Biogeosciences, 11, 857–871, 2014

Open Access
www.biogeosciences.net/11/857/2014/
doi:10.5194/bg-11-857-2014 Biogeosciences
© Author(s) 2014. CC Attribution 3.0 License.

Mangroves facing climate change: landward migration potential in

response to projected scenarios of sea level rise
D. Di Nitto1 , G. Neukermans1 , N. Koedam1 , H. Defever1 , F. Pattyn3,4 , J. G. Kairo5 , and F. Dahdouh-Guebas1,2
1 Biocomplexity Research Focus c/o Laboratory of Plant Biology and Nature Management, Mangrove Management Group,
Vrije Universiteit Brussel – VUB, Pleinlaan 2, 1050 Brussels, Belgium
2 Laboratoire d’Écologie des Systèmes et Gestion des Ressources, Département de Biologie des Organismes, Faculté des

Sciences, Université Libre de Bruxelles – ULB, CP 169, Avenue F.D. Roosevelt 50, 1050 Brussels, Belgium
3 Laboratory of Physical Geography, Vrije Universiteit Brussel, Pleinlaan 2, 1050 Brussels, Belgium
4 Unité de Recherche Sciences de la Terre,Université libre de Bruxelles, Brussels, Belgium
5 Kenya Marine and Fisheries Research Institute, P.O. Box 81651, Mombasa, Kenya

Correspondence to: D. Di Nitto ([email protected])

Received: 21 December 2012 – Published in Biogeosciences Discuss.: 25 February 2013

Revised: 29 September 2013 – Accepted: 24 October 2013 – Published: 13 February 2014

Abstract. Mangrove forests prominently occupy an intertidal 1 Introduction

boundary position where the effects of sea level rise will be
fast and well visible. This study in East Africa (Gazi Bay,
Kenya) addresses the question of whether mangroves can be Inhabiting the interface between land and sea, mangroves
resilient to a rise in sea level by focusing on their potential are amongst one of the most at-risk ecosystems when
to migrate towards landward areas. The combinatory analy- sea level rises (McLeod and Salm 2006). Throughout the
sis between remote sensing, DGPS-based ground truth and Quaternary, mangroves have shown high resilience to dis-
digital terrain models (DTM) unveils how real vegetation ruptions from large sea level fluctuations over historic
assemblages can shift under different projected (minimum timescales (Woodroffe 1990). However, adaptation probabil-
(+ 9 cm), relative (+ 20 cm), average (+ 48 cm) and maxi- ities strongly depend on the rates of sea level rise (SLR) and
mum (+ 88 cm)) scenarios of sea level rise (SLR). Under sediment supplies in combination with subsurface processes
SLR scenarios up to 48 cm by the year 2100, the landward that affect sediment elevation (Gilman et al., 2007; Gilman
extension remarkably implies an area increase for each of the et al., 2006; McLeod and Salm 2006; Wolanski and Chappell
dominant mangrove assemblages except for Avicennia ma- 1996; Woodroffe 1990). Ellison and Stoddart (1991) sug-
rina and Ceriops tagal, both on the landward side. On the one gested that mangroves are stressed by SLRs of between 9
hand, the increase in most species in the first three scenar- and 12 cm over 100 yr and concluded that faster rates could
ios, including the socio-economically most important species seriously threaten mangrove ecosystems. This view has been
in this area, Rhizophora mucronata and C. tagal on the sea- challenged by Snedaker et al. (1994), who cited historical
ward side, strongly depends on the colonisation rate of these records showing changes nearly twice that high in mangrove
species. On the other hand, a SLR scenario of + 88 cm by the expansion under relative sea level; however hard scientific
year 2100 indicates that the area flooded only by equinoctial data or SLR simulations are not available.
tides strongly decreases due to the topographical settings at As mangrove ecosystems are very dynamic, the ability of
the edge of the inhabited area. Consequently, the landward these forests to migrate to more landward zones is a very im-
Avicennia-dominated assemblages will further decrease as a portant aspect when considering the effect of SLR on man-
formation if they fail to adapt to a more frequent inundation. groves. If the possibility presents itself, mangroves will ad-
The topography is site-specific; however non-invadable areas just to a SLR by expanding landward or laterally into ar-
can be typical for many mangrove settings. eas of higher elevation, or even by growing upward in place
(McLeod and Salm 2006). However, the mangrove areas that

Published by Copernicus Publications on behalf of the European Geosciences Union.

858 D. Di Nitto et al.: Mangroves facing climate change

are most vulnerable are those situated in a physiographic set- Nagelkerken et al., 2008; Walters et al., 2008; Wells et al.,
ting that limits landward migration due to obstacles or steep 2006).
gradients and with a net decrease in sediment elevation or This study focuses on the critical factor “tidal range” in
sediment accretion that is insufficient to keep up with SLR order to investigate the potential for landward migration of
(Gilman et al., 2008). Landward obstructions, artificial or mangrove vegetation assemblages in Gazi Bay (Kenya) un-
natural, have an effect on ecosystems that would normally der different SLR scenarios. As mangrove species have their
move landward in response to erosive forces. Where there preferred hydroperiod, the vegetation distribution in the dif-
is a rise in sea level relative to the land, a coastal squeeze ferent inundation classes at present is extrapolated towards
takes place (Doody, 2004). On the species level, adaptation future SLR scenarios based on a static mangrove surface el-
can occur through landward migration at different speeds as evation. Digital terrain modelling is derived from differen-
mangrove species maintain their preferred hydroperiod or by tial GPS field measurements and used to simulate water lev-
sediment accretion (Gilman et al., 2008). Mangrove species els in a GIS environment. In combination with a mangrove
composition can strongly affect a mangrove’s resistance and species map, preliminary results are generated regarding the
resilience to SLR given that, on the one hand, individual effect of SLR in the study site in Gazi Bay (Kenya). The
species have varying tolerances of the period, frequency and focus resides on individual mangrove species and their pos-
depth of inundation, and, on the other hand, different vege- sible colonisation of back-mangrove areas that become ac-
tation zones have different rates of change in sedimentation cessible when sea level rises. We deliberately adopt a re-
elevation (Krauss et al., 2003; McKee et al., 2007; Rogers et ductionistic approach by taking abstraction of alterations
al., 2005). Furthermore, several scientists have also investi- in sedimentation and elevation and other consequences of
gated how different functional root types of several mangrove global change such as increases in temperature, CO2 con-
species respond to changes in elevation in order to determine centration and storm frequency, as well as possible shifts in
the vulnerability to SLR (Ellison and Stoddart 1991; Vin- seasonal periods (Pernetta, 1993; UNEP, 1994; Woodroffe,
cente 1989). 1990; Woodroffe and Grime, 1999). However we feel that, in
Species-specific competition may allow some species to this context, relevant conclusions can be made. First of all,
outcompete others and to become more dominant within the this study represents the first attempt to simulate the effect of
newly formed species composition (Lovelock and Ellison, SLR based on a large amount of detailed information on to-
2007). Establishment and dispersal play a significant role pography and vegetation covering the whole bay. Secondly,
in these processes. They are, however, different for various many researchers have already gathered valuable informa-
species and strongly dependent on many biotic factors such tion within this study area on diverse subjects like regen-
as buoyancy, period of obligate dispersal, longevity and pe- eration, vegetation structure dynamics, human impacts and
riod of establishment (Allen and Krauss, 2006; Clarke et al., propagule dispersal (e.g. Abuodha and Kairo, 2001; Bosire et
2001; Drexler, 2001; Tomlinson, 1986), whilst wind and hy- al., 2003, 2008b; Dahdouh-Guebas Farid and Koedam, 2006;
drodynamics of tides and currents can be equally important Dahdouh-Guebas et al., 2002a; Di Nitto et al., 2008; Kairo et
abiotic factors (Stieglitz and Ridd, 2001). Additionally, fac- al., 2001; Kirui et al., 2008; Neukermans et al., 2008). The
tors like microtopography, the soil type at the top and root latter gives us the opportunity to draw preliminary conclu-
structures can also have a significant effect on the fate of sions on the potential for landward migration of mangroves
propagules once released from their parental tree, as can in Gazi Bay and to create some views on the future vegetation
human-induced degradation, like tree cutting (Di Nitto et al., structure dynamics, which can contribute to their resilience
2008). to SLR. Resilience is understood here as the survival of the
To date, mangroves have been subjected to non-climate- formation, even if displaced in space.
related anthropogenic stressors which have accounted for
most of the global average annual rate of mangrove loss,
estimated to be 1–2 %, with losses during the last quarter 2 Material and methods
of a century ranging from 35 to 86 % (Alongi, 2002; Duke
et al., 2007; FAO 2003, 2007; Valiela et al., 2001). So far, 2.1 Study area
relative SLR has been a smaller threat to mangroves. How-
ever, it may constitute a substantial proportion of predicted Gazi Bay (4◦ 260 S, 39◦ 300 E) is a shallow tropical water sys-
losses (about 10–20 % of total estimated losses) as several tem situated circa 40 km south of the historic port (Kilin-
studies have already shown that many mangrove areas have dini) of Mombasa (Fig. 1). The mangrove forest covers an
not been keeping pace with current rates of relative SLR (Ca- area of approximately 6.5 km2 and is drained by two tidal
hoon et al., 2006; Gilman et al., 2007; McKee et al., 2007). creeks. The tidal regime within the bay is semi-diurnal with
We would like to emphasise the importance of understand- a macro tidal range of 3.5 m and an ebb-dominant asym-
ing mangrove responses to SLR as these ecosystems provide metry (Kitheka 1996, 1997). Ten East African mangrove
tremendous social, economic and ecological value (Barbier, species are present within this bay fringed with mangrove
2003; Dahdouh-Guebas et al., 2005; Mumby et al., 2004, forests, seagrass beds and coral reefs, more specifically Avi-

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D. Di Nitto et al.: Mangroves facing climate change 859

Fig. 1. Representation of (A) the Kenyan coast (Dahdouh-Guebas et al., 2000) and (B) Gazi Bay. The satellite image (QuickBird) shows the
whole bay of Gazi ; however this research focuses on the western part as encompassed by the overlaid vegetation map. S. alba = Sonneratia
alba, R. mucronata = Rhizophora mucronata, C. tagal Lw = Ceriops tagal on the landward side, C. tagal Sw = Ceriops tagal on the seaward
side A. marina Lw = Avicennia marina on the landward side and A. marina Sw = Avicennia marina on the landward side. Classification of
the mangrove species coverage was obtained by Neukermans et al. (2008).

cennia marina (Forsk.) Vierh., Bruguiera gymnorrhiza (L.) R. mucronata and C. tagal (Sw and Lw)) are mapped with an
Lam., Ceriops tagal (Perr.) C. B. Robinson, Heritiera lit- overall accuracy (OA) of 72 %.
toralis Dryand., Lumnitzera racemosa Willd., Rhizophora
mucronata Lam., Sonneratia alba Sm., Xylocarpus grana- 2.2 Topographical field survey and construction
tum Koen, a second yet unidentified Xylocarpus species, of a DTM
and Pemphis acidula Forst. (Gallin et al., 1989) (nomencla-
ture according to Tomlinson, 1986). Topographical measure-
The aim of the topographical field surveys was to construct
ments (see Sect. 2.2) were conducted throughout the western
a digital terrain model (DTM) in order to simulate water lev-
part of the bay during two dry periods (July–August 2003
els at present and for different Intergovernmental Panel on
and 2005).
Climate Change (IPCC) scenarios of SLR (for explanation
Mangrove species distribution within this study area was
on IPCC scenarios, see Sect. 2.3). Measurements were car-
obtained by Neukermans et al. (2008). A classification of
ried out using a Leica GPS-AT302, which is a centimetre-
a Standard QuickBird multispectral satellite image was per-
precise differential global positioning system (DGPS) with
formed in combination with ground truthing based on vegeta-
a fixed reference station and a mobile rover station. Since
tion transects by the point-centred quarter method (PCQM+)
a dense mangrove cover disrupts the DGPS signal, a strati-
of Dahdouh-Guebas and Koedam (2006). The two socio-
fied design was applied targeting the low-cover mangroves,
economically most important species within this study area,
back-mangrove areas, tidal mudflats and creeks. Resolution
R. mucronata and C. tagal (Dahdouh-Guebas et al., 2000,
of the DTM varies from 1m in the topographically “rough”
2004a), are mapped with user’s accuracies above 85 %,
areas to 50 m in areas characterised by a relatively flat and
whereas all four dominant mangrove species (A. marina (on
even surface. All DGPS points were post-processed in SKI
the seaward side (Sw) and the landwards side (Lw)), S. alba,
(Static Kinematic Program), and after converting these geo-
graphical coordinates into projected coordinates (WGS 1984,

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860 D. Di Nitto et al.: Mangroves facing climate change

UTM zone 37S) and assigning their absolute height, a thor- not available; however we do not underestimate the impor-
ough knowledge of the field was used to add extra points tance of sediment in mangrove vegetation dynamics in view
and breaklines in order to eventually optimise the constructed of SLR.
DTM. As the height measurements of these points are rel- The modelling exercise started with an assessment of the
ative, we followed the high-water line of a chosen spring current species-related zonation or spatial structure present
tide on two consecutive days and collected the xyz data of in Gazi Bay. First of all, the height boundaries for each in-
116 points using the DGPS. Based on the Kilindini tide ta- undation class according to Watson (1928) (Table 1) was de-
bles (Kenya Ports Authority, KPA) the approximated abso- fined based on the combination of the tide tables (July 2003–
lute height of the water was calculated, and the relative ele- July 2004) published by the KPA and the monthly inunda-
vations in the DTM converted to approximate absolute field tion frequencies per class (Table 1). In further analysis, in-
topography. We recognise a temporal delay in tides between undation frequencies higher than those of “class 1” will be
Mombasa and Gazi Bay; however this does not influence our referred to as “class 0”. Using ArcGIS 8.2, these boundaries
study. were classified into inundation classes based on the DTM for
The final coordinates resulting from the topographical the current scenario versus different IPCC scenarios of eu-
measurements were inserted into a geographical information static SLR. The relative scenario of + 20 cm coincides with
system (GIS) and served as an input to create a triangular the current trend of SLR within the long-term data set (1985–
irregular network (TIN) of the area. The TIN was based on 2003) obtained from gauge measurements by the Kenya Ma-
the (non-constrained) Delauney triangulation of the original rine and Fisheries Research Institute at the Kilindini Port in
set of points by use of Voronoi diagrams, a theory for which Mombasa. This initiative is part of the Global Sea Level Ob-
we refer to Raper (1990). In this paper it is not the intention serving System (GLOSS) founded by the Intergovernmental
to investigate in depth the impact of these elevation errors Oceanographic Commission (IOC) of UNESCO.
through principal component analysis (Lopez, 1997), but we Secondly, an overlay between the vegetation map and the
give an estimation of the absolute mean error and the stan- current inundation classes (Fig. 2b) gives an estimation of
dard deviation in densely covered and less densely covered the vegetation surface of each species within each inunda-
areas. After extracting 30 points from each of the latter areas, tion class. To review the accuracy of the DTM and/or the
the TIN was reconstructed and height values were reassessed classification of the vegetation, it is important to investigate
for these particular points. whether the distribution of the species within the inunda-
tion classes deviate from a random distribution. To perform
2.3 Spatial analyses the statistical analyses, the complete area was divided into
10 equally sized blocks. Within each block the areal cover-
IPCC has predicted several SLR scenarios (+ 9 cm (mini- age (ha) was calculated for each species in all inundation
mum), + 20 cm (relative), + 48 cm (average) and + 88 cm classes of the current situation. Secondly, a Kolmogorov–
(maximum)) by the year 2100 (IPCC 2001)1 based on Smirnov test was performed to compare the observed cumu-
atmosphere–ocean general circulation models and emission lative distribution function to a theoretical normal distribu-
scenarios incorporating uncertainties regarding changes in tion, whereafter Kruskal–Wallis tests were completed to in-
terrestrial ice, permafrost and sediment deposition. The main vestigate whether the vegetation distribution within the inun-
purpose of the spatial analyses is to predict possible changes dation classes is random. Since the species concerned are not
in vegetation assemblages under these different scenarios of randomly distributed, extrapolations of the vegetation struc-
SLR. ture towards future IPCC scenarios of SLR were performed.
This modelling exercise mainly focuses on the potential of The area increase (%) of each inundation class within each
mangroves to migrate towards landward areas, but it is solely scenario was calculated in relation to the current situation,
based on sea level rise relative to a static mangrove surface whereafter these percentages were multiplied by the current
elevation. In this stage, data on sediment-related changes are vegetation area (ha).
1 We based our analysis on SLR scenarios of the IPCC Third
2.4 Sensitivity analysis
Assessment Report (TAR) (2001) and not on those of the Fourth
Assessment Report (AR4) (2007), which forecast a range from 9 to A source of uncertainty in the input data is the DTM’s abso-
88 cm by 2100 and a range from 18 to 59 cm by 2090–2099, respec- lute height, which was calibrated using Kilindini port gauge
tively. The reason is the following: due to lacking of published liter-
measurements. To address the sensitivity of the model to the
ature, AR4 models do not include uncertainties in climate–carbon-
cycle feedback, nor do they include the full effects of changes in ice
absolute height uncertainty of the DTM, we investigated the
sheet flow. The AR4 projections do, however, include a contribu- impact of changes in the height boundaries of the inunda-
tion due to increased ice flow from Greenland and Antarctica at the tion classes. Upper and lower height boundaries are slightly
rates observed for 1993–2003, but these flow rates could increase or altered at a time and in a systematic manner, more specifi-
decrease in the future. The AR4 could have similar ranges to those cally by an increase and decrease of these boundary intervals
of TAR if uncertainties were to be treated in the same way. with 5, 10 and 15 % corresponding to 4, 6 and 8 cm. The

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D. Di Nitto et al.: Mangroves facing climate change 861

Table 1. Inundation classes and monthly inundation frequency according to Watson (1928). Height boundaries (metres above datum) of
present and future inundation classes are presented: a minimum (+9 cm), relative (+20 cm), average (+48 cm) and maximum (+88 cm)
scenario is based on IPCC eustatic SLR scenarios for the year 2100. In further analysis, inundation frequencies higher than those of “class
1” will be referred to as “class 0”.

Inundation Flooded by Monthly Present Minimum Relative Average Maximum

classes inundation situation scenario, scenario, scenario, scenario,
frequency (m) +9 cm (m) +20 cm (m) +48 cm (m) +88 cm (m)
1 All high tides (AHT) 56–62 2.10–2.60 2.19–2.69 2.30–2.80 2.58–3.08 2.98–3.48
2 Medium high tides (MHT) 45–56 2.60–3.10 2.69–3.19 2.80–3.30 3.08–3.58 3.48–3.98
3 Normal high tides (NHT) 20–45 3.10–3.50 3.19–3.59 3.30–3.70 3.58–3.98 3.98–4.38
4 Spring high tides (SHT) 2–20 3.50–3.80 3.59–3.89 3.70–4.10 3.98–4.28 4.38–4.68
5 Abnormal (equinoctial tides) (EHT) 0–2 3.80–4.20 3.89–4.29 4.10–4.40 4.28–4.68 4.68–5.08

10

11

12

13

14

Figure 2
Fig. 2. (A) Presentation of the DTM. (B) 3-D presentation of the combination between (B1) inundation classes, (B2) vegetation map and
(B3) QuickBird image. (C) Presentation of the inundation classes: (C1) current situation, (C2) scenario +9 cm, (C3) scenario +20 cm, (C4)
scenario +48 cm and (C5) scenario +88 cm. AHT stands for all high tides; MHT, medium high tides; NHT, normal high tides; SHT, spring
high tides; and EHT, equinoctial high tides. Inundation frequencies higher than those of “class 1” will be referred to as “class 0”.

comparison between the reference map (Fig. 2C1) and the 3 Results
output maps after altering the height boundaries was assessed
with an error matrix, giving overall (OA), user’s (UA) and 3.1 Construction and validation of the digital
producer’s accuracies (PA) (for calculations, see Appendix terrain model
A).
The DTM of the study area is shown in Fig. 2a. After post-
processing in SKI, 4105 points were accepted with an aver-
age error on x, y and z of 1.16, 2.08 and 0.89 cm, respec-
tively, whereafter several breaklines and 82 extra points were
manually added to optimise the DTM. Breaklines along the
creek banks are, however, crucial and had to be added as

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862 D. Di Nitto et al.: Mangroves facing climate change

Table 2. Presentation of the total area (ha) occupied by each man-

grove species with the whole studied area (TMA) and the total area
occupied by each mangrove species within the inundation classes at
present (TMAI). Sw = seaward side and Lw = landward side.

Species TMA TMAI Difference (%)

Avicennia marina Sw 46.59 41.65 11.86
Avicennia marina Lw 32.34 29.53 9.52
Ceriops tagal Lw 37.99 36.53 3.99
Ceriops tagal Sw 13.50 13.27 1.76
Rhizophora mucronata 109.42 105.50 3.71
Sonneratia alba 7.96 4.95 60.89

estimates (based on measurements within the creek) due to

high mangrove coverage. Absolute mean error and standard
deviation for densely covered and less densely covered areas
are 0.013 m ± 0.106 and 0.089 m ± 0.374, respectively.

3.2 Simulation of sea level rise scenarios

The current situation covers a total (studied) area of

423.43 ha, of which the regularly flooded area and the non-
flooded area encompass 386.53 and 36.90 ha, respectively.
When looking at the inundation classes within the different
scenarios (Fig. 2C1 to C5), we can conclude that there is an
overall trend of transgression into the terrestrial areas. The
maximum scenario in particular (+ 88 cm) represents a sig- Fig. 3. (A) Graph of the total area (ha) per species within the
nificant area increase of class 0 and class 1 (AHT) (for ab- four SLR scenarios, (B) future prediction of the dynamics of man-
breviations, see Fig. 2c). More specifically, the percentage groves, non-flooded area and class 0.
area increase of these two classes from the current situation
towards the maximum scenario of SLR is 245 and 103 %,
respectively. After calculating the extent of each mangrove between TMA and TMAI of 61 %. The high discrepancy be-
species within each current inundation class, Kolmogorov– tween TMA and TMAI for S. alba could be explained by a
Smirnov tests were completed with results showing signif- possible lower accuracy of the DTM at the breaklines mark-
icance values < 0.05 for each species. The vegetation dis- ing the creek bank.
tribution is therefore not normal and nonparametric tech- All other species appear to have an adequate distribution
niques have to be used for further analyses. The following within the whole study area: Avicennia marina Sw (seaward)
Kruskal–Wallis test proved that the distribution of the vege- mainly resides in class 1 (AHT) (26 %) and class 2 (MHT)
tation within the inundation classes is not random; all signif- (45 %), Rhizophora mucronata mainly appears in class 2
icance values are < 0.05. Each species evaluated within the (MHT) (53 %) and class 3 (NHT) (22 %), whilst Ceriops ta-
area has a preference for certain inundation classes, confirm- gal, which is an inner mangrove, occupies the areas in several
ing the occurrence of a specific zonation or spatial structure mid-classes. A. marina Lw (landward) dominates the land-
in Gazi Bay and therefore also an adequate accuracy of the ward classes with 35 % in class 4 (SHT). An extrapolation of
field measurements. changes in vegetation assemblages towards future scenarios
Due to the errors on the classification of the vegetation (Fig. 3a) demonstrates that, in comparison to the average sce-
map (see Sect. 2.1 and Neukermans et al., 2008) and the to- nario of SLR (+ 48 cm), all species will decrease in the max-
pographical measurements, the total area (ha) occupied by imum scenario (+ 88 cm), resulting in a decline of 13 % in
each mangrove species within the whole study area (TMA) 100 yr. Although most species show a possible area increase
does not fully coincide with the total area (ha) occupied throughout the minimum, relative and average scenario, this
by each mangrove species within the inundation classes at is not the case for A. marina Lw, as this species will dimin-
present (TMAI). This, however, does not exceed values be- ish throughout all scenarios, with a highest decrease of 60 %
tween 2 and 12 (Table 2), except for Sonneratia alba, which in the maximum scenario. When considering the two socio-
mainly occurs in class 0 (38 %) and class 1 (AHT) (35 %), economically most important species R. mucronata and C.
consequently being the only species with a high difference tagal in the most probable relative scenario of + 20 cm SLR,

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D. Di Nitto et al.: Mangroves facing climate change 863

an area increase of 15 % occurs in comparison to the cur- dient from land to sea, extrapolations can be made to areas
rent situation. Finally, the area proportions between the total with similar characteristics.
mangrove area, the non-flooded area and class 0 are shown
in Fig. 3b as percentage increase or decrease compared to 4.1 Vegetation dynamics of mangrove assemblages
the current situation. The maximum scenario shows a con- under different scenarios of SLR
siderable decrease in total mangrove area of 13 %, whereas
for the relative scenario this area increases by 4 %. The most Bearing in mind the reductionistic approach, the extent of
marked increase is for the area of class 0, namely 245 % in the most common assemblages, apart from Avicennia marina
comparison to the current situation. and Ceriops tagal on the landward side (Lw), are forecasted
to increase in surface under the different scenarios of SLR
3.3 Sensitivity analysis and error matrix for map (except for the maximum scenario of + 88 cm). This forecast
comparison or accuracy assessment is in line with a few earlier reports that current sea level rise
rates do not pose a threat to mangrove ecosystems (e.g. Mc-
Table B1 (see Appendix B) shows the results of the error Kee et al. 2007; Snedaker et al., 1994; Tan and Zhang, 1997),
matrices for map comparison or accuracy assessment. When but contradicts many others (e.g. Ellison and Stoddart, 1991;
comparing the vegetation distribution within adjusted height Fujimoto and Miyagi 1990; Parkinson et al., 1994; Pernetta,
boundaries for each inundation class, the outcome appears to 1993). However, considering the uncertainties regarding the
be relatively sensitive to an increase or decrease of 15 %. The impact of global change on mangrove growth and develop-
overall accuracy, with a comparable outcome for Khat , fluc- ment, such contradictions are not unexpected. In addition,
tuates between 87.34 and 65.88 % when considering an in- our reductionistic approach focuses on tidal range and the
crease or decrease up to 10 %, yet strongly declines towards possible dispersal range of propagules, but it does not take
53.61 to 48.02 % when height boundaries of each inundation into account the biogeomorphological capacity to maintain
class are adjusted by 15 %. As the applied vegetation classifi- or to protect a mangrove forest.
cation confirms the occurrence of a specific zonation or spa- Landward migration of mangroves in Gazi Bay appears to
tial structure in Gazi Bay, which is highly related to inunda- be limited under the maximum scenario as the highest in-
tion patterns, we can conclude that sensitivity to alterations in tertidal inundation class strongly decreases due to the topo-
topography can be significant from a certain limit and should graphical settings at the edge of the inhabited area. Conse-
therefore be aligned to vegetation distributions when data are quently, the coastal squeeze will signify a decrease in the Avi-
available. Furthermore, the wind setup may have affected the cennia-dominated assemblages if they fail to adapt to a more
high-water-line measurements at spring tide. In addition, the frequent inundation or if competition with other species pre-
inundation classification according to Watson (1928), which vails. Dahdouh-Guebas et al. (2004a) made a prediction of
is based on inundation frequency, may not always yield fully future vegetation structure in Gazi Bay based on retrospec-
satisfactory results, especially in regions with an irregular tive remote sensing, social surveys and tree distribution, and
tidal regime and/or irregular elevation profile, where the du- results show that the surface extent of A. marina on the land-
ration of inundation seems equally important, as was shown ward side has been reducing since 1972. Furthermore, the
by Van Loon et al. (2007). current situation in Gazi Bay is characterised by large bare
and sandy sites on the landward side which have remained in
the same state for a substantial time; that is, we have observed
4 Discussion no colonisation for at least 16 yr). When landward areas are
accessible during SLR, dispersal and early growth become
This study was to investigate whether mangrove assemblages important stages in a plant life that fundamentally determine
in Gazi Bay have the potential to migrate to more land- community structure and population dynamics (Clarke et al.,
ward areas, which can contribute to their resilience to SLR 2001; Sousa et al., 2007). These processes are very complex.
(Fig. 4), understood as the survival of the formation within A dense mangrove forest can provide an adequate propagule
the site. Although the focus of this study was mainly on tidal supply for dispersal towards newly colonisable areas, but (1)
range, we emphasise the importance of sediment supply, es- as Clarke et al. (2001) stated, establishment of young trees
pecially for scenarios of SLR higher than 20 cm 100 yr−1 is mainly related to the presence of parental trees, while this
(relative scenario). Whether mangroves can be resilient to is not so much the case for juveniles and the hydrochorous
SLR strongly depends on the physiographic setting in which dispersal of propagules, and (2) suitability for stranding or
these ecosystems occur, human activities that are carried out self-planting of propagules is strongly dependent on the pres-
in the wetland and on how species-specific competition and ence of root structures (which can facilitate the entanglement
adaptation will unfold. There is no clear-cut answer that can of propagules) and the compactness of the soil (clay- or silt-
be applied to global mangrove coverage; however, by study- dominated) (Di Nitto et al., 2008).
ing this particular mangrove area with a mesotidal regime As in other transitional systems, plant establishment and
and a common vegetation zonation along a gentle slope gra- community succession is driven by tolerance to physiological

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864 D. Di Nitto et al.: Mangroves facing climate change

Fig. 4. Overview scheme summarising the discussion on resilience of mangroves facing sea level rise, more specifically concerning the case
study in Gazi Bay (Kenya).

stress and plant–plant interactions (Bertness, 1991; Mil- The reported forecasts can also have an important socio-
brandt and Tinsley, 2006); hence species-specific competi- ecological implication. Although the forest adjacent to the
tion could signify a natural blockage for landward migra- village has long been overexploited for wood and decreased
tion of mangroves. However, in several cases, facilitation is in area, anthropogenic disturbance has diminished over the
a common mechanism of succession in terrestrial habitats, last years and some mangrove assemblages have even ex-
meaning that an early coloniser changes the abiotic condi- panded (Dahdouh-Guebas et al., 2004a). An increase in man-
tions in a way that allows an entry and finally a displacement grove area under different scenarios of SLR, provided that it
of a second species to a previous intolerable habitat (Con- does not go at the expense of qualitative degradation, may
nell and Slayter, 1977). This was, for instance, the case for imply an increase in anthropogenic threats such as tradi-
(1) saltwort (Batis maritima L.), as it was identified as an tional utilisation (McLeod and Salm, 2006). Clear felling of
abundant initial coloniser of an extensive black mangrove mangroves species can have severe consequences for future
(Avicennia germinans L.) die-off area (Milbrandt and Rins- vegetation dynamics. Furthermore, most mangrove creeks
ley, 2006), and (2) salt marsh cordgrass (Spartina alterni- (like the case in Gazi Bay) are characterised by the occur-
flora Loisel.), being a potential initial soil stabiliser creating rence of time–velocity asymmetry in which ebb flow is more
successional stages firstly for Laguncularia racemosa (L.) dominant than flood flow (Kitheka 1997, 1998; Kitheka et
C. F. Gaertn, which is secondly outshaded and replaced by al., 2002). Sediment trapping occurs during incoming flood
Avicennia schaueriana Stapf and Leechm. ex Mold. (Cunha- tides, and there is no significant export of sediments during
Lignon et al., 2009). ebb tide (Furukawa and Wolanski, 1996; Wattayakorn et al.,
1990); however degradation of mangroves can lower trap-
ping efficiency (Kitheka et al., 2002), consequently increas-
ing vulnerability to sea level rise.

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D. Di Nitto et al.: Mangroves facing climate change 865

Fig. 5. Decision tree to aid resilient site selection for mangroves according to McLeod and Salm (2006). This three can be applied once
candidate sites of high biodiversity have been selected using biological criteria, for instance *, factors that indicate strong recovery potential
(see Appendix Table C1). This decision tree was adjusted (**) to implement the possibility of a shift in vegetation structure. MPA = marine
protected area.

4.2 Vegetation dynamics of individual species under tation zones that are daily inundated and are never submitted
different scenarios of SLR to large salinity variations (Tomlinson, 1986). When sea level
rises, this species is forecasted to increase in area (except un-
When landward areas become accessible for the migration der the maximum scenario), yet as investigated by Dahdouh-
and colonisation of mangrove species, we have to ask the Guebas et al. (2004a), the juvenile layer within these S. alba
same question as Alongi (p4, 2008): “Are trends in man- stands is limited and propagule establishment is hampered
grove forest replacement in response to catastrophic distur- by currents that are generally known to be strongest along
bances the result of somewhat deterministic sequences as in the seaward side (Diop et al., 2001). The distribution of the
terrestrial forests, or are they the result of a stochastic “first young individuals of S. alba is more related to the adult
come, first served” opportunistic response or neither?” Em- trees, whereas juveniles are generally spread over a wider
pirical data support the idea that recovery is stochastic with area (Dahdouh-Guebas et al., 2004a). The latter also ap-
distinct succession stages, yet early sequences of species re- plies for the species Avicennia marina on the seaward side
placement are greatly influenced by species present at ini- (Sw). Furthermore, Imai et al. (2006) verified that S. alba
tial recovery (Alongi, 2008; Clarke et al., 2001; Sousa et al., seedlings and saplings, which require sunny conditions for
2007). Within this study the extrapolation of the present veg- their growth, were more abundant in gaps than in the under-
etation distribution towards scenarios under a rising sea level storey. Competition with a more landward species such as
is based on species-specific preference for certain inunda- Rhizophora mucronata might demonstrate that an area in-
tion frequencies. The survival of these species in their shift crease of S. alba could be overestimated by our analyses.
in a more landward direction is strongly dependent on their However, colonisation by S. alba on seaward sand banks
colonisation rate and interspecific competition. The most has occurred throughout the years. Additionally, bearing in
seaward mangrove species Sonneratia alba appears in vege-

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866 D. Di Nitto et al.: Mangroves facing climate change

mind the site-specific rates of sea level rise and sediment et al., 1998; McKee, 1995), yet research is needed on in-
input rates, Ellison and Stoddart (1991) claimed that man- terspecies interactions influencing mangrove forest regener-
grove ecosystems can keep pace with SLR of 8–9 cm per ation in post-disturbance mangrove communities.
100 yr, making seaward expansion and colonisation of these
daily inundated areas possible. Rates of 9–12 cm per 100 yr 4.3 Recommendation for further research and
will cause stress, yet the ability for mangroves to adjust to management strategies
even higher rates is unlikely. The minimum scenario of SLR
(+ 9 cm) could in fact provide an additional and suitable In the light of mangrove ecosystem stresses caused by cli-
habitat for S. alba and A. marina (Sw). mate change, managers face the dual challenge of selecting
R. mucronata and Ceriops tagal are two economically and implementing conservation strategies in order to main-
valuable pioneer species that will most likely increase as pre- tain and restore resilient mangrove forests.
dicted unless anthropogenic impact rises. Multivariate vege- In this study the emphasis resides on tidal range and not
tation structure analysis showed that C. tagal is very abun- on sediment supply; however, we give a preliminary vulner-
dant in the understorey of assemblages dominated by other ability assessment of this mangrove area based on a slightly
mangroves, which could mask a dynamic shift (Dahdouh- adjusted decision tree (Fig. 5) to aid resilient site selection
Guebas et al., 2004a). R. mucronata and C. tagal already for mangroves by McLeod and Salm (2006). This decision
occupy the mid-zone within the mangrove area and knowl- tree was applied after appointing Gazi Bay as a high biodi-
edge on the dispersal of their propagules indicates that prop versity candidate site based on biological and environmental
roots and pencil roots clearly have the ability to entangle criteria (Table C1; see Appendix C). Decisions were made
propagules and that preference of propagule dispersal goes based on the available literature involving the mangrove area
to flat areas and substrates with a more compact soil struc- in Gazi Bay and the relative SLR scenario of + 20 cm, which
ture (clay, silt) (Di Nitto et al., 2008). One disadvantage for coincides with the current trend along the coast of Kenyan.
R. mucronata could, however, be represented by a further sil- Following this decision tree, the mangrove area in Gazi
tation along the seaward sand bank creating a patch of arid Bay appears to be adequately resilient for at least 100 yr
conditions and higher light intensity more favourable for A. and can most likely be appointed as a marine protected area
marina (Dahdouh-Guebas, et al. 2004a). (MPA). However, although we do not intend to focus only
Avicennia marina (Lw) will have to adapt to greater in- on MPAs, we want to anticipate a future scenario of sea
undation frequencies. It is known that this species can toler- level rise and indicate gaps in, on the one hand, scientific
ate high salinity variation, so could the double zonation of and, on the other hand, site-specific knowledge that neces-
this species on the landward side versus the same species sitate further research. Given (1) the mesotidal regime and
on the seaward side support the idea of dynamic adaptation? permanent rivers and creeks that provide freshwater and sed-
Genetic analyses based on 48 RAPD (randomly amplified iment (mainly during wet season), (2) the knowledge that the
polymorphic DNA) loci have demonstrated that four DNA drainage basin of both Mkurumuji and Kidogoweni rivers,
fragments show a slight differentiation in allelic frequency which extend into the coastal ranges of the Shimba Hills Na-
between the two A. marina stands in spite of their short dis- tional Reserve, has limited anthropogenic pressures with re-
tance separation (Dahdouh-Guebas et al., 2004b). This in- spect to the intactness of the hydrological regime, and given
dicates that there is less genetic exchange between the dis- that (3) landward migration in Gazi Bay is possible under the
junctive stands than within one stand, consequently suggest- relative scenario of sea level rise, the decision tree leads us
ing that an ecological or physical barrier might exist. Tidal towards the question whether recruitment is strong. The an-
range might facilitate the dispersal of propagules in both di- swer is definitely “yes”; however we feel that the possibility
rections; however obstruction by complex root structures can of a shift in vegetation structure needs to be implemented,
prevent this exchange. Additionally, interspecific competi- rendering Gazi Bay a site that is “Maybe OK for MPA”. Ac-
tion with the adjacent species C. tagal could disadvantage A. cording to McLeod and Salm (2006) the decision tree would
marina as McCusker (1977) confirms that a salinity increase have led towards “Good choice for MPA”.
causes a reduction in water use efficiency for the seedlings The recommendations for further research and manage-
of Rhizophora, but not for Ceriops. Furthermore, an elevated ment strategies, which can be applied globally, are the fol-
CO2 level will enhance the efficiency of water use (UNEP, lowing: (1) identifying an early coloniser to promote early
1994); however this advantage is lost when salinity becomes establishment of mangrove seedlings, (2) measuring changes
too high, for instance, at low inundation frequency areas on in elevation by means of surface elevation tables (SETs), (3)
the landward side. Another drawback for A. marina is an in- assuring the possibility of landward migration and (4) inves-
crease of temperature, since this species has lowest optimal tigating propagule dispersal by combined hydrodynamic and
temperature for leaf development (Hutchings and Saenger, ecological behaviour modelling.
1987).
There are several well-established physiologic mecha-
nisms influencing mangrove community composition (Duke

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D. Di Nitto et al.: Mangroves facing climate change 867

Appendix A Table B1. Results of the error matrices for map comparison or accu-
racy assessment when comparing the vegetation distribution within
Calculation of an error matrix for map comparison or adjusted height boundaries for the inundation classes. Values repre-
accuracy assessment sent the overall accuracy and Khat in percentages.

– Map 1 is a raster grid of n classes as a model output Input Adjustments Comparison of vegetation distributions
parameter in input criteria within the adjusted inundation classes
– Map 2 is a raster grid of n classes from an alternative Overall accuracy Khat
model or comparison reference layer. (%) (%)
Inundation +5% 87.34 85.34
Producer’s accuracy (PA) classes +10% 76.67 75.3 1
Producer’s accuracy (PA): Takes into account the accuracy (height +15% 48.02 49.61
of individual classes and therefore indicates the probability boundaries) –5 % 78.09 75.28
of the cell value in Map 2 being the same as in Map 1. –10 % 65.88 65.39
–15 % 53.61 50.72
PA = xii /x+i · 100 %
N = total number of cells in the error matrix.
xii = total number of correct cells in a class
x+i = sum of cell values in the column
Khat : Measure of agreement or accuracy based on KAPPA
User’s accuracy (UA): Takes into account the accuracy of in- analysis to compare maps of similar categories in order to
dividual classes but indicates the probability of the cell value determine if they are significantly different
in Map 1 being the same as in Map 2. Pr Pr
!
i=1 xii − i=1 (xi+ · x+i )
UA = xii /xi+ · 100 % Khat = N
N 2 − ri=1 (xi+ · x+i )
P

xii = total number of correct cells in a class
r = number of rows in the matrix
xi+ = sum of cell values in the row
xii = total number correct cells in a class
Overall Accuracy (OA): Summarizes the total agree- (i.e. value in row i and column i)
ment/disagreement between the maps and incorporates the xi+ = total for row i
major diagonal while excluding the omission and the com-
x+i = total for column i
mission errors.

OA = D/N · 100 %
D = total number correct cells as summed along
the major diagonal
N = total number of cells in the error matrix

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868 D. Di Nitto et al.: Mangroves facing climate change

Table C1. Mangrove resilience factors that contribute to site selection (according to McLeod and Salm, 2006). Case study: Gazi Bay, Kenya.

Factors that allow for peat building to keep up with sea level rise Applicable to Gazi Bay Yes/No Literature available per factor
Association with drainage systems including permanent rivers and Yes e.g. Dahdouh-Guebas et al.
creeks that provide freshwater and sediment (2004a), Kitheka (1996, 1997),
Njambuya (2006), Obade et al.
(2004), Ohowa et al. (1997)
Sediment-rich macrotidal environments to facilitate sediment redistri- Yes
bution and accretion
Actively prograding coast and delta Yes
Natural features (bays, barrier islands, beaches, sandbars, reefs) that re- Yes
duce wave erosion and storm surge
Factors that allow for landward migration
Mangroves backed by low-lying retreat areas (for example, salt flats, No/Yes in certain places e.g. Di Nitto et al. (2008), Neuk-
marshes, coastal plains) which may provide suitable habitat for coloni- ermans et al. (2008), Obade et al.
sation and landward movement of mangroves as sea level rises (2004)
Mangroves in remote areas and distant from human settlements and Yes
agriculture, aquaculture, and salt production developments
Mangroves in areas where abandoned alternate land use provides op- Yes, unmanaged coconut plantations
portunities for restoration, for example, flooded villages, tsunami-prone
land, unproductive ponds
Factors that enhance sediment distribution and propagule dispersal
Unencumbered tidal creeks and areas with a large tidal range to im- Yes e.g. De Ryck (2009), Di Nitto
prove flushing, reduce ponding and stagnation, and enhance sediment et al. (2008), Kitheka (1996,
distribution and propagule dispersal 1997), Ohowa et al. (1997)
Areas with a large tidal range that may be better able to adjust to in- Yes
creases in sea level due to stress tolerance
Permanent strong currents to redistribute sediment and maintain open Yes
channels
Factors that indicate survival over time
Diverse species assemblage and clear zonation over range of elevation Yes e.g. Beeckman et al. (1989),
(intertidal to dry land) Bosire et al. (2008a), Bosire et al.
(2006), Dahdouh-Guebas et al.
(2002a), Dahdouh-Guebas et al.
(2004a), Dahdouh-Guebas et al.
(2002b), Kairo (2001), Kairo
et al. (2001), Neukermans et al.
(2008), Tack et al. (1992), Van
Tendeloo (2004)
Range in size from new recruits to maximum size class (location and Yes
species dependent)
Tidal creek and channel banks consolidated by continuous dense man- Yes
grove forest (which will keep these channels open)
Healthy mangrove systems in areas which have been exposed to large No
increases in sea level due to climate-induced sea level rise and tectonic
subsidence
Factors that indicate strong recovery potential
Access to healthy supply of propagules, either internally or from adja- Yes
cent mangrove areas
Strong mangrove recruitment indicated by the presence, variety, and Yes
abundance of established mangrove propagules
Close proximity and connectivity to neighbouring stands of healthy Yes
mangroves
Access to sediment and freshwater Yes
Limited anthropogenic stress Yes, no major residential area in the
vicinity, selected as a fairly pristine
East African site in the EU PUMPSEA
project: http://www.pumpsea.icat.fc.ul.
pt/main.php
Unimpeded or easily restorable hydrological regime Yes
Effective management regime in place such as the control of usual Yes
threats like dredging and filling, conversion to aquaculture ponds, and
construction of dams, roads, and dikes that disrupt hydrological regime,
etc.
Integrated coastal management plan or protected area management plan Yes/No
implemented

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D. Di Nitto et al.: Mangroves facing climate change 869

Acknowledgements. Many thanks are due to the people of Gazi tory traits correlate with patterns of adult distribution?, J. Ecol.,
Bay, more specifically Latifa S. Ba’alawy and her relatives for the 89, 648–659, 2001.
hospitable family environment and R. Abdul for the assistance on Connell, J. H. and Slayter, R. O.: Mechanisms of succession in nat-
the field. This research was funded the Flemish Interuniversity ural communities and their role in community stability and orga-
Council (VLIR) and the Fonds David & Alice Van Buuren. D. D. is nization, Am. Natural., 111, 1119–1144, 1977.
the recipient of a VLIR PhD scholarship. This work was presented Cunha-Lignon, M., Mahiques, M. M., Schaeffer-Novelli, Y., Ro-
in part at (1) the International Symposium of Aquatic Vascular drigues, M., Klein, D. A., Goya, S. C., Menghini, R. P., To-
Plants (ISAVP) (January 11–13, 2006, Brussels, Belgium) (2) the lentino, C. V., Cintrón-Molero, G., and Dahdouh-Guebas F.:
7th International Symposium on GIS and Computer Cartography Analysis of mangrove forest succession using cores: a case study
for Coastal Zone Management (CoastGIS) (July 12–16, 2006, in the Cananéia-Iguape Coastal System, São Paulo, Brazil, Braz.
Wollongong, Australia) and the (3) MMM3 Meeting on Mangrove J. Oceanogr., 57, 2009.
ecology, functioning and Management (2–6 July 2012, Galle, Sri Dahdouh-Guebas, F. and Koedam, N.: Empirical estimate of the
Lanka). reliability of the use of the Point-Centred Quarter Method
(PCQM): Solutions to ambiguous field situations and description
Edited by: B. Satyanarayana of the PCQM+ protocol, Forest Ecol. Manage., 228, 1–18, 2006.
Dahdouh-Guebas, F., Mathenge, C., Kairo, J. G., and Koedam,
N.: Utilization of mangrove wood products around Mida Creek
(Kenya) amongst subsistence and commercial users, Economic
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