MANDAL & NASKAR

Tropical Ecology 49(2): 131-146, 2008

International Society for Tropical Ecology
www.tropecol.com

131

ISSN 0564-3295

Diversity and classification of Indian mangroves: a review

R.N. MANDAL*1 & K.R. NASKAR2
1

Central Institute of Freshwater Aquaculture P.O. Kausalyaganga, Bhubaneswar 751002,

Orissa, India
2
Central Inland Fisheries Research Institute, Regional Station, CGO Complex, Kolkata, India
Abstract: Indian mangrove vegetation covers about 6,749 km2 along the 7516.6 km long
coast line, including Island territories. The entire mangrove habitats are situated in three
zones: (1) East Coast, about 4700 km2, (2) West Coast, about 850 km2, and (3) Andaman &
Nicobar Islands about 1190 km2, with Lakshadweep Atoll. These three zones have been further
categorized into Deltaic, Coastal, and Island habitats following Thoms classification of
estuarine habitats. Estimates of the number of species considered mangrove in the world,
range from 48 to 90, and in India from 50-60. We estimate 82 species of mangroves distributed
in 52 genera and 36 families from all the 12 habitats in India. The relative mangrove diversity
(RMD) of each of the 12 habitats is calculated as, RMD=100 [(Fn+Gn+Sn)/N], where Fn, Gn
and Sn are respectively, numbers of families, genera and species, and N = 170 (sum of numbers
of families, genera and species in mangrove vegetation of all the 12 habitats in India).
Sundarbans recorded the maximum RMD (90%) and Lakshadweep Atoll the minimum (9.4%).
The inter tidal vegetation is classified into three categories: Major mangroves, Mangrove
associates, and Back mangal, on the basis of their morpho-anatomical characters representing
adaptation to halophytic condition.
Resumen: En la India la vegetacin de manglar cubre cerca de 6,749 km2 a lo largo de
7,516.6 km de lnea costera, incluyendo territorios isleos. Todos los hbitats de manglar
estn situados en tres zonas: (1) la Costa Oriental, con cerca de 4,700 km2, (2) la Costa
Occidental, con cerca de 850 km2, y (3) las islas Andamn y Nicobar, con cerca de 1,190 km2,
incluido el atoln Lakshadweep. Estas tres zonas fueron a su vez categorizadas en tres
hbitats: deltaico, costero e insular, de acuerdo con la clasificacin de Thom para los hbitats
estuarinos. Las estimaciones del nmero de especies consideradas de manglar en el mundo
varan entre 48 y 90, y en la India van de 50 a 60. Nosotros estimamos que hay 82 especies
de manglar distribuidas en 52 gneros y 36 familias en los 12 hbitats en la India. La
diversidad relativa del manglar (DRM) de cada uno de los 12 hbitats se calcul como
DRM=100 [(Fn+Gn+Sn)/N], donde Fn, Gn y Sn son, respectivamente, los nmeros de
familias, gneros y especies, y N = 170 (suma de los nmeros de familias, gneros y especies
en la vegetacin de manglar de los 12 hbitats en la India). En Sundarbans se registr la
mxima DRM (90%) y en el atoln Lakshadweep se registr la mnima (9.4%). La vegetacin
intermareal fue clasificada en tres categoras: Principalmente manglar, Asociada al
manglar y Manglar posterior, con base en caracteres morfoanatmicos que representan
adaptaciones a la condicin haloftica.
Resumo: A vegetao dos mangais indianos cobrem cerca de 6,749 km2 ao longo da linha
de costa, incluindo os territrios insulares. O conjunto dos habitats do mangal encontram-se

Corresponding Author; e-mail: rnmandal2003@yahoo.com

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DIVERSITY & CLASSIFICATION OF MANGROVES

situados em trs zonas: (1) Costa Oriental, cerca de 4700 km2, (2) Costa Ocidental, cerca de 850
km2 e (3) Ilhas de Andaman & Nicobar cerca de 1190 km2 com o Atoll de Lakshadweep. Estas
trs zonas foram posteriormente categorizadas nos habitats Estuarino, Costeiro e Insular de
acordo com a classificao dos habitats estuarinos. Estimativas do nmero das espcies
englobadas como de mangal, oscilam entre as 48 e as 90, e na ndia entre 50-60. De todos os 12
habitats na ndia estimou-se em 82 as espcies de mangal distribudas por 52 gneros e 36
famlias. A diversidade relativa do mangal (RMD) de cada dos 12 habitats foi calculada como
RMD=100 [(Fn+Gn+Sn)/N], onde Fn, Gn e Sn so, respectivamente, nmeros de famlias,
genro e espcies, e N=170 (soma do nmero de famlias, gneros e espcies de vegetao de
mangal de todos os 12 habitats na ndia). Sundarbans apresentou a maior RMD (90%) e o Atoll
de Lakshadweep o mnimo (9,4%). A vegetao entre-mars, na base dos caracteres morfoanatmicos representando as condies de adaptao halfita classificada em trs categorias:
Mangal principal, Associadas do mangal e Retaguarda do mangal.

Key words: Classification, diversity, India, mangroves, mangrove habitats.

Introduction
Biodiversity is prevalent in the tropical
estuarine system, particularly in the inter tidal
forested vegetation known as Mangrove (Mooney et
al. 1995), which covers about 240 x 103 km2 (Lugo
et al. 1990; Twilley et al. 1992). This vegetation
occurs in river deltas, lagoons and estuarine
complexes (Thom 1982), and also colonizes
shorelines and islands in sheltered coastal area,
with locally variable topography and hydrology,
leading to relatively specific structure and function
(Lugo & Snedaker 1974).
Mangrove has been variously defined in
literature. The oxford dictionary mentioned the
words mangrove and mangrowe since 1613,
indicating tropical trees or shrubs found in coastal
swamps with tangled roots that grow above the
ground, whereas the Americans, the Spanish, and
the Portuguese used the term Mangle and
Mangue indicating trees and shrubs of the genus
Rhizophora (Mepham & Mepham 1984). Later, the
term mangrove was referred to the individual
plant or tidal forest or both, as Mangrove plants
and Mangrove ecosystem (MacNae 1968).
Chapman (1976) used the term mangrove for
inter tidal plants, and considered plant
communities of inter tidal forest as mangrove
ecosystem called mangal. The term mangal was
also commonly used in French and in Portuguese
to refer to both forest communities and to
individual plants. Several workers have opined

that plants growing in between the highest and

the lowest tidal limits may be considered
mangrove (Aubreville 1964; Blasco 1975, 1977;
Clough 1982; Davis 1940; Grzimek et al. 1976;
MacNae 1968; Naskar & GuhaBakshi 1987;
Tomlinson 1986). The tidal limits of various
habitats, however, can vary.
India with a long coastline of about 7516.6 km,
including the island territories (Anonymous 1984),
has a mangrove cover of about 6,749 km2, the
fourth largest mangrove area in the world (Naskar
& Mandal 1999). These mangrove habitats (69E89.5E longitude and 7N-23N latitude) comprise
three distinct zones: East coast habitats having a
coast line of about 2700 km, facing Bay of Bengal,
West coast habitats with a coast line of about 3000
km, facing Arabian sea, and Island Territories
with about 1816.6 km coastline. The long
coastlines and their mangrove vegetation have
immense role in protecting coastal biodiversity.
This paper highlights the diversity of mangrove
habitats in relation to environmental factors,
mangrove species diversity, along with their
distribution, regeneration and growth, and
classification of vegetation growing in and around
mangrove habitats in India.

Habitat diversity
Based on Thoms (1982) classification of
coastal habitats, the Indian mangrove habitats are
categorized as below:

MANDAL & NASKAR

133

(a) Deltaic mangrove habitat

(c) Island mangrove habitat

It is characterized by high tidal range with

associated strong bi-directional tidal currents.
These currents are responsible for the dispersion
of sediments brought to the coasts by rivers and
the main river channels, which are funnel shaped
and are fed by numerous tidal creeks. It includes
the major estuaries of East coast and two gulfs of
West coast, Gujarat.

It is commonly found at the mouths where

rivers are seen to border the open sea. Neither
marine nor river deposition has been sufficient to
fill up what is an open estuarine system.
Sometimes the bedrock of onshore area has been
drowned by a rising sea level. It includes shallow
but protected inter tidal zones of Bay Islands and
Lakshadweep Atoll.
Considering geomorphological characters, the
mangrove habitats of India can be viewed as
follows:

(b) Coastal mangrove habitat

It is characterized by low tidal range. River
discharge of fresh water and sediments lead to
rapid deposition of terrigenous sands, silts and
clays to form deltas. These deltas build seawards
over flat offshore slopes composed of fine-grained
pro-delta sediments. It includes the inter-tidal
coastlines, minor river mouths, sheltered bays and
backwater areas of the West coast.

(a) East coast mangrove habitats

Sundarban mangrove forest (SF), Ganga delta,
West Bengal; Subarnarekha mangrove forest (SM),
Bhitarkanika mangrove forest (BF), Mahanadi
delta, Orissa; Godavari and Krishna delta
(G&KD), Andhra Pradesh; Pichavaram Estuary

Fig. 1. Distribution of mangroves in 12 different habitats in India, along

with that in Bangladesh. SF= Sundarbans mangrove forest, West Bengal;
SM= Subarnarekha mangrove forest, Orissa; BF= Bhitarkanika mangrove
forest, Orissa; G&KD= Godavari & Krishna delta, Andhra Pradesh; PE=
Pichavaram mangrove estuary, Tamilnadu; CE= Cochin estuary, Kerala;
C&MA= Coondapur/Malpe area, Karnataka; ZE= Zuary estuary, Goa; BM=
Bombay mangrove creeks, Maharashtra; BE= Bhabnagar estuary, Gujarat;
A&N= Andaman & Nicobar islands; LA= Lakshadweep Atoll; BDF=
Bangladesh mangrove habitat..

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DIVERSITY & CLASSIFICATION OF MANGROVES

(PE), Cauvery Estuary, Tamilnadu Fig. 1, are

considered to fall under deltaic mangrove forest,
situated at the mouth of respective major rivers
carrying freshwater, facing Bay of Bengal by North
to South and North-West to South directions
(Jagtap et al. 1993; Mandal 1996; Sanyal et al.
1998; Untawale & Jagtap 1992). They altogether
cover about 4700 km2 or 70% of the total mangrove
area, and are rich in mangrove species. In East
coast habitats, SF of Indian part alone covers
about 4200 km2 (Govt. of India 1987), while the
undivided Sundarbans (including Bangladesh
part, BDF) is the largest single mangrove block in
the world (Blasco 1977).

(b) West coast mangrove habitats

Cochin estuary (CE), Kerala; Coondapur/
Malpe area (C&MA), Karnataka; Zuary estuary
(ZE), Goa; Bombay mangrove creeks (BM),
Maharashtra; and Bhabnagar estuary (BE),
Gujarat Fig. 1, are considered to fall under coastal
mangrove habitats in the inter tidal zones along
with mouths of minor rivers or minor estuaries
and back waters, facing Arabian sea by North-East
to West and East to West directions. They
altogether occupy about 850 km2 (12% of the total
Indian mangrove area), though Sidhu (1963) once
mentioned the cover at about 1140 km2. The
mangrove vegetation is sparse, less extended, and
confined to patches due to scanty upstream
freshwater supply, excessive amount of silt-clay
deposition, low average rainfall and relatively low
tidal fluctuation (Blasco & Aizpuru 1997; Naskar
& GuhaBakshi 1987; Untawale 1984).

(c) Andaman & Nicobar and Lakshadweep

islands
Andaman & Nicobar Islands (A&N) and
Lakshadweep Atoll (LA), comprising more than
200 and 37 islands, respectively, harbour island
mangroves (Fig.1). This is the second largest
mangrove coverage, estimated at about 1199.6 km2
or about 18% of the total Indian mangrove. Tall
dense mangrove forest occurs in A&N Islands due
to favourable climatic conditions such as short dry
season and high tidal fluctuation, heavy amount of
rainfall (Blasco & Aizpuru 1997; Dagar et al. 1991;
Naskar & Mandal 1999). On the other hand, small
and sparse mangroves with stunted growth are
reported from LA (Newberri & Hill 1981; Spicer &
Newberri 1979).

Habitat diversity based on tidal

fluctuation
Lear & Turner (1977) observed that mangrove
vegetation is found between MSL (Mean sea level)
and HATL (Highest Astronomical Tide Level).
They recommended that mangrove might be
subdivided into two groups: (i) Major mangroves
zone - between the MSL and MHWSTL (Mean
High Water Spring Tide Level), and (ii) Minor
mangroves zone - between MHWSTL and HALT.
Blasco (1975), following Aubrevilles view (1964),
divided Indian mangrove habitats into two groups
based on the tidal range: (i) Swampy mangroves situated below the high tidal level and submerged
by sea water twice a day, and (ii) Tidal mangroves
- inundated only during spring tide and sometimes
by unprecedented sea surges.

Factors responsible for mangrove

species regeneration and growth
Generally, tidal waters bring nutrients along
with other essential minerals to the on-shore region
where they become available to mangroves. This
tidal water was earlier considered to be the only
factor playing a major role in the regeneration and
growth of mangroves. But, it has been observed that
other factors also contribute; rainfall (200 cm - 300
cm), atmospheric humidity (60%-90%), and moderate
temperatures (19C - 35C) have been considered
ideal for mangroves growth (Blasco 1977; Naskar &
Mandal 1999). In spite of having maximum tidal
fluctuations, Bhabnagar estuary (BE), therefore,
does not have high mangrove species diversity
because of its low average rainfall (60 cm annum-1),
and inadequate upstream freshwater supply (Blasco
& Aizpuru 1997). Venkatesan (1966) has argued that
the mangrove habitats would remain productive as
long as they get inundated with tidal water, receive
high rainfall annually, and are benefited with
continuous upstream freshwater which usually carry
silt, sediments and organic matter. Mandal (1996)
supported the above views while investigating seed
germination and seedling development of mangroves.
The major mangroves initially require fresh water to
continue their physiological process until they
develop salt secretary organs, such as salt glands,
corkwart, gall, and other related mechanisms
(Naskar et al. 1997; Naskar & Mandal 1999).

MANDAL & NASKAR

Mangrove species diversity

Till date, the usage of the term mangrove and
the number of mangrove species assigned on that
basis vary remarkably among different workers.
The species growing in the same region are
designated differentially; some consider them as
mangroves, but others do not. For instance, Jones
(1971) reported 27 mangrove species from the
Eastern Australia, whereas Walsh (1974)
mentioned 55 mangrove species from the same
region, and Bunt et al. (1982) selected 45 species
among them as mangrove. World-wide, Chapman
(1976)
reported
90
species
representing
mangroves, Saenger et al. (1983) recorded 83
species, UNDP/UNESCO (1986) reported 65
species, and Tomlinson (1986) mentioned only 48
mangrove species, out of which 40 were considered
true mangroves found in the Old world Tropics
(Indo-West pacific region) and 8 true mangroves
from the New world Tropics. In India also, the
reported number of mangrove species varies
among researchers: 50-60 species (Blasco et al.
1975); 33 species from West coast and 47 species
from East coast, but about 55 species from all the
habitats
(Untawale 1987); 35 true mangrove
species, 28 mangrove associates and 7 back
mangals (Naskar & GuhaBakshi 1987); 59 species,
out of which 34 species from East coast and 25
species from West coast (Banerjee et al. 1989); and
50 species (Jagtap et al. 1993).

Indian scenario
Usually, the coastal estuarine vegetation that
occurs in inter-tidal zones i.e. between the highest
and the lowest tidal limits is considered to be
mangrove. A total of 82 mangrove species (Table
1), distributed in 52 genera and 36 families, has
been recorded by different workers. Sundarban
mangrove forest (SF), West Bengal, shows the
highest taxa diversity: 69 species, 49 genera, 35
families, including two species, viz. Scyphiphora
hydrophyllacea and Atalentia corea reported for
the first time from Indian Sundarbans (Mandal et
al. 1995). In addition, Acanthus volubilis, which
was regarded as extinct, has been recorded again
with its very limited population existing in
confined location. This species has not been
reported from any other mangrove habitat in
India. Importantly, it has been found that both the
mangrove palms: Nypa fruticans and Phoenix

135

paludosa are largely restricted to Sunderban

mangrove area and to Andaman & Nicibar Islands;
the latter species also occurs in Bhitarkanika
mangrove forest.
Other habitats have the
following
diversity
(Appendix
Table
1):
Subarnarekha mangrove forest (SM), 18 species,
14 genera, 11 families; Bhitarkanika mangrove
forest (BF), 57 species, 37 genera, 29 families,
including one new species Heritiera kanikensis
(Banerjee 1984) and with relatively higher number
of true mangroves; Godavari & Krishna delta
(G&KD), 36 species, 26 genera, 21 families;
Pichavaram mangrove estuary (PE), 35 species, 26
genera, 20 families, including one new species
Rhizophora annamalayana Kathir., the hybrid of
R. apiculata x R. mucronata (Kathiresan 1995).
Naskar (1993) has reported more than 60
mangrove species, including 25 major mangroves
in East coast zones. In the West coast, Cochin
estuary (CE) with 32 species, 24 genera, 19
families, despite negligible area coverage, has the
second highest number of species after Zuary
estuary, Goa, which has the highest numbers: 33
species, 25 genera, 19 families, among habitats in
the West coast. Coondapur & Malpe area (C&MA)
has 22 species, 16 genera, 12 families; Bombay
mangroves (BM) has 29 species, 21 genera, 16
families, including Salvadora persica, a mangrove
associate herb growing only in two habitats (BM
and BE). Bhabnagar estuary (BE) has 26 species,
19 genera, 15 families. Andaman & Nicobar
Islands (A&N) harbour 61 species distributed in 39
genera and 30 families, including two new species,
R. lamarkii and R. stylosa (Singh et al. 1987), and
has the maximum species of the genus Rhizophora
(four species). Lakshadweep Atoll (AL) has 8
species distributed in 5 genera and 3 families.

Relative mangrove diversity of habitats

An index has been developed to represent the
relative mangrove diversity (RMD) of different
habitats. Relative mangrove diversity (RMD) = 100
x [(Fn+Gn+Sn)/N], where, Fn, Gn, and Sn are
respectively, numbers of families, genera and
species of a habitat, and N = 170 (sum of reported
numbers of families, genera and species from all
12 mangroves habitats in India, i.e. 36+52+82
(Appendix Table 1). Among the 12 habitats, SF
records 90% RMD, followed by A&N- 76.5%, BF72.3%, G&KD- 48.8%, PE- 47.6%, ZE- 45.3%, CE44.1%, BM-38.8%, BE- 35.3%, C&MA- 29.4%, SM-

136

DIVERSITY & CLASSIFICATION OF MANGROVES

25.3%, and LA- 9.4%. Thus SF shows the highest

RMD and LA the lowest among all the habitats.
Higher the RMD, greater is the resemblance of the
habitat to overall taxa composition of mangrove
vegetation of India.

Classification
Different opinions
Saenger et al. (1983) listed 60 mangrove
species growing exclusively in the tidal zones, as
true mangroves. Mepham & Mepham (1984)
suggested that any arborescent species growing in
the tidal zones might be referred to as Potential
mangrove or Frequent mangrove. Tomlinson
(1986) recommended that mangrove species were
basically of two types, viz., (1) Major element of
mangals or true mangroves with complete
fidelity to the mangrove environment, and (2)
Minor element of mangals not conspicuous in
mangrove habitats, rather might prefer the
peripheral habitats of mangrove regions. Later, Li
& Lee (1997) used two terms, viz., true mangrove
and semi mangrove. Subsequently, several
workers, e.g. Watson (1928), Tomlinson (1980),
Chai (1982), Mepham & Mepham (1984), and
Naskar (1993) coined the term Mangrove
associate for the flora representing nonarborescent, herbaceous, sub-woody and climber
species, found growing mostly in regions bordering
the tidal periphery of mangrove habitats.

Indian perspective
The present study enumerated the modified
morphological and anatomical characters of leaves,
stems, roots, and reproductive organs of 63 species
and evaluated them in relation to halophytic
adaptation. Leaves: lamina (1) fleshy, flattened,
shiny, (2) coriaceous in dorsal surface, thus it
reflects bright sunlight and checks transpiration
rate, marked with symbols lm I, 2; anatomy (1)
thick cuticle & waxy epidermis, (2) dense hair,
unicelled/multicelled glands, (3) sunken stomata,
(4) hypodermal aqueous tissue, (5) central aqueous
tissue, (6) sclereids/stone cells, (7) salt glands/cork
wart, lenticels, each contributes to reduction of
transpiration rate, storing of water, maintenance
of succulence, provision of mechanical support, and
excretion of excess salt, marked with symbols la 1,
2, 3, 4, 5, 6, 7. Stem: (1) swollen trunk base, (2)

gall, provide mechanical support and facilitate

aeration, respectively, marked with symbols sm 1,
2; anatomy - (1) greater number of vessels mm-2
and multiseriate fibre, (2) secondary anomalous
growth, perform functions like uplift of a huge
amount of water and protection of conductive
tissue, marked with symbols sa 1, 2. Roots: (1) all
types of aerial roots, (2) pneumatophores, (3) cable
roots, (4) pseudo taproots, provide mechanical
support and facilitate breathing, marked with
symbols rm 1, 2, 3, 4; anatomy (1) lenticels, (2)
number of air cavities, (3) sclereids, (4) circular
frequent sclereid ring, aid in breathing and
provision of mechanical support, marked with
symbols ra 1, 2, 3, 4. Reproductive organ: (1)
vivipary, (2) cryptovivipary, facilitate seed
germination while attached with mother plant,
marked with symbols rom 1, 2; anatomy (1)
presence of numerous air cavities, (2) conspicuous
vacuum between seed and pericarp, support
breathing and salt excretion, and buoyancy,
respectively, marked with symbols roa 1, 2. In
calculation, one modified part has been assigned
one point and the sum of the total points has been
put against each taxon (Appendix Table 2).
Families such as Rhizophoraceae, Avicenniaceae,
Sonneraceae,
Combretaceae,
Arecaceae,
Sterculiaceae,
Meliaceae,
Euphorbiaceae,
Rubiaceae,
Agialitidaceae,
Poaceae,
and
Acanthaceae score points between 7-14. They
possess characters which belong to at least three
modified organs out of four, viz., leaves, stems,
roots, and reproductive organs. Members of all
these families are considered major mangrove.
Other families such as Acanthaceae, Tiliaceae,
Asclepediaceae,
Malvaceae,
Chenopodiaceae,
Loranthaceae,
Aizoacae,
Boraginaceae,
Verbenaceae, Amaryllidaceae, Convolvulaceae,
Rubiacae, and Pteridaceae score points betweens 2
- < 7. They have characters, which belong to at
least one modified organ out of four. All such
species have been considered mangrove associate.
None of the members belonging to the families
Fabaceae, Solanaceae, and Rutaceae scores any
point because of absence of character modification
essential for halophytic adaptation; and so, such
species are considered back mangals. Under this
assignment, 30 species fall under major
mangroves, 21 species mangrove associates and
12 species back mangal.

MANDAL & NASKAR

Loss of mangrove area

Inspite of their immense role in protecting
human resource as well as biodiversity, these
unique mangrove habitats of India have been
facing tremendous threats due to indiscriminate
exploitation of mangrove resources for multiple
uses like fodder, fuel wood, timber for building
material, alcohol, paper, charcoal and medicine
(Upadhyay et al. 2002). Apart from those,
conversion of forest area to aquaculture and
agriculture, construction of port and harbour,
extension of human inhabitation, over-grazing,
urbanization, industrialization, and chemical
pollution are major common occurrences that
dwindle mangrove area (Blasco & Aizpuru 1997;
Naskar 2004; Upadhyay et al. 2002). Owing to
these threats > 33% of the Indian mangrove areas
has been lost within the last 15 years. Of this, East
coast area has lost about 28%; West coast area
about 44%; and Andaman & Nicobar Islands
about 32% (Jagtap et al. 1993; Naskar 2004).

Conclusions
The proposed Relative Mangrove Diversity
(RMD) of the Indian habitats can help in assessing
the diversity of Indian mangrove species in
different habitats: gain or loss in course of time.
Study of morpho-anatomical characters in relation
to adaptation to halophytic conditions can help in
formulating strategic plans to afforest mangroves;
the classification system based on these characters
may also be a guideline to select the appropriate
zone for respective flora.

Acknowledgements
We are grateful to the Director, CIFRI,
Barrackpore for providing us laboratory facilities for
carrying out some investigation of mangroves. We
record our sincere thanks to those who helped us a
lot during field study. We duly acknowledge the
help extended by Dr. K. K. Sharma, Dr. N. K.
Maity, Dr. M. Samanta and B. K. Sarker for their
valuable suggestions for preparing this manuscript.

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