Journal of Plant Ecology Advance Access published February 3, 2012

Journal of
Plant Ecology Invaders control on
PAGES 110

doi: 10.1093/jpe/rtr050 post-disturbance succession in

coastal mangroves
Shekhar R. Biswas1, Md. Saiful Islam Khan2,3 and
Azim U. Mallik4,*
1
Department of Ecology and Evolutionary Biology, University of Toronto, 3359 Mississauga Road, Mississauga, Ontario L5L
1C6, Canada
2
United Nations Environment Programme-World Conservation Monitoring Centre (UNEP-WCMC), Cambridge CB3 0DL, UK
3
Forest and Landscape, LIFE, Copenhagen University, 1958 Frederiksberg C, Denmark
4
Department of Biology, Lakehead University, 955 Oliver Road, Thunder Bay, Ontario P7B 5E1, Canada
*Correspondence address. Department of Biology, Lakehead University, 955 Oliver Road, Thunder Bay, Ontario,
Canada P7B 5E1. Tel.: +1-807-343-8927; Fax: +1-807-346-7796; E-mail: amallik@lakeheadu.ca

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Abstract

Aims biological invasion associated with large disturbance. We argue that

In recent years, coastal mangroves have been frequently affected by
large disturbances (cyclones, hurricanes, flooding and tsunamis) and
post-disturbance vegetation is often dominated by small stature
mangrove, mangrove-associate and non-mangrove species poten-
tially affecting ecosystem functioning. Knowledge on the processes
of mangrove vegetation development and recovery (succession)
following normal and large disturbances will benefit practitioners
in designing robust ecosystem management/restoration plans. Here
we propose a conceptual model of disturbance-mediated succession
in mangroves.
Methods
Based on field observations and species life history traits, we
develop conceptual models of mangrove succession under normal
disturbance regime and recently experienced increased frequency
of large disturbances. We evaluate our conceptual models by
conducting a scenario testing experiment.
Important Findings
We suggest two predominant processes affecting mangrove
succession after disturbance: propagule limitation due to damage
of seed producing mature trees and dispersal barrier resulting from
large disturbances affect mature trees more than the small-stature
non-tree (shrubs, herbs and climbers) species creating a larger
propagule shortage for mangrove tree species than non-tree species.
Secondly, large disturbances facilitate invasion of free-floating
aquatics, which may interfere with the flow-facilitated propagule
dispersal and seedling establishment of mangrove species. In a sce-
nario testing experiment, we have shown that similar levels of dis-
turbance impact vegetation development and recovery differently
depending on the presence or absence of invasive species. We con-
clude that since biological invasion is one of the major drivers of
post-disturbance mangrove succession, the dimension of biological
invasion should be included in prediction, management and
restoration of mangrove forests.
Keywords: biological invasion d cryptic ecological
degradation d dispersal barrier d large disturbances d mangrove
restoration d mangrove succession
Received: 12 October 2011 Revised: 10 November 2011 Accepted:
26 December 2011

et al. 2011). Mangroves, a unique type of plant community

INTRODUCTION
Disturbances affect dynamics and composition of ecological
communities (White and Pickett 1985). The degree to which
ecological communities recover following a disturbance
depends on the severity and frequency of the disturbance
and the resiliency of the ecosystem (Holling 1973; Peters
along the tropical and subtropical coastlines, are species poor
but a resilient system where natural disturbances and subse-
quent community recovery through succession is a norm
(Alongi 2008). However, recently, the typical pattern of
natural disturbances has been changed largely due to
climate change; large disturbances (the disturbance that

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2 Journal of Plant Ecology
affects spatially large area with its high magnitude and ampli-
tude, e.g. cyclones, hurricanes, catastrophic flooding, etc.;
Dale et al. 1998) are now more intense and frequent than oc-
curred anytime in the past (Solomon et al. 2007). This altered
pattern of disturbances is likely to affect mangrove succession
(Grindrod et al. 2002; Lugo 2008). Thus, it is not surprising that
the species composition in post-disturbance mangroves differs
from that prior to disturbance (sensu Lugo 2008). Importantly,
the species composition in post-disturbance mangroves
appears to be characterized by high abundance of small stature
mangrove, mangrove-associate and salt-tolerant non-man-
grove species of herbs, shrubs and climbers (Baldwin et al.
1995; Bardsley 1985; Brown 2007; Ferwarda et al. 2007; Lugo
2000; Paling et al. 2008; Roth 1992; Smith et al. 1994, 2009;
Urquhart 2009). Although occurrence of mangrove-associate
and non-mangrove species along the forest edges is a normal
phenomenon in mangrove forests, an increasing abundance of
these species in the forest interior is rather unusual (Brown
2007; Harun-or-Rashid et al. 2009). Without any change in
the spatial extent of forest cover, changes in species composi-
tion in terms of increasing abundance of low stature man-
grove-associate or non-mangrove species in place of
mangrove species may lead to cryptic ecological degradation
in the mangrove forest (Dahdouh-Guebas et al. 2005a). This
is a cause for concern as change in species composition is often
associated with a change in ecosystem function (Field et al.
1998; Osborn and Polsenberg 1996). Unlike the large-stature
mangrove trees, increased abundance of small-stature man-
grove-associate or non-mangrove species may compromise
the efficiency of mangroves to buffer large hydrodynamic dis-
turbances (Dahdouh-Guebas et al. 2005a). In addition, changes
in species composition may have cascading effects on ecolog-
ical associations such as mangrove-associated faunal commu-
nity (e.g. Ellison et al. 1996; Lee 2008), food web (Brusati and
Grosholz 2009), and ecologysociety linkage such as livelihood
of mangrove-dependent human communities (Biswas et al.
2009). Thus, management actions need to counteract, and
for that, an in-depth understanding of the processes of man-
grove vegetation development and recovery, i.e. successional
processes following large disturbances is essential.
Mangroves occur in the highly dynamic environment
where land accretion and erosion are the norms, and thus,
mangrove succession occurs in diverse physical and geomor-
phic contexts (Friess et al. 2011). Formation of mangroves
in the newly raised lands (primary succession) and recovery
of old mangroves in established land affected by large
disturbances (secondary succession) are the two major types
of mangrove succession in the context of large disturbances.
It has been suggested that large disturbances affect mature
mangrove vegetation constraining propagule availability nec-
essary for both primary and secondary succession (Kauffman
and Cole 2010; Smith et al. 1994, 2009). However, the degree
to which a species will be affected by disturbance severity and
frequency is related to its life history traits (e.g. stem height
and diameter, branching pattern, mode of regeneration and
growth habit). Mangrove trees are greatly affected by large dis-
turbances due to their large stature, but mangrove associates or
non-mangrove species are relatively less affected by such
disturbances because of their smaller stature (Bardsley 1985;
Baldwin et al. 1995; Smith et al. 1994). This inequality in
vegetation damage from a disturbance translates into higher
propagule shortage for mangrove tree species than non-tree
mangrove associates and non-mangrove species. Presumably,
mangrove-associate or non-mangrove species capitalizing on
the lower vegetation damage from disturbance and altered
ecosystem condition gain an upper hand in the recovered
mangroves (Dahdouh-Guebas et al. 2005b). Inequality of
propagule availability among the two (tree and non-tree
species) will also influence the species composition of newly
raised lands.
However, disturbance in an ecosystem often creates cascade
of events (Peters et al. 2011), so that the effects of large distur-
bances on mangroves may not be limited to only propagule
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limitation. Among other factors, large disturbances facilitate
biological invasion (Elton 1958; Bellingham et al. 2005;
Harun-or-Rashid et al. 2009; Lugo and Helmer 2004; Williams
and Grosholz 2008). Many invasive species in mangroves are
free-floating aquatics, and they may affect flow-facilitated
propagule dispersal and seedling establishment along river-
banks (Biswas et al. 2007; Gordon 1998; Harun-or-Rashid
et al. 2009; Schmitz et al. 1993). Disturbance-related interfer-
ence of invasive species in propagule dispersal and seedling
establishment of mangrove, non-mangrove and mangrove-
associate species have not been explicitly studied, but our field
observation in the Sundarbans mangrove offers useful
insights: dispersal of mangrove tree propagules are more
affected than non-mangrove or mangrove-associate species.
In the Sundarbans, we observed that large quantities of a float-
ing invasive species, Eichhornia crassipes (Mart.) Solms, enter in
the mangrove system after large flooding (large disturbance).
The floating mats of E. crassipes blocks large mangrove tree
propagules (e.g. Heritiera fomes Buch.-Ham., Bruegueria
gymnorhhiza (L.) Lam., Rhizophora spp.) at a larger proportion
than the small propagules of mangrove-associate species such
as Derris trifoliata Lour. (Troup 1921) in water channels, result-
ing in increased abundance of D. trifoliata in the forest interior;
this pattern of species filtering seems irreversible (Fig. 1).
Studying the mangroves of Florida, Schmitz et al. (1993) also
reported a similar role of the invasive species E. crassipes in
habitat modification and physical barrier to propagule
dispersal, causing reduced community-level establishment of
large-seeded mangrove species (see Gordon 1998). Surpris-
ingly, the dimension of biological invasion remains largely un-
accounted for in understanding post-disturbance vegetation
succession in mangroves.
The goal of this contribution is to highlight the effects
of biological invasion on post-disturbance succession in
mangroves. We propose conceptual models of mangrove
succession under normal disturbance regime (regular tidal
flooding and cyclones and infrequent large flooding and
Biswas et al. | Post-disturbance succession in coastal mangroves 3

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Figure 1: stages of vegetation development in the Sundarbans mangrove. The left hand panel shows succession in normal conditions: mature
mangrove forest (a); propagule supply (b); Sonneratia apetala stand and in the inset Derris trifoliata at riverbank (c). The middle panel shows post-
disturbance succession: affected vegetation (d) and invasive flow (e); newly raised land colonized and dominated by D. trifoliata (f, g). The right
hand panel shows: Eichhornia crassipes created a mat along the riverbank (h), floating E. crassipes blocked a small creek (i), E. crassipes colonized at
riverbank (j) and sediment trapping by E. crassipes (k). Mangrove species = S. apetala, mangrove-associate species = D. trifoliata and invasive
species = E. crassipes.

cyclones) and recently experienced increased frequency of
large disturbances (frequent large flooding and cyclones).
We argue that: (1) under normal disturbance regime, man-
grove regeneration follows typical successional patterns,
e.g. pioneer mangrove species (and sea grasses) / mid serial
mangrove species / climax mangrove species. (2) Following
high frequency of large flooding and cyclones, the effect of
dispersal barrier due to biological invasion causing propagule
limitation may facilitate shift in dominance of mangrove
species to mangrove-associate or non-mangrove species. To
test our conceptual model, we conducted a scenario testing
experiment with varying degrees of dispersal barrier
(1090%) and propagule limitation (1090%) and explored
their effects on dominance of mangrove species. We also
explored the scenarios of mangrove formation when man-
grove-associate species experience equal or less (one-half,
one-third, one-fourth) dispersal barrier than large mangrove
species.
4 Journal of Plant Ecology

A CONCEPTUAL MODEL OF MANGROVE
SUCCESSION
Vegetation development in normal conditions
We will consider a mature mangrove stand comprising a man-
grove tree species, Sonneratia apetala Buch.-Ham (hereafter
referred to as mangrove species) and a mangroves-associate
species, D. trifoliata (hereafter referred to as mangrove-
associate species), and limit the scope of this model to processes
relevant to increasing abundance of mangrove-associate
species in mangroves (Fig. 2).
Let us consider that the mangrove stand has Mt mature
individuals of a mangrove species and At mature individuals
of a mangrove-associate species at time t (see Table 1 for a list
of symbols). If fecundities of mangrove and mangrove-
associate species are Fm and Fa, the net propagule productions
are MtFm and AtFa, respectively. These propagules will disperse
to newly raised lands or suitable vacant habitats in pre-existing
(Fransworth 2000; Harun-or-Rashid et al. 2009), say St
1.
Viable propagules germinate and seedlings of mangrove and
mangrove-associate species occupy by cm and ca proportions
of the vacant habitat, respectively. Seedlings may experience
density-dependent mortality (survivorship: Cm and Ca).
Although seedling banks are important in post-disturbance
mangrove regeneration (Roth 1992), it is most likely that large
disturbances affect seedling banks of both mangrove and
mangrove-associate species equally because of their small
stature. Therefore, we did not include this factor in the
conceptual model. Similarly, we assumed constant and equal
density-dependant mortality for both species because we are
primarily interested in dispersal barrier and propagule
limitation.
Taken together, the dominance of mangrove species over
mangrove-associate species in the new (post-disturbance)
habitat at time t + 1 can be expressed by their ratio (equation 1)
or scaled dominance (equation 2, scale 01; >0.5 means

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stands by tidal flow. During dispersal, a portion of the propa-
gules can be washed-out from the mangrove system to the sea
by tidal current and some propagules can be lost by predation
(total loss: am and aa). Similarly, a portion of the propagules
can encounter physical barrier from the invasive aquatics
(E. crassipes) at their dispersal routes (bm and ba).
Mangrove species do not persist in the soil seed bank
for extended period but mangrove-associate species do
mangroves dominance). For simplicity of interpretation, we
consider scale-free ratio from this point forward (equation 1),
so that if M:A > 1, mangrove species dominates over
mangrove-associate species.
Mt Fm vm f1
am + bm g1
ca Cm
Mt + 1 : At + 1 = : 1
At Fa va f1
aa + ba g + St
1 1
cm Ca

Figure 2: simplified schematics of a normal (A) versus an alternate pathway of mangrove succession (B). The grey and dark hatched areas in-
dicate occupancies of mangrove and mangrove-associate species, respectively. Arrows indicate propagule flow (solid arrow = uninterrupted flow,
broken arrow = interruption). The shaded areas at dispersal route indicate presence of invasive species. The bottom row indicates: mangroves are
more affected than mangrove associates in relation to disturbances (a), disturbance increases biological invasion (b), with increasing invasion,
propagule availability of mangrove species drops sharply but little effects on mangrove-associate species (c) and with increasing invasive-barrier,
occupancy of mangrove-associate species increases (d). The solid lines in the bottom panel indicate mangrove species and the dotted lines indicate
mangrove-associate species.
Biswas et al. | Post-disturbance succession in coastal mangroves 5

Table 1: symbols used in the conceptual model and their definition

Hypothetical values used in the

Symbol Definition (unit) Mangrove vs. mangrove-associate species scenario testing experiment

Mt Mature individuals at times t (number per area) For mangrove species 100
At For mangrove-associate species 20
Fm Fecundity (number of individuals per year) For mangrove species 500
Fa For mangrove-associate species 250
vm Propagule viability (proportion) For mangrove species 0.8
va For mangrove-associate species 0.8
Sm Propagules in the soil seed bank (proportion) For mangrove species 0
Sa For mangrove-associate species 0.1
Cm Per capita survivorship following density-dependent For mangrove species 0.9
Ca seedling mortality (proportion) For mangrove-associate species 0.9
am Propagule loss due to tidal washout and propagule For mangrove species 0.1
aa predation (proportion) For mangrove-associate species 0.1
bm Propagules blocked by invasive species (proportion) For mangrove species 0.10.9
ba For mangrove-associate species 0.10.9

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cm Newly raised land colonized (proportion) For mangrove species 1= 3000
ca For mangrove-associate species 1
cm
dm Mature species affected by large disturbances (proportion) For mangrove species 0.10.9
da For mangrove-associate species 0.10.9

Mt Fm vm f1
am + bm g1
ca Cm
Mt#+ 1 : A#t + 1 = : 2
Mt Fm vm f1
am + bm g1
ca Cm + At Fa va f1
aa + ba g + St
1 1
cm Ca

Under normal disturbance regime, mangrove species colo-
nize first followed by mangrove associates (Das and Siddiqi
1985), so that colonization of a mangrove-associate species
may be constrained by habitat availability (1 cm), but colo-
nization of a mangrove species is unconstrained. In this con-
dition, propagule losses for both species are approximately
equalbecause proportion of washed-out and predated prop-
agules may be equal for both species (am = aa). There is no dis-
persal barrier of invasive species that could affect propagule
dispersal or seedling establishment of these two species differ-
ently (bm = ba 0). In addition, soil seed bank contribution is
very little, and we can exclude this factor. Under this condi-
tion, the dominance of mangrove species over mangrove
associate will be as follows (equation 3).
Mt Fm vm Cm
Mt + 1 : At + 1 typical = : 3
At Fa va 1
cm Ca
A mangrove species often produce large amounts of propa-
gule, and in a mangrove forest, the abundance of mangrove
species are generally higher than that of mangrove-associate
species. In addition, the numerator in equation 3 is uncon-
strained while denominator is constrained (1
cm). Taken to-
gether, M:A is likely to be >1 and thus mangrove species
dominates over mangrove-associate species.
Vegetation development and recovery following
large disturbances
Large disturbances (cyclones, hurricanes and high flooding)
cause death and damage of mature mangrove (dm) and
mangrove-associate species (da). Only survived adults (1

dm and 1
da) can produce seeds/propagules. Because high
tidal surges associated with large disturbances washout
more propagules than that after normal (low) tidal surges
(Lugo 2000), soil seed bank can be important and species that
possess soil seed bank can have an advantage. Therefore,
following large disturbances, the dominance of mangrove
species over mangrove-associate species can be written as
follows.
Mt Fm vm 1
dm f1
am + bm g1
ca Cm
Mt + 1 : A t + 1 = :
At Fa va 1
da f1
aa + ba g + St
1 1
cm Ca
4
Previous studies have found that large disturbances
cause higher dm than da (Bardsley 1985; Baldwin et al.
1995; Smith et al. 1994), but they did not consider b (prop-
agule blockage by invasive species). We suppose that the
proportion of propagules blocked by invasive species is
higher for mangrove species (because of larger seeds) than
6 Journal of Plant Ecology
mangrove-associate species (bm > ba). If the previous state-
ment holds, then, the portion of habitats colonized by the
mangrove species (cm) is likely to be relatively small and
mangrove-associate species may occupy a higher portion of
vacant habitat for colonization (1
cm). Now the dominance
of mangrove species over mangrove-associate species will be
as follows.
Mt Fm vm 1
dm f1
am + bm g1
ca Cm
Mt + 1 : At + 1 =
fAt Fa va 1
aa + St
1 g1
cm Ca
Note that in equation (5) (under large disturbance), the
numerator is more constrained (dm, bm) than equation (3)
under normal disturbance regime. However, the denomina-
tor is mostly unconstrained or getting a benefit (St
1).
Thus, the values of M:A under this scenario is likely to be
smaller than that in equation (3). If large disturbances are
frequent and mangrove species do not get sufficient time
to recover (Lugo 2008; Smith et al. 2009), there will be few
seed producing adult mangrove species left. In that context,
M:A is likely to be <1 and mangrove-associate species may be
dominating.
In a scenario of mangrove-associate species creating dense
seed dispersal barrier at riverbanks, propagules of mangrove
species may encounter further constraint in reaching potential
colonization sites (French et al. 2011). In other words,
a mangrove-associate species may indirectly limit the coloni-
zation opportunity of a large-seeded mangrove species, and
ultimately, mangrove-associate species may dominate the
vegetation (equation 6).
Mt Fm vm 1
dm f1
am + bm g1
ca Cm
Mt + 1 : At + 1 feedback =
fAt Fa va 1
aa + St
1 gCa
METHODS: A SCENARIO TESTING
EXPERIMENT
We considered a hypothetical initial condition (Table 1) and
estimated proportion of mangrove species in recovered
mangroves in the following year of a large disturbance. We
considered two different scenarios of affected vegetation:
(1) mangrove and mangrove-associate species are equally
affected (dm = da) and (2) mangrove species are two,
three and four times more affected than mangrove-
associate species (dm = 2da, dm = 3da and dm = 4da). Within
each scenario of affected vegetation (d), we considered two
different patterns of invasion barrier (b): (1) equal
blockage to propagules of both species (bm = ba) and (2)
disproportional blockage of propagules of mangrove and
mangrove-associate species (bm = 2ba, bm = 3ba and bm =
4ba). Finally, we computed the proportion of mangrove
species in recovered mangroves when mangrove species
simultaneously experience higher (twofour times) propor-
tion of affected vegetation (d) and higher dispersal barrier
(b) than mangrove-associate species.
RESULTS
When mangrove and mangrove-associate species were
:
equally affected by large disturbances (dm = 2d), mangrove
5 species dominated (proportion of mangrove; >0.90), and this
pattern was consistent for most ranges of dispersal barrier by
invasive species (Fig. 3a). However, when mangrove
species were more affected than mangrove-associate species
(dm > da), the proportion of mangrove species decreased
but remained high enough to maintain the dominance (pro-
portion > 0.5; Figs 3bd). Only in an extreme scenario of 90%
affected mangrove vegetation against 22.5% affected
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mangrove-associate vegetation, the proportion of mangroves
fall below 0.5 (Fig. 3d). Similarly, when both species were
equally affected by large disturbances, disproportionate
dispersal barrier (mangrove propagules experiencing two to
four times more blockages than mangrove associate) could
not change dominance of mangrove species, unless >80%
mangrove propagules were blocked (Fig. 4). However, when
both factors acted simultaneously, i.e. a greater proportion of
affected vegetation (dm > da) and propagule barrier (bm > ba)
for mangroves than mangrove-associate species, a relatively
small disturbance affected the dominance of mangroves
greatly compared to when each of these factors acted alone
(Fig. 5).
:
DISCUSSION
6
So far, the prediction of post-disturbance mangrove vegeta-
tion recovery has been made based primarily on the extent of
physical damage to vegetation and the associated change in
habitat conditions, suggesting that frequent large disturban-
ces markedly alter community structure and composition
(Lugo 2008; Smith et al. 1994, 2009). Our study supports this
general view. But our study for the first time demonstrates
that biological invasion is an important dimension in post-
disturbance succession and must be considered in the predic-
tion of mangrove vegetation development and recovery after
disturbance.
The susceptibility of mangroves to invasion is context
dependent (Foxcroft et al. 2011), and therefore, the relative
importance of biological invasion in mediating vegetation
succession will vary among mangroves. Connectivity of
mangroves to terrestrial or freshwater habitats and the
presence of invasive species in that habitats and anthropo-
genic activities in and around mangroves are examples of
contexts that may confound susceptibility of mangroves to
invasion (Anastasiu et al. 2011; Biswas et al. 2007; Howard
Biswas et al. | Post-disturbance succession in coastal mangroves 7

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Figure 3: proportion of mangrove species in recovered mangroves when vegetation of mangrove and mangrove-associate species are equally
affected (a) and when mangrove species are two (b), three (c) and four (d) times more affected than mangrove-associate species. Different lines
within each plot represent different proportion of propagules blocked by invasive species. Lines are ordered from lowest (top: 0.1) to highest
(bottom; 0.9).

and Harley 1998; Levin et al. 2001). For example, due to
high aquatic-terrestrial connectivity and intense upstream
anthropogenic activities in the Sundarbans, susceptibility
of this forest to invasion is high and a large amount of
invasive species may enter into this forest following large
disturbances (Fig. 1e; Biswas et al. 2007). Although there
are numerous reports on the presence of floating invasive
species in mangroves (Howard and Harley 1998; Lugo
1998; Schmitz et al. 1993), the extent of invasion may not
be as large as in the Sundarbans. The timing of a disturbance
or an invasion event may be another determinant of post-
disturbance invasion: if a disturbance or an invasion event
occurs during monsoon (in the context of tropical climate),
excessive rainfall may lower the hurdle of salinity that an
invasive species otherwise encounters. For example, in the
Sundarbans during the monsoon period, salinity drops close
to fresh water level (Wahid et al. 2007) and the exotic-
invasive species E. crassipes does not experience the usual
saline condition that restricts its invasion. We acknowledge
that well-designed empirical studies on disturbance-medi-
ated patterns and processes of biological invasion and subse-
quent vegetation development are critical for further
verification of our findings.
It is known that physical factors and dispersal barrier
moderate mangrove species distribution at a continental scale
(Duke et al. 1998). Our study implicitly emphasizes the
importance of dispersal barrier and physical factors in
structuring mangrove species at local scale and suggests that
an invasive species by affecting propagule dispersal of
mangrove and non-mangrove species can change post-
disturbance species composition and community dominance
(Harun-or-Rashid et al. 2009; Schmitz et al. 1993). However,
the relative importance of propagule dispersal may vary
among types of succession (primary vs. secondary) (Friess
et al. 2011) and the places of colonization (river mouths vs.
riverbanks; forest edge vs. interior); thus, the role of biolog-
ical invasion in mediating succession is complex and context
dependent.
8 Journal of Plant Ecology

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Figure 4: proportion of mangrove species in recovered mangroves when mangrove propagules are two (b), three (c) and four (d) times more
blocked than propagules of mangrove-associate species. Different lines within each plot represent different proportion of physically affected veg-
etation. Lines are ordered from lowest (top; 0.1) to highest (bottom; 0.9).

Our proposed conceptual model is an enormous simplifica-
tion of a complex reality in the land-sea interfaces and needs
careful interpretation. For example, post-disturbance
hydrologic alteration (Lewis 2005) and spatial and temporal
dimensions of disturbances are important factors in deter-
mining mangrove succession, but we did not consider these
factors here. We believe that consideration of temporal
dimension will amplify rather than reducing the interactive
effects of invasive species affecting mangrove succession if
large disturbances are frequent. It would be worthy to
develop a full dynamic model to predict case specific patterns
in the future.
In conclusion, our study highlights the complexity of
post-disturbance vegetation development and recovery in
coastal mangroves and puts forward an important missing
dimension, biological invasion, into the context. Our results
imply that similar levels of disturbances may have different
consequences on mangrove vegetation recovery in the
presence of an invasive species. These findings have impor-
tant implications for practitioners in mangrove management
and restoration. Depending on the mangrove system and
its susceptibility to invasion, management of invasive
species in terms of removal of propagule dispersal barriers
and seedbed filters such as floating and ground-deposited
debris may be required in concert with management of re-
sidual vegetation (removal of affected vegetation) (Dale
et al. 1998) and/or plantation of native mangrove species af-
ter large disturbances. We cannot overemphasize the need
for empirical studies on the complexity of biological
invasion and its role on mangrove formation. We need this
understanding urgently in order to take informed manage-
ment decisions for the protection of our coastal greenbelts
that are so critical for the livelihood of coastal communities
and their very existence in the face of large disturbances that
are predicted to be increasingly common due to climate
change.
Biswas et al. | Post-disturbance succession in coastal mangroves
Figure 5: contour plots showing proportion of mangrove species in
recovered mangroves when mangrove species encounter two (a),
three (b) and four (c) times more affected vegetation and dispersal bar-
rier simultaneously.
FUNDING
Natural Science and Engineering Research Council of Canada
Research (46483-2009 to A.U.M.).
ACKNOWLEDGEMENTS
We thank Fatima-tuz Zohora and IUCN Bangladesh country office for
providing several photographs used in Fig. 1. The opinions expressed
9
in this paper are those of the authors, and not necessarily the opinions
of the UNEP-WCMC.
Conflict of interest statement. None declared.
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