6.

Mangrove Structure and Classification

The mangrove spreads widely.

its roots looping. braru:hing. grappling. entwining.
revelling in the sludge;
from its braru:hes it drops aerial roots
into the static convulsions below.
And over all is a dense roof of glossy leafage
which effectively excludes sunlight
from the stinking. impenetrable chaos beneath.
One can do nothing with mangroves but avoid them.
Edgar Beale (1977:172)

6.1 Classification of Mangrove Communities

The physico-chemical and biotic interactions discussed in the previous chapters
produce a range of mangrove communities, which differ in their function and
structure. These functional and structural attributes can be used to classify mangrove
communities, a process which, through its data reduction, can provide an overview
of the types of mangrove communities and how dominant interactions shape them.
Any of the various attributes (physico-chemical, functional, structural or
floristic) of mangrove communities may be useful in classifying them, and the
appropriate selection of attributes depends upon the purpose of the classification.
The classificatory schemes discussed here, however, appear to be of some universal
value in comparing mangrove communities over a range of scales.

6.1.1 Phytosociological Classification

The consistent co-occurrence (and mutual absences) of particular pairs of plant
species, together with their dominance (cover abundance), were used by the Ziirich-
Montepellier (sometimes referred to as the Braun-Blanquet) School of European
ecologists to establish a system of discrete plant communities (Braun-Blanquet
1964). Each of the communities (now termed syntaxonomical units) recognized were
given names, governed by rules contained in the 'International Code of
Phytosociological Nomenclature' (3rd Edition) (Weber et at. 2(00). The scientific
names of the dominant species are used and under Articles II of that Code, the
syntaxa have standardized endings, depending on their rank (Table 6.1).
Table 6.1: Codified tenninations for variously ranked syntaxa according to the International Code of
Phytosociological Nomenclature.

Rank Tennination
Association -etum
Alliance -ion
Order -etalia
Class -etea
Subassociation -etosum
Suballiance -enion
Suborder -enalia
Subclass -enea

While such community labels are very convenient and likely to facilitate a
degree of standardization, there can be no doubt that discrete, consistent and
persistent plant communities are a fiction (Bastow and Chiarucci 2(00).

P. Saenger, Mangrove Ecology, Silviculture and Conservation

© Springer Science+Business Media Dordrecht 2002
184 Mangrove Ecology, Silviculture and Conservation

In his salt marsh studies, Chapman (1960) used a phytosociological

classification; consequently, it is not surprising that he applied this approach to
mangroves (Chapman 1970). He identified and named 8 Alliances, 15 Orders and 40
Associations, a number that almost matched the number of mangrove species (55)
that he recognized. With only minor modifications, Chapman (1975, 1976, 1977)
used this classificatory scheme and attempted to link it to successional sequences.
From time to time, this approach has been used by others in salt marsh aOO
mangrove studies, particularly when comparing reasonably distinct forest
community associations at broad regional scales (Bridgewater 1975, 1985, 1989,
Suzuki and Mochida 1982, Miyawaki et al. 1983, Suzuki and Saenger 1996).
Although the phytosociological approach has never been widely used outside
Europe, and sometimes bemused American ecologists (Snedaker and Brown 1982), it
nevertheless survives. Thus, as recently as 2001, Bouzulle et al. used this approach
to describe syntaxonomical units colonizing abandoned salt pans in western France.

6.1.2 Classification Using Structural Attributes

Specht (1970) developed a structural classification of evergreen plant communities
that uses those properties which reflect the amount of photosynthetic tissue
(contributing to energy input) and the biomass of respiring aerial plant tissue
(involved in energy output). The properties used are (1) the height and life form of
the tallest stratum (which provides an estimate of the biomass) and (2) the 'foliage
projective cover' (FPC) of the tallest stratum. The FPC is the areal proportion of
photosynthetic tissue vertically above the landscape. Ideally, it should be measured
using some crosswire device to determine the presence or absence of foliage
vertically above a large number of randomly selected points in the community. More
often it is estimated using photographic techniques.
Using these two properties, the identification of structural formations can be
achieved. These formations (such as forest or woodland) can be defined further by
including the name ofthe dominant genus or species (such as Avicennia woodland).
The general absence of well-developed understorey and shrub strata in mangrove
communities (Janzen 1985) and the marked tendency towards dominance by one
species of canopy tree mean that it is rarely necessary to seek further precision.
Using this classification, the range of mangrove structural formations that have been
encountered are shown in Table 6.2. The most common of these are closed
communities of variable height (Blasco and Aizpuru 1997). Only rarely d>
mangroves exceed 30 metres in height. Open canopies are associated with high
salinity sites, often at or near high-water spring levels, where rainfall or run-off are
low or moderately seasonal. Open canopies also may occur where persistent
waterlogging is a feature of the environment and, in some other instances, in dwarfed
communities whose structure results from seasonally high salinities (Lin aOO
Sternberg 1992b).
The assumption is made in this classification that the communities are mature,
that is, fully reflecting the constraining effects of water balance, soil fertility,
temperature and light. In practice, this assumption is not difficult to meet as
successional response by FPC is rapid (Specht and Morgan 1981), minimizing any
error arising from this parameter; the age/size structure of the population in relation
to neighbouring sites permits a reasonable assessment of the developmental
(successional) phase of the ecosystem. This scheme can be used validly also where