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Biology of Mangroves and Magrove Ecosystems (Kathiresam and Bingham, 2001)

BIOLOGY OF MANGROVES AND MANGROVE ECOSYSTEMS                                    1

           Biology of Mangroves and Mangrove Ecosystems

          ADVANCES IN MARINE BIOLOGY VOL 40: 81-251 (2001)

                    K. Kathiresan1 and B.L. Bingham2

       Centre of Advanced Study in Marine Biology, Annamalai University,
                Parangipettai 608 502, India
      Huxley College of Environmental Studies, Western Washington University,
    Bellingham, WA 98225, USA e-mail (correponding author)

1. Introduction .............................................................................................. 4
    1.1. Preface........................................................................................ 4
    1.2. Definition ................................................................................... 5
    1.3. Global distribution ..................................................................... 5
2. History and Evolution ............................................................................. 10
    2.1. Historical background ................................................................ 10
    2.2. Evolution .................................................................................... 11
3. Biology of mangroves
    3.1. Taxonomy and genetics.............................................................. 12
    3.2. Anatomy..................................................................................... 15
    3.3. Physiology ................................................................................. 18
     3.4. Biochemistry ............................................................................. 20
    3.5. Pollination biology..................................................................... 21
    3.6. Reproduction, dispersal and establishment ................................ 22
    3.7. Biomass and litter production .................................................... 24
4. Mangrove-associated flora
    4.1. Bacteria ...................................................................................... 27
    4.2. Fungi and fungus-like protists.................................................... 29
    4.3. Microalgae.................................................................................. 33
    4.4. Macroalgae................................................................................. 34
    4.5. Seagrasses .................................................................................. 36
    4.6. Saltmarsh and other flora ........................................................... 37
5. Mangrove-associated fauna
    5.1. Zooplankton ............................................................................... 38
    5.2. Sponges and Ascidians............................................................... 39
    5.3. Epibenthos, infauna, and meiofauna .......................................... 41
    5.4. Prawns, shrimp and other crustaceans ....................................... 43
    5.5. Crabs .......................................................................................... 45
    5.6. Insects......................................................................................... 49
    5.7. Mollusks..................................................................................... 50
    5.8. Fish............................................................................................. 52
BIOLOGY OF MANGROVES AND MANGROVE ECOSYSTEMS                                   2

   5.9. Amphibians and Reptiles ........................................................... 56
   5.10. Birds ......................................................................................... 56
   5.11. Mammals.................................................................................. 57
6. Responses of mangroves and mangrove ecosystems to stress ................ 58
   6.1. Responses to light ...................................................................... 58
   6.2. Responses to gases ..................................................................... 59
   6.3. Responses to wind...................................................................... 61
   6.4. Responses to coastal changes..................................................... 61
   6.5. Responses to tidal gradients and zonation ................................. 63
   6.6. Responses to soil conditions ...................................................... 64
   6.7. Responses to salinity.................................................................. 66
   6.8. Responses to metal pollution ..................................................... 67
   6.9. Responses to organic pollution .................................................. 69
   6.10. Responses to oil pollution ........................................................ 70
   6.11. Responses to pests.................................................................... 71
   6.12. Responses to anthropogenic stress ........................................... 73
   6.13. Responses to global changes.................................................... 75
7. Ecological role of mangrove ecosystems
   7.1. Litter decomposition and nutrient enrichment ........................... 76
   7.2. Food webs and energy fluxes..................................................... 78
8. Concluding remarks ................................................................................ 80
   Acknowledgements ........................................................................... 82
   References .........................................................................................

    Mangroves are woody plants that grow at the interface between land and sea in
tropical and sub-tropical latitudes where they exist in conditions of high salinity, extreme
tides, strong winds, high temperatures and muddy, anaerobic soils. There may be no other
group of plants with such highly developed morphological and physiological adaptations
to extreme conditions.
     Because of their environment, mangroves are necessarily tolerant of high salt
levels and have mechanisms to take up water despite strong osmotic potentials. Some also
take up salts, but excrete them through specialized glands in the leaves. Others transfer
salts into senescent leaves or store them in the bark or the wood. Still others simply
become increasingly conservative in their water use as water salinity increases.
Morphological specializations include profuse lateral roots that anchor the trees in the
loose sediments, exposed aerial roots for gas exchange and viviparous water-dispersed
     Mangroves create unique ecological environments that host rich assemblages of
species. The muddy or sandy sediments of the mangal are home to a variety of epibenthic,
infaunal, and meiofaunal invertebrates. Channels within the mangal support communities
of phytoplankton, zooplankton, and fish. The mangal may play a special role as nursery
habitat for juveniles of fish whose adults occupy other habitats (e.g., coral reefs and
seagrass beds)
     Because they are surrounded by loose sediments, the submerged mangroves roots,
trunks, and branches are islands of habitat that may attract rich epifaunal communities
including bacteria, fungi, macroalgae, and invertebrates. The aerial roots, trunks, leaves

and branches host other groups of organisms. A number of crab species live among the
roots, on the trunks or even forage in the canopy. Insects, reptiles, amphibians, birds and
mammals thrive in the habitat and contribute to its unique character.
    Living at the interface between land and sea, mangroves are well adapted to deal
with natural stressors (e.g., temperature, salinity, anoxia, UV). However, because they
live close to their tolerance limits, they may be particularly sensitive to disturbances like
those created by human activities. Because of their proximity to population centers,
mangals have historically been favored sites for sewage disposal. Industrial effluents have
contributed to heavy metal contamination in the sediments. Oil from spills and from
petroleum production has flowed into many mangals. These insults have had significant
negative effects on the mangroves.
    Habitat destruction through human encroachment has been the primary cause of
mangrove loss. Diversion of freshwater for irrigation and land reclamation has destroyed
extensive mangrove forests. In the past several decades, numerous tracts of mangrove have
been converted for aquaculture, fundamentally altering the nature of the habitat.
Measurements reveal alarming levels of mangrove destruction. Some estimates put global
loss rates at one million ha y-1, with mangroves in some regions in danger of complete
collapse. Heavy historical exploitation of mangroves has left many remaining habitats
severely damaged.
     These impacts are likely to continue, and worsen, as human populations expand
further into the mangals. In regions where mangrove removal has produced significant
environmental problems, efforts are underway to launch mangrove agroforestry and
agriculture projects. Mangrove systems require intensive care to save threatened areas. So
far, conservation and management efforts lag behind the destruction; there is still much to
learn about proper management and sustainable harvesting of mangrove forests.
     Mangroves have enormous ecological value. They protect and stabilize coastlines,
enrich coastal waters, yield commercial forest products and support coastal fisheries.
Mangrove forests are among the world’s most productive ecosystems, producing organic
carbon well in excess of the ecosystem requirements and contributing significantly to the
global carbon cycle. Extracts from mangroves and mangrove-dependent species have
proven activity against human, animal and plant pathogens. Mangroves may be further
developed as sources of high-value commercial products and fishery resources and as sites
for a burgeoning ecotourism industry. Their unique features also make them ideal sites for
experimental studies of biodiversity and ecosystem function. Where degraded areas are
being revegetated, continued monitoring and thorough assessment must be done to help
understand the recovery process. This knowledge will help develop strategies to promote
better rehabilitation of degraded mangrove habitats the world over and ensure that these
unique ecosystems survive and flourish.


1.1. Preface

    Mangrove forests are among the world’s most productive ecosystems. They enrich
coastal waters, yield commercial forest products,
protect coastlines, and support coastal fisheries
(Figures 1 and 2). However, mangroves exist under
conditions of high salinity, extreme tides, strong
winds, high temperatures and muddy, anaerobic soils.
There may be no other group of plants with such
highly developed morphological, biological,
ecological and physiological adaptations to extreme
    Mangroves and mangrove ecosystems have
been studied extensively but remain poorly
understood. With continuing degradation and
destruction of mangroves, there is a critical need to
understand them better. Aspects of mangrove biology
have been treated in several recent reviews. Tomlinson
                     (1986) described
                     the basic botany    Figure 2. A) General view of coastal
                                edge of a mangrove forest. B) A black
                     of mangroves.
                                mangrove thicket, Avicennia, showing
                     Snedaker and      aerial roots (pneumatophores). C) Closer
                     Snedaker (1984)    view of the pneumatophores of
                     reviewed earlier
                     mangrove research and made recommendations for
                     further research. An overview of tropical mangrove
                     community ecology, based primarily on Australian
                     work, can be found in Robertson and Alongi (1992).
                     Li and Lee (1997) reviewed much of the Chinese
                     mangrove literature published between 1950 and
                     1995. Ellison and Farnsworth (2000) have recently
                     published a general review of mangrove ecology.
                         As researchers continue to discover important
                     facts about mangroves and the role they play in the
                     global ecosystem, the volume of published
Figure 1. A) The seward edge of a
mangrove forest, showing red      information has grown enormously and increasing
mangroves, Rhizophora. B) A young    numbers of workers are drawn to these unique
plant of Rhizophora, showing prop
                     environments. Thus, there is a need for periodic
roots carrying epifauna, including
                     reviews of the rapidly expanding literature. In this
barnacles and oysters. C) The
                     review, we emphasize work on mangrove ecosystems
propagules of Rhizophora, developed
from the fruit, before release (photos: completed between 1990 and 2000, though for space
A, A.J. Southward; B, K. Kathiresan;  reasons we can list only a fraction of the studies. Our
C, B.L. Bingham)
                     intent is to make information more readily available to

researchers around the world in hopes of facilitating and stimulating further study of the
mangrove environment.

1.2. Definition

    Mangroves are woody plants that grow at the interface between land and sea in
tropical and sub-tropical latitudes (Figures 1 and 2). These plants, and the associated
microbes, fungi, plants, and animals, constitute the mangrove forest community or
mangal. The mangal and its associated abiotic factors constitute the mangrove ecosystem
(Figure 3). The term “mangrove” often refers to both the plants and the forest community.
To avoid confusion, Macnae (1968) proposed that “mangal” should refer to the forest
community while “mangroves” should refer to the individual plant species. Duke (1992)
defined a mangrove as, “…a tree, shrub, palm or ground fern, generally exceeding one half
metre in height, and which normally grows above mean sea level in the intertidal zone of
marine coastal environments, or estuarine margins.” This definition is acceptable except
that ground ferns should probably be considered
mangrove associates rather than true mangroves.
The term “mangrove” is also used as an
adjective, as in ”mangrove tree” or “mangrove
fauna.” Mangrove forests are sometimes called
“tidal forests”, “coastal woodlands”, or “oceanic
rain forests.”
    The word “mangrove” is usually
considered a compound of the Portuguese word
“mangue” and the English word “grove.” The
corresponding French words are “manglier” and
“paletuvier” (Macnae, 1968) while the Spanish
term is “manglar”. The Dutch use            Figure 3. Physical and biological
“vloedbosschen” for the mangrove community       components of mangrove ecosystems.
and “mangrove” for the individual trees. German
use follows the English. The word “mangro” is a common name for Rhizophora in
Surinam (Chapman, 1976). It is believed that all these words originated from the
Malaysian word, “manggi-manggi” meaning “above the soil.” This word is no longer used
in Malaysia, but is used in eastern Indonesia to refer to Avicennia species.

1.3. Global distribution

     Mangroves are distributed circumtropically, occurring in 112 countries and
territories. Global coverage has been variously estimated at 10 million hectares (Bunt,
1992), 14-15 million hectares (Schwamborn and Saint-Paul, 1996), and 24 million hectares
(Twilley et al., 1992). Spalding (1997) gave a recent estimate of over 18 million hectares,
with 41.4% in south and southeast Asia and an additional 23.5% in Indonesia (Figure 4).
Mangroves are largely restricted to latitudes between 30° north and 30° south. Northern
extensions of this limit occur in Japan (31°22’N) and Bermuda (32°20’N); southern
extensions are in New Zealand (38°03’S), Australia (38°45’S) and on the east coast of
BIOLOGY OF MANGROVES AND MANGROVE ECOSYSTEMS                                          6

South Africa (32°59’S; Spalding, 1997, Yang
                            South and
et al., 1997). Mangroves are not native to the    Southeast Asia

Hawaiian Islands, but since the early 1900’s, at   The Americas

least 6 species have been introduced there.       West Africa

     Mangrove distributions within their       Australasia
ranges are strongly affected by temperature
                           East Africa and
(Duke, 1992) and moisture (Saenger and        the Middle East

Snedaker, 1993). Large-scale currents may also             0   2         4         6    8
                                      Area covered by mangrove forests (million ha)
influence distributions by preventing             Figure 4. Global coverage of
propagules from reaching some areas (De Lange and De mangrove forests (modified from
Lange, 1994). Individual mangrove species differ in the    Spalding, 1997).
length of time their propagules remain viable, their
establishment success, their growth rate, and their tolerance limits. These factors, which
appear quite consistent around the world, interact to produce characteristic distributional
ranges for most species (Duke et al., 1998a; Table 1).
BIOLOGY OF MANGROVES AND MANGROVE ECOSYSTEMS                                                                                               7

Table 1. Mangrove species, their taxonomic authorities, and global distributions.

                                                                              Malay Archipeligo

                                                                                                    Southwest Pacific
                                          Southeast USA

                                                                     Southeast Asia

                                                                                                              West Pacific
                                                               South Asia

                                                                                        East Asia

Family       Species

                                                                 !       !         !       !
Avicenniaceae   Avicennia alba Blume
          Avicennia balanophora Stapf and Moldenke ex Molodenke
          Avicennia bicolor Standley
          Avicennia eucalyptifolia (Zipp. ex Miq.) Moldenke
                                             !      !
          Avicennia germinans (L.) Stearn
          Avicennia lanata Ridley
                                                            !     !       !         !       !      !        !
          Avicennia marina (Forsk.). Vierh.
                                                                         !         !       !      !
          Avicennia officinalis L.
          Avicennia schaueriana Stapf and Leechman ex Moldenke
          Avicennia africana Palisot de Beauvois

                                                                 !       !         !
Bignoniaceae    Dolichandrone spathacea (L. f.) K. Schumann

                                                                                  !       !
Bombacaceae    Camptostemon philippinensis (Vidal) Becc.
          Camptostemon schultzii Masters

                                                                 !       !         !             !
Caesalpiniaceae  Cynometra iripa Kostel
                                                                 !       !         !
          Cynometra ramiflora L.

                                             !      !
Combretaceae    Conocarpus erectus L.
                                             !      !             !       !
          Laguncularia racemosa (L.) Gaertn. f.
                                                                 !       !         !
          Lumnitzera littorea (Jack) Voigt.
                                                            !     !       !         !       !
          Lumnitzera racemosa Willd.
          Lumnitzera X rosea (Gaud.) Presl. (hybrid of
           L. racemosa and L. littorea)
BIOLOGY OF MANGROVES AND MANGROVE ECOSYSTEMS                                   8

                                           !  !  !  !  !  !
Euphorbiaceae  Excoecaria agallocha L.
                                           !  !  !
         Excoecaria indica (Willd.) Muell. - Arg.
         Excoecaria dallachyana (Baill.) Benth.

                                         !  !  !  !  !  !
Lythraceae    Pemphis acidula Forst.
         Pemphis madagascariensis (Baker) Koehne

Meliaceae    Aglaia cucullata (Pellegrin ) Roxb.
                                           !  !  !  !  !    !
         Xylocarpus granatum Koen.
                                           !  !  !    !
         Xylocarpus mekongensis Pierre
                                           !  !  !
         Xylocarpus moluccensis (Lamk.) Roem.

                                           !  !  !  !  !
Myrsinaceae   Aegiceras corniculatum (L.) Blanco
                                             !  !  !
         Aegiceras floridum Roemer and Schultes

                                               !  !
Myrtaceae    Osbornia octodonta F. Muell. loc. cit.

Pellicieraceae  Pelliciera rhizophoreae Triana and Planchon

Plumbaginaceae  Aegialitis annulata R. Brown
                                           !  !
         Aegialitis rotundifolia Roxburgh

                                           !  !  !  !
Rhizophoraceae  Bruguiera cylindrica (L.) Bl.
         Bruguiera exaristata Ding Hou
                                         !  !  !  !  !  !  !  !
         Bruguiera gymnorrhiza (L.) Lamk.
                                             !  !
         Bruguiera hainesii C. G. Rogers
                                           !  !  !  !  !
         Bruguiera parviflora Wight and Arnold ex Griffith
                                         !  !  !  !  !
         Bruguiera sexangula (Lour.) Poir.
                                           !  !  !  !
         Ceriops decandra (Griff.) Ding Hou
                                       !  !  !  !  !  !  !  !  !
         Ceriops tagal (Perr.) C. B. Robinson
                                           !  !  !  !
         Kandelia candel (L.) Druce
                                           !  !  !  !  !    !
         Rhizophora apiculata Bl.
                                     !  !
         Rhizophora mangle L.
                                         !  !  !  !  !  !  !  !
         Rhizophora mucronata Poir.
         Rhizophora racemosa Meyer
         Rhizophora samoensis (Hochr.) Salvoza
                                           !  !  !  !  !  !
         Rhizophora stylosa Griff.
                                           !    !    !
         Rhizophora X lamarckii Montr. (hybrid of R. apiculata
          and R. stylosa)

         Rhizophora X annamalayana Kathir. (hybrid of
          R. apiculata and R. mucronata )
         Rhizophora X selala (Salvoza) Tomlinson (hybrid of
          R. stylosa and R. samoensis)
                                    !  !
         Rhizophora x harrisonii Leechman (hybrid of R.mangle
          and R. stylosa )

                                        !  !  !  !
Rubiaceae    Scyphiphora hydrophyllacea Gaetn. f.

                                      !  !  !  !  !  !  !
Sonneratiaceae  Sonneratia alba J. Smith
         Sonneratia apetala Buch.-Ham.
                                        !  !  !  !
         Sonneratia caseolaris (L.) Engler
                                        !  !  !
         Sonneratia griffithii Kurz
         Sonneratia lanceolata Blume
                                          !  !
         Sonneratia ovata Backer
         Sonneratia X gulngai Duke (hybrid of S. alba
          and S. caseolaris)

                                        !  !  !
Sterculiaceae  Heritiera fomes Buch.-Ham.
         Heritiera globosa Kostermans
                                        !  !  !  !
         Heritiera littoralis Dryand. In Aiton

    Mangroves have broader ranges along the warmer eastern coastlines of the
Americas and Africa than along the cooler western coastlines. Mangroves prefer a humid
climate and freshwater inflow that brings in abundant nutrients and silt. Mangroves grow
luxuriantly in alluvial soils (loose, fine-textured mud or silt, rich in humus). They are
abundant in broad, sheltered, low-lying coastal plains where topographic gradients are
small and tidal amplitudes are large. Repeatedly flooded but well-drained soils support
good mangrove growth and high species diversity (e.g., Azariah et al., 1992). Mangroves
do poorly in stagnant water (Gopal and Krishnamurthy, 1993).


2.1. Historical background

    Mangroves have been known and studied since ancient times. Descriptions by
Nearchus (325 B.C.) and Theophrastus (305 B.C) of Rhizophora trees in the Red Sea and
the Persian Gulf are the earliest known records. Plutarch (70 A.D.) and Abou’l Abass
(1230) wrote about Rhizophora and its seedlings (Macnae, 1968; Chapman, 1976). The
bibliography of mangrove research compiled by Rollet (1981), however, shows only 14
references before 1600, 25 references from the 17th century, 48 references in the 18th
century, and 427 in the 19th century. In contrast, there were 4500 mangrove references
between 1900 and 1975 and approximately 3000 between 1978 and 1997, illustrating the
explosion of interest in mangroves.
    Mangroves have a long historical link with human culture and civilization. In the
Solomon Islands, the bodies of the dead are disposed of and special rites are performed in
the mangrove waters (Vannucci, 1997). In the third century, a Hindu temple to the
mangrove Excoecaria agallocha was erected in south India. Rock carvings show the plant
being worshipped anciently as a “sacred grove” and even today it is believed that a dip in
the holy pond of the temple cures leprosy. The city where this temple is found bears the
name of the mangrove. In Kenya, shrines built in the mangrove forests are worshipped by
the local people, who believe spirits of the shrine will bring death to those who cut the
surrounding trees.
    The Portuguese, probably the first Europeans to visit the mangrove forests of the
Indian Ocean (around the 14th century), learned the traditional Indian technique of rice-
fish-mangrove farming, as demonstrated by letters from the Viceroys to the King of
Portugal. Some six centuries ago, this Indian technology was also transferred by Jesuit and
Franciscan Fathers to the African countries of Angola and Mozambique (Vannucci, 1997).
In the 19th century, the British used the practical knowledge gained over centuries by the
Indians to manage mangroves at Sunderbans for commercial timber production (Vannucci,
1997). An unusually creative use of mangroves is described in a traditional story from
India about two countries at war. The larger country planned to invade their small
neighbors during the night. The smaller nation, which had mangrove forests on its
coastline, plotted to discourage their enemies by placing lighted lamps on the aerial roots
of mangroves. What appeared to be a large flotilla of ships discouraged the invaders and
ended the hostilities.

2.2. Evolution
    The evolutionary history of mangroves remains problematic with a number of
competing theories. Mangroves evolved from terrestrial rather than marine plants.
Mangrove pollen fossils have been found below marine foraminiferan assemblages (i.e., in
the lower deposits of estuarine environments) suggesting the evolution of these plants from
a non-marine habitat to an estuarine habitat (Srivastava and Binda, 1991). In the distant
past, these land plants adapted to brackish water and became the “core” mangrove flora.
The diversity of mangroves is much higher in the Indo-West Pacific than in the Western
Atlantic and Caribbean. Two competing hypotheses have been presented to explain this
pattern. The center-of-origin hypothesis suggests that all mangrove taxa first appeared in
the Indo-West Pacific and subsequently dispersed to other regions. The vicariance
hypothesis, on the other hand, states that all mangroves originated around the Tethys Sea.
Continental drift then isolated the flora in different regions of the earth where
diversification created distinct faunas.
    Ellison et al. (1999) evaluated these two hypotheses using 1) a review of the
mangrove fossil record, 2) a comparison of modern and fossil distributions of mangroves
and mangrove-associated gastropods, 3) an analysis of species-area relationships of
mangroves and gastropods, 4) an analysis of nestedness patterns of individual plants and
gastropod communities, and 5) an analysis of nestedness patterns of individual plants and
individual gastropod species. The evidence from all 5 analyses supported the vicariance
hypothesis, suggesting a Tethyan origin of mangroves. This argues that the much higher
diversity of mangroves in the Indo-West Pacific relates to conditions there that favored
diversification. For example, the continual presence of extensive wet habitat may have
allowed more species to make the transition from terrestrial to brackish-water habitats. The
Atlantic, Caribbean or and East Pacific all saw periods of drying which could have
prevented such adaptation. Ricklefs and Latham (1993) suggest that limited dispersal,
combined with the closure of the Tethys connection to the Atlantic Ocean in the mid-
Tertiary, restricted most mangrove taxa to the Indo-Pacific.
    Studies of mangrove biochemistry and genetics should provide further evidence
concerning mangrove evolution and dispersal. For example, Dodd et al. (1998) found
significant genetic differentiation between mangroves in eastern and western Atlantic
provinces. Three species from western Africa showed significantly greater lipid diversity
and longer carbon chains than conspecifics from eastern South America, suggesting that
the western Atlantic mangroves show derived characteristics. The authors concluded that
this evidence suggests it is unlikely that Atlantic mangroves dispersed from the Tethys via
the Pacific.
    Mangroves are quite old, possibly arising just after the first angiosperms, around
114 million years ago (Duke, 1992). Avicennia and Rhizophora were probably the first
genera to evolve, appearing near the end of the Cretaceous period (Chapman, 1976). Pollen
records provide important information about subsequent radiation. Fossil pollen from
sediments in the Leizhou Peninsula, China suggest that mangroves expanded from south to
north, reaching their northern limit on the Changjiang Delta by the mid-Holocene (Y.
Zhang et al., 1997). A similar study of pollen from late Holocene samples in Bermuda
suggests that mangroves were established there in the last 3000 years, when sea level rise
decreased from 26 to 7 cm per century (J.C. Ellison, 1996).

     A detailed study of pollen records from Mexico, the Antilles, Central America and
northern South America (Graham, 1995) show that neotropical environments were first
occupied by Acrostichum, Brevitricolpites variabilis, Nypa and Pelliceria in the early
Eocene, about 50 million years ago. Avicennia appeared in this region in the late Miocene
(about 10 million years ago). Six mangrove species and three associated genera were
present by the middle Pliocene (3.5 million years ago), and fifteen plant genera were
present by the Quaternary period. Twelve additional species were added during the
Cenozoic to produce the present-day assemblage of about 27 genera of mangroves and
associated plants (Rico-Gray, 1993; Graham, 1995).
     Continental drift produced massive mixing and dispersal of genes in geologically
recent times, greatly enhancing evolutionary processes. Though mangroves evolved in the
tropics, one species, Avicennia marina, is found in temperate latitudes, particularly in the
southern hemisphere (Saenger, 1998). This genus is of a western Gondwanan origin with
the subsequent radiation of several taxa facilitated by tectonic dispersal of southern
continental fragments (Duke, 1995). Mangrove fossils have clearly provided valuable
information about prehistorical mangrove evolution and dispersal. However, Burnham
(1990) cautions that reconstructions based on organic remains can differ substantially
depending on the mangrove parts studied (e.g., fruits and seeds vs. leaf litter).
     Mangrove ecosystems, in general, are dynamic, undergoing changes on time scales
of 10 - 104 y(Woodroffe, 1992). Indeed fossil mangroves are often found in regions where

they no longer exist: in Texas, USA (Westgate and Gee, 1990; Westgate 1994), west
Africa (Marius and Lucas, 1991), Hungary (Nagy and Kokay, 1991), India (Bonde, 1991;
Barni and Chanda, 1992), the Chao-Shan Plain of China (Z. Zheng, 1991), and Western
Australia (Kendrick and Morse, 1990), for example.
     Historical changes in mangrove distributions can reveal details about paleoclimates
and sea-level changes (Somboon, 1990; Khandelwal and Gupta, 1993; Y. Zhang and
Wang, 1994; Plaziat, 1995; Saito et al., 1995; Lezine, 1996; W. Zhang and Huang 1996; Y.
Zhang et al., 1997). For example, in the equatorial Pacific Ocean, there are alternating reef
and mangrove fossils in upper Miocene and lower Pliocene deposits (Cronin et al., 1991).
Similarly, Holocene sediments from the Maya Wetland of Belize indicate that mangrove
peat filled the lagoon by 4800 y ago (Alcala-Herrera et al., 1994). These patterns may
reflect fluctuating sea levels or large-scale climatic shifts. In Poverty Bay, New Zealand,
the presence of Avicennia marina var. resinifera during the early to mid-Holocene suggests
that the area then had a frost-free climate (Mildenhall, 1994). The mangrove fossil record
is clearly an area where continued research has the potential for providing significant
information, not only about the history of these unique plants, but also about the recent
history of the earth.

3.1. Taxonomy and genetics

3.1.1. Taxonomy
    Tomlinson (1986) recognized three groups of mangroves: major mangrove species,
minor mangrove species and mangrove associates. The major species are the strict or true
mangroves, recognized by most or all of the following features: 1) they occur exclusively
in mangal, 2) they play a major role in the structure of the community and have the ability

to form pure stands, 3) they have morphological specializations - especially aerial roots
and specialized mechanisms of gas exchange, 4) they have physiological mechanisms for
salt exclusion and/or excretion, 5) they have viviparous reproduction, and 6) they are
taxonomically isolated from terrestrial relatives. The strict mangroves are separated from
their nearest relatives at least at the generic level, and often at the sub-family or family
     The minor mangrove species are less conspicuous elements of the vegetation and
rarely form pure stands. According to Tomlinson (1986), the major mangroves include 34
species in 9 general and 5 families. The minor species contribute 20 additional species in
11 genera and 11 families for a total of 54 mangrove species in 20 genera and 16 families.
Duke (1992), on the other hand, identified 69 mangrove species belonging to 26 genera in
20 families. One family falls in the fern division (Polypodiophyta); the remainder are in the
Magnoliophyta (angiosperms). Families containing only mangroves are the
Aegialitidaceae, Avicenniaceae, Nypaceae and Pellicieraceae. Two orders (Myrtales and
Rhizophorales) contain 25% of all mangrove families. By reconciling common features
from Tomlinson (1986) and Duke (1992), we recognize 65 mangrove species in 22 genera
and 16 families (Table I).
     There are a number of problems with mangrove taxonomy (Duke, 1992) and many
of these are based on hybridization between described species. For instance, the systematic
distinction between Rhizophora mucronata in eastern Africa, R. stylosa in Australia, and
their putative hybrids is unclear. Rhizophora lamarckii, which occurs in New Caledonia,
Papua New Guinea and Queensland, Australia, is a sterile F1 hybrid between R. apiculata
and R. stylosa. Rhizophora x annamalayana, found in a south Indian mangrove forest, was
first identified as R. lamarckii but has since been reidentified as a new species hybrid
between R. mucronata and R. apiculata (Kathiresan, 1995a). Some hybrids, like
Rhizophora x harrissoni, can not be confirmed with wax chemistry (Dodd et al., 1995).
Molecular analyses may help eventually resolve the taxonomic problems. For example,
DNA sequence data from the chloroplast gene rbcL indicate that the Rhizophoraceae
belongs not to the Myrtales, but to a rosid clade that includes the families Euphorbiaceae,
Humiriaceae and Malphighiaceae (Conti et al., 1996).

3.1.2. Genetic variation
    There is significant inter- and intraspecific variability among mangroves. For
example, physiological differences have been identified between West African and
Western Atlantic Avicennia germinans (Saenger and Bellan, 1995) and distinct
chemotypes have been described for A. germinans and Rhizophora (Corredor et al., 1995;
Dodd et al., 1995; Rafii et al., 1996). Variability may result from genotypic differences or
from phenotypic responses to local environments. Mean leaf area of Rhizophora mangle in
Mexico, for example, is positively correlated with annual precipitation and negatively
correlated with latitude. This morphological response to local conditions may allow the
trees to maximize their photosynthetic efficiency (Rico-Gray and Palacios-Rios, 1996a).
Similarly, leaf area indices can be used to differentiate Rhizophora mangle from basin and
dwarf forest types in southeast Florida, USA (Araujo et al., 1997). In contrast, variation in
Rhizophora mangle flower morphology appears to have a genetic basis. Dominguez et al.
(1998) found significant differences between populations on the Pacific and Atlantic coasts
of Mexico, among populations on each coast, and within individual populations. They

hypothesized that frequent extinctions, followed by recolonization of a few individuals, has
produced genetic differentiation.
     Genetic variability has been clearly demonstrated through biochemical markers like
iridoid glycosides (Fauvel et al., 1995), foliar leaf waxes (Dodd et al., 1995, 1998; Rafii et
al., 1996), and isoenzymes (Duke, 1991). It is also evident in differences in length and
volume of chromosomes (Das et al., 1994). Lakshmi et al. (1997) measured intraspecific
genetic variability in Acanthus ilicifolius through DNA-based molecular markers that are
insensitive to environmental influences (i.e., random amplified polymorphic DNAs and
restriction fragment length polymorphisms). They found 48 genotypes in eight distinct
populations. There were no differences in chromosome number (2n = 48). Genetic
polymorphism is even higher in Excoecaria agallocha. The E. agallocha polymorphism is
independent of morphological and sexual differences (Parani et al., 1997).
     Changes in gene frequency, such as those produced by inbreeding, can lead to
genetic differentiation. Inbreeding may result if pollen are shed before the flower opens
(Lowenfeld and Klekowski, 1992). If inbreeding is prevalent, a mangrove forest may be a
virtually monospecific stand with little genetic diversity. Pollination by bees produces
geitonogamous selfing in Kandelia candel. However, there is little genetic differentiation
among 13 populations along the coastlines of Hong Kong, indicating that dispersion of
propagules is sufficient to maintain high levels of gene flow in this species (Sun et al.,
1998). In contrast, genetic differentiation, has led to subspeciation in Avicennia marina
(Duke, 1991, 1995). It has been assumed that Avicennia propagules commonly move long
distances. However, allozyme studies suggest that Avicennia species in the Indo-West
Pacific and eastern North America have limited gene flow. This may indicate that true
dispersal distances are much shorter than has been commonly believed (Duke et al.,
     Gene mutations can also cause species divergence. One or 2 gene mutations are
needed for biochemical differences, 5-10 for physiological changes, >10 for morphological
variations and >100 for taxonomic changes (Saenger, 1998). A single recessive gene
causes albinism in Rhizophora seedlings. This albino mutation is in the nuclear genome but
has a profound effect on ultrastructure of the chloroplasts (Klekowski et al., 1994a).
Pigment fingerprint studies of chlorophyll-deficient mutants show that most albino
genotypes are deficient in chlorophylls, xanthophylls, and carotenes (Corredor et al.,
1995). Recent studies of post-zygotic mutations reveal that fewer than 0.1% of the
Rhizophora in Puerto Rico exhibit somatic mutations. These mutations are often manifest
in shoot apices as complete or partial periclinal chimeras (Klekowski et al., 1996). Rates of
both mutation and outcrossing vary among mangrove populations. For instance, the Puerto
Rican Rhizophora are more outcrossed and have lower mutation rates for chlorophyll-
deficiency than Florida Rhizophora.

3.13. Tissue Culture
    There have been few studies of tissue culture in mangroves. This is because
explants frequently turn brown or black shortly after isolation, with tissue death usually
following (Kathiresan, 1990, 1994). The high tannin and phenol content of mangroves may
be responsible for the browning problem (Kathiresan and Ravi, 1990; Ravi and Kathiresan,
1990). Antioxidants can prevent phenolic browning in explants collected during the
monsoon season (Kathiresan and Ravikumar, l997).

    Callus induction has been achieved in Sonneratia apetala and Xylocarpus
granatum by supplementing the medium with double strength vitamins (Kathiresan and
Ravikumar, 1997). Baba and Onizuka (1997) have improved techniques for callus
induction and initiation of redifferentiation in the callus of Bruguiera gymnorrhiza,
Kandelia candel, Pemphis acidula and Rhizophora stylosa. Adventitious roots were
produced in P. acidula, but neither adventitious buds nor roots could be induced in the
remaining species.
    Researchers are currently working to identify and micropropagate unique plant
genotypes for commercial purposes. Mangals may provide good raw material for such
work. For instance, in vitro multiplication of the salt-marsh Sesuvium portulacastrum,
associated with Indian mangroves, has been achieved by axillary bud culture (Kathiresan,
1994; Kathiresan et al., 1997). In vitro cell cultures of this plant synthesize antibacterial
substances in higher quantities than do the intact plants, demonstrating the potential of
these systems for production of valuable metabolites (Kathiresan and Ravikumar, 1997).
    Cell protoplast fusion techniques may allow us to transfer salinity tolerance from
mangrove plants to non-salt-tolerant species (Swaminathan, 1991). Methods for extracting
and preparing protoplasts from tissue cultures of Bruguiera gymnorrhiza have been
developed by Eguchi et al. (1995). Sasamota et al. (1997) have done similar work with the
cotyledons of Avicennia marina and A. lanata. Such creative tissue culture work may
allow researchers to better understand, and make use of, the unique characteristics of

3.2. Morphology and anatomy

3.2.1. Root anatomy
    Mangroves are highly adapted to the coastal environment, with exposed breathing
roots, extensive support roots and buttresses, salt-excreting leaves, and viviparous water-
dispersed propagules. These adaptations vary among taxa and with the physico-chemical
nature of the habitat (Duke, 1992). Perhaps the most remarkable adaptations of the
mangroves, however are the stilt roots of Rhizophora, the pneumatophores of Avicennia,
Sonneratia and Lumnitzera, the root knees of Bruguiera, Ceriops and Xylocarpus and the
buttress roots of Xylocarpus and Heritiera. The roots of many mangroves do not penetrate
far into the anaerobic substrata. Instead, the trees produce profuse lateral roots for support.
Their effectiveness is well illustrated by the tallest mangrove trees, found in Ecuador,
which attain heights of more than 60 m and may be 100 yold (Emilio, 1997).
    The specialized roots are important sites of gas exchange for mangroves living in
anaerobic substrata. The exposed surfaces may have numerous lenticels (loose, air-
breathing aggregations of cells; Tomlinson, 1986). Avicennia possesses lenticel-equipped
pneumatophores (upward directed roots) through which oxygen passively diffuses. The
lenticels may be closed, partially opened or fully opened, depending on environmental
conditions (Ish-Shalom-Gordon and Dubinsky, 1992). The spongy pneumatophores are
generally short (< 30 cm), but grow much larger and become more numerous in Avicennia
marina living in anaerobic and oil-polluted conditions. This phenotypic response
apparently increases surface area for gas exchange (Saifullah and Elahi, 1992). In

Sonneratia, the pneumatophores may be 3 m long and stout from heavy secondary
thickening (Tomlinson, 1986).
     Oxygen may also pass through non-lenticellular portions of the pneumatophores.
Horizontal structures (subrisules) may be important in air exchange, particularly in rapidly
growing pneumatophores where the newly formed tip lacks lenticels (Hovenden and
Allaway, 1994). Pneumatophores are normally unbranched. However, following the 1991
Gulf War, mangroves in the Arabian Gulf began developing branched pneumatophores and
adventitious roots (Boeer, 1993).
     The general structure of mangrove roots is similar to that of most other vascular
plants. They typically have a root cap, lateral roots arising endogenously, exarch
protoxylem, and alternating strands of primary phloem and xylem. Many also have an
enlarged polyarch stele with a wide parenchymatous medulla. Aerial roots are modified for
life above ground. Compared to the underground roots, they have an exaggerated zone of
elongation behind the apical meristem (Tomlinson, 1986). They also have significant
secondary thickening (similar to the stems). When the aerial roots reach the ground, they
shift to having a short elongation zone and little to no secondary growth. They also become
spongy to adapt to sub-soil existence. In Rhizophora, the roots become thinner and form
“capillary rootlets” with a simple diarch stele and a narrow cortex. Like aquatic plants, true
mangroves lack root hairs. Hence, the endodermis is an effective absorbing layer
(Tomlinson, 1986).

3.2.2. Wood anatomy
    Tomlinson (1986) has summarized the unique anatomical features of mangrove
woods. Growth rings are conspicuously anomalous (as in Avicennia; Das and Ghose, 1998)
or completely absent. Hence, aging trees is difficult. Duke and Pinzon (1992) suggest that
leaf scar nodal number is a better way to estimate the age of Rhizophora seedlings.
    Mangrove wood has special features that enable the trees to overcome the high
osmotic potential of seawater and the transpiration caused by high temperatures. There are
numerous narrow vessels running through the wood. These range in density from 32 •
mm-2 in Excoecaria to 270 • mm-2 in Aegiceras (Das and Ghose, 1998). The vessels help
create high tensions in the xylem since a slight decrease in vessel diameter produces a
disproportionally large increase in flow resistance (Scholander et al., 1964, 1965;
Tomlinson, 1986). The vessel elements, which form the vessels, normally have simple
perforation plates (Tomlinson, 1986). However, mangroves in the family Rhizophoraceae
(except Kandelia candel) have scalariform perforation plates.
    Water conduction through wood is strongly influenced by size and distribution of
the vessels. Water moves most quickly through ring-porous woods in which the largest
vessels are in the outermost growth layer. Conduction is much slower in diffuse-porous
woods where vessels are more uniform in size and distribution. The wood of most
mangroves is diffuse-porous but Aegialitis rotundifolia has ring-porous wood (Das and
Ghose, 1998).
    Wier et al. (1996) studied wound repair in Rhizophora mangle. A closing layer
isolates necrotic tissue within 17 d, and the wound is completely enclosed by periderm by
52 d. Isolation of the damage site and development of wound periderm may prevent spread
of pathogens to undamaged tissues.

3.2.3. Leaf anatomy
     Mangrove leaves are almost leathery with obscure leaf veins (there are no vein
sheaths). The cuticle is thick and smooth with small hairs, giving the plant a glossy
appearance. The leaves are of moderate size and are arranged in a modified decussate
(bijugate) pattern with each pair at an angle less than 180° to the preceding pair. This
arrangement reduces self-shading and produces branch systems that fill space in the most
photosynthetically efficient way (Tomlinson, 1986). The leaves generally show
dorsiventral symmetry though isolateral leaves are also found in Kandelia candel,
Sonneratia apetala and Phoenix paludosa (Das et al., 1996).
     Six types of stomata are known from mangrove leaves. These differ in their
arrangement of guard cells and subsidiary cells. In most species, a horn or beak-like
cuticular outgrowth covers either the outer side of the stomatal pore or both the inner and
outer sides. These structures reduce stomatal transpiration (Das and Ghose, 1993), which is
important given the high solute concentration of the water and the “physiological drought”
the trees experience. Heritiera fomes has deeply sunken stomata covered by trichomes. The
leaves in this species also have a palisade-spongy ratio that is small compared to other
halophytes (Das et al., 1995).
     Mangrove leaves have specialized idioblast cells including tannin cells
(Rhizophoraceae), mucous cells (Rhizophora, Sonneratia), crystalliferous cells
(Rhizophoraceae), oil cells (Osbornia) and laticifers (Excoecaria; Tomlinson, 1986). In
general, the leaves lack bundle sheath fibres and bundle sheath extensions, but possess
enlarged tracheids terminating in vein endings. Branched sclereids are abundant and well
developed in Aegiceras, Rhizophora, Sonneratia and Aegialitis. The sclereids may give
mechanical support to leaves or discourage herbivores. Both sclereids and tracheids may
also be involved in water storage (Tomlinson, 1986). Water is also stored in colourless,
non-assimilatory water-storage tissue that is hypodermal in dorsiventral leaves, but is deep-
seated in the extensive mesophyll region of isolateral leaves. In some species, the thick
layer of non-assimilatory tissue occurs in front of the assimilatory cells. This back scatters
incoming light, creating a gradient that may help the plant capture weak light, increasing
photosynthetic efficiency (Koizumi et al., 1998).
     Yoshihira et al. (1992) studied the distribution of pigments in mangrove leaves.
They found that different species concentrated the pigments in different parts of the leaves.
In Aegiceras corniculatum, the highest concentration of carotenoids and chlorophylls was
in the light-harvesting complex. In Rhizophora apiculata, however, chlorophyll was
concentrated in the chloroplast reaction center. The chlorophyll-binding proteins (including
the functional cytochrome B 6/f complex and the protein kinases) were found in the
thylakoid membranes in Bruguiera gymnorrhiza and Kandelia candel

3.2.4. Seed and seedling anatomy
    Avicennia marina forms endosperm haustoria during early embryonic
histodifferentiation. Once the growth phase is initiated, subsequent embryonic
development is extra-ovular. The mature seed, therefore, is enclosed by a pericarp that
originates entirely from the ovary wall. From the end of histodifferentiation until the
mature seeds are abscised, cotyledon cells become highly vacuolated and contain large
amounts of soluble sugars, which constitute the major nutrient reserves of the mature seed
(Farrant et al., 1992).

    Incipient phellogen usually develops toward the radicle end of mangrove seedlings
and masks the chlorophyllous tissue. Tannin cells are present in the aerenchymatous tissue,
stone cells are present in the outer cortex, and trichosclereids appear in the cortex and
medulla. Since the epidermis lacks stomata, numerous lenticels facilitate gas exchange.
    In experiments with six mangrove species, Youssef and Saenger (1996)
demonstrated that the seedlings have special features that allow them to tolerate flooding
and facilitate rhizosphere oxidation. Lacunae in the ground tissue constrict air flow
passages, conserving oxygen and enabling the mangrove to maintain aerobic metabolism
during periods of flooding. Variations in this anatomical feature are responsible for species
differences in tolerance to flood stress.

3.3. Physiology

3.3.1. Salt regulation
     Mangroves are physiologically tolerant of high salt levels and have mechanisms to
obtain fresh water despite the strong osmotic potential of the sediments (Ball, 1996). They
avoid heavy salt loads through a combination of salt exclusion, salt excretion, and salt
accumulation. For example, Rhizophora, Bruguiera, and Ceriops all possess ultrafilters in
their root systems. These filters exclude salts while extracting water from the soil. Other
genera (e.g., Avicennia, Acanthus, Aegiceras) take some salt up, but excrete it through
specialized salt glands in the leaves (Dschida et al., 1992; Fitzgerald et al., 1992). The salt-
excreting species allow more salt into the xylem than do the non-excretors, but still
exclude about 90% of the salts (Scholander et al., 1962, Azocar et al., 1992). Salt excretion
is an active process, as evidenced by ATPase activity in the plasmalemma of the excretory
cells (Drennan et al., 1992). The process is probably regulated by leaf hypodermal cells,
which may store salt as well as water (Balsamo and Thomson, 1995).
     Species of Lumnitzera and Excoecaria accumulate salts in leaf vacuoles and
become succulent. Salt concentrations in the sap may also be reduced by transferring the
salts into senescent leaves or by storing them in the bark or the wood (Tomlinson, 1986).
As water salinity increases, some species simply become increasingly conservative in their
water use, thus achieving greater tolerance (Ball and Passioura, 1993). In south Florida,
Rhizophora mangle decreases its salt stress by using surface water as its sole water source.
In the wet season, the fine root biomass increases in response to decreased salinity of the
surface waters, directly enhancing the uptake of low-salinity water (Lin and Sternberg,
     Most mangrove species directly regulate salts. However, they may also accumulate
or synthesize other solutes to regulate and maintain osmotic balance (Werner and Stelzer,
1990; Popp et al., 1993). For example, Aegiceras corniculatum, Aegialitis annulata and
Laguncularia racemosa accumulate mannitol and proline (Polania, 1990). Avicennia
marina accumulates glycine betaine, asparagine and stachyose (Ashihara et al., 1997).
Sonneratia alba synthesizes purine nucleotides that help it adapt to salt loads of 100 mM
NaCl (Akatsu et al., 1996). To facilitate the flow of water from root to leaves, the water
potential at the leaves is held lower (-2.5 to -6 MPa) than in the roots (-2.5 MPa;
Scholander et al., 1964).
     Because mangrove roots exclude salts when they extract water from soil, soil salts
could become very concentrated, creating strong osmotic gradients (Passioura et al., 1992).

However, viscous, polymeric substances in the sap limit flow rate and decrease
transpiration (Zimmermann et al., 1994). This, combined with high water-use efficiency,
slows the rate of water uptake and prevents salts from accumulating in the soil surrounding
the roots. This helps the mangroves conserve water and regulate internal salt
concentrations (Ball and Passioura, 1993; Ball, 1996). Low transpiration and slow water
uptake, however, are not characteristic of all mangrove species. Becker et al. (1997)
measured relatively high transpiration rates in both Avicennia alba and Rhizophora
    Transpiration rates vary with season, being higher in the dry season than in the wet
season in Bruguiera cylindrica (Herppich and Von Willert, 1995; Hirano et al., 1996). This
corresponds to changes in stomatal movement. The oscillatory behaviour of Avicennia
germinans stomata is affected by any factor that changes hydraulic flow through the plant.
This includes increases in vapour pressure deficit and osmotic potential of the substrata
(Naidoo and Von-Willert, 1994).
    Fukushima et al. (1997) studied the effects of salt on sugar catabolism in leaves and
roots of Avicennia marina. They showed that sugar catabolic pathways are different in
roots and leaves. Over 50% of the 14C-labeled sucrose the gave the plants was incorporated
into an unidentified sugar in the leaves. The remainder appeared in the roots as glucose,
fructose and sucrose. Neither pathway was significantly affected by salt levels.

3.3.2. Photosynthesis
     Mangroves show characteristic C3 photosynthesis. Basak et al. (1996) found
significant intra- and interspecific variation in photosynthetic activity of 14 mangrove
species, suggesting that the rates of photosynthesis may have an underlying genetic basis.
This possibility is supported by observations that the photosynthetic rate of Bruguiera is
under direct internal control and is not influenced by stomatal activity induced by changes
in salinity or light (Cheeseman et al., 1991; Cheeseman, 1994).
     In contrast, other researchers have shown that photosynthetic rates of some species
are strongly affected by environmental conditions. For example, low salinity conditions
reduce carbon losses in Avicennia germinans and Aegialitis annulata and lead to greater
CO2 assimilation (Naidoo and Von-Willert, 1995). Fluctuating soil salinities lead to
significantly lower intercellular CO2 concentration and reduced photosynthesis in scrub
forests of south Florida (Lin and Sternberg, 1992). The stunted mangroves in these habitats
have much lower canopies, more main stems and smaller leaves than mangroves in fringe
forests that experience less salinity variability. Steinke and Naidoo (1991) also
demonstrated experimentally that temperature affects the photosynthetic rate of Avicennia
marina. Temperature-induced changes in the relative rates of photosynthesis and
respiration, in turn, influence overall growth rates.
     Strong sunlight can also reduce mangrove photosynthesis through inhibition of
Photosystem II (Cheeseman et al., 1991). The photosynthetic rates of mangroves saturate
at relatively low light levels despite their presence in high sunlight tropical environments.
The fairly low photosynthetic efficiency may be related to the concentration of zeaxanthin
pigments in the leaves (Lovelock and Clough, 1992). To prevent damage to the
photosystems, the mangroves dissipate excess light energy via the xanthophyll cycle
(Gilmore and Bjorkman, 1994) and through the conversion of O2 to phenolics and
peroxidases (Cheeseman et al., 1997).

    Kathiresan and Moorthy (1994a) and Kathiresan et al. (1996c) demonstrated that
application of aliphatic alcohols can have a major stimulatory effect on mangrove
photosynthesis. Treatment with triacontanol (a long-chain aliphatic alcohol) increased the
photosynthetic rate of Rhizophora apiculata by 225%. A similar treatment with methanol
(a short-chain aliphatic alcohol) increased photosynthesis in R. mucronata by 612%.

3.4. Biochemistry

     Mangroves are biochemically unique, producing a wide array of novel natural
products. Excoecaria agallocha, for example, exudes an acrid latex that is injurious to the
human eye, hence its designation as “the blinding tree”. The latex is toxic to a variety of
marine organisms (Kathiresan and Thangam, 1987; Kathiresan et al., 1990b) and has
sublethal effects on the rice-field crab Oziotephusa senex senex, in which exposure
decreases whole-animal oxygen consumption and inhibits the ATPase system in gill and
hepatopancreas tissues (R. Ramamurthi et al., 1991). Soil bacteria and yeasts degrade the
toxic latex, preventing its accumulation in the mangal (Reddy et al., 1991).
     Researchers have isolated a variety of other mangrove compounds including
taraxerol careaborin and taraxeryl cis-p-hydroxycinnamate from leaves of Rhizophora
apiculata (Kokpol et al., 1990); 2-nitro-4-(2’-nitroethenyl phenol) from leaves of
Sonneratia acida (Bose et al., 1992); alkanes (46.7-97.9% wax) and triterpenoids (53.3%
wax) from leaves of Rhizophora species (Dodd et al., 1995); and iridoid glycosides from
leaves of Avicennia officinalis and A. germinans (Fauvel et al., 1995; Sharma and Garg,
1996). C.K.Rao et al. (1991) found arsenic in mangroves from the Goa Coast.
     Mangroves are also rich in polyphenols and tannins (Kathiresan and Ravi, 1990;
Ravi and Kathiresan, 1990; Achmadi et al., 1994). The levels of these substances may vary
seasonally (Basak et al., 1998), but older data should be interpreted cautiously since
standard methods for measuring tannins are very inaccurate for mangrove leaves (Benner
et al., 1990a).
     Substances in mangroves have long been used in folk medicine to treat disease
(Bandaranayake, 1998). Extracts have proven activity against human, animal and plant
pathogenic viruses including human immuno-deficiency virus (Premanathan et al., 1996),
Semliki forest virus (Premanathan et al., 1995), Tobacco Mosaic virus (Padmakumar and
Ayyakannu, 1997), Vaccinia virus (Premanathan et al., 1994a), Encephalomyocarditis
virus (Premanathan et al., 1994b), New castle disease virus (Premanathan et al., 1993), and
Hepatitis-B viruses (Premanathan et al., 1992). A few mangrove species, particularly those
belonging to the family Rhizophoraceae, show particularly strong antiviral activity
(Premanathan et al., 1992; Kathiresan et al., 1995a). Purified active fractions like acid
polysaccharides (galactose, galactosamine, glucose and arabinose) show potent anti-HIV
activity (Premanathan et al., 1999).
     Other unique mangrove biochemicals have potential commercial applications
(reviewed by Kathiresan, 2000). For example, mangrove extracts kill larvae of the
mosquitoes Anopheles stephensi (Thangam and Kathiresan, 1988), Culex tritaeniorhynchus
(Thangam and Kathiresan, 1989), Aedes aegypti (Thangam and Kathiresan, 1991, 1992a,
1994), and Culex quinquefasciatus (Thangam and Kathiresan, 1997). A pyrethrin-like
compound in stilt roots of Rhizophora apiculata shows strong mosquito larvicidal activity

(Thangam, 1990). Smoke from burned extracts repels and kills both Aedes aegypti
(Thangam et al., 1992) and Culex quinquefasciatus (Thangam and Kathiresan, 1992b) and
extracts applied directly to human skin repel adult Aedes aegypti (Thangam and
Kathiresan, 1993a).
    Phenols and flavonoids in mangrove leaves serve as UV-screening compounds.
Hence, mangroves tolerate solar-UV radiation and create a UV-free, under-canopy
environment (Moorthy, 1995). These substances also contribute to a black tea that can be
extracted from mangrove leaves (Kathiresan, 1995b). The “mangrove tea” is rich in
theaflavin, the substance responsible for the briskness and colour of tea. The tea, which
shows no mammalian toxicity, can be improved by UV irradiation (Kathiresan and
Pandian, 1991, 1993, Kathiresan, 1995b).
    Moorthy and Kathiresan, (1997a) proposed a physiological grouping of mangrove
species based on pigments, which may differ significantly among species (Basak et al.,
1996). Pigments concentrations may also vary with environmental conditions and season.
For example, Menon and Neelakantan (1992) found that total chlorophyll content was
positively related to light levels. Oswin and Kathiresan (1994) found that mangrove
chlorophyll and carotenoid levels, in general, are high during the summer but anthocyanin
levels are highest in the monsoon months. Flavonoids increase during the premonsoon

3.5. Pollination biology

     Mangroves have both self-pollinating and cross-pollinating mechanisms that vary
with species. For example, Aegiceras corniculatum and Lumnitzera racemosa are self-
pollinated. Avicennia officinalis is self-fertile, but can also cross-fertilize (Aluri, 1990). In
Avicennia marina, protandry makes self-pollination of an individual flower unlikely.
However, some fruits are set even when flowers are experimentally bagged to prevent
cross-pollination (between 4 and 41% of cross-pollinated flowers set fruit). Fruit abortion
is significantly higher in self-fertilized treatments, indicating some inbreeding depression
(Clarke and Myerscough, 1991a). There is a similar distinct trend for self-incompatibility
in Rhizophora, Ceriops and Sonneratia. This pattern is less clear in Bruguiera and
Kandelia (Ananda Rao, 1998).
     Mangroves are pollinated by a diverse group of animals including bats, birds, and
insects. Pollen is deposited on the animals as they deeply probe the flowers looking for
nectar; they subsequently transfer the pollen grains to the stigma of another flower. The
identity of the pollinators differs from species to species. Lumnitzera littorea, for example,
is pollinated primarily by birds while L. racemosa and small-flowered Bruguiera species
are pollinated by insects (Tomlinson, 1986). Sunbirds visit and may pollinate Acanthus
ilicifolius (Aluri, 1990) and large-flowered Bruguiera hainesii (Noske, 1993, 1995). Birds
are particularly important pollinators in the dry season when absence of terrestrial plant
flowers causes them to turn to mangroves as a food source.
     Bats are the major pollinators for Sonneratia, which opens its flowers to expose the
powdery stamens in the late night/early morning hours. If there are no bats, hawk moths
become the primary nighttime pollinators (Hockey and de Baar, 1991). Two lycaenid
butterflies may be important in the pollination of mangroves in Brisbane, Australia where

their abundance is directly correlated with the abundance of mangrove flowers (Hill,
1992). Bees regularly visit and pollinate species of Avicennia, Acanthus, Excoecaria,
Rhizophora, Scyphipora, and Xylocarpus. Some wasps and flies are highly dependent on
mangroves for nesting and are particularly important pollinators of Ceriops decandra,
Kandelia candel and Lumnitzera racemosa (Tomlinson, 1986). Rhizophora species
produce prolific amounts of pollen and are mainly wind-pollinated, though the stigma has
no special modifications to capture the wind-borne pollen (Tomlinson, 1986).

3.6. Reproduction, dispersal and establishment
    Bhosale and Mulik (1991) described four methods of mangrove reproduction:
viviparity, cryptoviviparity, normal germination on soil, and vegetative propagation.
Vivipary, the precocious and continuous growth of offspring while still attached to the
maternal plant, is a unique adaptation to shallow marine habitats (Thomas and Paul, 1996).
True viviparous species remain attached to the maternal plant for a full year while
cryptoviviparous offspring are only attached for 1-2 months (Bhosale and Mulik, 1991).
S.M. Smith and Snedaker (1995a) suggest that viviparous reproductive patterns allow
seedlings to develop some salinity tolerance before being released from the parent tree.
Figure 2c illustrates propagules of Rhizophora still attached to the parent.
    The timing of mangrove reproduction depends on local environmental conditions
and may differ broadly over the range of a species. For example, Duke (1990) found that
flowering in Avicennia marina occurred 6 months earlier in Papua New Guinea than in
Southern Australia and New Zealand. The period from flowering to fruiting was 2-3
months in the northern tropical site but stretched to 10 months in the southern temperate
locations. Flowering appeared to be controlled by daylength while air temperature set the
period for fruit maturation.
    Phytohormones are important in development, growth, and dispersal of mangrove
seeds, which may undergo no maturation drying, and remain metabolically active
throughout development (Farrant et al.; 1992, 1993). Phytohormones, like cytokinin
(particularly zeatin riboside) accumulate in both axes and cotyledons during reserve
accumulation. The level of abscissic acid (ABA) in the embryo stays low during this
period, making them sensitive to desiccation (though their dehydration tolerance increases
with development; Farrant et al., 1993). ABA levels in the pericarp increase throughout
seed development; the ABA in the pericarp may prevent precocious germination.
Farnsworth and Farrant (1998) suggest that ABA concentrations represent a trade-off
between salinity adjustment by the parental plant and developmental demands of the
embryo. Other biochemicals may be compartmentalized in the seeds. Mature propagules of
Rhizophora species exhibit high chlorophyll levels in the hypocotyl and high polyphenol
content in the radicle regions (Kulkarni and Bhosale, 1991).
    S.M. Smith et al. (1995) investigated the role of hormones in controlling flotation
and the development of roots and shoots in Rhizophora mangle propagules. Application of
gibberellic acid (GA3) caused the propagules to float horizontally, but painting with
naphthalene acetic acid (NAA) produced vertically floating propagules. NAA promoted
root elongation while GA3 enhanced stem elongation and leaf expansion (S.M. Smith et
al., 1996). A variety of hormones and chemicals (e.g., NAA, IBA, IAA, GA3, phenolics,
methanol, boric acids, triacontanol) promote root growth in propagules of other

Rhizophora and Avicennia species (Kathiresan and Thangam, 1990b; Kathiresan and
Moorthy, 1992, 1994a,b,c,d; Kathiresan et al., 1990a, 1994b, 1996b).
     Mangrove propagules have an obligate dispersal phase of several weeks before the
radicle extends for root development. If, however, the propagules do not contact the
sediment, they remain viable in seawater for several months (Clarke, 1993). Dispersal of
propagules depends on their buoyancy and longevity and on the activity of tides and
currents. The propagules of Kandelia candel are sensitive to light; high levels inhibit
rooting. Fan and Chen (1993) suggest that this is adaptive as it keeps the floating
propagules alive during potentially long dispersal periods. It is unclear, however, how
common it is for mangrove propagules to travel great distances. It has been experimentally
shown that most Avicennia marina propagules strand and establish close to their parents; it
is uncommon for them to move very far (Clarke and Myerscough, 1991b; Kathiresan and
Ramesh, 1991; Kathiresan, 1999). This conclusion is supported by the observation of
Saifullah et al. (1994) that dispersal only determines small-scale distributional patterns of
mangroves in Karachi, Pakistan. Larger-scale patterns are created by environmental
     Mangrove propagules may suffer high mortality during their dispersal. In field
studies, propagules of Ceriops tagal in northern Australia dispersed very short distances
(only 9% moved more than 3 m from the parent tree). Within that short distance, however,
a high percentage of them were damaged or eaten by predators (McGuinness, 1997a;
Figure 5). Farnsworth and Ellison (1997a) measured predation on mangrove propagules in
42 mangrove swamps in 16 countries and found rates ranging from 0 - 93% with a global
average of 28.3%. The major predators were grapsid crabs and insects in the Coeleoptera,
and Lepidoptera. In Kenya, grapsid crabs cleared nearly 100% of the seeds from landward
                                mangrove plantations (Dahdouh-Guebas
                                et al., 1998). Such high levels of seed
                                predation undoubtedly have significant
Percent of propagules

                                effects on population dynamics and stand
                   Not taken by predators

                                     Mortality is not restricted to
                                propagules. Mangroves are also
                                vulnerable during establishment and
              Not taken or damaged
                                early growth. In Belize, mortality of R.
                 by predators
                                mangle and A. germinans is highest
    0    20    40     60     80     100
                                during establishment. The mortality can
                                be attributed to (1) a failure to establish
                                before seed viability is lost, (2)
Figure 5. Loss of Ceriops tagal propagules to
predators in a northern Australia mangal. Propagules      predation, and (3) desiccation (Ellison
were marked and tethered then monitored for           and Farnsworth, 1993).
disappearance and damage. Crab predators removed or
                                     After establishment, survival is
damaged 83% of the propagules within the first 90
                                strongly influenced by physicochemical
days (after McGuinness 1997a).
                                stresses. For example, shading,
orientation of the seedling axis (e.g., upright vs. horizontal), soil fertility, and flooding can
all have significant impacts on survival (Hovendon et al., 1995; McKee, 1995a; Koch,
1997; McGuinness, 1997a). Post-establishment growth is also affected by a suite of
physical and chemical factors. Experimental work with Rhizophora species demonstrates

that propagule length, planting depth, soil type, salinity, concentration of leachates, pH and
light intensity are important determinants of growth (Kathiresan and Thangam, 1989,
1990a; Kathiresan and Ramesh, 1991; Kathiresan and Moorthy, 1993; Kathiresan et al.,
1993; Kathiresan et al., 1995b, 1996a; Kathiresan, 1999). Seedling growth can be
artificially stimulated by application of triacantanol and methanol. Both of these substances
increase the photosynthetic rate of the seedlings, the in vivo nitrate reductase activity, the
growth of roots and shoots, the protein and energy contents of leaves and roots, the
chlorophyll and carotenoid content in leaves, and the amount of chlorophyll in
photosystems I and II and in the light harvesting complex of the chloroplasts (Moorthy and
Kathiresan, 1993; Kathiresan and Moorthy, 1994a; Kathiresan et al., 1996a).
     New mangrove growth comes primarily from seeds and density of newly
established individuals can be very high (seedling densities reach 27,750 individuals • ha-1
in the Sunderbans of Bangladesh; Siddiqi, 1997). Vegetative regrowth from stump sprouts
(“copicing”) also occurs in some species (e.g., Excoecaria, Avicennia, Laguncularia,
Sonneratia; Tomlinson, 1986). Recently an air-layering technique has been used to
successfully induce vegetative propagation in Avicennia alba, A. officinalis, Sonneratia
apetala, Xylocarpus granatum and Rhizophora mangle. The technique was not successful
for A. marina or Kandelia (Kathiresan and Ravikumar, 1995a; Calderon and Echeverri,
1997; Ananda Rao, 1998). External application of auxins can stimulate growth of newly
planted mangrove cuttings. The auxins produce metabolic changes during initiation and
development of roots, enhancing levels of reducing sugars and increasing the mobilization
of nitrogen to the rooting zone (Basak et al., 1995; Das et al., 1997).

3.7. Biomass and litter production

     Mangroves and mangrove habitats contribute significantly to the global carbon
cycle. Mangrove forest biomass may reach 700 t ha-1 (Clough, 1992, Table 2) and Twilley
et al. (1992) estimate the total global mangrove biomass to be approximately 8.7 gigatons
dry weight (i.e., 4.0 gigatons of carbon). Accurate biomass estimates require measuring
volumes of individual trees. Da Silva et al. (1993) have developed equations for making
such measurements on living mangroves.
     Mangroves generally grow better in wet equatorial climates than they do in
seasonally monsoonal or arid climates (Clough, 1992) and the amount of litter they
produce is negatively correlated with latitude. Estimates of the annual global litterfall from
mangroves range from 130 to 1870 g m-2. In general, the litterfall is heaviest 1) in dry
summer months when thinning of the canopy reduces transpiration, and 2) in the wet rainy
season when fresh water input increases the nutrient supply (Roy, 1997; Wafar et al.,
1997). However, individual species may differ in the conditions that produce heavy litter.
For instance, Australian Rhizophora stylosa and Avicennia marina show heaviest litterfall
in hot climates with short dry seasons, but Ceriops tagal litterfall is heaviest in hot climates

Table 2. Mangrove standing biomass measurements.

Location          Species      Biomass    Amount (t •  Reference

Cuba (North America)    R. mangle    Roots        31.3   Fiala and Hernandez, 1993
              A. germinans  Roots        24.4

French Guiana (S.      Mixed forest  Total       31 – 315   Fromard et al, 1998

Mgeni Estuary (S. Africa)  Mixed forest  Above-ground     94.4   Steinke et al., 1995
              A. germinans  Below-ground     9.6

Sunderbans (India)     Avicennia sp.  Total        147.7   Choudhuri, 1991
(6 yr old trees)      B.                  11.2
              S. apetala              34.5
              C. tagal               4.8

Tritih, Java (Indonesia)  R. mucronata  Above-ground     93.7   Sukardjo  and   Yamada,

Matang mangal        Mixed forest  Total        202.4   Gong and Ong, 1990

Hainan Island, (China)   Mixed forest  Total       9.6-14.2   Liao et al., 1993
              S. caseolaris  Total        47.2   Liao et al., 1990

Near Brisbane (Australia)  A. marina    Above-ground   110-340   Mackey, 1993
                      Below-ground+   109-126

Mary River (Australia)   A.       Above-ground/   40/50    Saintilan, 1997
              corniculatum  below ground
              A. marina    Above-ground/   150/80
                      below ground
              E. agallocha  Above-ground/   140/40
                      below ground
              R. stylosa   Above-ground/   70/100
                      below ground
              C. australis  Above-ground/   110/50
                      below ground

with a long dry winter (Bunt, 1995). In India, Avicennia marina litter production is high in
the post-monsoon period and low in the pre-monsoon season (Ghosh et al., 1990).
Deviations from these general patterns of litterfall may result from habitat-specific stresses
(e.g., aridity, poor soils; Saenger and Snedaker, 1993; Imbert and Ménard, 1997).
     A number of researchers have measured mangrove litterfall. Results show a broad
range of litter volumes with production varying significantly from habitat to habitat. The
production appears to depend largely on local conditions, species composition, and

productivity of the individual mangal. Litter production has been variously measured at
0.011 t ha-1 y-1 in the mangroves of Kenya, 9.4 t ha-1 y-1 in Bermuda, and 23.69 t ha-1
y-1 in Australia (Table 3).
Table 3. Litter production in mangrove forests.

Location                 Species     Litter production   Reference
                             (t • ha-1 • yr-1 )

Guyana (South America)        A. germinans      17.71       Chale, 1996

Teacapan-Ague Brava Lagoon      Mixed forest      14.17       Flores-Verdugo et al, 1990

Bermuda (North America)        Mixed forest      9.40       Ellison, 1997

Bonny estuary (Nigeria)        R. racemosa       8.46       Abbey-Kalio, 1992
                   A. africana       6.41
                   Laguncularia sp.    8.18

South Africa             Mixed forest      4.50       Steinke and Ward, 1990

Gazi Bay (Kenya)           R. mucronata      0.02       Slim et al, 1996
                   C. tagal        0.01

Andaman Islands (India)        Mixed forest     7.10 - 8.50    Mall et al, 1991
                   B. gymnorrhiza    5.11 –7.09     Dagar and Sharma, 1993
                   R. apiculata     8.08 – 10.30    Dagar and Sharma, 1991

Mandovi-Zuari Estuary (India)     R. apiculata      11.70       Wafer et al, 1997
                   R. mucronata      11.10
                   S. alba        17.00
                   A. officinalis     10.20

Fly River Estuary (New Guinea)    Mixed forest     8.00 – 14.00    Twilley et al, 1992

Matang mangal (Malaysia)       Mixed forest      3.90       Gong and Ong, 1990

Jervis Bay, NSW (Australia)      A. marina        3.10       Clarke, 1994
                   A. corniculatum     2.10

Embley River (Australia)       R. stylosa       12.23       Conacher et al., 1996
                   C. tagal        5.39
                   A. marina        6.28

Australia               A. marina       15.98       Bunt, 1995
                   R. stylosa       23.69
                   C. tagal        12.90

    Litter from the mangroves is composed of leaves, twigs, branches, and seeds. Seeds
alone accounted for 25% of the total litterfall for Avicennia germinans and Rhizophora

mangle in a mangrove habitat in Martinique (Imbert and Ménard, 1997). In a temperate
mangal, the reproductive material was approximately 9% of the total for Avicennia marina
and 32% of the total for Aegiceras corniculatum. Clarke (1994) suggested that such
relatively high reproductive output may contribute to the low productivity and stunting of
mangroves at high latitude.
     Accumulated mangrove litter may wash into rivers and streams when rain or tides
inundate the forest. Consequently, mangrove litter may decompose either in the source
forest or in the river, with nutrients being retained or exported (Conacher et al., 1996).
Whether the litter (and its nutrients) remain in the habitat or are exported by water flow
may depend largely on the local animal community. On the east coast of Queensland, the
litter accumulation in a Ceriops forest was 6 g m-2 (0.06 t ha-1) while in an Avicennia
forest, it was closer to 84 g m-2 (0.84 t ha-1; Robertson et al., 1992). This enormous
difference in accumulation was attributed to the feeding activities of crabs.

4.1. Bacteria

    Mangroves provide a unique ecological environment for diverse bacterial
communities. The bacteria fill a number of niches and are fundamental to the functioning
of these habitats. They are particularly important in controlling the chemical environment
of the mangal. For example, sulfate-reducing bacteria (e.g., Desulfovibrio,
Desulfotomaculu, Desulfosarcina, and Desulfococcus; Chandrika et al., 1990; Loka-
Bharathi et al., 1991) are the primary decomposers in anoxic mangrove sediments. These
bacteria largely control iron, phosphorus, and sulfur dynamics and contribute to soil and
vegetation patterns (Sherman et al., 1998). Methanogenic bacteria are seasonally abundant
in sediments where Avicennia species dominate (T. Ramamurthy et al., 1990; Mohanraju
and Natarajan, 1992). Subsurface bacterial communities (along with epibenthic
microalgae) may sequester nutrients and hold them within nutrient-limited mangrove muds
(Alongi et al., 1993; Rivera-Monroy and Twilley, 1996).
    Bacteria are critical to the cycling of nitrogen in mangrove environments. Marine
cyanobacteria are a particularly important component of the microbiota, constituting a
source of nitrogen in every mangrove system (Sheridan, 1991, 1992; Hussain and Khoja,
1993; Krishnamurthy et al., 1995a; Palaniselvam, 1998). N2-fixing cyanobacteria isolated
from Avicennia pneumatophores in the Beachwood Mangrove Reserve, South Africa
supply 24.3% of the annual nitrogen requirements of that swamp. The N2-fixation rates are
controlled by light and temperature and show seasonal trends (low in the winter and high
in the summer; Mann and Steinke, 1993). Fixation rates are higher when the cyanobacteria
are on the mangrove than when they are held on an artificial growth medium (Toledo et al.,
    N2-fixing bacteria are efficient at using a variety of mangrove substrates despite
differences in carbon content and phenol concentrations (Pelegri and Twilley, 1998).
However, their abundance may be dependent on physical conditions and mangrove
community composition. N2-fixing Azotobacter, which show potential as biofertilizers, are
abundant in the mangrove habitats of Pichavaram, south India. Their abundance in the
mangal exceeds that in marine backwaters and estuarine systems (S. Ravikumar, 1995).
Sengupta and Choudhuri (1991) studied N2-fixing bacteria in a Ganges River mangrove

community. They found high numbers in the rhizospheres of plants in inundated areas but
plants on occasionally inundated ridges and in degraded areas had fewer rhizosphere
bacteria. Ogan (1990) found similar distinct differences in nodulation and nitrogenase
activity among sites and among species in a Nigerian mangal.
     Two halotolerant N2- fixing Rhizobium strains have been isolated from root nodules
of Derris scandens and Sesbania species growing in the mangrove swamps of Sunderbans
(Sengupta and Choudhuri, 1990). If the non-N2-fixing bacteria are removed from the
rhizosphere, N2-fixing activity drops, indicating that other rhizosphere bacteria contribute
to the fixation process (Holguin et al., 1992). The non-N2 fixer, Staphylococcus sp.,
isolated from mangrove roots, promotes N2-fixation by Azospirillum brasilense. This can
be achieved by growing the two species in mixed culture or simply by adding a cell-free
dialysate of the Staphylococcus sp. to the A. brasilense culture. Aspartic acid is the
compound responsible for the effect (Holguin and Bashan, 1996).
     In addition to processing nutrients, mangrove bacteria may also help process
industrial wastes. Iron-reducing bacteria are common in mangrove habitats in some mining
areas (Panchanadikar, 1993). Eighteen bacterial isolates that metabolize waste drilling fluid
have been collected from a mangrove swamp in Nigeria (Benka-Coker and Olumagin,
1995). Interestingly, four additional bacterial strains isolated from the same swamp depress
growth rates of Staphylococcus and Pseudomonas species and could, therefore, decrease
normal rates of organic decomposition (Benka-Coker and Olumagin, 1996).
     Bacteria play a number of other roles in the mangal. Some live symbiotically with
other organisms. For example, rod bacteria can be commonly found in the hindguts of
mangrove detritivores (Harris, 1993) and deeply branched sulfur-oxidizing bacteria occur
as endosymbionts within members of the bivalve family Lucinacea in sulfide-rich, muddy
mangrove areas. Bauer-Nebelsick et al. (1996) and Ott et al. (1998) have described sulfur-
oxidizing bacteria that live as obligate ectosymbionts on colonial sessile ciliates
(Zoothamnium niveum) in a Belizian mangal.
     Other mangrove bacteria are parasitic or pathogenic. Bdellovibrios capable of
parasitizing Vibrio spp. are common in an Australian mangrove habitat. Their abundance
there (36.6 ml-1) is much higher than in nearby Great Barrier Reef habitats (9.5 ml-1;
Sutton and Besant, 1994). Also in Australia, Bacillus thuringiensis, which shows
insecticidal activity against mosquito larvae of Anopheles maculatus, Aedes aegypti and
Culex quinquefasciatus, has been isolated from mangrove sediments (Lee et al., 1990a;
Lee and Seleena, 1990). Actinomycetes (fungi-like bacteria) that occur in many mangrove
habitats (Kala and Chandrika, 1993; Vikineswary et al., 1997) may show antifungal
activity (Vikineswary et al., 1997).
     Bacterial populations show distinct spatial distribution patterns. Many live
epiphytically on the surfaces of mangroves, but different species appear to prefer different
parts of the tree. Leaves of Avicennia marina and Sesuvium portulacastrum harbour large
numbers of Flavobacterium while roots and stems have large populations of Vibrio spp.
(Abhaykumar and Dube, 1991). In many species, the aerial roots, especially
pneumatophores, harbour particularly dense bacterial cyanopopulations that may show
sharp vertical zonation. Coccoid forms occur in the upper zone of the pneumatophores.
Filamentous non-heterocystous forms predominate in the middle zone, and filamentous
heterocystous forms are largely restricted to the lower zones (Toledo et al., 1995a;
Palaniselvam, 1998). In the forests of Aldabra Lagoon, heterocystous forms like
BIOLOGY OF MANGROVES AND MANGROVE ECOSYSTEMS                                29

Scytonema sp. also form conspicuous growths on pneumatophores, but non-heterocystous
species are restricted to the sediment surface (e.g., Alongi and Sasekumar, 1992).
     Cyanobacteria in the mangal colonize any submerged surface including sediments,
roots, aerial roots, branches and trunks (Sheridan, 1991). Microbial mats in mangrove tidal
channels often have an outer layer of cyanobacteria
and a reddish inner layer of anoxygenic            100

phototrophic bacteria (Lopez-Cortes, 1990). A          80
cyanobacterium (Calothrix viguieri) isolated from

                                   % hairiness
the surface of mangrove roots show a peculiar
morphological response to salinity variation. In low      40

salinity, it develops hairs (Figure 6). The hairs are      20
shed if salinity is increased . The hairs may be an
adaptation to hydrolyze pulses of organic             0   6      12      18   24

phosphorus that occur in the habitat after heavy        200

rains (Mahasneh et al., 1990).

                                   Mean hair length (µm)
     Bacterial counts are generally higher on       150

attached mangrove vegetation than they are on         100
fresh leaf litter. This is probably because attached,
undamaged leaves leak amino acids and sugars but        50

do not release much tannin (Kathiresan and
Ravikumar, 1995b). Shome et al. (1995) isolated         0
                                  0   6      12      18   24
thirty-eight distinct bacteria from mangrove leaf           Time after transfer to fresh water (h)

litter and sediments in south Andaman and           Figure 6. Effects of salinity on hair
                                formation in the mangrove bacterium
characterized the bacterial community. The bacteria
were generally gram-positive (76.3%), motile (87%),
fermentative (6.9-82.1%), pigmented (31%), and antibiotic resistant (100% against
polymixin B and 50% against chloramphenicol). Photosynthetic bacteria, including purple
sulfur bacteria (Chromatium spp.) and purple non-sulfur bacteria (Rhodopseudomonas
spp.), have been isolated from mangroves in Pichavaram, south India (Vethanayagam,
1991; Vethanayagam and Krishnamurthy, 1995). Nine species of purple non-sulfur
bacteria have also been found in mangroves of Egypt (Shoreit et al., 1994). Growth of the
purple sulfur bacteria in these habitats is limited by low light and sulfide. In contrast, high
light and sulfide limit growth of green sulfur bacteria (Chandrika et al., 1990).

4.2. Fungi and fungus-like protists

    Mangals are home to a group of fungi called “manglicolous fungi.” These
organisms are vitally important to nutrient cycling in these habitats (Hyde and Lee, 1995;
Kohlmeyer et al., 1995). Kohlmeyer and Kohlmeyer (1979) were the first to review this
group. They recognized 43 species of higher fungi, including 23 Ascomycetes, 17
Deuteromycetes, and 3 Basidiomycetes. Hyde (1990a) listed 120 species from 29
mangrove forests around the world. These included 87 Ascomycetes, 31 Deuteromycetes,
and 2 Basidiomycetes.
    Work in individual habitats has revealed surprisingly diverse fungal communities
(e.g., Hyde, 1990b; Hyde, 1996). Chinnaraj (1993a) identified 63 species of higher fungi in

mangrove samples from Andaman and Nicobar Islands alone. Similar samples from
Lakshadweep Island yielded 32 species (Chinnaraj, 1992) and 39 species were found in
mangrove samples from the Maldives (Chinnaraj, 1993b). D.R. Ravikumar and Vittal
(1996) found 48 fungal species in decomposing Rhizophora debris in Pichavaram, south
India. On the Indian Ocean coast of South Africa, Steinke and Jones, (1993) identified 93
species of marine fungi, including 55 from mangrove wood (particularly Avicennia
marina). Table 4 lists some of the fungal species identified in these studies.
Table 4. Some fungal species isolated from mangrove habitats.

Species                         Author

Aigialus striatispora                  Hyde (1992c)
Aniptodera longispora                  Hyde (1990b)
Aniptodera salsuginosa                 Nakagiri and Ito (1994)
Calathella mangrovei                  Jones and Agerer (1992)
Cryptovalsa halosarceicola               Hyde (1993)
Eutypa bathurstensis                  Hyde and Rappaz (1993)
Falciformispora lignatilis               Hyde (1992d)
Halophytophthora kandeliae               Ho et al (1991)
Halophytophthora kandeliae               Newell and Fell (1992b)
Halophytophthora vesicula                Newell and Fell (1992b)
Halophytophthora. spinosa                Newell and Fell (1992b)
Halosarpheia minuta                   Leong et al (1991)
Hapsidascus hadrus                   Kohlmeyer and Kohlmeyer (1991)
Hypoxylon oceanicum                   Whalley et al (1994)
Julella avicenniae                   Hyde (1992a)
Khuskia oryzae                     Pal and Purkayastha (1992a)
Lophiostoma asiana                   Hyde (1995)
M. ramunculicola                    Hyde (1991b)
Massarina armatispora                  Hyde et al (1992)
Massarina velatospora                  Hyde (1991b)
Payosphaeria minuta                   Leong et al (1990)
Pedumispora                       Hyde and Jones (1992)
Phomopsis mangrovei                   Hyde (1991a)
Saccardoella                      Hyde (1992b)
Trematospaeria lineolatispora              Hyde (1992d)

    Surveys are revealing a number of range extensions, new species, and even new
genera. Collections of mangrove fungi in Macau and Hong Kong, for instance, yielded 45
species. Twenty-eight of these were new records for Macau and 21 were new records for
Hong Kong (Vrijmoed et al., 1994). These discoveries are leading to significant taxonomic
revision of these groups (Jones et al., 1994, 1996; Alias et al., 1996; Goh and Yipp 1996;
Ho and Hyde, 1996; Vrijmoed et al., 1996; Honda et al., 1998).
    The fungus-like thraustochytrids are important endobionts in dead or living plants
and in calcareous shells (S. Raghukumar, 1990). A number of these occur in mangrove
swamps where they help decompose mangrove leaf litter (S. Raghukumar et al., 1994;
Bremer, 1995). Both thraustochytrids and chytridiomycetes (including Schizochytrium,

Thraustochytrium and Ulkenia) have been isolated from Costa Rican mangrove swamps
(Ulken et al., 1990). Two thraustochytrids, Thraustochytrium striatum and Schizochytrium
mangrovei, have been isolated from an Indian mangal at Goa. Both produce amoebae-like
structures, move using pseudopodia, and phagocytose bacterial cells (S. Raghukumar,
     Marine oomycetes (fungus-like protists) also occur in mangrove communities. T.K.
Tan and Pek (1997) found five Halophytophthora species in Singapore mangroves. This is
the first time three of them have been seen in tropical mangroves and the first time one has
been reported outside Australia. Oomycetes in the genus Halophytophthora have special
importance in mangrove habitats (Newell and Fell, 1992a). They greatly facilitate the
decomposition of mangrove material. Newly fallen Rhizophora mangle leaves are quickly
infested with mycelial growths of Halophytophora vesicula and H. spinosa. Rapid lateral
extension of the mycelia within the leaves apparently follows establishment of a single
zoospore (Newell and Fell, 1995). In laboratory cultures, the established Holophytophora
are subsequently colonized by bacteria and labyrinthulas (Newell and Fell, 1994).
Holophytophora species are generally good competitors against true fungi but have
difficulty colonizing leaves that already have bacterial films (Newell and Fell, 1997).
     Newell and Fell (1996) speculate that Halophytophora completes its colonization
of submerged leaves, from attachment of zoospore cysts to release of new zoospores, in the
early stages of leaf decomposition, before there is substantial entry into the leaves
themselves. Mild drying, low salinity and low temperatures may enhance zoospore release.
The release rates are low in older, decaying leaves and high in newer, less-decayed leaves
(Newell and Fell, 1996). Leaño et al. (1998) showed that the zoospores of other mangrove
fungi are chemically attracted to plant material and extracts. This undoubtedly aids in the
colonization of new substrata.
     A few researchers have studied the physiology and biochemistry of manglicolous
fungi. Many of the species produce interesting compounds. For example, most of the soil
fungi produce lignocellulose-modifying exoenzymes like laccase (C. Raghukumar et al.,
1994). Preussia aurantiaca synthesizes two new depsidones (Auranticins A and B) that
display antimicrobial activity (Poch and Gloer, 1991). Cirrenalia pygmea produces
melanin pigments that appear to protect the hyphae from sudden changes in osmotic
pressure; when melanin synthesis in cultures is inhibited with tricyclazole, the fungus
becomes sensitive to osmotic shock (Ravishanker et al., 1995). High salinities also
increase the number and types of amino acids this species produces (Ravishankar et al.,
     Ascus and ascospore ultrastructure have been studied in the fungi Swampomyces
armeniacus and Marinosphaera mangrovei (Read et al., 1995) and in Dactylospora
haliotrepha (Au et al., 1996). Ascocarp formation has been tested in single and mixed
cultures of Aigialus parvus, Lignincola laevis and Verruculina enalia growing on the wood
of Avicennia alba, Bruguiera cylindrica and Rhizophora apiculata. Sporulation was
delayed and fewer ascocarps were formed in mixed cultures, suggesting competition
among the fungi (T.K. Tan et al., 1995).
     A number of fungal species live directly on living mangroves but, in general, they
are not well known. Sivakumar and Kathiresan (1990) isolated ten fungal species from leaf
surfaces of seven mangrove species. The dominant phylloplane fungi were Alternaria
alternata, Rhizopus nigricans, Aspergillus and Penicillium spp. Abundances of these fungi

were negatively correlated with tannin content of the leaves. The fungi appear to prefer
leaf litter (which contain more amino acids) to fresh leaves (which contain more tannins
and sugars; S. Ravikumar and Kathiresan, 1993). Other fungi are harmful to the living
mangroves. Two new parasitic species (Pestalotiopsis agallochae and Cladosporium
marinum) have been isolated from the leaves of Excoecaria agallocha and Avicennia
marina (Pal and Purkayastha, 1992b). Pathogenic fungi may have contribute to diebacks of
Rhizophora mangle stands in Costa Rica (Tattar et al., 1994).
     A number of fungal species colonize subsurface mangrove roots. Nair et al. (1991)
found 25 fungal species from 15 genera in the rhizosphere of Avicennia officinalis;
adjacent non-rhizosphere soil held only 16 species from 10 genera. Sengupta and
Choudhuri (1994) found Rhizoctonia and VA-mycorrhiza-like fungi in the mangrove
community at Sunderbans. When Cajanas seedlings in nutrient-poor conditions were
inoculated with the VA-mycorrhizal isolates, there was a significant increase in growth.
This was due, in part, to mobilization of insoluble phosphate by the fungus.
     Distributions of fungal species within the mangrove habitat may reflect physical
conditions and/or habitat preference. It may also reflect age of the stand. Working in Belize
(Central America), Kohlmeyer and Kohlmeyer (1993) found that fungal diversity depends
on age of the mangrove stand. They discovered 43 species in established Rhizophora
stands but only 7 in recently introduced Rhizophora. Some species may be quite specific in
their habitat preferences. For example, of the 48 fungal species Ravikumar and Vittal
(1996) found in a south Indian mangal, 44 were on prop root while seedling and wood
samples only held 18 and 16 species respectively. The fungal species appeared to partition
the mangrove habitat. Verruculina enalia was most abundant on prop roots and seedlings
while Lophiostoma mangrovei was most common on wood. Physical conditions, or genetic
differentiation created by isolation, may lead to differences in fungal morphology and
physiology. Pestalotiopsis versicolor strains, isolated from Ceriops decandra growing in
different regions of the Sunderbans, vary in mycelial mat texture, growth rate and
sporulation intensity (Bera and Purkayastha, 1992).
     Differences in physical requirements may lead to vertical zonation of the fungi.
Hyde (1990b) found 57 intertidal fungal species on Rhizophora apiculata at Brunei
mangal. Most of these occurred above the mean tidal level. A similar study of senescent
Acanthus ilicifolius at Mai Po, Hong Kong revealed that the apical portions of the trees are
colonized by typical terrestrial fungi but the basal portions are colonized by marine species
(Sadaba et al., 1995). The authors attributed this to the nature of the substratum and the
frequency of tidal inundation. Other fungal species live directly on the sediment surface
but are still entirely restricted to mangrove habitats (Soares et al., 1997).
     Wood degrading fungi are well-known in mangrove habitats. Thirty species of such
lignicolous fungi have been recorded in Malaysian mangals. The most abundant are
Halosarpheia marina, Lulworthia sp., Lignincola laevis, Halosarpheia retorquens, Eutypa
sp., Kallichroma tethys, Marinosphaera mangrovei, Phoma sp. and Julelia avicenniae.
Diversity and abundance are greatest on Avicennia wood (T.K. Tan and Leong, 1992; Alias
et al., 1995). Test panels of different woods placed in mangrove waters along the Goa
coast of India showed four common lignicolous fungi (Periconia prolifica, Lignincola
laevis, Aniptoder sp. and Lulworthia sp.). Panels treated with copper chrome arsenic were
more resistant to fungal infestation than those treated with chrome boric (Santhakumaran et
al., 1994).

    Nakagiri and Ito (1994) found a new lignicolous fungus (Aniptodera salsuginosa)
with unique ascospore appendages and an unusual ascus apical apparatus on decomposing
mangrove wood. The ascospore appendages are functional only when they are submerged
in brackish water. The ascospores are discharged through a fissure in the ascus wall at the
margin of the apical disc; the ascus pore in this disc does not function in ascospore release.

4.3. Microalgae

    Phytoplankton and benthic microalgal communities make important contributions
to the functioning of mangrove environments. However, their contribution to total
estuarine production is relatively small in most regions of southeast Asia, Australia,
Central America and tropical South America. Robertson and Blaber (1992) suggested that
the contribution of plankton to total net production in mangrove habitats ranges from 20-
50%. Careful measurements are verifying that predication for large systems. Phytoplankton
are responsible for 20% of the total production in mangrove estuaries in the Fly River
Delta in Papua New Guinea (Robertson et al., 1991, 1992) and 20-22% of the total
production in the Pichavaram mangroves of south India (Kawabata et al., 1993).
    Phytoplankton contributions to productivity in localized mangrove areas may be
much smaller. Lee (1990) found that phytoplankton and benthic macroalgae together
contribute less than 10% of the net primary production in Hong Kong mangals and Boto
and Robertson (1990), using nitrogen measurements, estimated that benthic cyanobacteria,
microalgae and macroalgae together contribute only 6% of the gross primary production in
mangrove ecosystem of northeastern Australia. Robertson and Blaber (1992) state that
phytoplankton productivity is significantly lower in estuarine mangrove areas than it is in
lagoons or open embayments fringed by mangroves.
    High turbidity, large salinity fluctuations and a generally small ratio of open
waterway to mangrove forest area contribute to the low light levels and shading that limit
productivity of the microalgae, especially the benthic forms (Alongi, 1994; Harrison et al.,
1994). High summer temperatures may also limit production (Lee, 1990). Rates of primary
production, which are generally low in the dry season, increase on ebb tides and decrease
on flood tides (Kitheka, 1996).
    In the Fly River delta of Papua New Guinea, Robertson et al., (1992) measured
very low production rates of only 0.022 to 0.0693 g C m-3 d-1. However, localized
conditions may lead to much higher rates. For example, daily production in the coastal
lagoons of Mexico may reach 2.4 g C m-3 d-1 (Robertson et al., 1992). Increased
productivity may relate to elevated nutrient levels. Production in lagoons of the Ivory
Coast reach 5 g C m-3 d-1. However, the effect is largely a result of nitrogen and
phosphorus input from nearby human population centers.
    Naturally occurring substances may also regulate phytoplankton growth. Selvam et
al. (1992) found phytoplankton productivity to be four times higher in mangrove waters
than in adjacent marine waters in south India. Refractive materials like humic acid, which
are abundant in the mangroves, stimulate phytoplankton growth there (Schwamborn and
Saint-Paul, 1996). In the Celestun Lagoon (northern Yucatan Peninsula, Mexico) low

concentrations of natural phenolics stimulate phytoplankton growth, but higher wintertime
levels depress the growth rates (Herrera Silveira and Ramirez Ramirez, 1996).
    While microalgae may make only small contributions to total productivity in
estuarine mangrove systems, they may be critical to supporting higher trophic levels
(Robertson and Blaber, 1992). This may be particularly true because of the high nutritional
quality of phytoplankton relative to mangrove detritus. Phytoplankton biomass,
productivity, and size are closely tied to diversity and abundance of higher trophic levels.
Teixeira and Gaeta (1991) determined the composition of the phytoplankton community in
a Brazilian mangal. Nanoplankton (cells from 2 - 20 µm) constituted over 80% of the total
phytoplankton. Laboratory testing showed that the smaller cells were responsible for a
significant part of the total productivity. Picoplankton (cells < 2 µm) accounted for 3-29%
of the total 14C uptake. The effects of this skewed phytoplankton size distribution on the
zooplankton community composition has not been studied.
    Despite relatively low productivity, mangrove phytoplankton communities can be
quite diverse. However, composition and density of the plankton community are strongly
affected by local environmental conditions (Lee, 1990). For example, low phytoplankton
diversity in Rhizophora habitats is related to the release of tannins by roots, decomposing
wood, and leaves (Robertson and Blaber, 1992). Phytoplankton populations also respond to
temperature and salinity variation. Thus, communities may show marked seasonal
variation (Mani, 1994). Phytoplankton studies at West Bengal, India revealed 46 species of
Bacillariophyceae, Dinophyceae and Cyanophyceae (Santra et al., 1991). Coscinodiscus,
Rhizosolenia, Chaetoceros, Biddulphia, Pleurosigma, Ceratium and Protoperidinium were
the dominant genera, existing almost year round. At least 82 phytoplankton species (72%
diatoms,15% dinoflagellates) occur in the Pichavaram mangroves of south India (Kannan
and Vasantha, 1992). The diatoms Nitzschia closterium, Pleurosigma spp., Thalassionema
nitzschioides and Thalassiothrix frauenfeldii are most abundant. Thirty-one of those
species may form seasonal blooms (Mani, 1992). Chaghtai and Saifullah (1992) reported
such a bloom of the diatom Navicula in the Karachi mangroves of Pakistan.
    Dinoflagellate assemblages have been particularly well studied in Belizean
mangrove habitats where a diverse collection of benthic and epiphytic species exists
(Faust, 1993a,b,c,d; Faust and Balech, 1993). Many are new species (e.g., Prorocentrum
maculosum, P. foraminosum, P. formosum, Plagiodinium belizeanum, Sinophysis
microcephalus). Faust and Gulledge (1996) found many microalgal species associated with
floating mangrove detritus. Dinoflagellates constituted the greatest proportion (50-90%),
followed by diatoms (5-15%), cyanobacteria (3-25%) and dinoflagellate cysts (1-7%).
Ciliates and nematodes were the major dinoflagellate consumers in the detritus.

4.4. Macroalgae

    The macroalgal flora is rich in mangrove habitats where it contributes to production
while also providing habitat and food for a number of invertebrate and fish species. Red
algae, especially in the genera Bostrychia, Caloglossa and Catenella, are most commonly
associated with mangroves and may be quite abundant. For instance, the total annual
biomass of Bostrychia tenella in a south Nigerian estuary reaches 1.84 mg • cm-2, which is
38% of the total algal production there (Ewa-Oboho and Abby-Kalio, 1993). The biomass
of algae in the mangrove lagoons of Puerto Rico is similar to the total annual leaf litterfall

from the Rhizophora fringe, leading to an algal-dominated foodweb (Rodriguez and
Stoner, 1990).
    Algal diversity can also be quite high in mangrove environments. Recent surveys
have revealed diverse macroalgal communities in Papua New Guinea (25 species; King,
1990), the Nicobar Islands in the Andaman Sea (61 species; Jagtap, 1992), and the coast of
Mauritius (127 species; Jagtap, 1993). King and Puttock (1994) and King (1995) provide
exhaustive reviews of the very diverse Australian mangrove macroalgal flora. Algal
assemblages tend to be richest in shallow areas with a mixture of hard and soft substrates.
Lowest diversity occurs where there is low light, soupy muds, or homogeneous, large-grain
sands (as in the Netherlands Antilles, Kuenen and Debrot, 1995).
    Algal surveys have produced new records for a number of species including
Stictosiphonia kelanensis from Atlantic mangroves (Fujii et al., 1990); Bostrychia pinnata,
Bostrychia simpliciuscula, Caloglossa angustalata (Rhodophyta) and Boodleopsis
carolinensis (Chlorophyta) from Singapore (West, 1991a); Bostrychia pinnata, Caloglossa
ogasawaraensis, C. stipitata and Halochlorococcum operculatum from Peru (West,
1991b); Bostrychia pinnata and Caloglossa ogasawaraensis from the Atlantic coast, USA
(West and Zuccarello, 1995); Bostrychia calliptera from the Central Gulf of Mexico
(Collado-Vides and West, 1996) and C. ogasawaraensis, C stipatata, C. lepricuriiI, B.
moritziana, B. pinnata, B. radicans and Catenella caespitosa in Southern Mexico and
Guatemala (Pedroche et al., 1995).
    Recent work has investigated genetic differentiation of some of the widely
distributed red algae. Male Caloglossa ogasawaraensis from a Peruvian mangle readily
hybridize with female C. ogasawaraensis from Brazil, producing viable tetrasporophytes
(West, 1991b). Similar studies with Bostrychia radicans from the Pacific and Atlantic
coasts of North America have been done. Almost all isolates from the northern Pacific
coast of Mexico are compatible and produce cystocarps that release viable carpospores.
However, isolates from the Atlantic coast of the United States show greater incompatibility
(Zuccarello and West, 1995).
    Algal abundance and diversity are largely determined by the physico-chemical
characteristics of the mangal (Mazda et al., 1990a) and these may be extremely variable.
As with the mangroves themselves, the most successful macroalgae have special
adaptations that help them tolerate extreme conditions. Work on the physiology of algae
associated with mangroves includes a study of salinity and the polyol (D-dulcitol, D-
sorbitol) content of Bostrychia (West et al., 1992). The success of B. simpliciuscula in the
mangrove swamps of Singapore may be attributed to its physiological adaptations to
salinity extremes. The polyols serve as osmoprotectors that B. simpliciuscula synthesizes
and sequesters as salinity increases (Karsten et al., 1994, 1996). Floridoside compounds,
which may be essential for survival of the algae, also change with salinity (Karsten et al.,
1995). Caloglossa leprieurii, which is also common in mangrove environments, has a
novel metabolic pathway that may be a similar biochemical adaptation to environmental
extremes (Karsten et al., 1997). Salinity gradients create distinct ecotypes of Caloglossa
leprieurii in mangals along the Brisbane River, Australia (Mosisch, 1993).
    Salinity, temperature, desiccation, tidal inundation, wave action, wetting frequency
and light intensity are all environmental factors likely to produce patterns of horizontal and
vertical distribution seen in many mangrove algae (e.g., Phillips et al., 1994; Farnsworth
and Ellison, 1996b). In the Gazi Bay of Kenya, there is distinct macroalgal zonation. The

upper intertidal is covered by Boodleopsis pusilla while the mid-intertidal is dominated by
Halimeda opuntia, Gracilaria salicornia and G. corticata. The low water mark has
primarily Halimeda macroloba and Avrainvillea obscura (Coppejans et al., 1992). A
distinct zonation has also been described for algae growing on the pneumatophores of
Avicennia marina (Steinke and Naidoo, 1990). There are generally three zones: an upper
Rhizoclonium zone; a middle Bostrychia zone, and a lower Caloglossa zone (Phillips et al.,
    The composition of the mangrove algal community may depend largely on the
nature of the early colonizers. Eston et al. (1992) monitored colonization of artificial
substrata by mangrove macroalgae and found that Bostrychia radicans and several other
species settled early. There was no evidence that later species could displace the early
colonists. This macroalgal community showed no succession; the pioneer community was
also the final community. Established macroalgae can also affect distribution of the
mangroves directly. For example, in southeastern Australia, the alga Hormosira banksii
inhibits intertidal establishment of grey mangrove (Avicennia marina) seedlings (Clarke
and Myerscough, 1993).
     A number of algae from mangrove habitats have potential commercial value. For
example, the red alga Gracilaria changii from Malaysian mangrove habitats is an excellent
source of agar; the agar content is between 12 and 25% of its dry weight (Phang et al.,
1996). Monostroma oxyspermum, Catenella impudica and Caloglossa lepriurii are all
edible food resources. The latter two species are also potential sources of dyes. Caulerpa
sp. has yielded bioactive substances that may hold promise as pharmaceutical agents (e.g.,
Ananda Rao et al., 1998).

4.5. Seagrasses

    Seagrasses are closely associated with mangrove habitats in many parts of the
world. In the Andaman Sea, there are three mangrove-associated sea-grasses, Thalassia
hemprichii, Enhalus acoroides and Halophila ovalis (Poovachiranon and Chansang, 1994).
Intertidal mangrove areas in the Gazi Bay, Kenya are colonized by Thalassia hemprichii,
Halophila ovalis and Halodule wrightii (Coppejans et al., 1992) while Halophila baccarii
occurs on intertidal mudflats of Indian mangals (Jagtap, 1991).
    The seagrass biomass in mangrove areas may be quite high. In an Andaman Sea
mangal, Poovachiranon and Chansang (1994) measured seagrass biomass ranging from 55-
1941 g wet wt • m-2, corresponding to 32-297 g dry wt • m-2. As with the macroalgal
communities, seagrass diversity and abundance are largely regulated by a combination of
light level and substrate type. In the Spaanse waters of the Netherland Antilles, the richest
assemblages of seagrasses occur in shallow areas with high light and a mix of hard and soft
substrates. Diversity is much lower where light is low and the substrates are loose muds or
homogeneous, coarse-grained sands (Kuenen and Debrot, 1995).
    Seagrasses generally require high light levels to grow and survive. Planktonic
primary producers require only about 1% of the surface irradiance to maintain a net
positive carbon balance. In contrast, seagrasses may require 10-20% of the daily average
surface irradiance to survive (Fourqurean and Ziemean, 1991). Growth rate may decrease
naturally in the winter months as a result of low temperatures and shortened daylengths.
However, in recent years, there have been precipitous declines of seagrass beds in

mangrove environments. Seagrass mortality has often been linked to reduced water quality
and increased turbidity that decrease light penetration (Giesen et al., 1990; Larkum and
West, 1990). In turbid waters, flocs from the mangroves themselves contribute to shading
of the seagrass (Wolanski et al., 1997).
     Though seagrass beds often occur in close proximity to mangroves, the two habitats
may not be closely coupled. Tussenbroek (1995) found that seagrass growth, biomass and
primary production were all higher in the vicinity of mangrove discharges than they were
in other habitats. Respiratory CO2 derived from mangrove particulate organic matter
(POM) could be a carbon source for seagrass and could promote faster growth. Ebb flows
are generally stronger than flood flows in mangrove creeks, which should promote a net
export of nutrients and POM. In general, however, fluxes from mangrove forests seem to
have little effect on adjacent seagrass beds (Fleming et al., 1990). For example, Hemminga
et al. (1994) failed to detect any input of mangrove POM in a seagrass bed only 3 km
away. POM was exported from the mangrove forest, but deposition was rapid and little
material reached the seagrass bed. Similarly, in the Gazi Bay of Kenya, leaf production and
nitrogen:phosphorus ratios of Thalassodendron ciliatum were unrelated to the input of
mangrove carbon and 13C studies confirmed that the mangroves contribute little reduced
carbon to adjacent seagrass beds (Lin et al., 1991). Nor does it appear that dissolved
nutrients move from the mangal to nearby grassbeds. The few dissolved nutrients
generated by the mangroves are likely to be used for primary production within the
mangrove zone itself (Kitheka et al., 1996).
     Mangroves and seagrasses serve parallel functions in the habitats they share. Both
trap sediments and help capture chemical elements, including trace metals (Costa and
Davy, 1992; Lacerda, 1998). Both also help support fish population by serving as food for
fish, as critical habitat for fish, and as growth surfaces for epizonts that fish eat. A number
of fish species may use seagrass/mangrove habitat as a nursery area. In Guadeloupe,
French West Indies, fish diversity is higher in Thalassia testudinum beds near mangroves
than in the adjacent coral reefs (Baelde, 1990). Similarly, in Belize, Central America, fish
abundance and biomass were highest in a mangrove creek, followed by a seagrass bed and
the sand-rubble zone of an adjacent lagoon (Sedberry and Carter, 1993). Arancibia et al.
(1993) found more than 80 fish species using the mangrove/seagrass habitat; seven species
were found only in these areas.

4.6. Saltmarsh and other flora

     Saltmarsh plants replace mangroves at their northern limit on the Gulf and Atlantic
coasts of North America but the southern limit of the saltmarsh distribution may be set by
competition with the mangroves. For example, the common saltmarsh grass Spartina
cannot survive high salinities and fast sediment accretion. As a result, it grows poorly in
areas where mangroves thrive (Kangas and Lugo, 1990). This usually leads to its
replacement by mangroves, as in Paranagua Bay, Brazil (Lana et al., 1991).
     Though saltmarsh species are generally not common in mangrove habitats, a large
number of other non-mangrove plant species may be found coexisting with the mangroves.
A floristic survey of the tidal mangrove flora in the Sunderbans, India, documented 1175
angiosperm species in 680 genera and 154 families (Nasakar and Bakshi, 1993). Working
in the tropical mangrove forests of the Yucatan Peninsula, Olmsted and Gomez (1996)

found approximately 100 epiphytic species in the families Orchidaceae, Bromeliaceae,
Cactaceae, Araceae, Piperaceae and Polypodiaceae scattered through the canopy and on
trunks of mangrove trees. The orchid Brassavola nodosa is an epiphyte on red mangroves
(Rhizophora mangle) in Belize, Central America, where it grows anywhere from 1-300 cm
above the ground. The largest specimens occur high above the ground where plentiful light
enables them to flower continuously through the summer (Murren and Ellison, 1996).
Lichens may also be abundant on the bark of the mangroves in some habitats (J.C. Ellison,

5.1. Zooplankton

    Diverse communities of zooplankton exist in mangrove habitats and abundances
can be extremely high, reaching 105 individuals m-3 with biomasses up to 623 mg m-3.
These numbers are significantly higher than what is often recorded in offshore waters
(reviewed by Robertson and Blaber, 1992) and the planktonic organisms may contribute to
regional food webs. Such high abundances, however, do not occur in all mangrove
environments. On the west coast of India, for example, Goswami (1992) found lower
zooplankton biomass in the mangroves than in contiguous estuarine and neritic habitats.
    Zooplankton in mangrove waters can be grouped into three size classes. The
smallest organisms are the microzooplankton (organisms between 20 and 199 µm). This
group includes tintinnids, radiolarians, foraminiferans, ciliates, rotifers, copepod nauplii,
barnacle nauplii, and mollusk veligers. Krishnamurthy et al. (1995b) found 81 such species
in the Pichavaram mangroves of south India. Tintinnids were the dominant
microzooplankters with 50 species and densities ranging from 60 to 44,990 individuals m-
 . The most important genera were Tintinnopsis and Favella (Godhantaraman, 1994;
Krishnamurthy et al., 1995b). They also found 40 rotifer species in 17 genera. Except for
rotifers, whose populations peaked in the premonsoon and monsoon months, the microzoo-
plankters were most abundant in the summer, corresponding with highest phytoplankton
    Copepods are the most abundant group in the mangrove mesoplankton (organisms
between 200 µm and 2 mm). In the Pichavaram mangroves of south India, copepod
densities reach 80,740 individuals • m-3 (Godhantaraman, 1994); the genera Acartia and
Acrocalanus (Calanoida), Macrosetella and Euterpina (Harpacticoida) and Oithona
(Cyclopoida) are the most abundant. In Kenyan mangrove waters, copepods constitute
48.5-92.4% of the zooplankton. Zooplankton counts are high in the creek mouth compared
to the inner creek. Abundances peak around May when heavy rains increase nutrient input
(Osore, 1992). Species in the cyclopoid genus Oithona are particularly abundant in many
studies of mangrove plankton. Harpacticoids (e.g., Pseudodiaptomus spp.) and calanoids
(e.g., Acartia spp., Paracalanus spp. and Parvocalanus spp.) are also important (Ambler et
al., 1991). Barnacle nauplii occur in mangrove canals throughout Raby Bay, Australia, but
the copepod Acartia tranteri is found only in the innermost canals (King and Williamson,
    Dioithona oculata is a particularly interesting member of the copepod assemblage
in some mangrove habitats. Individuals congregate to form swarms in light shafts among
mangrove prop roots. The swarms maintain their position in currents up to 2 cm • sec-1

(Buskey et al., 1996). Buskey et al. (1995) showed that the swarms form in response to an
endogenous rhythm. They cannot, therefore, be induced to swarm in artificial light shafts
created at night.
     Copepods and other mesoplanktonic organisms are food for the macrozooplankton
(organisms larger than 2 mm). Jellyfish are the most important macrozooplanktonic
species. The medusa Tripedalia cystophora is attracted to light shafts where non-breeding
individuals actively feed on copepods (reproductive males and gravid females do not feed;
R.W. Stewart, 1996). Planula larvae of Cassiopea species show a strong preference for
mangrove substrata, specifically settling and undergoing metamorphosis on submerged,
deteriorating mangrove leaves (Hofmann et al., 1996). The larvae are apparently attracted
to a soluble protein (molecular weight > 5000 daltons) leaching from the mangrove leaves
(Fitt, 1991, Fleck and Fitt, 1999).
     Meroplankton (planktonic larval stages of benthic invertebrates) may constitute up
to 70% of the zooplankton and span a range the full range of zooplankton sizes.
Brachyuran zoeae can be especially abundant. For example, decapod larval densities
reached 1000 individuals • m-3 in a mangrove area of Costa Rica. These early larval stages
are exported from the mangrove areas on outgoing tides; incoming tides bring the older
stages back to the habitat (Dittel and Epifanio, 1990).
     Bingham (1992) studied larval recruitment of invertebrates (e.g., sponges, oysters,
barnacles, bryozoans, ascidians) living epifaunally on Rhizophora mangle prop roots in the
Indian River Lagoon, Florida (USA). The major factor controlling adult distributions was
transport and recruitment of planktonic larvae as influenced by water flow through the
habitat. Physical factors also contributed to community structure, but on much larger
scales. Farnsworth and Ellison (1996a) reached similar conclusions for R. mangle root
communities in Belize, Central America. To better understand larval recruitment processes
and their importance to the structure and dynamics of mangrove marine communities,
Wolanski and Sarsenski (1997) have developed computer models that simulate the
dispersal of fish and shrimp larvae through mangrove habitats.

5.2. Sponges and Ascidians

     Because they are often surrounded by muddy or sandy sediments, submerged
mangroves roots, trunks, and branches are islands of habitat that attract rich epifaunal
communities. The epifauna may include a diverse array of invertebrate groups including
sponges, hydroids, anemones, polychaetes, bivalves, barnacles, bryozoans, and ascidians.
Encrusting sponges and ascidians are particularly important in many environments and
may be specially adapted to life there. A number of ascidian and sponge species are largely
restricted to mangrove surfaces (Goodbody, 1993, 1994, 1996; Bingham and Young
1991a; de Weerdt et al., 1991) and epifaunal species that do occur in other habitats may
show distinctly different growth forms when they are attached to mangrove roots
(Swearingen and Pawlik, 1998).
     As with mangrove bacterial, fungal, and algal communities the invertebrate
epifauna can show distinct distributional patterns correlated with desiccation, wave action,
temperature and salinity. Rützler (1995) described vertical zonation of sponges on the prop
roots of Rhizophora mangle in Belize. Differential desiccation tolerance produced the

zonation, with the most resistant species occurring higher on the roots. Farnsworth and
Ellison (1996a) found a particularly rich ascidian epifauna on mangroves in leeward areas
of another Belizian mangal.
     Epifaunal organisms may play important roles in the structure and function of the
mangal. Sponges, for example, may be food resources for other invertebrates and fish.
Many sponges have anti-predator defenses including siliceous or calcareous spicules and
noxious or toxic chemicals (McClintock et al., 1997). However, mangrove species are
generally not as well defended chemically as sponges from reef habitats (Pawlik et al.,
1995; Dunlap and Pawlik, 1996). Surprisingly, the palatable species also seem to lack any
particular structural or nutritional features that would discourage predators (Swearingen
and Pawlik, 1998). In light of this vulnerability, the mangrove habitat itself may, to some
extent, be a refuge for less protected species. Species here may also rely on faster growth
or greater reproductive output to compensate for predation losses (Chanas and Pawlik,
1995). In contrast to the sponges, some of the mangrove ascidians may have unusual
chemicals that are potent feeding deterrents (Vervoort et al., 1997).
     Mangrove sponges may also lack the allelochemicals that protect them from
overgrowth by other species in space-limited coral environments. Bingham and Young
(1991b) tested 8 sponges commonly found on submerged roots of Rhizophora mangle in
the Indian River, Florida, the Florida Keys, and Belize, Central America. None of the
sponges appeared to use allelochemicals to reduce settlement or survival of potential
competitors. In fact, several epifaunal invertebrate species recruited more heavily in the
presence of the sponges.
     Despite a seeming lower level of anti-predator and anti-competitor chemicals in
mangal than in coral reef communities, epifauna invertebrates in the habitat may still be
sources for interesting, and valuable, compounds. Ecteinascidia turbinata, for instance, is a
colonial ascidian that grows primarily on the submerged prop roots of Rhizophora mangle
in many areas of the Caribbean. It was recently discovered that E. turbinata produces
compounds (ecteinascidins, Figure 7) that show strong activity against a variety of
carcinomas, melanomas, and lymphomas (Rinehart et al., 1990; Wright et al., 1990, Sakai
et al., 1992). This discovery has led to large scale collection of this species for extraction,
isolation, purification and testing of the compounds. Depending on the method used, these
collections adversely affect the wild populations (in addition to damaging the mangrove
trees on which they grow; Pain, 1996). Long
distance dispersal of E. turbinata appears to
depend on rafting of adult colonies; larval
dispersal is highly localized. Collection
techniques that damage the mangroves or
remove large patches of the population,
therefore, could have severe consequences of this
species (Bingham and Young, 1991a).
     The close association of invertebrate
epifauna and mangroves may have led to
                            Figure 7. Bioactive compound (ecteinascidin)
mutualisms between them. For example,         extracted from the mangrove ascidian
sponges and ascidians may protect the         Ecteinascidia turbinata. The ecteinascidins
mangroves on which they grow. Ellison and       have shown strong in vivo activity against a
                            variety of cancer cells.
Farnsworth (1990, 1992) found that epifaunal

sponges and ascidians decreased the amount of damage wood-boring isopods did to the
roots of Rhizophora mangle. Roots without the sponge/ascidian cover showed significantly
more damage and 55% lower growth. In estuarine regions where physical conditions
prevented establishment of epifaunal sponges and ascidians, nearly 100% of the R. mangle
roots were damaged by the isopods.
    The invertebrate/mangrove mutualism may also take the form of a symbiotic
nutrient exchange. Sponges attached to submerged roots of Rhizophora mangle induce the
roots to produce fine rootlets that penetrate and grow throughout the sponge tissue.
Measurements indicate that the roots obtain dissolved inorganic nitrogen from the sponges.
The sponges, in turn, obtain carbon from the roots. Ellison et al. (1996) experimentally
transplanted sponges to bare R. mangle roots in a Belizean mangrove habitat. Within 4
weeks, adventitious rootlets had appeared over the surface of the root. The sponges
attached to the roots grew 1.4 – 10 times faster than did control sponges attached to PVC
pipes in the same habitat. Miller-Way and Twilley (1999) suggest that nitrogen-fixing
bacteria living symbiotically with Ulosa
rutzleri and Lissodentoryx isodictyalis on            E. turbinata     H. magniconulosa
mangrove roots release significant amounts
of NO3 to surrounding waters.
                          Percent Cover
    The epifaunal communities on
                            40        L. isodictyalis
mangrove roots may show strong
fluctuations. In the Florida Keys, USA,        20
Rhizophora mangle root communities
change dramatically over short time           0
intervals (1-2 months, Figure 8). Physical




disturbance from tidal flows, species-
specific predation and fragmentation of the Figure 8. Fluctuations in cover of two epifaunal sponges
dominant sponges produce the variability.    (Lissodendoryx isodictyalis and Haliclona magniconulosa)
                        and a colonial ascidian (Ecteinascidia turbinata) on
The perturbations prevent competitive
                        submerged Rhizophora mangle prop root (Florida Keys,
processes from producing the more stable    USA). Photographic measurements were made at 2-3 month
equilibrium assemblages seen in some      intervals for 32 months (after Bingham & Young, 1995).
other mangrove epifaunal communities
(Bingham and Young, 1995).

5.3. Epibenthos, infauna, and meiofauna
    The muddy or sandy sediments of the mangal may be home to a variety of
epibenthic, infaunal, and meiofaunal invertebrates. The composition and importance of
these communities varies enormously from habitat to habitat depending on the sediment
characteristics of the individual mangal.
    Mangrove sediments generally support higher densities of benthic organisms than
do adjacent non-vegetated sediments (Edgar, 1990, Sasekumar and Chong, 1998). Sheridan
(1997) identified over 300 benthic taxa in red mangrove (Rhizophora mangle), seagrass
and mud habitats in southern Florida. Densities, which ranged from 22,591 - 52,914
individuals • m-2 were always higher in the red mangrove peat than in the other habitats.
The fauna was composed primarily of annelids and tanaids with maximum densities of
31,388 and 35,127 individuals • m-2 respectively (Sheridan, 1997).

     The epibenthos may include hydrozoans. For instance, the hydrozoan Vallentinia
gabriellae, which feeds on a variety of zooplankters, is common in some south Floridan
mangals (Rey et al., 1992). Calder (1991) found that hydroids in a Belizean mangal
respond to water flow. The hydroid fauna is richer and more diverse in areas exposed to
waves and tidal currents than in sheltered, still-water areas of the mangal. Polychaetes are
the dominant macrobenthos in mangrove flats at Inhaca Island, Mozambique where their
distributions are controlled by sediment grain size, salinity and ground water (Guerreiro et
al., 1996). Oligochates may also be abundant in shallow mangal waters. Diaz and Erseus
(1994) found one oligochaete family, the Limnodriloidinae, entirely restricted to mangrove
     The most successful benthic species in the mangal are those that can adapt to the
salinity and temperature stresses that are characteristic of these environments (Ferraris et
al., 1994). Extreme fluctuations in these physical features may prevent colonization by
benthic species. For example, Lana et al. (1997) found that benthic infaunal abundance
and diversity were significantly lower in mangrove sites than in more seaward zones of
Paranagua Bay, Brazil.
     Mangrove meiofaunal communities may also include annelids (especially
oligochaetes) and crustaceans. However, they are generally dominated by nematodes. As a
result, nematodes have been better studied than any other members of the mangrove
meiofauna (Olafsson, 1996). In the dry tropical mangroves of northeastern Queensland,
nematode abundances may reach 2117 individuals • cm-2 with seasonal fluctuations
contributing to variability in the community (Alongi, 1990a). The study of mangrove
meiofaunal communities has led to descriptions of several new nematode species. These
include Parapinnanema ritae, P. alii and P. rhipsoides from Guadeloupe (Gourbault and
Vincx, 1994); Chromaspirina okemwai, Pseudochromadora interdigitatum and
Eubostrichus africanus from Ceriops sediments along the Belgian coast of the North Sea
(Muthumbi et al., 1995); and Papillonema danieli and Papillonema clavatum from Ceriops
sediments of Kenya (Verschelde et al., 1995).
     The distribution of the nematode fauna has been intensively studied in a temperate
mangrove mudflat of southeastern Australia (Nicholas et al., 1991). Approximately 85% of
the nematodes occurred in the top layer of the soft mud, but 5-7 species penetrated the
deeper anoxic muds down to 10 cm. Abundances were affected by tidal zonation.
Nematode biomass was approximately 888 mg dry wt m-2 (≈ 383 mg C m-2) in the low
tide zone but was only 19 mg dry wt m-2 (≈ 8 mg C m-2) in the upper tide zone.
     Nematode populations may vary with food content, grain size and organic content
of the mangrove sediment (Hodda, 1990). The meiofaunal community is undoubtedly part
of the detrital food web. Tietjen and Alongi (1990) found a significant correlation between
biomass of Avicennia marina litter, bacterial abundance, and nematode abundance. The
relationship disappears as detritus ages. However, a direct role of nematodes in organic
matter cycling could not be demonstrated experimentally. Nor does the meiobenthic
community appear to have much direct predator/prey interactions with the epibenthos.
Schrijvers et al. (1995, 1997) showed this experimentally through exclusions of
meiobenthic species from Kenyan Ceriops tagal and Avicennia marina habitats. There is
still much to learn about the role of these less-easily studied members of the mangal

5.4. Prawns, shrimp and other crustaceans
5.4.1. Prawns and shrimp
     Mangrove habitats and prawn/shrimp populations are tightly linked in many
regions. Analyses of commercial prawn catches have repeatedly shown strong correlations
between abundance and biomass of prawns and extent of the surrounding mangrove areas
(Sasekumar et al., 1992; Kathiresan et al., 1994c; Vance et al., 1996b).
     Robertson and Blaber (1992) proposed three explanations for this relationship.
First, organic detritus exported directly from the mangroves provides food and habitat for
juvenile penaeids in offshore areas (Daniel and Robertson, 1990). Second, the waters in
the numerous channels and small creeks of the mangrove receive high levels of terrestrial
runoff, rich in nutrients. Export of these nutrients (controlled largely by groundwater
flows; Mazda et al., 1990b; Ovalle et al., 1990) contribute to productivity. This
productivity, in turn, may support offshore penaeid populations. Third, the mangrove
waterways directly serve as nursery grounds for juvenile penaeids that move offshore and
enter the commercial fishery as they mature. This hypothesis is strongly supported by
surveys of larval, postlarval and juvenile penaeids in nearshore habitats (Vance et al.,
1990, 1996b, 1997; Mohan et al., 1997; Primavera, 1998; Rajendran and Kathiresan,
     Sheridan (1992) found low shrimp abundance among Rhizophora mangle prop
roots in Rookery Bay, Florida. Only 4% of the collected animals were in the roots,
compared with 74% in adjacent seagrass beds. This, however, seems to be unusual, and
numbers and biomass of prawns and shrimp are generally higher in mangrove areas than in
adjacent nearshore habitats (Chong et al., 1990; Sasekumar et al., 1992). Study of these
diverse shrimp communities is revealing new species (Miya, 1991; Bruce, 1991).
     In a six-year study, Vance et al. (1997) determined the primary factors controlling
juvenile prawn abundance in mangroves to be larval supply and postlarval settlement. The
young of many shrimp species appear to use the mangal. Juveniles of eight penaeid prawn
species (primarily Metapenaeus monoceras and Penaeus indicus) are common in the
Pichavaram mangroves. Catches of the juveniles in core mangrove areas are greater than in
open waters (Rajendran, 1997). In Oman, R. Mohan and Siddeek (1996) similarly found
abundant postlarval and juvenile shrimp in the detritus-rich, muddy substrates of a mangal
they studied. Distributions of the juveniles within the mangal are strongly influenced by
salinity; densities are highest at intermediate salinities (R, Mohan et al., 1995).
     As the shrimp grow, they may eventually leave the mangal. In the Matang
mangroves of Malaysia, Chong et al. (1994) measured prawn densities of 4092 individuals
• ha-1 in the mangal but only 2668 individuals ha-1 in the adjacent mudflats. However,
biomass was approximately the same in both areas, suggesting that larger individuals move
out of the mangal. Using size distribution data, Rajendran and Kathiresan (1999a),
concluded that postlarvae of prawns recruit into the Pichavaram mangroves in the
postmonsoon period; subadults then leave during the premonsoon and monsoon periods.
The annual offshore commercial catch of adult P. merguiensis is significantly correlated
with number of prawn emigrating from the estuary during the wet season (Vance et al.,
1997). There is also a strong positive relationship between rainfall and subsequent offshore

commercial catch of adult shrimp (Staples et al., 1995), probably due to flushing from the
mangrove habitat as a result of heavy rains.
    Although the mangal may be a sink for settlement and early growth of shrimp and
prawns, it may also be a source for larvae that are transported to other habitats. Mangrove
waters in the Klang Strait of Malaysia may collect 65 billion penaeid prawn larvae before
their annual transport and settlement in coastal nursery grounds. Tidal currents and lateral
trapping in mangrove-lined channels cause this aggregation (Chong et al., 1996).
    There may be a number of benefits for juvenile shrimp and prawns living in
mangrove habitats. The habitat is complex and provides a variety of niches within which
species can exist. For example, in the mangroves of Muthupet, India, Penaeus indicus, P.
merguiensis and Metapenaeus dobsoni show clear preference for detritus-rich muddy
substrates in which they feed. In contrast, P. monodon shows no such preference. Other
shrimp feed directly on the mangroves. Cholesterol extracted from Rhizophora leaves
promotes growth of juvenile Penaeus indicus and increases their conversion efficiency
(Ramesh and Kathiresan, 1992). However, not all mangrove products are beneficial.
Excoecaria agallocha latex is toxic to larvae of the freshwater prawn Macrobrachium
lamarrei (Krishnamoorthy et al., 1995) and to penaeid prawns (Kathiresan and Thangam,
    The mangrove forest, with its small creeks and channels, its hanging roots, and soft
substrates may also provide refuge from predators. Prawns in these habitats tend to be most
active near high tide and at night (Stoner, 1991; Vance, 1992; Vance and Staples, 1992;
Rajendran, 1997). This presumably allows them to forage when food is most accessible
and predation danger is lowest.
    Some mangrove shrimp may avoid predation by burrowing in the muddy
sediments. Primavera and Lebata (1995) found that Metapenaeus were particularly active
burrowers. Penaeus monodon is also a burrower, but burrowing activity is size dependent
and increases as the animals grow. Shrimp may also escape predators by migrating with
the tides. Vance et al. (1996a) observed that juvenile P. merguiensis are very mobile,
moving substantial distances into the mangrove forest at high tide. An extreme example of
shrimp migration is the semi-terrestrial Merguia oligodon, a species common in some
Kenyan mangroves. This species lives among the aerial roots of Rhizophora mucronata. It
is active at night, grazes on mangrove bark, and climbs mangrove roots and trunks up to 80
cm above the ground (Vannini and Oluoch, 1993). Vance et al. (1997) have used a stake-
netting method to study distribution and movements of prawn in intertidal mangrove
forests. This technique shows promise as a way to provide better information about the
shrimp and prawns and their roles within the mangal.

5.4.2. Other Crustaceans
    While shrimp and prawns do not generally harm the mangroves, and may actually
be beneficial (e.g., through bioturbation of muddy sediments), other crustaceans do
significant damage. For example, barnacles can grow abundantly on mangrove roots and
pneumatophores (Foster, 1982; Anderson et al., 1988; Bayliss, 1993; Ross and
Underwood, 1997). Balanus amphitrite and other fouling organisms, for instance, kill
42.5% of the mangrove seedlings in Goa, India (Santhakumaran and Sawant, 1994). Some
species of barnacles belonging to the genera Euraphia, Elminius and Hexaminius appear
to prefer mangroves over other substrates. The settling barnacle larvae may even show

strong preferences for mangrove species and discriminate among parts of the trees. The
barnacle densities are controlled by the physical environment of the mangal (primarily
desiccation and temperature). Populations are greater on seaward than on landward areas
of the forest. Densities are also greater on lower surfaces than on upper surfaces of trunks
and leaves (Ross and Underwood, 1997). In some mangrove areas barnacle numbers are
also greater in the mid-intertidal than in the upper or lower intertidal zones (Kathiresan et
al., 2000), and the species show marked zonation (Baylisss, 1993), with chthamalids
occuring above Balanus amphitrite. The recruitment of barnacles, and other sessile
invertebrates within the mangal is largely controlled by larval abundance, tidal currents,
duration of larval life and density of the adult populations (Bingham, 1992; Farnsworth and
Ellison, 1995; Young, 1995). It was at one time thought that the selection by barnacles of
their mangrove habitat was so extreme that one species of Hexaminius (H. folorium)
occurred only on twigs and leaves, while another (H. popeiana) was restricted to the bark
(Anderson et al., 1988; Ross and Underwood, 1997). Recent studies indicate that
phenotypic variation is responsible, and that the leaf-occurring form is comparable
morphologically and by DNA to the form on mangrove bark (Ross, 1996; Ross and
Pannacciulli, personal communication). This situation can be compared with the changes
in form and colour seen in the bivalve Enigmonia aenigmatica and the snail Littoraria
pallescens when inhabiting different parts of the mangrove, trees as noted on page 81.
     Burrowing isopods (e.g., Sphaeroma terebrans and S. peruvianum) also do
tremendous damage to mangroves in many regions of the Atlantic, the Caribbean, and the
eastern Pacific. Numerous juveniles and adults can be found living inside a single root or
stem. Their burrowing can significantly affect root growth and development (Ellison and
Farnsworth, 1990; Santhakumari, 1991).
     Other crustaceans use mangrove waters temporarily during certain phases of their
life history. One of the better known is the Caribbean spiny lobster (Panulirus argus),
juveniles of which use the mangroves as nursery habitat (Monterrosa, 1991). Like the
shrimp and prawns, however, the lobsters migrate out of the mangal as they grow. Adult
lobsters remain in the mangal only if their preferred habitat (under coral heads) is
unavailable (Acosta and Butler, 1997). Migration to other habitats may reflect a search for
better food resources.

5.5. Crabs
    Crabs are characteristic members of the invertebrate mangrove fauna and have
received much attention. Some indication of the diverse array of mangrove-associated
crabs can be found in annotated checklists from India (Sethuramalingam and Ajmal Khan,
1991), Malaysia and Singapore (C.G.S. Tan and Ng, 1994) and Brazil (Vergara-Filho et
al., 1997).
    Within the complex mangrove environment, crabs fill a variety of niches. For some
species, the relationship with mangroves is obligatory; they depend directly on the
mangroves for survival (Vergara-Filho et al., 1997). Others simply have ranges that
overlap the mangal. The mud crab Scylla serrata inhabits seagrass and algal beds in the
mangroves of Pichavaram, south India (Chandrasekaran and Natarajan, 1994). Floating
leaves in a Costa Rican mangal harbor a unique community dominated by Uca crabs
(77.8% of all organisms counted; Wehrtmann and Dittel, 1990). Perhaps one of the most
striking associations is seen with the hermit crab Clibanarius laevimanus. Individuals of

this species climb the mangrove roots and rest on them during the entire low water period,
forming dense clusters of up to 5,000 individuals (Gherardi et al., 1991; Gherardi and
Vannini, 1993). Gherardi et al. (1994) have studied size, sex and shell characteristics of
this unique mangal species.
     Mangrove crabs are morphologically, physiologically, and behaviorally
well-adapted to their environment. For example, the semaphore crab, Heloecius cordifor-
mis, is active at low tide when it is completely exposed to air. Its branchial chambers are
modified for respiration both in the air and under water (Maitland, 1990). A number of
other crab species (particularly in the Family Grapsidae) live directly on the mangrove
trees. Species in this group generally have a square, flattened carapace, a relative
shortening of the dactylus on the walking legs and a lengthening of the propodus (Vannini
et al., 1997a). These structural specializations appear to be adaptations for their tree-
dwelling existence.
     Crabs living in the mangal must adjust to significant temperature and salinity
fluctuations. Some, like the grapsid Metopograpsus messor, retreat to burrows where
temperatures are less variable and consistently lower than the sediment or air temperatures.
When it is out of the burrow, M. messor uses evaporative cooling to keep its body
temperature lower than the surrounding air (Eshky et al., 1995).
     Other crabs have adopted a nocturnal lifestyle, possibly to escape high temperatures
and/or predators (Micheli et al., 1991). The hermit crabs Coenobita rugosus and Coenobita
cavipes are active 24 hours a day but are most active when they are among the mangrove
roots. Barnes (1997) suggests that they do this because wind speeds (and desiccation
potential) are lower there. Desiccation can significantly affect ion balances and mangrove
crabs are physiologically adapted to resist major changes. In Ucides cordatus and Carcinus
maenas total Na+ efflux is markedly reduced during emersion. The reduction in ion and
water loss results from decreased urine output (Harris et al., 1993). When U. cordatus is
placed in low salinity water, active sodium uptake increases 4-5 fold (Harris and Santos,
     Crabs in mangrove habitats show distinct distributional patterns related to substrate
characteristics, salinity, degree of tidal inundation, and wave exposure. In the Indian
Sunderbans, these conditions produce a vertical zonation of crab species (Chakraborty and
Choudhury, 1992, Kathiresan et al., 2000). Machiwa and Hallberg (1995) also found a
horizontal zonation of crabs in East Africa,. The terrestrial edge of the mangal was
occupied by grapsids while mixed associations of ocypodids dominated the open areas of
sand and mud.
     Different crab species respond differently to disturbance and this affects species
distributions. In Kenya, Sesarma guttatum prefers shaded habitats and is most common in
regions with an established mangrove canopy. In contrast, Uca urvillei and
Microphthalmus depressus prefer clear-cut areas. In the more landward Avicennia zone,
the species composition of the crab community remains constant whether the vegetation is
intact or clear-cut (Ruwa, 1997).
     Mangrove crabs can be divided into distinct guilds based on their feeding mode.
Some species (e.g., Uca and Macrophthalmus spp.) are detritivores that extract their food
from the sediments while others (e.g., the portunid Scylla serrata) are opportunistic
scavengers (Micheli et al., 1991). There are also a number of active predators. The
swimming portunid Thalamita crenata lives on the extreme seaward fringe of the

mangrove swamp where it preys heavily on bivalves and slow-moving crustaceans
(Cannicci et al., 1996c). This species is active during high tides, but only when the water is
between 10 and 30 cm deep, suggesting that their foraging behavior is controlled by
hydrostatic pressure changes associated with tidal flux (Vezzosi et al., 1995).
    Epixanthus denatus, which is very abundant in mangrove creeks along the Kenyan
coast, is another active predator. It forms dens among the mangrove roots and feeds on
almost any slow-moving invertebrate (including other crabs) that comes within a 3-m
radius. Their intense predation may be responsible for the climbing behaviour of many
potential prey species (Cannicci et al., 1998). Crabs in the mangrove face significant
predation risks and may show specific anti-predator adaptations. Diaz et al. (1995) suggest
that postlarval and juvenile crabs may avoid predation by responding to specific light cues.
Aratus pisonii, which lives among roots and branches, is attracted to narrow dark
rectangles but avoids large dark rectangles. The authors speculate that the narrow
rectangles resemble roots that represent refuges while the larger rectangles indicate
predators. In contrast, Chlorodiella longimana, a subtidal species, moves toward all dark
rectangles regardless of their size.
    In addition to scavenger and predator guilds, there is a guild of herbivorous
mangrove crabs that feed directly on mangrove litter. In Ao Nam Bor, Thailand, up to 82%
of the diet of sesarmid crabs consists of mangrove material (Poovachiranon and
Tantichodok, 1991). A number of these herbivores show clear feeding preferences. For
example, Sesarma meinertii generally prefers Bruguiera gymnorrhiza to Avicennia marina
leaves (Micheli et al., 1991). However, Steinke et al. (1993a) showed that the age of the
litter was more important than its source in determining preference. The crabs chose
yellow B. gymnorrhiza and A. marina leaves over green leaves of either species. Sesarma
messa and S. smithii both prefer decaying leaves to those that are simply senescent,
irrespective of leaf species. Neosarmatium meinerti does not choose among mangrove
species but does, however, strongly prefer fresh leaves. Its heavy, non-selective feeding on
mangrove seedlings and propagules could make it a significant threat to afforestation
efforts (Dahdouh-Guebas et al., 1997).
    It is unclear what factors are responsible for these feeding preferences. Micheli
(1993a, b) found that preferences were not affected by tannins, water content, % organics,
C:N ratio, or leaf toughness. Many of the herbivorous crabs store the leaves in their
burrows for some time. However, the nutritional value of the leaves does not increase
during the time they are stored, indicating that the crabs are simply storing the leaves and
not gardening them to encourage bacterial or fungal growth (Micheli, 1993a, b). Given that
the mangrove leaves, in general, have low nutritional value and the crabs do not have a
mechanism to promote bacterial or fungal growth, it may be very important for them to get
the maximum food value out of the leaves they eat. This may contribute to their specific
    Although mangrove leaves are not particularly nutritious, they do produce
sufficient energy to influence survival, growth and reproduction of the crabs and it seems
reasonable to assume that is a sufficient selective force to produce feeding preferences.
Survival of two mangrove crabs, Chiromanthes bidens and Parasesarma plicata, is
directly related to litter type. They do best when fed brown Avicennia marina leaves,
followed by brown Kandelia candel, yellow A. marina, and finally, yellow K. candel
(Kwok and Lee, 1995). Mangroves at different sites in Venezuela produce leaves with

different nutritional value. The crabs are smaller in sites where the stunted trees produce
leaves of low nutritional value (Conde and Diaz, 1992; Conde et al., 1995).
                                      Some of the herbivorous
                                  crabs do not simply graze on
                                  fallen leaves. Some actively
                                  forage in the canopy of the tree.
                                  In the mangrove swamps of East
                                  Africa, Sesarma leptosoma is an
                                  active climber that can reach the
                                  tops of the tallest trees. It never
                                  descends into the water nor
                                  venture out on the mud at low
                                  tide. This behaviour provides
                                  protection from predators
 Figure 9. Average daily migration patterns of the crab
                                  (Cannicci et al., 1996a). It spends
 Sesarma leptosoma into and out of the mangrove canopy (after
 Vannini and Ruwa, 1994).                     most of the night among the
                                  mangrove roots but, in the
morning, moves up into the canopy to feed on fresh leaves. Increasing temperatures and
the danger of desiccation eventually drive the crabs down to the bases of the trees where
they spend the hottest hours of the day. In the evening, they return to the canopy for
another short feeding period (Figure 9; Vannini and Ruwa, 1994; Vannini et al., 1997b).
The crabs tend to return to the same feeding
spot each time they visit the canopy and even
follow the same path to get there (Figure 10).
Cannicci et al. (1996b) suggest that site fidelity
is important as it takes the crabs near leaf buds
where they can find water trapped among the
scales. Reduced light delays the migration. It is
unclear why migration of this intertidal animal
is regulated by light instead of tides (Vannini
et al., 1995).
     Feeding by crabs hastens composting of
mangrove material and contributes to cycling
of nutrients through the mangal (Lee, 1998). In
the Gazi Bay, Kenya, crabs (along with large
snails) process over 18% of the fallen litter
(Slim et al., 1997). The large mangrove
grapsid Sesarma meinertii consumes Avicennia
marina leaves at a rate of 0.78 g m-2 d-1,
accounting for 43.58% of the leaf-fall in a        Figure 10. Branch fidelity in the mangrove
warm temperate mangrove swamp in southern         crab Sesarma leptosoma. Arrows indicate
                             movement of three individuals. Individuals B
Africa (Emmerson and McGwynne, 1992).
                             and C always returned to the same branch in
     Digging by crabs, in conjunction with
                             the canopy and even followed the same path
other benthic fauna like nematodes,            to get there. Individual A returned to one of 2
polychaetes, and mudskippers (Kristensen et        branches (after Cannicci et al, 1996b).
al., 1995) can also have a profound effect on
BIOLOGY OF MANGROVES AND MANGROVE ECOSYSTEMS                                        49

nutrient cycling and the physical and chemical environment of the mangal (Lee, 1998).
Burrows enhance aeration, facilitate drainage of the soils, and promote nutrient exchange
between the sediments and the overlaying tidal waters (Ruwa, 1990). Crab burrows
generally have two or more openings and may form extensive labyrinths of interconnected
tunnels. Using dye injections and flow measurements, Ridd (1996) estimated that, in a 1
km2 area of a North Queensland mangal, 1,000 to 10,000 m3 of water move through crab
burrows on each tidal cycle. T.J. Smith et al. (1991) removed burrowing crabs from a
mangal and observed significant increases in soil sulfide and ammonium levels relative to
control sites. These chemical changes led to decreased mangrove growth and reproduction.

5.6. Insects
    Insects constitute a significant portion of the fauna in many mangrove
communities. They may be permanent residents of the mangal or only transient visitors. In
either case, they often play important roles in the ecology of the system and contribute to
the unique character of these habitats. Surveys of mangrove insects are revealing complex
assemblages of species filling a wide variety of niches. For example, Veenakumari et al.
(1997) found 276 insect species in the mangals of Andaman and Nicobar Islands of India;
197 of these were herbivores, 43 were parasites and 36 were predators. Similar levels of
diversity and abundance have been found in the insect fauna of Thailand’s Ranong
mangroves (Murphy, 1990a). Many of the insects reported in mangals are only temporary
visitors; their ranges included many other habitat types. As a result, they provide linkages
between the mangal and other environments (Ananda Rao et al., 1998).
    Terrestrial organisms living in mangrove environments are faced with harsh
conditions of strong sunlight, high temperatures and desiccation. Many of the insects (and
other terrestrial arthropods) avoid these conditions by emerging only at night, or by living
entirely within the plants. In the mangals of Belize, wood-boring moths and beetles
excavate tunnels through the mangroves. The
tunnels then become home to more than 70 other                           Leaves always exposed
species of ants, spiders, mites, moths, roaches,                          Leaves submerged at high tide

termites, and scorpions (Rützler and Feller, 1996;
                                       Percent damaged


Feller and Mathis, 1997). A number of organisms
(including isopods, amphipods, myriapods, and
spiders in addition to insects) escape high
temperatures and desiccation by living in the
intertidal portions of the mangal. During periods       0
                                 Total leaf Holes through Damage to Internal leaf   Damage to
of high tide, these organisms retreat to air-filled        damage   the leaf  leaf margin  damage   underside of
                                                           primary vein

cavities where they remain until they are again      Figure 11. Effect of tidal submergence on
exposed by the falling water (Murphy, 1990b).       Laguncularia racemosa leaf damage
                              (Guanacaste Province, Costa Rica).
    Herbivorous insects can cause significant
                              Submergence limits access by terrestrial
damage to the mangroves, attacking leaves and
                              herbivores and may also cause chemical and
boring through the wood. Seedlings may be         structural changes in the foliage. Standard
particularly vulnerable to attack and are strongly     errors are shown. Statistical analysis showed
affected by proximity to adult trees. In a Belizean    that exposed leaves had significantly more
                              damage in all cases (after Stowe, 1995).
mangal, seedlings growing under the intact adult

canopy suffered twice as much herbivore damage as seedlings in areas without an
established canopy (Farnsworth and Ellison, 1991). Immersion in seawater may help
protect the trees. Portions of the mangrove canopy that are submerged by tidal waters
suffer significantly less herbivore damage than those that remain exposed (Stowe, 1995;
Figure 11).
    Recent records of insects in mangrove include 28 species of dragonflies in India
(Mitra, 1992), a water strider, Mesovelia polhemusi in Belize (Spangler, 1990), an unusual
psyllid, Telmapsylla, in Florida and Costa Rica (Hodkinson, 1992), and termites,
Nasutitermes nigriceps, in Jamaica (Clarke and Garraway, 1994). However, some of the
more important and best studied mangrove insects are bees, ants, and mosquitoes. Honey
bees produce significant quantities of honey from the mangroves of India, Bangaladesh,
the Caribbean and southwest Florida. The honey is an important food resource for humans
in some regions (e.g., Padrón et al., 1993). In India, the dominant bee species (Apis
dorsata), may travel hundreds of miles to forage in the mangrove forests during periods of
peak blooming (March and July). It builds honeycombs on several mangrove species, but
prefers Excoecaria (Krishnamurthy, 1990). In contrast, the same bee species in southern
Vietnam forages on mangrove vegetation primarily during the rainy season and rarely
builds combs (Crane et al., 1993).
    Twenty-two ant species are known from Brazilian mangrove habitats. Camponotus
and Solenopsis are most common genera (Cortes-Lopes et al., 1996). Clay and Andersen
(1996) found 16 ant species in an Australian mangal. Two of these, both in the genus
Polyrhachis, are apparently restricted to this habitat. In northern Australia, Polyrhachis
sokolova nests directly in the soft mud of the mangal (Nielsen, 1997). Adams (1994)
studied niche partitioning in four ant species common in Panamanian mangroves. These
species partition the mangrove canopy in non-overlapping territories that are maintained
through a combination of pheromonal signals and tactile displays.
    Holes in the mangrove trees (particularly Avicennia species) and crab-burrows
provide ideal sites for mosquito breeding (Thangam, 1990). Mosquitoes are ubiquitous in
mangrove habitats and may act as vectors for diseases of vertebrates. Populations are often
dense and species diversity can be high (eighteen species occur in the Pichavaram
mangroves of south India alone; Thangam and Kathiresan, 1993b). Predation by fishes
may reduce successful mosquito oviposition. Hence, mosquito populations are lower in
sites with high fish densities (Ritchie and Laidlaw-Bell, 1994). Regardless of its
composition, any mangrove forest that is flooded by < 14% of the highest daily tide can
potentially produce the mosquito Aedes taeniorhynchus (Ritchie and Addison, 1992).
Addison et al. (1992) identified A. taeniorhynchus oviposition sites and quantified larval
production by locating and counting egg shells.
     Mangals tend to be reservoirs for a number of pathogenic viruses including
Dengue, Haemorrhage Fever, Bakau and Ketapang. Mosquitoes are the most common
vectors for these viruses. However, several families of other diptera associated with faecal
contamination in mangroves of Singapore and Malay also contribute to the spread of
human disease (Murphy, 1990c).

5.7. Mollusks
    Mollusks are found throughout most mangrove habitats. They live on and in the
muds, firmly attached to the roots, or forage in the canopy. They occupy a number of

niches and contribute to the ecology of the mangal in important ways. The nature of the
molluskan community is strongly influenced by physical conditions. For example, in the
mangroves of China, Jiang and Li (1995) found that density and biomass of the mollusks
(including 52 species) were consistently highest in the high tide zones and decreased with
depth. In addition, species abundance increased with salinity. Such a pattern is likely to be
found in other mangals. This sensitivity of mollusks to their physical/chemical
environment may make them good bioindicators. Skilleter (1996) has used the composition
of the molluskan assemblage to assess the health of urban mangrove forests.
    The molluskan fauna in mangrove habitats is composed primarily of bivalves and
snails and most study has focused on these groups (e.g., Balasubrahmanyan, 1994). Other
mollusk groups (e.g., nudibranchs, chitons, scaphopods) are less obvious and have been the
subject of only a few studies (e.g., Sigurdsson, 1991). Much of the work with bivalves and
snails has concerned individual species and their specific adaptation to the mangrove
environment. For example, Dious and Kasinathan (1994) studied the high desiccation,

salinity, and temperatures tolerances of two pulmonate snails, Cassidula nucleus and
Melampus ceylonicus, from a south Indian mangal. Special conditions in the mangal may
result in local adaptation. Crow (1996) compared movement of the snail Bembicium
auratum in mangrove habitats and on rocky shores. Movement patterns in the mangroves
were very different (despite similar distributions). This suggests that models developed in
rocky intertidal communities may not be directly applicable to mangrove communities.
     Color variability may represent a special adaptation to the mangrove environment
or possibly a reaction to the complex chemical defenses of the plants. For example, the
leaf-inhabiting mangrove snail, Littoraria pallescens, has distinct color morphologies that
are sometimes associated with other shell differences. The color variation may reflect
predation pressures (Cook, 1990; Cook and Kenyon, 1993).
     The unique tree-climbing bivalve, Enigmonia aenigmatica, which occurs mostly on
Avicennia and Sonneratia also shows color variation. The shells are normally red to deep
purple. However, the shells of individuals attached directly to the mangrove leaves are
golden yellow. (Sigurdsson and Sundari, 1990). This enigmatic bivalve is one of the gems
of the mangal, and belongs to the Anomiidae, of which most species stay cemented to the
substratum. It uses its highly mobile foot to reach the desired level in the mangroves and
fastens temporarily with transparent byssus threads (Yonge, 1957; Berry, 1975; 1976)
     Other bivalves are adapted to the chemical environment of the mangal. Two
corbiculids, Geloina erosa and G. expansa, from Iriomote Island, Japan, occasionally
secrete thin organic sheets on the inner shell. Formation of these sheets may be a response
to shell dissolution in the acidic mangal environments; the sheets occur only in specimens
that have suffered extensive shell damage (Isaji, 1993, 1995).
     Frenkiel et al.. (1996) reported the bivalve Lucina pectinata from muddy mangrove
sediments. Like most other lucinids inhabiting sulfidic sediments, including also seagrass
beds and salt marshes, this species carries endosymbiotic chemoautotrophic sulfur-
oxidising bacteria in the gill, and the blood is rich in haemoglobin (see Somero et al., 1989
and Fisher, 1990 and references therein). These bivalves get their organic matter from the
bacteria, but the symbiosis requires proximity to both sulfide and oxygen. It has been
suggested that in seagrass beds these bivalves might benefit from the proximity to plant

roots carrying oxygen (Fisher & Hand, 1984); a similar relationship could be suggested for
mangrove roots.
    Some mollusks are critical to the basic ecology of some mangals. For example, the
mangrove snail Thais kiosquiformis plays a central role in maintaining the function and
productivity of mangroves in Costa Rica by “cleaning” their root systems of encrusting
barnacles (Koch and Wolff, 1996). Ellison and Farnsworth (1992) measured similar effects
in the mangals of Belize. Detritivorous snails (e.g., Terebralaia palustris in Gazi Bay,
Kenya) aid nutrient cycling in the mangal by processing mangrove litter (Slim et al.,
1997). Bivalves may contribute significantly to the organic biomass in the habitat and may
be a link between phytoplankton communities and higher trophic levels (e.g., Ingole et al.,
1994; Deekae and Idoniboye-Obu, 1995).
    Researchers have collected detailed information on mangrove oysters, largely
because they can be valuable food (e.g., Tack et al., 1992; Ruwa and Polk, 1994). Around
Tuticorin, India, mangroves provide ideal conditions for production of edible oysters
(Crassostrea madrasensis) and oyster beds are an important part of the habitat
(Rajapandian et al., 1990). Newkirk and Richards (1991) have found that exposing the spat
of Crassostrea rhizophorae to air increases growth and enhances the yield of marketable
oysters. This response may reflect an adaptation to the tidal regime of their mangrove
    Teredinids (shipworms) and pholads are specialized bivalves that burrow through
wood. Some species within these groups do extensive damage to mangroves by destroying
submerged roots and branches. Seven such species (Bankia campanellata, B. carinata,
Dicyathifer manni, Lyrodus pedicellatus and Teredo furcifera, Martesia striata and M.
nari) live in the Pichavaram mangroves of south India (Sivakumar and Kathiresan, 1996).
Morton (1991) recently discovered the first mangrove shipworms in Hong Kong (Lyrodus
singaporeana). A survey for teredinids in the mangroves of Sao Paulo, Brazil revealed four
species (Nausitora fusticula, Bankia fimbriatula, B. gouldi and B. rochi). Differences in
salinity tolerance affect distributions of these species within the mangal (Lopes and Narchi,

5.8. Fish
     Mangroves have a rich and diverse
assemblage of fish (Figure 12), some with
commercial value. Other fish species are
important links in the mangrove food web.
Still others are only temporary residents
that spend most of their life history
elsewhere. Whatever their role, all are
important to the character of the mangal.
     Extensive studies of the fish
                       Figure 12. The canopy of Rhizophora mangle provides
community have been made in
                       habitat for terrestrial birds and insects. The submerged
Alligator Creek, northeastern
                       prop roots provide solid surfaces for attachment of a
Queensland, Australia (Robertson and     variety of marine invertebrates. In addition, many fish
Duke, 1990a, b); in the Embley River     species use the habitat as a nursery area; the complex
estuary (Blaber et al., 1990 a, b; Salini  tangle of roots provides a refuge from predators (from
                       Rutzler & Feller, 1988).

et al., 1990; Brewer et al., 1991) and the Leanyer Swamp of the Northern Territory; and in
the Dampier region of Western Australia (Robertson and Blaber, 1992). The fish fauna is
generally very rich; 197 species occur in the mangroves of the Embley River alone. Such
high diversity is not restricted to Australian mangals. One hundred and seventeen fish
species, in 49 genera, have been recorded in the Matang mangrove waters of Malaysia
(Sasekumar et al., 1994; Yap et al., 1994) while Hong and San (1993) reported 260 fish
species in the mangroves of Vietnam.
     Abundances of the fish can also be very high. In Mexican mangroves, fish
biomasses up to 10 g m-2 have been recorded (Flores-Verdugo et al., 1990; Arancibia et
al., 1993). In Moreton Bay, Australia, the biomass reaches 20 g m-2 and ninety-six percent
of the biomass (46% percent of the species, 75% of the total fish) is from species important
in regional fisheries (Morton, 1990). Robertson and Blaber (1992) measured fish
biomasses up to 29 g m-2, with densities up to 161 individuals m-2.
     In a Queensland, Australia mangal, sampling suggests that fish regularly move
through the habitat with the tidal flows. Density and biomass at high tide were 3.5
individuals m-2 and 10.9 g m-2 respectively. On the ebb tide, the fish moved to small,
shallow creeks where density and biomass reached 31.3 individuals m-2 and 29.0 g m-2
(Robertson and Duke, 1990a). Fish distributions and abundances may also change on diel
or seasonal cycles (Chandrasekaran and Natarajan, 1993). In southwestern Puerto Rico,
fish present in the mangal during the day may completely disappear at night (Rooker and
Dennis, 1991). Accurately assessing populations of highly mobile species in such a
complex environment requires special sampling techniques (Lorenz et al., 1997).
     A comparison of catches in various habitats suggests that some species specifically
choose to reside in the mangal. For example, the number of fish species in the coastal
mangroves of Malaysia (119) exceeds that in all other habitats (inshore waters held 92
species, mudflats held 70 species, and near inshore waters held only 58 species; Chong et
al., 1990). A similar result has been found for mangrove habitats in Belize (Sedberry and
Carter, 1993). The relative importance of the mangal as habitat, however, may decrease if
nearby environments include coral reefs. Acosta (1997) found much higher fish diversity
on the reefs of La Parguera, Puerto Rico than in adjacent mangroves.
     Fish in mangrove habitats are important predators, consuming amphipods, isopods,
shrimp, nematodes, insects, gastropods, other fish, and algae (e.g., Erondu, 1990; Brewer
and Warburton, 1992; Williamson et al., 1994; Rooker, 1995; Columbini et al., 1995,
1996). In the Matang mangroves of Malaysia, a suite of fishes feed on shrimp. Croakers
(Family Sciaenidae) specialize on penaeid shrimp, consuming about 1.2 kg of shrimp ha-1
d-1 or about 17% of the total shrimp biomass (Yap et al., 1994). In the Philippines, shrimp
predation is significantly higher in bare sand areas than among mangrove pneumatophores
(Primavera, 1997).
     Feeding activities of mangrove fish can be strongly affected by local conditions. In
northeastern Florida Bay, salinity influences feeding behavior of mojarras and gold-spotted
killifish. In upstream areas with high salinity variation, the fish eat nutritionally poor algae;
in less variable downstream areas, they eat a much better diet of benthic invertebrates. Ley
et al. (1994) suggested that fish gut content measurements can be a tool to assess
environmental conditions and habitat quality. As such, it could be useful in comprehensive
monitoring and restoration programs.

    Mangals may play a special role as nursery habitat for juvenile fish. The juvenile
stages of adults that occur in other habitats (e.g., coral reefs and seagrass beds) may
migrate to the refuge on the mangal (Pinto and Punchihewa, 1996). It is common to find
large numbers of larvae and juvenile fish in net samples from mangrove habitats (Dennis,
1992; Tzeng and Wang Yu, 1992; Alvarez-Léon, 1993; Matheson and Gillmore, 1995) and
densities of juvenile fish in mangrove habitats are often higher than in adjacent habitats
(Robertson and Blaber, 1992). Thollot (1992) found that samples from the mangroves of
Southwest Lagoon, New Caledonia held 262 fish species, including the young of 30% of
the reef species. Most fish collected in the Lagos Lagoon (Nwadukwe, 1995) and in the
mangrove waters of Martinique Island (Louis et al., 1995) were small and sexually
immature. In Belize, most of the fish collected from mangrove waters are juveniles of
species that live out on the marine reefs as adults (Sedberry and Carter, 1993). Despite this
linkage to coral reefs, mangroves also have their own unique fish assemblages. Gill net
sampling in a tropical mangrove creek in SW Madagascar produced 60 species of juvenile
fish. Only six of those occurred on an adjacent coral reef (Laroche et al., 1997).
   Robertson and Blaber (1992) present three explanations for the high density of
juvenile fish in mangrove waters. First, mangrove estuaries supply an enormous amount of
food appropriate for juvenile fish (Chong et al., 1990). Second, reduced visibility in the
turbid mangrove waters may reduce predation by large fish. Third, the structural
complexity of the mangroves provides excellent shelter and protection for the juveniles.
    There is correlative evidence for the third possibility. In the Solomon Islands,
mangrove estuaries clogged with woody debris harbour pomacentrids and some species of
Apogonidae and Gobiidae. These groups are largely absent from mangrove estuaries that
are clear of the woody material (Blaber and Milton, 1990). Daniel and Robertson (1990)
also found a highly significant relationship between amounts of mangrove detritus and fish
densities or biomass in mangrove creeks. In Sri Lanka, fishermen increase their catches in
lagoons by placing thickets of dead mangrove twigs on the lagoon floor. Netting around
these thickets produces much higher catches than in adjacent bare mud areas (Robertson
and Blaber, 1992). Experiments with this technique in south India show that Avicennia
debris works better than Rhizophora debris, producing much higher catches (Rajendran,
1997; Rajendran and Kathiresan, 1998). The stilt and prop roots of some mangroves
provide a complex environment that would seem to provide an ideal refuge. However,
Mullin (1995) found more fish species in the open waters adjacent to Rhizophora mangle
stands than among the prop roots themselves, but it was unclear why the fish avoided the
    While mangroves in general may serve as nursery habitat for many fish species,
individual mangals may not. For example, Ley et al. (1999) found that mangrove habitats
in northeastern Florida Bay did not function as nursery-grounds. The authors suggest that
this particular mangrove estuary may be atypical for two reasons: (1) it has no lunar tides
and lacks typical tidal circulation, (2) it has little submerged vegetation; and (3) it
experiences severe hypersaline conditions. Conditions in this particular environment may
be sufficiently stressful to prevent its use by juvenile fish.
    Fish living in the mangal must adjust to temporal and spatial variability in physical
and chemical conditions, and some species possess specific adaptations to deal with this.
For example, the widely distributed hermaphrodite killifish (Rivulus marmoratus) is well
adapted to mangrove microhabitats (Taylor et al., 1995). Its specializations include an

ability to survive in moist detrital substrates during periods of low water or drought and
reproduction by internal self-fertilization (Davis et al., 1995).
     Fish without such specific adaptations may respond behaviorally to physical cues
that indicate physically or chemically stressful microhabitats. This can lead to distinct
distributional patterns (Blaber et al., 1994). For example, Heath et al. (1993) demonstrate
experimentally that thermal cues affect fish distributions within mangrove ponds.
Cyprinodon prefers higher temperatures than other fish species tolerate. Temperature
differences within the mangal could, therefore, spatially separate fish populations. Fish
may also prefer certain areas of the mangal based on the nature of the substrate. Kimani et
al. (1996) sampled fish populations in an estuarine mangrove bay in Kenya for 12 months.
Diversity, density, and biomass were all lower in a silty area than in three other mangrove
areas with established seagrass beds.
     Hypoxia, which affects plasma osmolality, plasma chloride ion concentration, and
hematocrit in fish (Peterson, 1990; Peterson and Gilmore, 1991) can also influence their
distributions. For example, juvenile snook (Centropomus undecimalis) move toward
oxygenated surface waters when deeper waters become anoxic (Peterson and Gilmore,
1991). Habitat degradation can lead to changes in fish distribution. In the Virgin Islands,
differences in fish abundance and diversity between degraded and natural mangrove sites
are directly related to water quality (Boulon, 1992). The simple installation of culverts
creates better water exchange in some mangals, promoting reestablishment of marsh
vegetation, and increased fish production (Lin and Beal, 1995).
     Mudskippers are a group of unusual amphibious fish (Family Gobiidae: Subfamily
Oxudercinae) that are characteristic residents of many mangals. A variety of anatomical,
physiological, and behavioral adaptations help them tolerate the stressful mangrove
environment (Chew and Ip, 1990; Colombini et al., 1995, 1996; Ikebe and Oishi, 1996,
1997; Ip et al., 1991; Ishimatsu et al., 1999; Ogasawara et al., 1991; reviewed by Clayton,
     Researchers have long believed that one of the primary adaptations of mudskippers
is an ability to endure extremely hypoxic conditions. However, recent evidence suggests
that this may not be true for all species; many mudskippers are less tolerant of hypoxia
than has been assumed. Takeda et al. (1999) demonstrated that Periophthalmodon
schlosseri could recover from post-exercise oxygen debt, but only in air. Furthermore,
laboratory experiments demonstrate that P. schlosseri may rarely use aquatic gill
ventilation at all, apparently prefering aerial respiration (Aguilar et al.., in press). A limited
capacity for aquatic oxygen uptake has also been proposed for mudskippers in the genus
Periophthalmus. Aguilar (2000) suggests that Periophthalmus spp. lack physiological
mechanisms to tolerate hypoxia and instead have a range of adaptive behaviors that allow
them to avoid low-oxygen conditions.
     A unique behavioral mechanism may enable some species to completely escape
hypoxia. A number of mudskippers dig extensive burrows into anoxic mangrove
sediments and might be expected to experience extremely hypoxic conditions, particularly
during periods of low tide. Ishimatsu et al. (1998) found that Periophthalmodon schlosseri,
for example, creates burrows as deep as 125 cm. However, the burrows have special
excavated chambers that hold pockets of air. The P. schlosseri fill the pockets by gulping
air at the surface, carrying it down to the chamber and releasing it there. The fish,
therefore, escape rather than tolerate hypoxia. Such burrowing and air trapping,

particularly where mudskipper populations are dense, may significantly affect oxygenation
and chemistry of the mangrove sediments (Kristensen et al., 1995).
5.9. Amphibians and Reptiles
    Reptiles, including crocodiles, alligators, lizards, snakes and turtles live in many
mangroves. About 35 reptile species are known from the Sunderbans of Bangladesh alone.
The most notable ones are saltwater crocodiles (Crocodylus porosus), monitor lizards
(Varanus bengalensis, V. salvator and V. flavascens), King cobras (Ophiophagus hannah),
Green Pit vipers (Vipera trimeresurus), Rock pythons (Python molorus), and the olive
Ridley turtles (Lepidochelys olivacea, Hussain and Acharya, 1994). In Liberia, Nile
crocodiles occur only in the brackish water of mangrove swamps and at river mouths
(Kofron, 1992). Indo-Pacific crocodiles Crocodylus porosus are abundant in the upper
mangrove sections of the Klias River, Sabah, Malaysia (Stuebing et al., 1994).
    The amphibian fauna of the Sunderbans mangal includes four genera of frogs
(Rana, Bufo, Microhyla and Rhacophorus). The ground frog Rana hexadactyla, the tree
frog Rhacophorus maculatus, and the toad Bufo melanostictus are particularly common
(Gopal and Krishnamurthy, 1993; Hussain and Acharya, 1994). The amphibian fauna has
not been well studied in most other mangals.
    Human activities that impact mangroves have cascading effects on the reptile and
amphibian fauna. There have been drastic declines in the population of crocodiles in the
mangals of Liberia (Kofron, 1992) and of both crocodiles and snakes in the mangroves of
Bangladesh (Hussain and Acharya, 1994). Habitat loss through human encroachment is a
primary cause of the decline. These impacts are likely to continue, and worsen, as human
populations expand further into the mangals.

5.10. Birds
    Mangroves provide important habitat for landbirds, shorebirds and waterfowl, and
they are home to a number of threatened species including spoonbills (Ajala ajala), large
snowy egrets (Cosmorodium albus), scarlet ibis (Eudocimus ruber), fish hawks (Pandion
haliaetus), royal terns (Sterna hirundo), West Indian whistling-ducks (Dendrocygna
arborea), and Storm's Storks (Danielsen et al., 1997; Panitz, 1997; Staus, 1998). The birds
in the mangal may be permanent residents that forage and nest in the mangroves and the
mangrove waters or they can be temporary visitors. Lefebvre et al. (1994) measured bird
abundances and grouped species according to their diet and the frequency with which they
use mangrove habitats. Distributions and abundances of the feeding guild were consistent
with the abundance and distribution of their invertebrate prey (Lefebvre and Poulin, 1997).
    Lefebvre et al. (1992) studied distributions of passerine birds in the mangroves of
Venezuela and found that they form highly stable territories. In Singapore, sand pipers,
plovers, herons and egrets all regularly use mangrove habitat (Murphy and Sigurdsson,
1990). About 315 species of birds are known from the Sunderbans of Bangladesh. The
most common ones are white-bellied sea eagles (Haliaetus leucogaster) and Pallas’s fish
eagles (Haliaetus leucorhyphus; Hussain and Acharya, 1994). Alves et al. (1997) counted
32 bird species (2 marine species, 18 terrestrial species, and 12 waterfowl) in the
mangroves of Jequiaman, Brazil.
    Migratory birds visiting the mangroves may fly long distances to find food and
nesting places there. This may be particularly true in the Neotropics (Parrish and Sherry,
1994; Confer and Holmes, 1995; Lefebvre and Poulin, 1996; Panitz, 1997). Seventy-seven
BIOLOGY OF MANGROVES AND MANGROVE ECOSYSTEMS                                             57

bird species have been recorded in the Pacific mangroves of Colombia. Forty-three percent
of these are permanent residents, 22% are regular visitors and 18% are temporary winter
residents (Naranjo, 1997). One migratory species, the black-crowned night heron
(Nycticorax nycticorax) is an important vector for disease. This mangrove-breeding bird is
the principal host for Japanese Encephalitis Virus, which it widely disseminates during its
migrations (Murphy and Sigurdsson, 1990).
    Some of the resident bird species are highly dependent on mangroves for their
survival. The yellow warbler (Dendroica petechia) and the mangrove vireo (Vireo pallens)
are nearly confined to mangroves (Parkes, 1990; Buden, 1992). The mangrove gerygone
spends 80% of its time on Avicennia marina (Noske, 1996) while A. germinans provides
important breeding habitat for Florida Prairie Warblers (Dendroica discolor paludicola)
and Cuban Yellow Warblers (D. petechia gundlachi; Prather and Cruz,1995).
    Because of this dependence, disturbances to the mangal may reverberate through
the bird populations. This may be particularly true where the bird species show strong site
fidelity (Warkentin and Hernandez, 1996). The habitat disturbances may be natural, such
as the frequent cyclonic storms that strongly affect myna populations in the Pichavaram
mangroves of south India (Nagarajan and Thiyagesan, 1995). More frequently, however,
they are caused by human activities.
    Mangrove forest destruction and fragmentation, usually due to development, reduce
effective habitat and threaten mangrove-dependent species. Bancroft et al. (1995) found
reduced populations of mangrove cuckoos
(Cocyzus minor) in fragmented mangrove
areas of Florida, USA (Figure 13).                    Original      1991

Similarly, the mangrove finch (Cactospiza                                        Northern Flicker
heliobates), once present in six mangrove      80
                                     1 km

areas on two of the Galapagos Islands, are
                            Occurrence (%)

                                                  Mangrove Cuckoo
now restricted to four mangrove pockets on                White-eyed Vireo

one island. Habitat destruction has                                         Yellow-billed Cuckoo

completely eliminated the finches from the      20

other island (Grant and Grant, 1997).
Ironically, one potential threat to these        0-1  2-5    5 - 10         10 - 20 > 20

populations is the birdwatchers who explore                  Forest size (ha)

hoping to see the birds in their natural       Figure 13. Abundance of mangrove-associated
environment (Klein et al., 1995; Ellison and     land birds as a function of forest size. Inset
                           shows changes in the mangrove forests of Upper
Farnsworth, 1996a).
                           Matecumbe Key (Florida Keys, Florida, USA)
    Protection of the mangrove-dependent
                           from an original 252.2 ha forest to 105 forest
birds will require effective management of the    fragments totalling only 61.4 ha. Records of 4
entire mangrove habitat. This may be complex     bird species show that abundance is strongly
and require evaluation of habitat needs on a     affected by forest size (after Bancroft et al, 1995).
species by species basis. For example, in
Florida Bay, bald eagles (the U.S. national bird) nest almost exclusively in mangrove trees
(Avicennia germinans and Rhizophora mangle). Many of the nest sites are in snags (dead
trees) suggesting that a comprehensive eagle management plan will require preservation of
both living and dead mangroves (Curnutt and Robertson, 1994).

5.11. Mammals

    A variety of mammals make their homes in the mangal. Their ecology within the
mangal and their associations with the mangroves themselves have been little studied and
are poorly known. Some of the noteworthy species present include dolphins (Platenista
gangetica), mangrove monkeys (Macaca mulatta) and otters (Lutra perspicillata ) in India
(Gopal and Krishnamurthy, 1993); flying fox (Pteropus conspicillatus and Pteropus
alecto) in northern Australia (Richards, 1990; Loughland, 1998) capuchin (Cebus apella
apella) in Brazil (Fernandes, 1991). In southeastern Brazil, distributions of some cetacean
species can also be related to the distribution of mangroves (Martuscelli et al., 1996).
Small clawed otters (Lutrinae) shelter amongst Acrostichum ferns during the dry season in
the mangroves of Singapore and Malay (Sivasothi and Burhanuddin, 1994).
    Thirty-two mammal species once lived in the Sunderban mangals of Bangladesh.
Four of these, Javan rhinoceros (Rhinoceros sondaicus), wild buffalo (Bubalus bubalis),
swamp deer (Cervus duvauceli) and hog deer (Axis porcinus), have gone extinct since the
beginning of this century (Hussain and Acharya, 1994). Two additional species, the Royal
Bengal Tiger (Panthera tigris) and the Chital deer (Axis axis), are currently endangered.
Studies have shown that the chital deer browses directly on the mangroves. Its feeding
damages Avicennia officinalis, Xylocarpus mekongensis, Bruguiera sexangula, and
Aegiceras corniculatum but has no effect on Heritiera fomes, Excoecaria agallocha or
Ceriops decandra (Siddiqi and Hussain, 1994).
    Loss of mammalian species in the world's mangrove environments is probably the
result of habitat fragmentation. This is particularly true for some of the larger species that
have large home ranges. This may largely explain the loss of species from the Sunderbans.
Habitat loss, however, can also have a major effect on smaller species. Forys and
Humphrey (1996) studied distribution and movement of an endangered marsh rabbit
(Sylvilagus palustrishefneri) in the Lower Florida Keys, USA. They found that the rabbits
use mangrove tracts as dispersal corridors between marsh habitats. Preservation of this
species will require protection of both the marsh and the mangrove corridors. Careful study
will be required to implement effective conservation plans for the mammals faced with
shrinking, and fragmenting, mangrove habitat.


6.1. Responses to light
    Although mangroves occur in tropical habitats where they are exposed to high light
intensities, their photosynthetic rates tend to plateau at relatively low light levels. The trees
possess mechanisms to deal with the high sunlight (see section 3.3.2). For example,
Avicennia marina shows good resistance to high sunlight, hot and dry conditions and is
well adapted to arid zones (ElAmry, 1998). However, there is evidence that intense light
can damage the mangroves despite these adaptations. For example, Cheeseman et al.
(1991) demonstrated that rates of photosynthesis drop in mangroves exposed to excessive
sunlight. This could explain why Rhizophora seedlings establish and sprout most readily
under the shady canopy of larger trees (Kathiresan and Ramesh, 1991; Kathiresan, 1999).
Kathiresan and Moorthy (1993) also demonstrated that the seedlings grow faster in the
shade, use NO3 more efficiently, and show more efficient photosynthesis.

    The negative effects of sunlight may, in some cases, be due to the high doses of
UV-B radiation that the mangroves receive (Moorthy, 1995). To date, however, there have
been few studies of UV-B and mangroves (Lovelock et al., 1992; S.M. Smith and
Snedaker, 1995b). Moorthy (1995) and Moorthy and Kathiresan (1997a) studied mangrove
responses to UV-B in the Pichavaram mangroves of south India (total sunlight and UV-B
intensities here may exceed 1300 W m-2 and 0.31 W m-2 respectively). Species in the
Rhizophoraceae showed greater UV-B tolerance than did Avicennia species or other
succulent plants.
    To better understand potential effects of global increases in UV-B, Moorthy and
Kathiresan (1997b) grew Rhizophora apiculata seedlings under the UV-B regimes
predicted for 10, 20, 30 and 40% stratospheric ozone depletions. Net photosynthetic rates
of seedlings increased 45% under the 10% UV-B treatment (stomatal conductance
increased 47%). Raising the UV intensity to the 40% level, however, produced a 59%
decrease in net photosynthesis and a 73% increase in intercellular CO2 concentrations
(Moorthy, 1995).
    Increasing UV-B exposure also produced biochemical changes. Phenol and
flavonoids levels increased with UV-B dose, but anthocyanin concentrations dropped.
Small UV-B doses enhanced amino acid and protein levels but the effect was reversed at
higher UV-B levels. The UV-B, in general, enhanced saturated fatty acids (maximum
increase of 88%) and reduced unsaturated fatty acids (maximum decrease of 26%). Any
UV-B exposure also inhibited the activity of nitrate reductase while simultaneously
enhancing total tissue nitrate (Moorthy and Kathiresan, 1998). Both growth and
biochemical responses indicate that the mangroves are stressed by the high intensity UV-B.
    While excessive sunlight can damage mangroves, shading can also have negative
effects. Mangrove seedlings under a closed canopy showed lower growth in south Florida
(Koch, 1997) and Belizean mangals (Ellison and Farnsworth, 1993). The appearance of
gaps in the canopy produced rapid growth of the previously shaded trees. In dense
mangrove forests, shaded saplings have lower shoot biomass than those exposed to the sun.
The saplings may compensate for this by increasing development of the pneumatophores
(Turner et al., 1995).
    Shade tolerance differs among mangrove species. Clarke and Allaway (1993) found
that shading had no effect on growth or survival of Avicennia marina. But, McKee (1995b)
noted that shading with brief periods of light exposure increased biomass and growth in
Avicennia germinans and Laguncularia racemosa. The same treatment had little effect on
Rhizophora mangle. Ellison and Farnsworth (1993) found that R. mangle seedlings
performed better overall in larger canopy gaps. In R. mangle, ontogenetic developments
produce changes in light adaptation. Seedlings are apparently adapted for the shaded
understory environment while mature trees do better in the sunlit canopy (Farnsworth and
Ellison, 1996b).

6.2. Responses to gases
    Because of the environment they live in, mangroves may experience episodic, or
chronic oxygen stress. The consequences of low oxygen vary among species and appear
related to the physiological and morphological adaptations of each (McKee et al., 1996).
For example, McKee (1993) found that flooding and anoxia reduced the total biomass of
Avicennia germinans seedlings 20-40% relative to drained controls. However, under
BIOLOGY OF MANGROVES AND MANGROVE ECOSYSTEMS                                60

identical conditions, the biomass of Rhizophora mangle seedlings increased 9-24%. The
differential tolerance of these species for flooding and low oxygen may partly result from
differences in root aeration. R. mangle maintains high oxygen concentrations in its roots
even under reducing soil conditions; A. germinans does not. Skelton and Allaway (1996)
showed that a congener (A. marina) also does not maintain gas pressures under low-
oxygen conditions. Pressures in the aerial roots drop during high tide, probably due to
removal of respiratory CO2 from gas spaces during flooding. As the waters recede on the
low tide, a rapid influx of air may take place.
      High methane levels can be associated with anoxia in mangrove environments.
The methane flux from the sediments is strongly influenced by freshwater loading and
nutrient input (Sotomayor et al., 1994; Giani et al., 1996). Fluxes may also vary along tidal
gradients being generally low on the landward fringe and high in the seaward transition
zone between Avicennia and Rhizophora communities (e.g., Ye et al., 1997). Mangrove
species with pneumatophores may be best equipped to deal with high methane loads. The
pneumatophores themselves may be conduits for release of methane gas (Sotomayor et al.,
1994). Pneumatophore-bearing species also release more methane through their leaves
than do those lacking pneumatophores (Lu et al., 1998).
      Predicted global changes in atmospheric carbon dioxide are likely to have strong
                                 effects on mangroves. Farnsworth et al.
                                 (1996) grew Rhizophora mangle under
   25                400
                Ambient CO2
                                 double ambient CO2 for 1 year. Growth
                Elevated CO2
                              Total stem length (cm)
      Number of branches

                                 rate, net assimilation, and photosynthetic

                                 rate all increased significantly. Seedlings in
                                 the enhanced CO2 treatment had greater

                                 biomass, longer stems, more branching, and
                                 more leaf area than control seedlings
   0                  0
                                 (Figure 14). They also became reproductive
    0 100 200 300 400         0  100 200 300 400

                                 after only 1 year (2 years sooner than under
                                 normal conditions). Ellison (1994) found
Total plant biomass (g)

                        Total leaf area (cm2)

                                 that, in addition to stimulating productivity,

                                 increased CO2 led to more efficient use of
                                 water as a result of reduced stomatal
                                 conductance (Ellison, 1994).
   60                1000

                                     The effects of increased CO2 may
   20                400
                                 vary with other physical and chemical
    280 320  360 400         280  320  360 400

                                  conditions (Ball and Munns, 1992). For
           Days elapsed since planting
                                  instance, Rhizophora apiculata and R.
Figure 14. Effects of increased CO2 on Rhizophora
mangle growth and morphology. Predicted changes          stylosa both benefit from increased CO2,
in the global climate could result in significant         but the stimulatory effect is much greater
increases in CO2 levels. Experimentally doubling the
                                  under low salinity conditions (Ball et al.,
ambient CO2 levels produced significant increases in
                                  1997). The effects of increased CO2 may
number of branches, total stem length, plant biomass,
                                  also differ among habitats and species.
and total leaf area (after Farnsworth et al, 1996).
                                  Snedaker and Araújo (1998), for example,
studied the effects of a 6 - 34 % increase in CO2 concentration on four mangrove species in
south Florida (Rhizophora mangle, Avicennia germinans, Laguncularia racemosa, and
Conocarpus erectus). Elevated CO2 reduced stomatal conductance and transpiration and

significantly increased instantaneous transpiration efficiency in all of these species.
However, it did not increase net primary productivity in any species and actually reduced
the productivity of Laguncularia racemosa.

6.3. Responses to wind
     Tropical storms (hurricanes and cyclones) cause enormous damage to mangrove
forests, particularly in the Caribbean and the Bay of Bengal. The immediate consequence
is loss of the mangrove trees themselves. In 1988, for example, a severe cyclone in
Bangladesh destroyed 8.5 million trees with a loss of 66.3 million m3 of commercial saw
timber (e.g. Mastaller, 1996). Large numbers of trees have similarly been defoliated and
destroyed by storms in South Florida, Guadeloupe, Nicaragua, Belize, and Puerto Rico.
     Damage to the mangroves may also have indirect consequences. For example,
Hurricanes Gilbert and Joan, which both hit the Caribbean in the last quarter of 1988,
caused mass mortality of invertebrates growing on the roots of Rhizophora mangle
(Orihuela et al., 1991). Hurricane Hugo, while passing through the coastal environment of
Guadeloupe, French West Indies, killed large numbers of fish, producing changes in the
fish community (Bouchon et al., 1991, Imbert et al., 1996).
     A far-reaching consequence of mangrove mortality can be serious erosion of the
coastal habitat. Hurricane Andrew made landfall in 1992 on the mangrove-fringed coasts
of south Florida. Uprooted mangrove vegetation left behind unprotected intertidal and
subtidal sediments that were subsequently eroded by currents and waves. (Davis, 1995;
Doyle et al., 1995; Swiadek, 1997). In Bangladesh, a mangrove afforestation project was
initiated in 1966. A primary goal of the project was to provide a mangrove buffer that
would protect the coast from frequent cyclone damage. By 1993, 0.12 million ha had been
afforested (Saenger and Siddiqi, 1993).
     Some effects of storm damage may not be seen until some time after the event.
Often, trees that are broken or severely damaged by the storm later die. T.J.Smith et al.
(1994) found that mangrove mortality in south Florida continued for many weeks after
hurricane Andrew. Many mature trees later died from injuries sustained in the storm.
Propagules and seedlings in the habitat were also killed, largely by sedimentation and high
porewater sulfides.
     Differential species mortality associated with a major storm can change community
structure. For example, smaller Rhizophora mangle were not greatly affected by Hurricane
Andrew whereas large Laguncularia racemosa were heavily damaged (McCoy et al.,
1996). These differences in survival have produced a shift in species distributions in some
areas (Baldwin et al., 1995). A similar shift can be seen with Avicennia germinans and
Rhizophora mangle. Because A. germinans, cannot tolerate long periods of pneumatophore
submergence, storms and hurricanes in Florida may promote replacement of A. germinans
by R. mangle (Rey et al., 1990a). Even where the regenerated forest is composed of the
same species present before the hurricane, different regeneration rates may shift the
proportions of those species (Roth, 1992, 1997).

6.4. Responses to coastal changes
    Mangroves are tightly bound to the coastal environments in which they occur. Not
only are they influenced by chemical and physical conditions in their environment, they

help create those conditions. As a result, perturbations to the system can have cascading
long-term effects.
     Many changes seen in coastal mangals can be attributed to changes in hydrology.
Some of these changes are favorable. For example, in Singapore, increases in the
frequency of tidal inundation has created new mangal (comprised of Avicennia and
Sonneratia alba) adjacent to established mangrove stands (Lee et al., 1996). More often,
however, hydrological changes result in destruction of mangroves. In Pichavaram, south
India, changes in topography and tidal flushing have caused large-scale degradation of
mangroves. The mangroves are healthy and diverse where the land is flat. If water flow is
reduced, flat areas become shallow basins. The poor flushing and resultant hypersalinity
stunt the mangroves or replace them with saltmarsh (Suaeda spp.) or barren soil devoid of
vegetation. The process can be reversed by simply increasing the free flow of tidal waters
(Selvam and Ravichandran, 1998).
     In Senegal, decreasing rainfall and increasing evaporation have markedly changed
mangrove populations. The changes have been accelerated by altered tidal conditions
resulting from the breaching of a protective sand dune (Diop et al., 1997). Riverine
mangroves affect the dynamics of tidal currents (Wolanski et al., 1992), producing
asymmetrical tidal currents that may be 50% stronger on the ebb than on the flood. Erosion
from these flows creates deep channels through the habitat (Medeiros and Kjerfve, 1993).
Deforestation changes the tidal asymmetry and leads to changes in channel structure
(Wolanski et al., 1992).
     Human attempts to modify the physical character of the mangal by erecting hard
structures or by dredging can also drastically alter the system. In Florida, a culvert
connecting a mangrove marsh to a tidal lagoon was closed in 1979. This led to
overflooding and hypersalinity ( > 100 ppt) that eliminated the marsh. The culvert was
reopened in 1982, but the mangroves did not recover (Rey et al., 1990a, b).
     Another potential consequence of flow modification is change in sedimentation
patterns (e.g., Q. Zhang et al., 1996). On spring flood tides under normal conditions, about
80% of the sediment transported into the Middle Creek (Cairns, Australia) is trapped by
the mangroves. This corresponds to 10-12 kg of sediment m-1 on each spring tide and
could produce sediment accretions of 0.1 cm y-1 (Furukawa et al., 1997). Such levels of
sediment trapping can produce major changes to the habitat. Chakraborti (1995) traced the
evolutionary history of coastal quaternary deposits on the Bengal Plain of India and found
that mangroves were the dominant geomorphic agents in the evolution of tidal shoals and
their eventual accretion to the mainland.
     Damage to the mangroves strongly affects sediment budgets and promotes coastal
erosion (Kamaludin and Woodroffe, 1993). The eroded sediments may then cause further
damage the mangroves. For example, Young and Harvey (1996) showed that sediment
accretion interferes with root aeration in Avicennia marina var. australasica. Similarly,
movements of sand in mangrove habitats on Portuguese Island, Mozambique have caused
high mortality of Ceriops tagal. This has changed the mangrove species composition and
had significant secondary effects; all crustaceans and mollusks have also disappeared from
the mangal (Hatton and Couto, 1992).
     Major changes in mangrove distribution and abundance in coastal regions could
result from habitat loss associated with rising sea level (Fujimoto and Miyagi, 1990;
Woodroffe, 1990; Ellison, 1993; Parkinson et al., 1994). The vulnerability of individual

mangals can be evaluated through annual measurements of soil elevation deficit (elevation
change minus sea-level rise). Cahoon and Lynch (1997) suggest that this is a better
measure than the more commonly used accretion deficit (accretion minus sea-level rise).
Historical patterns of sea level rise in a mangrove environment can be evaluated through
measurements of the mangrove trees themselves since the δ18 oxygen fraction in the wood
is an effective seawater tracer (Ish-Shalom-Gordon et al., 1992).
     Ellison and Stoddart (1991) suggested that mangroves are stressed by sea level rises
between 9 and 12 cm • 100 y-1 and concluded that faster rates could seriously threaten
mangrove ecosystems. This view has been challenged by Snedaker et al. (1994) who cite
historical records showing mangrove expansion under relative sea level changes nearly
twice that great. X. Tan and Zhang (1997) conclude that, given current rate estimates, sea
level rises pose no significant threat to most mangrove forests in China. The effects of sea-
level rise on any mangrove habitat will be influenced by local wetland type, geomorphic
setting, and human activities in the wetland. There is a need for better models predicting
effects of sea level and other coastal changes on individual mangals (e.g., Bacon,1994).

6.5. Responses to tidal gradients and zonation

     Zonation can be a structural feature of
mangrove forests in some parts of the world
(T.J. Smith, 1992; Woodroffe, 1992).
However, unlike open coast habitats where
zonation patterns are distinct, mangrove
distributions are extremely variable and
extensive surveys may be necessary to fully
document patterns, particularly if diversity is
high (Bunt 1996). The “zones” may be
obscured by broad overlap in species
distributions (e.g., Bunt and Bunt, 1999; Bunt
and Stieglitz, 1999; Figure 15) or they may
simply be absent in some mangals (Ellison et
al., in press). Bunt (1999) has developed
methods specifically for evaluating and
describing mangrove zonation.
     Where zonation does occur,
contributing factors may include plant       Figure 15. Distribution of mangroves south of Townsville,
succession, geomorphology, physiological      North Queensland, Australia. Stippled areas of the map
adaptation, propagule size, seed predation     represent mangal. Details of transect A-B are shown in the
and interspecific interactions (e.g., Bunt et   lower panel. The dominant mangrove species (i.e.,
                          Avicennia marina, Rhizophora stylosa, Ceriops tagal,
al., 1991; Woodroffe, 1992; Ellison and
                          Bruguiera gymnorhiza, and Osbornia octodonta) are
Farnsworth, 1993; Schwamborn and Saint-      represented by different symbols. Note that the mangroves
Paul, 1996). The relative importance of      are closely associated with rivers and creeks, resulting in
these factors, however, depends on the       broadly overlapping species distributions (after Macnae,
individual habitat (e.g., McKee, 1995c) and    1967).
there is disagreement about the general

importance of some of them. For example, Robertson et al. (1991) argue that succession
plays a minor role in mangrove zonation and that simple erosion and sedimentation control
the distribution of mangroves along the seaward edge of the mangal. The complexity and
uniqueness of these communities may make it difficult even to define successional stages
(Roth, 1992). The term “old-growth”, for instance, cannot be applied easily to mangrove
forests (Lugo, 1997).
    One potential cause of mangrove zonation is the differential ability of propagules to
establish at different tidal heights. This is directly related to propagule size. It has been
suggested that small propagules drift further inland and establish better in shallow water
than do large propagules (thus producing a species zonation dependent on propagule size;
Kathiresan, 1999). The importance of this process to the creation of mangrove zones has
been clearly demonstrated for Avicennia bicolor and Rhizophora racemosa on the Pacific
coast of Costa Rica (Jiménez and Sauter, 1991). However, the more general importance of
this process has been contested (T.J.Smith, 1992).
    Interspecific differences in tolerance for physiological stress is perhaps the best
demonstrated cause of mangrove zonation. However, while physiological responses to
physicochemical conditions undoubtedly influence mangrove distributions in some
habitats, conclusions must be made cautiously since field measurements do not always
support laboratory conclusions (Schwamborn and Saint-Paul, 1996). Despite this
limitation, it is clear that mangrove species respond differently to different tidal regimes.
For example, in the Indian Sunderbans, a mangrove forest that experiences total diurnal
inundation is dominated by Avicennia marina and A. alba while Excoecaria agallocha,
Ceriops decandra and Acanthus ilicifolius dominate sites that are not completely inundated
(Saha and Choudhury, 1995). Nypa fruticans also seems to prefer sites with low level of
tidal inundation (Siddiqi, 1995). Kathiresan et al. (1996a) studied growth of Rhizophora
apiculata seedlings in different tidal zones of a south Indian estuary. Individuals in the
low intertidal grew 2.5 times faster and sprouted 4 times as many leaves as individuals in
the highest zone. Similar patterns of differential survival and growth have been seen in the
mangroves of Qatar (Abdel-Razik, 1991) and New Zealand (Osunkoya and Creese, 1997).
    Ellison and Farnsworth (1993) studied survival and growth of Rhizophora mangle
and Avicennia germinans seedlings at tidal heights corresponding to lowest low water
(LLW), mean water (MW), and highest high water (HHW). R. mangle seedlings survived
in the MW (69%) and LLW (56%) treatments; Avicennia survived at MW (47%), but not
at LLW. Neither species survived at HHW. Seedlings of both species suffered twice as
much insect damage in the MW treatment as in the LLW treatment. The combination of
insect herbivory and differential flood tolerance may create the zonation of these two
species in the Caribbean. Further experimental studies of this nature should help clarify the
causes and consequences of zonation in mangrove communities.

6.6. Responses to soil conditions
    Soil properties have a major impact on mangrove nutrition and growth. Some of the
most important characteristics are siltiness, electrical conductivity, pH, and cation
exchange capacity (Kusmana, 1990; V.B. Rao et al., 1992; Pezeshki et al., 1997). The
most important factor, however, appears to be nutrient concentrations. Mangals are finely
balanced, highly effective nutrient sinks with net imports of dissolved nitrogen,

phosphorus, and silicon. Nutrient fluxes in these environments are closely tied to plant
assimilation and microbial mineralization (Alongi, 1996; Middelburg et al., 1996).
    Nutrients availability may limit growth and production in many mangals. Varying
nutrient concentrations can also change competitive balances and affect species
distributions (Chen and Twilley, 1998). As a result, nutrient pulses can create immediate,
and impressive, changes in the vegetation. For example, on the southeast coast of India,
high nutrient concentrations and low salinity from monsoons produce rapid growth in the
mangroves. Seedlings grow 5X as much and produce 4X as many leaves in the post-
monsoon season as they do in the dry season (Kathiresan et al., 1996a).
    The limiting nutrients may vary with individual mangrove habitats. For example,
potassium levels may be important in some regions. Rhizophora apiculata seedlings do
significantly better in plantation sites with enriched potassium (Kathiresan et al., 1994a).
In general, however, mangroves in low-nutrient carbonate soils are limited by phosphorus.
What phosphorus is present may be bound with calcium, effectively holding it within the
sediments (Silva and Mozeto, 1997). In mesocosm and field experiments with Rhizophora
mangle seedlings, phosphorus enrichment produced nearly a 7 fold increase in stem
elongation rates and a 3 fold increase in leaf area. Nitrogen addition produced no such
response (Koch and Snedaker, 1997). Low phosphorus availability similarly limits growth
of dwarf R. mangle in a Belizean mangal (Feller, 1995) and promotes development of hard,
long-lived leaves called sclerophylls. The sclerophylls may be an adaptation to conserve
nutrients in these oligotrophic habitats (Feller, 1996). Mangroves may have other
mechanisms to hold valuable nutrients. For example, mature photosynthetically active
leaves have much higher nitrogen and potassium concentrations than senescent leaves.
This is apparently a consequence of nutrients being translocated out of the aging leaves and
into other plant parts before the leaves fall (Soto, 1992).
    Damage to a mangal may compromise its ability to retain nutrients. For examples,
at severely damaged mangrove sites in North Queensland, Kaly et al. (1997) measured a
significant loss of both nitrogen and phosphorus from the soils. This may have been related
to declines in the crab populations and a dramatic decrease in density of burrows. The
effects of disturbance will differ from habitat to habitat and will depend on the sediment
characteristics and flow regimes of each site. For example, Triwilaida and Intari (1990)
found no differences in soil nutrient concentrations between degraded and healthy
mangrove stands in the Pedada Strait, Riau.
    Sulfides are a characteristic feature of mangrove sediments that influences
mangrove distributions and is influenced by their presence. Tidal mixing, bioturbation, and
the mangroves themselves (Holmer et al., 1994) control the distributions and
concentrations of the sulfides. For example, reoxidation of sulfides is facilitated by roots;
soils are often less reduced near the aerial roots of some species. This leads to lower
sulfide levels. In a neotropical Florida mangal, the zone dominated by Rhizophora mangle
(with its numerous aerial prop roots) has moderately reducing soils with low sulfide levels.
In contrast, the Avicennia germinans zone has strongly reducing soils with high sulfides
(McKee, 1993). Surprisingly, this pattern is not repeated in a similar mangal in Brazil. In
that location, the R. mangle soils are highly reducing with high sulfide concentrations. The
sulfide content of the A. germinans soil is highly variable as the rhizosphere changes from
oxygenated to anoxic conditions (Lacerda et al., 1995). The Avicennia soils also contain
more exchangeable trace metals (Lacerda et al., 1993).

    Reduction of sulfate to sulfide is generally slower in young forests, resulting in
higher nutrient levels and lower sulfide toxicity (Alongi et al., 1998). High sulfide levels
can damage mangrove seedlings, causing stomatal closure, reduced gas exchange, reduced
growth, and high mortality (Youssef and Saenger, 1998). Disturbance can increase rates of
sulfate reduction. Clearing of mangrove forests, or simple formation of canopy gaps can
change the physical and chemical characteristics of the underlying soils (Ewel et al.,
1998a), leading to anaerobiosis and increased sulfide in the sediments (Ibrahim, 1990).
Heavy organic input can also increase sulfide production. In Ghana, pyrite (ferric sulfide)
accumulates directly in the upper layer of the mangrove soils. The rate of accumulation is
directly related to vegetation thickness (Nonaka et al., 1994).
    Under normal conditions, sulfides combine with metals in the sediment and
precipitate out as metal sulfides. When the metals available for sulfide precipitation are
exhausted, H2S is formed (Kryger and Lee, 1995). Kryger and Lee (1996) found that the
H2S from anaerobic processes accumulates in cable roots of Avicennia species as the
sediments age. Concentrations of H2S in the roots may be 30-40 times higher than in the
surrounding sediments. The H2S accumulation can kill the mangroves if their
pneumatophores are covered by silt and cannot transport oxygen to the rhizospheres.
Because they have aerial roots, Rhizophora species can better survive on aged mangrove
soils high in H2S. They may, therefore, be a natural successor to the less-tolerant Avicennia

6.7. Responses to salinity
    Salinity, as controlled by climate, hydrology, topography and tidal flooding, affects
the productivity and growth of mangrove forests (Sylla et al., 1996; Twilley and Chen,
1998). It can also strongly influence competitive interactions among species (Ukpong,
1995; Ukpong and Areola, 1995; Cardona and Botero, 1998). The distributions of plant
species within the mangal, in many cases, can be explained primarily by salinity gradients
(Ukpong, 1994; Ball, 1998).
    In general, mangrove vegetation is more luxuriant in lower salinities (Kathiresan et
al., 1996a). However, low salinity associated with long periods of flooding contributes to
mangrove degradation through reduced cell turgor and decreased respiration (Triwilaida
and Intari, 1990). On the Pacific coast of Central America, freshwater availability (largely
from rainfall and surface runoff) controls reproductive phenology, growth and mortality of
Avicennia biocolor (Jiménez, 1990).
    Even in mangals with strong riverine input, the combined effects of evaporation
and transpiration may remove much of the fresh water entering the system (Simpson et al.,
1997). The plants must, therefore, have some salinity tolerance. True mangroves (e.g.,
Avicennia spp. and Rhizophora spp.) tolerate higher salinity than do non-mangroves, but
tolerance also varies among the true mangroves. For example, Rhizophora mucronata
seedlings do better in salinities of 30 g l-1, but R. apiculata do better at 15 g l-1
(Kathiresan and Thangam, 1990a; Kathiresan et al., 1996b). Sonneratia alba grows in
waters between 5 and 50 % seawater, but S. lanceolata only tolerates salinities up to 5%
seawater (Ball and Pidsley, 1995). Mangrove seedlings require low salinity (S.M. Smith et
al., 1996), but their salt tolerance increases as they grow (Bhosale, 1994).

     Short periods of high salinity may trigger events in the mangrove life history. For
instance, high salinity at the end of dry period, followed by an extended rainy period
controls establishment of Rhizophora seedlings (Rico-Gray and Palacios-Rios, 1996b).
Chronic high salinity, however, is always detrimental to the mangroves. Hypersalinity
stunts tree growth in A. marina stands (Selvam et al., 1991), reduces biomass in
hydroponically grown Bruguiera gymnorrhiza (Naidoo, 1990), and causes denaturing of
terminal buds in Rhizophora mangle seedlings (Koch and Snedaker, 1997). Saline
interstitial water reduces leaf area, increases leaf sap osmotic pressure, increases the leaf
area/weight ratio and decreases total N, K, and P (Medina et al., 1995). Simple salinity
fluctuations also have significant negative effects on photosynthesis and growth (Lin and
Sternberg, 1993). In Senegal, hypersalinity (from a decade of low rainfall and high
evaporation) has caused salt flats to grow into mangrove areas, completely destroying the
vegetation (Diop et al., 1997).
     Extremely high salt concentrations in the groundwater of tropical salt flats are
responsible for the complete absence of macrophytes (including mangroves). There are
often very sharp changes in groundwater salt concentrations at the interface between salt
flats and mangroves, suggesting that the mangroves are modifying the salinity of the
groundwater (Ridd and Sam, 1996). Mathematical models of groundwater flow show that
human activity hundreds of kilometers inland can destroy vast mangrove areas by changing
groundwater flow and modifying salinity levels (Tack and Polk, 1997).

6.8. Responses to metal pollution
    Because of their proximity to population centers and industrialized regions,
mangrove habitats have often received inputs of heavy metals and the sediments may show
significant metal contamination (Mackey et al., 1992; Larcerda et al., 1993; Rivail et al.,
1996; Lacerda, 1998; Tam and Yao, 1998). The mangroves themselves, however, generally
have low concentrations of heavy metals. Consequently, they are very poor indicators of
trace metal contamination. For example, in Sepetiba Bay, Rio de Janeiro, the sediments
contain 99% of the Mn and Cu and almost 100% of the Fe, Zn, Cr, Pb and Cd in the total
mangrove ecosystem. The tissues of Rhizophora mangle contain less than 1% of the total
of these metals (Silva et al., 1990). On the Saudi coast of the Arabian Gulf, there is no
correlation between the concentrations of metals in sediments and in the leaves of
mangroves living on the contaminated soil (Sadiq and Zaidi, 1994).
    The low level of metals in the mangroves themselves may be due to 1) low
bioavailability in the mangal sediments 2) exclusion of the metals by the mangroves or 3)
physiological adaptations that prevent metal accumulation in the plants. Mangrove roots
appear to be barriers that prevent metals from reaching the more sensitive parts of the plant
(Tam and Wong, 1997). Oxygen exuded by underground roots forms iron plaques that
adhere to the root surfaces and prevent trace metals from entering the root cells. Where the
metals do enter, there are apparent mechanisms to keep them from circulating freely
through the plant. Heavy metal concentrations in Rhizophora apiculata seedlings decrease
from root to stem and from stem to leaves (Moorthy and Kathiresan, 1998a).
    The chemical and physical environment of the mangal may efficiently trap trace
metals in non-bioavailable forms. For example, rapid precipitation of stable metal sulfides
under anoxic conditions decreases the bioavailability of trace metals in the mangrove
sediments (Di-Toro, 1990; Mackey and Mackay, 1996; Figure 16). All but the most mobile

                            elements (e.g., Mn and Zn) may be
                            in strongly bound fractions. Thus,
                            the mangal may help control trace
                            metal pollution in tropical coastal
                            areas (Lacerda, 1997).
                                Trace metals may also be
                            bound in organic complexes that
                            show low bioavailability (Clark et
                            al., 1998; Lacerda, 1998). For
                            example, Cr, which does not form
                            sulfide minerals, is immobilized in
                            refractory organic compounds in
                            mangrove sediments (Lacerda et
                            al., 1991). Mercury can be similarly
                            bound. However, it may be bound
                            as the highly toxic dimethyl-
                            mercury. Under oxic conditions,
                            dimethyl-mercury is volatile and
                            unstable; in reducing mangrove
                            sediments, however, it may persist
                            and accumulate (Quevauviller et
                            al., 1992).
                                 While mangrove sediments
                             generally have a high capacity for
 Figure 16. Summary of major metabolic processes
 involving metals in anoxic mangrove sediments (after  absorbing and holding trace metals,
                             heavy loads may exceed the
binding capacity of the sediment (Stigliani, 1995). Tam and Wong (1996a, b) irrigated
mangrove soil samples with metal-laden artificial wastewater. They found that the upper
centimeter of the soils bound Cu, Cd, Mn and Zn. However, there were also higher
concentrations of Mn, Zn and Cd in the water-soluble, exchangeable fraction of the treated
sediments than in the untreated, native sediments.
     Disturbances may also cause the mangrove soils to lose their metal-binding
capacity, resulting in mobilization of the metals. The mangal then shift from a heavy metal
sink to a heavy metal source (Lacerda, 1998). Disturbances may be in the form of
prolonged dry periods (Clark et al., 1997), changes in the frequency and duration of tidal
flooding (Chiu and Chou, 1991) or changes in salinity (Spratt and Hodson, 1994). Often,
these disruptions are associated with human activities (Lacerda, 1998).
     S. Zheng et al. (1997) suggest that mangrove afforestation projects should not be
done on Cu or Zn-polluted soils since seedlings secrete organic acids that may increase
solubility of the metals. Rhizophora apiculata seedlings planted in an area formerly used
for tin mining showed high mortality (approximately 47% in the first four years). The
mortality, however, was attributed to altered microtopography and soil particle distribution
rather than metal contamination (Komiyama et al., 1996).
     Metals in mangrove sediments do not appear to strongly affect bacterial
populations, even under heavy loads (Tam, 1998). However, if the metals are bioavailable,
they may accumulate in the macroinvertebrate fauna. In Yingluo Bay, He et al. (1996)

found high Zn and Cd levels in mollusks living in the mangal. Crustaceans had elevated Cu
levels and sipunculids concentrated Pb. Meyer et al. (1998) similarly found that mangrove
oysters (Crassostrea rhizophorae) in a northeastern Brazil mangal accumulated mercury
and are good biomarkers for mercury contamination. Bioaccumulation of such substances
can carry substantial human health risks. Mangrove sediment chemistry and the fate of
heavy metals are subjects that merits much more study.

6.9. Responses to organic pollution
    Three characteristics have long made mangrove habitats favored sites for sewage
dumping: (1) flow through the habitat disperses wastes from a point source over vast areas,
(2) the vegetation itself filters nutrients from the water, and (3) the mangrove soil, algae,
microbes, and physical processes absorb large amounts of the pollutants (Wong et al.,
1995, 1997b).
    Nutrients (primarily nitrogen and phosphorus) are often major components of the
pollution. Researchers have studied the ability of mangals to absorb nutrients and the
effects of the pollutants on the mangal community as a whole. In general, mangrove soils
efficiently trap wastewater-borne phosphorus, but are less effective at removing nitrogen
(Tam and Wong, 1995). Tam and Wong (1996a, b) experimentally tested the ability of
mangrove soils to absorb nutrients when treated with synthetic wastewater. The soils
retained both nitrogen and phosphorus. The bulk of these were trapped in the upper 1 cm
of the sediment where they could be processed by bacterial communities (Corredor and
Morell, 1994).
    Wong et al. (1995, 1997a) found that two full years of sewage discharge did not
adversely affect mangrove growth in the Funtian mangal of China. Nor did sewage affect
biomass, density, or community structure of the benthic macrofauna (Yu et al., 1997).
Furthermore, wastewater input did not seem to increase litter production or litter decay
rates (N.F.Y. Tam et al., 1998).
    While these studies suggest that mangroves are tolerant of organic pollution, results
should be viewed cautiously since they may not hold in other habitat. The effects of
sewage dumping will depend on the quantities of sewage, the duration of dumping, and the
unique characteristics of each mangal. Particularly important are the patterns of water flow
through the habitat since this will determine flushing rates and residence times of the
pollutants (Ridd et al., 1990; Uncles et al., 1990; Wolanski et al., 1990; Wattayakorn et al.,
    High levels of organic pollution can contribute to disease, death, and changes in
species compositions within the mangal (Tattar et al., 1994). Mandura (1997) found that
sewage discharge killed pneumatophores of Avicennia marina in the Red Sea. The loss of
the pneumatophores decreased surface area for respiration and nutrient uptake and retarded
the growth of the trees. The pollution can also have cascading effects on invertebrate
populations (e.g., Sanches and Camargo, 1995).
    Beyond simple nutrients, organic pollution in mangrove environments may include
other anthropogenic chemicals and debris. Mangrove sediments in Cienaga Grande de
Santa Marta and Chengue Bay (Colombian Caribbean) contain significant organochlorine
pesticide residues. The concentrations of some of these vary seasonally (Espinosa et al.,
1995). Large amounts of plastic and non-mangrove wood are present in the mangroves of

Jamaica. The volume of these solid wastes correlates strongly with total rainfall in a nearby
metropolitan area (Green and Webber, 1996).

6.10. Responses to oil pollution
     Oil pollution from oil or gas exploration, petroleum production and accidental spills
severely damages mangrove ecosystems (Mastaller, 1996). Clean-up operations after such
calamities are costly and difficult (IUCN, 1993). Oiling of mangroves has a number of
significant consequences. One of the most immediate and obvious is defoliation of the
trees. The toxicity of the oil may depend on environmental conditions; oil has the greatest
effect on survival and growth of Rhizophora mangle when the trees are in hot, bright
outdoor conditions (Proffitt et al., 1995). Toxicity may also differ among mangrove
species. For example, along the coast of Sao Paulo, Brazil, an oil spill caused 25.9%
defoliation of Rhizophora mangle, 43.4% defoliation of Laguncularia racemosa, and
64.5% defoliation of Avicennia schaueriana (Lamparelli et al., 1997). Differential
mortality of the trees can potentially lead to long-term changes in the community structure.
     Oil in a mangrove habitat (whether from a spill or chronic input) can have other
less obvious effects on the mangroves. For example, sediments can have significant
hydrocarbon pollution long after a spill event, even when there is no evidence of petroleum
contamination on the trees or in water samples from surrounding water (Bernard et al.,
1995, 1996). Munoz et al. (1997) followed the breakdown of Arabian light crude oil in
mangrove peat for 8 full years. Sediments contaminated in the Galeta spill in Panama
continued to hold oil residue, including the full range of aromatic hydrocarbons, 5 years
after the spill (Burns et al., 1994). The authors suggest that it will take at least 20 years for
the toxicity to completely disappear.
     Grant et al. (1993) demonstrated that sediment oil can inhibit establishment and
decrease survival of mangrove seedlings for several years. This residual toxicity may
interfere with mangrove afforestation efforts (S. Zheng et al., 1997). Dutrieux et al. (1990)
planted Sonneratia caseolaris in soils that had been treated with oil. Many of the plants
were killed; the survivors were significantly stunted. The retained oil can also cause
mutation. Klekowski et al. (1994b, c) found a positive correlation between concentrations
of polycyclic aromatic hydrocarbons in mangal sediments, and the frequency of
Rhizophora mangle carrying chlorophyll-deficient mutations.
     The extent of mangrove damage from oil pollution will depend on the kind of oil,
and the magnitude and frequency of spilling. For example, fresh oil causes more leaf loss
in Avicennia seedlings than does aged oil (Martin et al., 1990; Grant et al., 1993). Boeer
(1996b) measured effects of a mineral oil spill in the Arabian Sea off Fujairah. The
mangroves were relatively unaffected and all signs of the spill were nearly gone only 7
months after the spill.
     Proffitt and Devlin (1998) tested the effects of sequential oilings on potted
Rhizophora mangle seedlings, first treating the seedlings with No. 6 fuel oil, followed, 34
months later, by crude oil. They found no evidence of cumulative or synergistic effects, but
this conclusion has been challenged because of unnatural laboratory conditions and low
statistical power (Ellison, 1999). Given the sensitivity of mangroves to soil conditions, it is
essential to study oil effects under conditions that reflect the natural environment as closely
as possible. For example, salinity should be held at field levels and realistic oil

concentrations should be used to model chronic exposure of plants in oil-contaminated soil
(Ellison, 1999).
    The most realistic measure of repeated oil exposure comes from field habitats
exposed to natural oiliness. Two large oil spills (the first in 1968 and the second in 1986)
have caused large-scale damage to mangrove forests in Panama. In addition to killing trees
outright, the oil retained in the sediments caused apparent sublethal effects (Duke et al.,
1997). The residual effects of oiling may make the mangroves more vulnerable to future
damage. More careful, long-term laboratory experiments under natural conditions are
necessary to understand the responses of mangroves to oil and the consequences of oiling.
    Oil contamination can damage animals living in the mangal, both in the sediments
and on submerged mangrove roots (e.g., Mackey and Hodgkinson, 1996). Five years after
the Galeta oil spill in Panama, there was a 60% decrease in the number of isopods on
submerged Rhizophora mangle prop roots and a 40-50% drop in the number of spiny
lobsters (Levings and Garrity, 1994; Levings et al., 1994). Oyster populations dropped
65% along mangrove channels and 99% in mangrove streams. The population decreases
are due, in part, to loss of root surface on which to attach (the surface area of submerged
roots decreased 38% in the channels and 74% in streams; Garrity et al., 1994).
    In addition to killing the mangrove fauna directly, oil can have indirect effects
resulting from habitat modification. Oil released during the 1991 Gulf War left a black tar
layer in the mangals along the Saudi Gulf. The tar layer created higher than normal
temperatures in the soil. The ecological consequences of the higher temperatures, and the
effect on epifauna and infauna, are not yet fully known (Boeer, 1996a).
    The general response of a mangrove forest to oiling can be divided into four
phases: 1) immediate effects, 2) structural damage, 3) stabilization and 4) recovery. The
third and fourth phases may take many years to occur, if they occur at all. In Brazil, a
mangrove area damaged by oil did not began to recover until approximately 10 years after
the event (Lamparelli et al., 1997). Assessing the effects of oil on mangrove environments
will require the development of creative methods for measuring impacts and accurate
modeling of the physical and chemical events associated with the spill (e.g., Jacobi and
Schaeffer Novelli, 1990; Lamparelli et al., 1997). These efforts, however, will only be
effective if they are supported by careful monitoring and long-term data sets.

6.11. Responses to pests
    A few of the many plants and animals that make their homes in the mangal are
serious pests that damage the mangroves, decreasing growth and productivity and, in
extreme cases, killing the trees. Some of the harmful species do not directly injure the
mangroves. Instead, they cause damage by competing for scarce resources. Allelopathic
interactions among mangrove species suggest that interspecific competition is a normal
process in the mangal. Toxic leachates from leaf litter of some mangroves (e.g. Lumnitzera
racemosa, Ceriops decandra and R. apiculata) inhibit the growth of roots and shoots of
Rhizophora apiculata and R. mucronata seedlings (Kathiresan and Thangam, 1989;
Kathiresan et al., 1993).
    In general, stressful osmotic conditions that lead to lignification and suberization
prevent the development of a luxuriant herbaceous undergrowth in mangrove forests so
there is not normally strong competition between mangrove and non-mangrove plants
(Schwamborn and Saint-Paul, 1996). However, damage to established stands can open

windows of opportunity for invasive species that may restructure the community (Kangas
and Lugo, 1990; Lugo, 1998). The mangrove fern Acrostichum, for example, is a weedy
pest that causes significant losses to mangrove forestry (Chan, 1996). The pest is currently
controlled by application of herbicides, but efforts to control it are being refocused on its
responses to shading and salinity (Medina et al., 1990).
     Mangroves themselves can become pests when they are introduced to new habitats.
At least 6 mangrove species have been introduced to the Hawaiian Islands since the early
1900’s. Rhizophora mangle has been a particularly successful transplant, but two other
species (Bruguiera gymnorrhiza and Conocarpus erectus) also have self-sustaining
populations. The mangroves were planted to help stabilize sediments in coastal mud flats.
As invaders, however, the mangroves have had negative effects. In particular, they
compete with native plants and modify habitats that are important to Hawaiian birds
(including endangered species). They also cause drainage problems in some areas (Allen,
     Other pest organisms damage the mangroves, not by competing with them, but
simply by living on their surfaces. For example, the spiders Tetragantha nitens and
Chiracanthium live on Rhizophora. They lay their eggs on the leaves, which induces leaf
rolling, chlorosis and wilting. Heavy infestations can kill the trees (Irianto and Suharti,
1994). The semi-parasitic mistletoe, Phthirusa maritima, has a more direct effect on the
trees. Infections in Conocarpus erectus and Coccoloba uvifera induce higher transpiration
rates, lower CO2 assimilation rates, and lower water-use efficiency (Orozco et al., 1990).
     By far the most extensive and serious damage to mangroves occurs through the
feeding activities of herbivorous animals. While most of the damage is done by animals
feeding in the canopy, several kinds of crustaceans and mollusks bore directly into
submerged mangrove wood and do significant damage. Spaeromatids are generally the
most common wood-borers (e.g., Sivakumar, 1992; Huang et al., 1996). Infestations of
these isopods are heavier in dead mangrove stumps than in live wood but the stumps and
woody debris provide a perennial source of larvae that also attack the living wood
(Sivakumar and Kathiresan, 1996). Distributions of these pests are controlled largely by
currents and tidal regimes.
     Of the animals feeding on the mangrove canopy, insects are undoubtedly the most
destructive. Murphy (1990d) described 102 insect herbivores that attack 21 mangrove
species in Singapore. Veenakumari et al. (1997) listed 197 species of herbivores on the
Andaman and Nicobar Islands. Some of the insect herbivores are serious crop pests that
simply use mangroves as alternative hosts. Others have apparent preferences for
mangroves (e.g., Mictis on Sonneratia, Glaucias on Lumnitzera, Calliphara on
Excoecaria, and Antestiopsis on Avicennia; Murphy, 1990d).
     Insect herbivores can completely defoliate mangrove stands. Rhizophora leaves that
have been attacked by scale insects (Aspidiotus destructor) first turn yellow at the site of
feeding, then brown and necrotic. In extreme cases, the leaves dry up, drop off, and the
entire seedling dies (Kathiresan, 1993). Periodic outbreak populations of the moth Achaea
serva defoliate large stands of Excoecaria agallocha (McKillup and McKillup, 1997). In
Singapore, feeding by Paralebeda and Selepa caterpillars can lead to total loss of shoots in
Excoecaria; Trabala krishna has the same effect on Sonneratia. Apical bud destruction
may reduce leaf production and change the architecture of the plant (Murphy, 1990d).

     Summer feeding by the caterpillar Nephpterix syntaractis in Hong Kong
completely defoliates Avicennia marina, severely reducing the reproductive output of the
trees (Anderson and Lee, 1995). Kandelia candel in the same region may experience a
35% defoliation (Lee, 1991). In Belize, Central America, an outbreak of the lepidopteran
Phocides pigmalion on Rhizophora mangle increases leaf abscission rates and reduces
above-ground net primary production by 5-20%. The lost production normally would have
been exported to surrounding marine environments (Ellison and Farnsworth, 1996b). The
insect defoliator, Pteroma plagiophleps (Lepidoptera: Psychidae), has been newly recorded
on the Indian west coast (Santhakumaran et al., 1995).
     Insect herbivores may show preferences among mangrove hosts. In an Ecuadorian
mangal, the bagworm, Oiketicus kirbyi removed 80% of the foliage of Avicennia
germinans, 10% of the Conocarpus erectus and < 5% of the Laguncularia racemosa (Gara
et al., 1990). The susceptibility of mangrove species, and individual mangrove plants, may
relate to their physico-chemical characteristics. High leaf toughness, measured as the ratio
of protein to fibre, reduces palatability and digestibility (Choong et al., 1992). Tannins also
deter herbivores. Avicennia species, which have low tannin levels, suffer more herbivore
damage than do Rhizophora species, which have more tannins (Kathiresan, 1992).
     Feeding preferences of the insects may also be influenced by the health of the
mangrove. Nutrient enriched trees tend to suffer higher herbivory. Herbivory by
Ecdytolopha (an endophytic insect that feeds in apical buds) and Marmara (which mines
stems) on Rhizophora mangle increased significantly when the trees were treated with P
and NPK. Fertilization with N alone did not increase herbivory (Feller, 1995). Damage
from feeding herbivores may also invite further attack. Farnsworth and Ellison (1993)
made small holes in the leaves of R. mangle and found that the artificial damage increased
natural damage from herbivorous insects; in 50 days, the size of the holes had increased
     Some herbivores feed specifically on the reproductive tissues and seeds of
mangroves. Crabs are particularly important seed predators (Osborne and Smith, 1990;
Robertson, 1991; McGuinness, 1997b; Dahdouh-Guebas et al., 1998). However, insects
can also attack mangrove seeds. Insect borers appear to impair the growth of Avicennia
marina propagules, but do not kill them (Robertson et al., 1990). A mite (Afrocypholaelaps
africana) feeds on mangrove pollen. Unopened flower buds are mite-free, but newly
opened flowers are infested by all post-embryonic stages of the mite. Egg-bearing female
mites are dispersed among the mangroves by the honeybee Apis mellifer. The mite
population declines as the mangrove flowering season ends (Seeman and Walter, 1995). It
is not clear what affect the mites have on the mangrove population.

6.12. Responses to anthropogenic stress
    In recent years, anthropogenic pressures have significantly damaged the world’s
mangroves, with alarming levels of habitat loss. For example, Ramirez-Garcia et al. (1998)
estimate a 32% decrease in mangroves in the Santiago River of Mexico in the past 23
years. Aksornkoae (1993) and Raine (1994) report more than a 50% reduction in the
mangrove forests of Thailand. Mndeme (1995) reports that the mangrove resources in the
Mafia District of Tanzania are in danger of collapse. In the Florida Keys, USA, Strong and
Bancroft (1994) report that 15% of the original mangrove forests have been cleared for
development; mean forest size has decreased 41%. Approximately 45% of the mangroves

in Indonesia have been heavily impacted by human activities (Choong et al., 1990). Some
estimates put global mangrove loss rates at one million ha y-1 (Mohamed, 1996). Such
levels of destruction and habitat fragmentation raise concerns about conservation of
biodiversity in the mangrove habitats and preservation of the mangals themselves.
     Ellison and Farnsworth (1996a) classified anthropogenic disturbances into four
types: extraction, pollution, reclamation, and changing climate. These disturbances are
listed in the order of their increasing spatial scale, their increasing temporal scales, and the
increasing time required for recovery. Research suggests that even relatively low impact
human activities can affect the mangrove environment. For instance, boardwalks placed in
the mangals around Sydney, Australia to provide access for educational and recreational
activities have modified sediment composition and changed benthic invertebrate
community structure (Kelaher et al., 1998a, 1998b). It may require fairly long periods for
the mangal to recover from even minor disturbances (Snedaker et al., 1992).
     Diversion of freshwater for irrigation and land reclamation has historically been a
major cause of wide-scale mangrove destruction (Conde and Alarcón, 1993, Twilley et al.,
1998). Throughout the world, mangroves and mangrove products have also been used for
timber, fuel, food, clothing, perfume, dyes, tannins, and medicine (Rasolofo, 1997;
reviewed by Bandaranayake, 1998). In the past several decades, extensive tracts of
mangrove have been converted for aquaculture. Shrimp ponds have become particularly
common in many former mangals (Twilley et al., 1993; Primavera 1995; de Graaf and
Xuan, 1998). Menasveta (1997) reports that nearly 55% of the mangroves in Thailand were
converted to shrimp ponds between 1961 and 1993. Pond culture now surpasses open
ocean fishing as the major source of shrimp there. Unfortunately, ponds in many regions
are unsustainable and up to 70% of them may be left idle after some period of production
(Stevenson, 1997). Because of changes in the sediments caused by pond construction, the
abandoned sites are difficult to revegetate with mangroves even after the shrimp farming
has ceased (de Graaf and Xuan, 1998).
     Intact mangals process heavy organic loads and could help oxidatively process
nutrients in shrimp pond effluents (Eguchi et al., 1997; Twilley et al., 1998). Robertson
and Phillips (1995) estimated that 2 to 22 hectares of mangrove forest could completely
filter excess nitrogen and phosphorus from a one-hectare shrimp pond. The effluents, in
turn, could promote growth of the mangroves. A 70% dilution of effluent from a semi-
intensive shrimp culturing pond in south India significantly increased growth of mangrove
seedlings (Rajendran and Kathiresan, 1996).
     In the Mekong Delta of Vietnam, living mangroves actually increase productivity
of shrimp aquaculture facilities. Binh et al. (1997) collected data suggesting that yields are
greater in shrimp ponds with 30-50% mangrove coverage. Farmers who integrate shrimp
and mangrove farming may, therefore, realize better economic returns (Hong and San,
1993). P.T. Smith (1996) found that sediments in the shrimp ponds are very similar to
those from nearby mangrove habitats, again suggesting that mangrove and shrimp
aquaculture should be compatible.
     Heavy historical exploitation of mangroves has left many habitats severely
damaged. The damage has consequences beyond loss of the trees themselves. For
example, because mangroves serve as nursery habitats for many crustaceans and fish,
damage can have a direct effect on fishery resources and the lives of those who depend on
them (John and Lawson, 1990; Twilley et al., 1991, 1998; Ruitenbeek, 1994; Fouda and

Al-Muharrami, 1995; Primavera, 1998). Recently, community-based approaches to
conservation and resource management have been launched with the participation of local
people (A.H. Smith and Berkes, 1993; Kairo, 1995; Semesi, 1998). Guidelines for
evaluation, restoration, and management of mangrove ecosystems are also being developed
(Field, 1996; Siddiqi and Khan, 1996; Ewel et al., 1998b; Gilbert and Janssen, 1998; Kaly
and Jones, 1998; Twilley et al., 1998).
    Efforts are being made to rebuild damaged mangrove ecosystems in many parts of
the world (Semesi, 1992; Chowdhury and Ahmed, 1994; Field, 1998). The programs are
called regeneration, reclamation, rehabilitation, or ecodevelopment. Finding adequate
supplies of viviparous seedlings for use in such afforestation projects is a challenge and
more effective methods are needed. Living seedlings can be cut and the cuttings induced to
produce roots and shoots. However, success of the cuttings depends on how they are done
(growth and survival depend on where the location and length of the cutting; Ohnishi and
Komiyama, 1998). In vitro micropropagation methods have been recently developed for
Excoecaria agallocha (C.S. Rao et al., 1998). These techniques hold promise for
mangrove regeneration.
    In South Sulawesi, Indonesia, where mangrove removal has produced significant
environmental problems, efforts are underway to launch mangrove agroforestry projects.
Planting of Rhizophora mucronata along the coast is mitigating coastal erosion and
preventing flooding (which otherwise damages aquaculture facilities). Controlled
harvesting of the mangroves produces income as the product is sold for fuel wood
(Weinstock, 1994). Efforts at mangrove agriculture are also underway in the Federated
States of Micronesia (Devoe and Cole, 1998). However, there is still much to learn about
proper management and sustainable harvesting of mangrove forests. Despite nearly 100
years of careful management, timber yields from the Matang Mangrove Forest Reserve in
Malaysia are declining significantly (Gong and Ong, 1995).

6.13. Responses to Global changes
     It is expected that increasing concentrations of atmospheric CO2 and other
"greenhouse gases" will bring changes in the global climate. It has been predicted that each
decade could bring a 0.3° rise in air temperature and a 6 cm rise in the global sea level
(Titus and Narayanan, 1996; Wilkinson, 1996; Gregory and Oerlemans, 1998). Because of
their location at the interface between land and sea, mangroves are likely to be one of the
first ecosystems to be affected by global changes. Most mangrove habitats will experience
increasing temperature, changing hydrologic regimes (e.g., changes in rainfall,
evapotranspiration, runoff and salinity), rising sea level and increasing tropical storm
magnitude and frequency (R.W. Stewart et al., 1990, Field, 1995; Michener et al., 1997).
Davis et al. (1994) have developed a framework for assessing risks to mangrove
ecosystems in the context of a changing global climate but the seriousness of the effects
will be strongly site-specific (Kjerfve and Macintosh, 1997).
     Small increases in air temperature may have little direct effect on the mangroves
(Field, 1995), but if temperatures exceed 35° C, root structures, seedling establishment and
photosynthesis will all be negatively affected. The broader effects of temperature increases
may be in modifying larger-scale distribution and community structure, increasing species
diversity in higher latitude mangals and promoting spread of mangroves into sub-tropical
saltmarsh environments (Ellison, 1994).

     Because they are so specialized, and may live so close to their tolerance limits,
mangroves are particularly sensitive to minor variation in hydrological or tidal regimes
(Blasco et al., 1996). Reduced rainfall and runoff would produce higher salinity and
greater seawater-sulfate concentrations. Both would decrease mangrove production
(Snedaker, 1995). The most important effects, however, would come from rising sea
levels, but responses will vary among locations and will depend on the local rate of the rise
and the availability of sediment to support reestablishment of the mangroves (Pernetta
1993; Parkinson et al., 1994; Semeniuk, 1994; Woodroffe, 1995, 1999). For example, in
the Caribbean, mangrove seedlings are very sensitive to low sediment availability,
suggesting that mangroves will not survive on Caribbean coral islands if sea levels increase
as predicted (Ellison, 1996).
     Ellison and Farnsworth (1997) studied the response of Rhizophora mangle to
increased inundation, mimicking the sea-level changes expected in the Caribbean in the
next 50-100 years. After 2.5 years of higher water, plants would have significantly lower
rates of photosynthesis and growth, be shorter and narrower, have fewer branches and
leaves, and more acid-sulfide in their soils. The authors suggested that increased mangrove
growth rates predicted for increasing atmospheric CO2 may be offset by decreased growth
resulting from changes in tidal regimes.
     Sayed (1995) tested the effects of higher water levels on Avicennia marina by
flooding potted seedlings. The treatment resulted in stomatal closure, loss of chlorophyll
fluorescence, and a slight reduction of leaf water potential. Post-flooding recovery,
however, was rapid, suggesting that sea level rises could lead to colonization of supratidal
flats by this species (Sayed, 1995). As sea level rises, mangroves, in general, would tend to
shift landward. Human encroachment at the landward boundary, however, may make this
impossible. Consequently, the width of mangrove systems would be likely to decrease as
the sea-level rose (Kjerfve and Macintosh, 1997).
     The mangrove-associated fauna would be affected both directly by climatic
changes and indirectly by changes in the mangroves. Species that are tolerant of increasing
temperatures (e.g., fish, gastropods, mangrove crabs and other crustaceans) may adjust
rapidly to the changes. In contrast, soft-bodied animals and bivalve mollusks would be
very sensitive to higher temperatures. Desiccation that would accompany increasing
temperatures would harm many marine species associated with mangroves (Kjerfve and
Macintosh, 1997). For mangrove-dependent species, however, the most serious
consequences of a changing climate would likely be the loss of habitat as the global
mangrove forests declined.


7.1. Litter decomposition and nutrient enrichment
    Mangrove ecosystems produce large amounts of litter in the form of falling leaves,
branches and other debris. Decomposition of the litter contributes to the production of
dissolved organic matter (DOM) and the recycling of nutrients both in the mangal and in
adjacent habitats. The organic detritus and nutrients could potentially enrich the coastal sea
and, ultimately, support fishery resources. The contribution of the mangroves could be
particularly important in clear tropical waters where nutrient concentrations are normally

     The nutrient cycling begins when leaves fall from the mangroves and are subjected
to a combination of leaching and microbial degradation (Lee et al., 1990b; Chale, 1993).
Leaching alone removes a number of substances and can produce high levels of DOM
(Benner et al., 1990b). Potassium is the most thoroughly leached element with up to 95%
of the total potassium being removed in a very short time (Steinke et al., 1993b).
Carbohydrates also leach quickly during early decomposition. Tannins, in contrast, leach
very slowly and the high tannin contents may slow establishment of bacterial populations
in the initial period of decomposition. As the tannins are eventually leached, the bacterial
populations rapidly increase (Steinke et al., 1990; Rajendran, 1997; Rajendran and
Kathiresan, 1999b).
     Bacteria and fungi contribute to decomposition of the mangrove material and to the
transformation and cycling of nutrients. Fungi are the primary litter invaders, reaching their
peak in the early phases of decomposition (Rajendran, 1997). The phylloplane fungi do not
attack live leaves; they begin to break the leaf material down only after it has been
submerged. There are two major phases of fungal decomposition. Cellulase-producing
fungi first attack the leaves between 0 and 21 days after submergence; xylanase producers
are active between 28 and 60 days. Pectinase, amylase and protease producers are present
throughout decomposition (Singh and Steinke, 1992; Raghukumar et al., 1994a).
     Bacterial colonies appear shortly after the litter has been colonized by fungi. The
bacteria grow quickly and can reach very high densities. Zhuang and Lin (1993) measured
bacterial densities from 2 x 105 to 10 x 105 • g-1 on Kandelia candel leaves that had
decomposed for 2-4 weeks. This was about 100 times higher than densities of
actinomycetes and filamentous fungi. The N2-fixing azotobacters are one of the important
groups in the decomposing litter (Rajendran, 1997) and their activities may increase the
nitrogen content of the leaves 2 - 3 times (Wafar et al., 1997; Rajendran, 1997).
     Chale (1993) measured a similar rapid nitrogen increase in leaves after six weeks of
decomposition and suggested that the litter 1) provides a surface for microbial nitrogen
synthesis and 2) acts as a nitrogen reservoir. The C:N ratio of decomposing Avicennia
marina leaves drops dramatically from approximately 1432 to 28, due primarily to a large
increase in their nitrogen content (Mann and Steinke, 1992; Singh and Steinke, 1992). In
another study, N.F.Y. Tam et al. (1990) saw the C:N ratio in decomposing leaves increase
for one week, then decrease, and finally stabilize at approximately 74. They hypothesized
that the initial increase resulted from the conversion of particulate and soluble nitrogen in
the litter to proteins in bacteria and fungi.
     A number of factors can affect the rate of litter decomposition and, therefore, the
rates of nutrient cycling. For example, litter decomposition rates vary among mangrove
species. Avicennia leaves, because they are thinner and have fewer tannins, decompose
faster than those of other species (Sivakumar and Kathiresan, 1990; Steinke et al., 1990;
Kristensen et al., 1995). Avicennia leaves also sink and begin to decompose immediately
whereas the leaves of other species (e.g., Sonneratia and Rhizophora) may float for several
days (Wafar et al., 1997). Lu and Lin (1990) found that litter of Bruguiera sexangula
decomposes quickly. Aegiceras corniculatum, in contrast, decomposes slowly (Tam et al.,
     Decomposition is influenced by tidal height, rainfall and temperature. In
subtropical mangrove forests, mangrove debris decomposes substantially faster in the rainy
season (e.g., Woitchik et al., 1997). Mackey and Smail (1996) studied decomposition of

Avicennia marina. They found significantly faster decomposition in lower intertidal zones
with greater inundation. They also found an exponential relationship between leaf
decomposition rate and latitude with leaves decomposing most quickly at low latitudes.
They attributed the pattern to temperature differences, and concluded that seasonality can
have important effects on organic cycling and nutrient export from mangrove systems.
     Breakdown and decomposition of mangrove litter is accelerated by the feeding
activities of invertebrates (Camilleri, 1992). The animals may process large volumes of
the litter, contributing significantly to nutrient dynamics. Litter turnover rates have been
estimated by measuring rates of leaf decomposition. However, estimates made this way are
generally 10-20 times lower than rates calculated from actual measurements of leaf fall and
litter standing crop. The difference in the estimates can be attributed to 1) tidal export and
2) the feeding activities of crabs. The
crab feeding may be the more
important of these in many regions.              A. marina
For example, in the Ao Nam Bor
                        Juvenile peneid shrimp • haul
mangrove forest in Thailand, crabs
process about 80% of the litter         4
deposited in the mid-intertidal zone
and nearly 100% of the leaves
deposited in the high intertidal         2

(Poovachiranon and Tantichodok,         1
1991). In field experiments, Twilley
et al. (1997) found that mangrove        0

crabs process the mangrove material           10    20   30    40    50 60   70

very quickly. They removed a full                   Days of decomposition

day’s accumulation of mangrove leaf       Figure 17. Number of penaeid shrimp associated with
litter in only 1 hr. Because the mangrove    decomposing leaves of Avicennia marina (in situ litterbag
                         experiment by Rajendran, 1997). Populations increase
material is quite refractory, it may need
                         dramatically, but only after several weeks of
to decompose for some time before it is     decomposition.
useful to other invertebrates. Wafar et al.
(1997) estimated that litter needs to
decompose for about two months before it can be used in most detritivores’ diets. In situ
observations verify that mangrove leaves attract shrimp, crabs, and fish (particularly
juveniles), but only after several weeks of decomposition (e.g., Rajendran, 1997;
Rajendran and Kathiresan, 1999a; Figure 17).

7.2. Food webs and energy fluxes

     Mangals contribute to complex food webs and important energy transfers.
However, it is not clear how, or whether, these processes affect the larger ecosystem.
While the living vegetation is a valuable food resource for insects, crustaceans, and some
vertebrates, most of the mangrove production is transferred to other trophic levels through
litterfall and detrital pathways (Figure 18). Mangrove forests produce organic carbon well
in excess of the ecosystem requirements. Duarte and Cebrian (1996) estimate that the
excess photosynthetic carbon approaches 40% of net primary production. While some of
this organic matter simply accumulates in the sediments, large amounts could potentially

                        be transported offshore (Alongi, 1990b;
                        Robertson et al., 1991, 1992; Lee, 1995). The
                        amount of material exported, however,
                        depends strongly on local conditions and
                        varies enormously among mangals (Twilley et
                        al., 1992).
                             Material exported from the mangroves
                        could potentially support offshore
                        communities (Marshall, 1994; Robertson and
                        Alongi, 1995; Van Tussenbroek, 1995), but
                        the connections between mangal and adjacent
                        habitats are complex, dynamic, and have been
                        difficult to demonstrate unequivocally (Alongi
                        et al., 1992; Twilley et al., 1992; Hemminga et
                        al., 1995; Alongi, 1998). For instance,
                         Jennerjahn and Ittekkot (1997) found that
                         organic matter in continental sediments in
 Figure 18. A stylised food web in a mangrove
                         eastern Brazil was very different from that in
 ecosystem. The food web may be highly
 localized without strong connections to other mangrove environments and concluded the
 habitats. The foundations of the web are    mangrove matter is largely retained and
 detritus, microbes, algae and seagrasses.
                         decomposed within the mangal itself. Studies
with stable isotopes also suggest that mangroves do not make major contribution to coastal
food webs (Primavera, 1996; Loneragan et al., 1997). In fact, the data suggest that carbon
may instead be flowing from oceanic systems into the mangrove habitat (Figure 19).
Oceanic carbon contributed up to 86% of the particulate organic carbon (POC) in water
samples from a Brazilian mangal (Rezende et al., 1990).
     It appears that mangroves, in general, make only a localized contribution to the
food web (Fleming et al., 1990; Mohammed and Johnstone, 1995). Sediment meiofauna,
for example, feed directly on mangrove detritus. The composition of the meiofaunal
community changes during the process of litter decay, suggesting that the community is
responding to chemical changes in the leaves (Gee and Somerfield, 1997). The meiofaunal
community, though large in some habitats, may largely be a trophic dead end that
contributes little to the larger food web (Schrijvers et al., 1998).
     The mangroves may have stronger trophic linkages with epibenthic invertebrates
and fish living in the mangal and in nearby habitats (e.g., seagrass beds). For example,
mangrove detritus contributes to the nutrition of juvenile Penaeus merguiensis living in
tidal creeks. The juveniles feed directly on mangrove detritus, on other small detritivorous
invertebrates, and on benthic microalgae growing in the mangal (Newell et al., 1995).
Shrimp in mangrove estuaries may also feed heavily on seagrass epiphytes (Loneragan et
al., 1997). Invertebrates may also feed on the variety of cyanobacteria and microalgae that
live on submerged portions of the mangroves and on leaf litter (e.g., Sheridan, 1991;
Farnsworth and Ellison, 1995; Pedroche et al., 1995).
     Pinto and Punchihewa (1996) found that syngnathid fish (pipefish) in the Negombo
Estuary of Sri Lanka fed primarily on mangrove litter. However, mangroves apparently
contribute little of the carbon assimilated by other fishes. This is true despite the movement

of a number of fish species between
mangrove habitats and nearby seagrass                              A
beds (Marguillier et al., 1997).                                S
                             -16      S
    Mangrove detritus is probably
more important as a substrate for                   SS
                             -18      S      S
                                   SS            S
microbial activity and nutrient                    S      A
regeneration than it is as a direct food                      S
                                    S      SS
source for detritivores. Wafar et al.                       SS
                        13                  S

(1997) analyzed energy and nutrient      (o/oo)         PP
                             -22            S
fluxes between mangroves and                            A
estuarine waters and concluded that                  P
                             -24                  M
mangroves contribute significantly to                             'D

the estuarine carbon budget. However,          -26            M
they contribute little to nitrogen and                 M
                                    M      M      P
                                   MM            M
phosphorus budgets. It is not clear                  M
                             -28           MM
whether any of these substances are                  M
exported from the mangal in sufficient                M

quantities to make significant
                               Sibunag River, Sementa Besar Laguna Joyuda,
contributions to energy flow and the              Philippines & Buloh River, Puerto Rico
ecology of the broader ecosystem
(Alongi et al., 1992; Alongi, 1998).       Figure 19. Ratios of stable carbon isotopes in
Mangrove sediments efficiently uptake, retain shrimp collected from mangrove habitats in the
and recycle nitrogen (Rivera-Monroy et al.,    Philippines (Primavera, 1996), Malaysia (Rodelli et
                         al., 1984), and Puerto Rico (Stoner and
1995). Resident bacteria and benthic algae
                         Zimmerman, 1988). Shrimp tissue δ13 values (S)
rapidly assimilate available ammonium and
                         are much closer to the δ13 val ues of plankton (P)
prevent its export (Kristensen et al., 1995;
                         and algae (A) than they are to those of mangrove
Middelburg et al., 1996). The mangrove      leaves (M) or detritus (D). This suggests that the
environment may, therefore, represent a      shrimp are deriving their carbon primarily from
nutrient and carbon sink rather than a source   algae and the plankton; the mangrove detrital
                         pathway contributes little to their nutrition (after
for adjacent habitats. Careful measurements
                         Primavera, 1996).
and creative experimentation will be
necessary to clarify the role these habitats
play in larger-scale food webs and energy fluxes.

    Mangrove ecosystems are receiving increasing attention, but we still lack much
basic information about their structure and function. There are still fundamental gaps in our
knowledge of the reproductive biology of mangroves, and mangrove evolution is poorly
understood. We are still far from understanding energy flow and food web dynamics in
mangrove environments and how the mangroves connect with other ecosystems. There is a
great need to better understand the effects of environmental change and pollution on
mangrove flora and fauna. Animals that are highly dependent on mangroves need
additional study, particularly with respect to larval supply and recruitment. Such
ecobiological research can be linked to management of mangroves and associated fishery
resources (e.g., Bacon and Alleng, 1992; Hudson and Lester, 1994; Fouda and Al-
Muharrami, 1995).

     Mangrove ecosystems are seriously threatened, mainly by human activities that
impact the habitat (Pons and Fiselier, 1991; Fouda and Al-Muharrami, 1995; Farnsworth
and Ellison, 1997a; Figure
20). The value of mangroves
has gone unrecognized for
many years (Farnsworth,
1998a) and the forests are
disappearing in many parts
of the world. The full extent
of the damage is not yet fully
known, but technological
advances (e.g., airborne
multispectral sensors and
satellite imagery) are
                   Figure 20. Factors impacting mangroves and ecosystems. These
allowing researchers to map
                   valuable systems are under pressure from a variety of physical,
and monitor mangrove
                   chemical, and biological processes. Many of the stresses on these
habitats (Ibrahim and Hashim;    environments result from human activities.
1990; Gang and Agatsiva,
1992; Lin et al., 1994; Aschbacker et al., 1995; Wei et al., 1995; Long and Skewes, 1996;
Green et al., 1997, 1998; N.F.Y. Tam et al., 1997; Blasco et al., 1998; De Jesus and Bina,
1998). The results of such studies are not encouraging; mangrove habitats continue to
shrink around the world.
     Even where efforts have been made to slow the destruction, remaining forests have
a number of problems. In some areas, the health and productivity of the forests have
declined significantly. In Indian mangrove ecosystems, 67% of the mangrove plants, 52%
of the macroalgae, 10% of the invertebrates and 4% of the vertebrates are endangered (e.g.,
Ananda Rao et al., 1998). Similar losses have occurred in the mangals of Singapore
(Turner et al., 1994) and are likely to be seen in other regions of the world. Mangrove
systems require intensive care to save threatened taxa from extinction. The causes of these
tragic losses differ from habitat to habitat but are generally tied directly or indirectly to
human activities. Individual study is required to determine the most effective remedial
measures. Where degraded areas are being revegetated, continued monitoring and thorough
assessment must be done to help us understand the recovery process (van Speybroeck,
1992). This knowledge will help us develop strategies to effectively rehabilitate degraded
mangrove habitats the world over.
     It has long been known that mangrove protect and stabilize coastlines. They are
more effective than concrete barriers in reducing erosion, trapping sediments, stabilizing
shorelines, and dissipating the energy of breaking waves (Pearce, 1996). We have learned
that they are critical nursery habitats for important marine species. Pioneering
investigations are now showing that mangroves and their associated fauna can be sources
of valuable products like black tea, mosquitocides, gallotannins, microbial fertilizers,
antiviral drugs, anti-tumor drugs and UV-screening compounds (Ravi and Kathiresan,
1990; Premanathan et al., 1992; Kathiresan and Pandian, 1991, 1993; Kathiresan, 1995b;
Kathiresan et al., 1995a; Ravikumar, 1995; Moorthy and Kathiresan, 1997b;
Bandaranayake, 1998; Palaniselvam, 1998; Kathiresan, 2000). Mangroves may be
developed as sources of high value commercial products and fishery resources and as sites

for a burgeoning ecotourism industry
(Thorhaug, 1990; Ruitenbeek, 1994;
Barton, 1995). Their unique features may
also make them ideal sites for
experimental studies of biodiversity and
ecosystem function (Osborn and
Polsenberg, 1996; Farnsworth, 1998b;
Field et al., 1998). All this will require
that the resource is understood, carefully
managed, and protected (Farnsworth and
Ellison, 1997b; Ammour et al.., 1999;
Figure 21). Involvement of local
communities in conservation and
education in wise use of our precious
mangrove resources will ensure that
these unique ecosystems survive and

                       Figure 21. Mangroves are highly dynamic and complex
                       systems that are still poorly understood. Continued
                       study, combined with concerted conservation efforts will
                       be necessary to preserve these fragile and unique
                       environments (from Rutzler & Feller, 1996).

We gratefully acknowledge the help of the editors of ‘Advances in Marine Biology’ in
bringing this review to publication. We thank the authorities of Annamalai University and
Western Washington University for providing facilities and resources to complete the
project. We thank Mrs. Sumathi Kathiresan, T. Smoyer, K. Short, R. Lopresti, D. Morgan,
S. Strom, B. Kjerfve and dedicated research students (Dr. N. Rajendran, Dr. V.
Palaniselvam, Ms. B. Kamakshi, Messrs. T. Ramanathan, M. Masilamani Selvam and K.
Sivakumar) for their help during the preparation of the manuscript. A.M. Ellison, S. Strom,
N.M. Aguilar and D. Morgan read and commented on portions of the manuscript. A.M.
Ellison kindly provided unpublished data and information. Two anonymous reviewers
provided constructive criticism


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by David Bael last modified 07-02-2007 13:43

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