Late 20th century mangrove encroachment in the coastal Australian monsoon tropics parallels the...

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Late 20th century mangrove encroachment in the coastal Australian monsoon tropics parallels the regional increase in woody biomass

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Grant J Williamson (Corresponding author) School of Plant Science, University of Tasmania, Private Bag 55, Hobart 7001, Australia. Email: grant.williamson@utas.edu.au Phone: +61 3 6226 1944 Fax: +61 3 6226 2698

Guy S Boggs Tropical Spatial Sciences Group, Charles Darwin University, Darwin NT 0909, Australia. Email: guy.boggs@cdu.edu.au

David MJS Bowman School of Plant Science, University of Tasmania, Private Bag 55, Hobart 7001, Australia. Email: david.bowman@utas.edu.au

Date of manuscript draft: 15 September 2009

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ABSTRACT 24

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In Kakadu National Park, a World Heritage property in the Australian monsoon

tropics 250 km to the east of Darwin, a number of recent studies have shown that

woody encroachment (expansion of woody communities) and densification

(increased biomass in woody communities) has occurred in the last 40 years. The

cause of this increase in woody biomass is poorly understood, but possibly associated

with the control of invasive Asian water buffalo, trend to higher rainfall, and

increased frequency of fires. Mangroves provide an important context to understand

these landscape changes given that they are unaffected by fire or feral water buffalo.

We examine change in mangrove distribution in a series of coastal tropical swamps

fringing Darwin, Northern Territory, Australia over a 30-year period using a series of

7 aerial photographs spanning 23 years from 1974 and a 2004 high-resolution satellite

image. In late 1974 Darwin was impacted by an intense tropical cyclone. Vegetation

at 3,000 randomly placed points was manually classified, and a multinomial logistic

model was used to asses the impact of landscape position (coastal, intertidal and

upper-tidal) and swamp on mangrove change between 1974 and 2004. Over the study

period there was instability and slight mangrove loss at the coast, stability in the

intertidal zone and mangrove gain in the upper-tidal zone, with an overall increase in

mangrove presence of 16.2% above the pre-cyclone distribution. A swamp that was

impacted by drainage works for mosquito control and the construction of a sewage

treatment plant, showed a greater mangrove increase than the two unmodified

swamps. The mangrove expansion is consistent with woody encroachment observed

in nearby but ecologically distinct systems. Plausible causes for this change include

changed local hydrology, changes in sea level and elevated atmospheric CO2

concentrations.

KEY WORDS

Mangroves, aerial photography, climate change, coastal ecology, landscape change,

vegetation dynamics, woody vegetation

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INTRODUCTION 53

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In many landscapes throughout the world there is a trend toward encroachment of

woody vegetation into adjoining treeless areas and increased cover of woody

vegetation in wooded communities, sometimes called densification. These processes

have been studied in great detail in Kakadu National Park, a World Heritage property

in the Australian monsoon tropics 250 km to the east of Darwin, using sequences of

orthorectified aerial photography. These studies have revealed that since the 1960s,

monsoon forest patches are expanding into tropical savanna (Banfai et al. 2007;

Banfai and Bowman 2007), tropical savannas are increasing their canopy cover

(Lehmann et al. 2008) and encroachment of fringing woody vegetation is occurring on

treeless freshwater floodplains (Bowman et al. 2008). There remains uncertainty and

debate about the cause of this expansion given that a number of changes have

occurred to potential drivers over the last 40 years. For example, feral Asian water

buffalo underwent a population irruption in the 1970s, which was not brought under

control until the 1990s (Petty et al. 2007). Fire regimes have changed from a

preponderance of fire in the late dry season in the late 1950s to sustained burning

throughout the late dry season (Bowman et al. 2007). Correlative studies have failed

to link the landscape scale woody expansion to either proxies of fire activity or

buffalo density, although these studies show that at the local scale these disturbances

can influence both encroachment and densification (Banfai and Bowman 2007;

Bowman et al. 2008; Lehmann et al. 2008). Collectively, researchers point to the

importance of the recent trend for increasing rainfall, or more controversially,

increased atmospheric CO2, in driving encroachment and densification in Kakadu

National Park.

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In this context, studies of mangrove communities provide an important perspective on

these processes given these systems are unaffected by landscape fire or Asian water

buffalo impacts. Nonetheless, mangroves are highly dynamic systems that respond to

slight changes in sediment accumulation, salinity, water depth and recurrence of tidal

inundation (Bridgewater and Cresswell 1999). For example, mangroves bordering salt

flats can show localized dieback or expansion in response to slight changes in the

hydrology of the system (Duke et al. 1998).

Aerial photography has been shown to be useful for assessing vegetation change over

medium-term time scales (Fensham and Fairfax 2002) and has been used to study

mangrove change (Jones et al. 2004; McTainsh et al. 1986; Saintilan and Wilton

2001). For example, a recent study conducted in Florida with aerial photographs

spanning 60 years has found a dramatic shift in mangrove vegetation up to 3.3km

inland, that appears to be associated with a combination of sea level rise and local

hydrological effects (Ross et al. 2000).

Our aim is to identify changes in mangrove distribution and examine correlates of this

change in the period 1974 to 2004 in three contiguous coastal swamps in the Northern

Territory of Australia. All three swamps were severely impacted by Cyclone Tracy, a

category 4 tropical cyclone that occurred in December 1974 and destroyed the city of

Darwin and damaged the surrounding native vegetation (Stocker 1976). Leanyer

Swamp has been impacted by urbanisation in its catchment and the construction of

drains in the early 1980s in order to increase the drainage of stormwater and to

prevent to pooling of water from high tides, a preferred habitat of saltwater

mosquitoes such as Aedes vigilax (Medical Entomology Section 1983). The other

swamps have been minimally impacted by urbanization. There is some evidence that

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nearby mangrove systems, for example, in the Mary River region (Mulrennan &

Woodroffe 1998) have responded to sea level change over the last 50 years.

Therefore, swamps provide an opportunity to chart mangrove recovery following

cyclone damage, assess the impacts of urbanization and contextualise background

changes associated potentially associated with global change including increased

rainfall, sea level rise and elevated CO2.

To examine changes in mangrove vegetation we use a sequence of high resolution

remotely sensed imagery (7 aerial photographs spanning 23 years that were

georeferenced to a QuickBird satellite image acquired in 2004).

Study Area

The study system comprises three contiguous swamps, Leanyer, Holmes Jungle and

Micket, developed around two tidal creeks, Buffalo Creek and Micket Creek, to the

east of Darwin in the Northern Territory of Australia (12°22´S, 130°55´E, Fig. 1).

This region experiences a monsoonal climate, with 90% of the average 1660 mm

annual rainfall occurring during the wet season from November to April, and a mean

daily maximum temperature of 32C that varies little throughout the year. The study

covers an area of approximately 30 km2 with a catchment area of 35 km2. The area

has a low elevation gradient and a maximum tidal range of close to 8 m, with the tide

reaching over 4 km from the coast along the tidal creeks at high tide. Vegetation

along each swamp is dominated by a gradient in salinity and water availability,

ranging from freshwater reed swamps, to brackish reeds and sedges, to mangrove

forest, with grassland and salt flats more distant from the tidal creeks. Comparable

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vegetation patterns, in nearby Howard’s Swamp, has been described by Wilson and

Bowman (1987).

MATERIALS AND METHODS

Aerial photographs taken at low tide and during the austral winter dry season (May to

August), covering the study area for the years 1974, 1976, 1979, 1983, 1985, 1990,

and 1997, were obtained from the Northern Territory Department of Planning and

Infrastructure. A high-resolution Quickbird Satellite image, also captured at low tide,

was used for the 2004 coverage. The photographs obtained included colour, black

and white, and colour-infrared prints for various years. The photographs were

registered to the 2004 satellite image in ArcMap 9.1, using a 3rd-order polynomial

transform, with RMS error values of around five metres. The 2004 satellite image

was manually digitised and classified into polygons representing a set of eight

vegetation or landscape units at a scale of 1:3000 following ground reconnaissance.

The creek network was digitised using the 2004 satellite image. The aerial

photographs were all taken on the low tide cycle thus enabling manual mapping of the

coastal line as evidenced by boundary between sand and mud flats.

Elevation data in the form of a digital contour map was obtained from the Northern

Territory Government, but it was found that the 1 m intervals were too coarse to

adequately represent the slope of the salt flat and low-lying mangrove areas. A field

survey was conducted using a differential GPS system at 120 randomly placed points

in the low-lying salt pan and brackish swamp areas, to fill in gaps in the contour map

to a vertical resolution of under 10cm. A digital elevation model with a cell size of

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100 m was produced from the combined data using ordinary kriging with a spherical

semi-variogram model using the Spatial Analysis extension in ArcGIS 9.1.

A total of 3,000 sample points were randomly placed across the vegetation map and a

number of habitat attributes were derived from the 2004 vegetation map and

preceding sequences of aerial photography. For each aerial photographic coverage the

points were manually classified into one of the 8 vegetation/landscape units by

examining the vegetation type inside circles of 10 m diameter at each point. For the

purposes of the analysis a generic mangrove formation was scored as present if > 50%

cover of the mangrove vegetation unit was observed at a point. Data for the elevation,

distance to creek, and distance to coast were spatially joined to the point samples.

Following field surveys on the ground and by helicopter during different times in the

tidal cycle, and following advice from the Medical Entomology Branch of the

Northern Territory Government who conduct frequent assessments of tidal extent for

the purposes of mosquito control, the swamps were a priori classified into three

landscape zones (coastal, intertidal, and upper tidal) based on distance from coast and

vegetation type and the sample points were assigned to these three landscape position

zones. While we recognize a continuous gradient in tidal and saline influence exists,

these three zones tend to represent distinct vegetation associations and tidal regimes.

The sample points were recoded as gain, loss or stable mangrove based on the

difference in mangrove presence at the point between 1974 and 2004, and a

multinomial logistic model, performed in R 2.8.1 (R Development Core Team 2008),

was used to determine the effect of swamp and landscape position on mangrove

change.

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Darwin tidal gauge and rainfall data used in interpretation was obtained from

Australian National Tidal Centre and the Australian Bureau of Meteorology

respectively.

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RESULTS 173

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A vegetation map and map of point classification into landscape position zones is

presented in figure 1. There was clear zonation in vegetation types from from the

coast inland. In the coastal zone the elevation was around 6 m Australian Height

Datum rising up to around 7 m in the upper tidal zone some 3 km inland (Table 1).

Distance from creek lines increases across this gradient from 100 to 300 m in the

coast and intertidal zones to almost 1 km in the upper-tidal zone (Table 1). The

coastal zone is dominated by mangrove vegetation, the intertidal zone by salt flats,

and the upper-tidal zone has the highest proportion of grass, sedge and reed vegetation

(Table 2). Vegetation patterns are related to salinity gradients and frequency and

depth of tidal inundation. For example, in the coastal zone mangroves (including

Bruguiera parviflora, Rhizophora stylosa and Avicennia marina) are dominant,

occupying areas subject to frequent tidal inundation. Mangroves also extend through

the intertidal zone along tidal creeks although much of this zone is comprised of

sparsely vegetated salt flats, that due to hypersaline conditions and extreme

evaporation rates only support mangrove vegetation close to frequently flushed creeks

The upper-tidal zone comprises a salinity gradient from low mangrove forests

dominated by Ceriops tagal, Lumnitzera racemosa, Avicennia marina and Sonneratia

lanceolata, through brackish Schoenoplectus littoralis reeds to Eleocharis dulcis in

areas subject primarily to freshwater inundation.

Across the 3,000 sample points, there was an overall increase in mangrove presence

of 16.2% between 1974 and 2004, amounting to an increase of approximately 123 ha

from the 1974 coverage of 761 ha. The upper-tidal zone showed a consistent increase

in mangrove cover over the 30 year study period, and little mangrove loss after the

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1974 cyclone. In contrast the coastal zone showed some mangrove loss over the study

period, particularly following the cyclone (Figure 2). The intertidal zone in Micket

and Holmes Jungle swamps showed little change in total mangrove area at the end of

the study period, though an increase in this zone was seen in Leanyer swamp that was

modified by drainage works (Figure 2).

From the beginning to end of the study period, the negligible transitions away from

mangrove vegetation were mostly to open water or salt flat (Table 3). The transition

matrix shows a large proportion of the points transitioning to mangrove vegetation

were originally brackish Schoenoplectus sedgelands characteristic of the upper tidal

zone, and fewer mangrove transitions were from salt flats or open water. In the upper-

tidal zone of Leanyer swamp, in particular, Eleocharis and Schoenoplectus

sedgelands, sample points showed a significant conversion to mangroves as a result of

drain construction for mosquito control (Table 2).

The swamp x landscape position interaction models of mangrove increase and

decrease between the start and end of the study period showed a trend towards greater

conversion to mangrove in the upper tidal zone (Figure 3a) and greater loss of

mangroves at the coastal zone (Figure 3b). There was a statistically significant

difference between swamps (p < 0.05), with Leanyer swamp having particularly high

rates of conversion to mangrove, and low rates of conversion away from mangrove.

Mean sea level and rainfall (Figure 4) in this region have fluctuated during the study

period, and although a number of years stand out as exceptional, no trend can be

discriminated at the available temporal resolution.

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DISCUSSION 219

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Despite a catastrophic cyclone in 1974 there has been a significant increase in the

coverage in mangrove distribution in the swamps adjacent to Darwin over the 30-year

study period. Immediately following the tropical cyclone, mangrove cover declined,

although there was recovery within 20 years back to the original distributional area of

the mangroves. There was both loss and gain of mangrove cover in the coastal zone

over the study period, consistent with the continuously shifting substrates found

previously in this system (Woodroffe and Grime, 1999). In contrast the upper-tidal

zone mangroves underwent expansion and densification, replacing treeless brackish

vegetation and, to a lesser extent, the hypersaline flats. The contrasting response of

mangroves with different positions along the elevation gradient from the coast to the

inland reflects the geomorphologies of the landscape settings: the coastal zone is

highly dynamic while in the intertidal zone favours more stable vegetation patterns

given the relatively fixed salinity gradients between creeklines and treeless saltflats

and floodplains (Hollins and Ridd 1997), unless impacted by drainage works.

Increased sedimentation as the result of urbanization and changing precipitation

regimes are often cited as drivers of mangrove expansion globally (Schwarz 2003,

Alongi 2005, Jupiter et al. 2007). The significantly different patterns of mangrove

increase in Leanyer swamp appear to be related to hydrological changes caused by

drainage works. This swamp was subject to the construction of a sewage treatment

plant, disrupting the flow of fresh and tidal water, and the construction of a system of

drains for mosquito control (Department of Construction and A.A.Heath 1978),

which allowed more rapid drainage of the swamp after heavy rainfall or high tides

(Medical Entomology Branch 1982). The pattern of mangrove expansion in Leanyer

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around the drains is visible in the satellite imagery (Figure 5), a change that has also

been observed in mosquito control channels in eastern Australia (Breitfuss et al. 2003;

Jones et al. 2004). Harty (2004) notes that urbanization, with the associated increased

nutrient and sediment loads and changes to hydrology through engineering works can

also promote mangrove establishment consistent with our finding for Leanyer swamp.

However, the fact that the two swamps not subject to human influence also showed

strong trends in mangrove in the upper-tidal zone signals some profound changes to

the ecology of these swamps.

At the boundary between temperate and subtropical climates, changing temperatures

and reduced frequency of frosts have been proposed as a mechanism for mangrove

expansion into treeless saltmarsh communities in the United States (Stevens et al.

2006). But changes in temperature are unlikely to impact significantly on mangrove

communities in northern Australia, which experiences a tropical climate with no

frosts. Herbert (2007) suggests that mangrove expansion into saltmarsh habitat at the

landward fringe in the Hunter estuary in New South Wales is due to increased tidal

range resulting from the construction of tidal barriers and channel dredging.

However, such engineering works have not taken place in the region of the Darwin

study, and any alteration in mean sea level or tidal amplitude is presumed to be the

result of global change rather than local anthropogenic impacts.

Elevated atmospheric CO2 concentration has frequently been proposed as a driver of

the expansion of woody species at the expense of non-woody vegetation (Bazazz

1990; Archer et al. 1995; Bond and Midgley 2000; Eamus and Palmer 2007), as

higher CO2 concentrations are expected to increase photosynthetic rate and water use

efficiency more in woody plants than non-woody plants. Mangrove species have

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shown responses in growth to elevated CO2 (Farnsworth et al. 1996) although this is

mediated by salinity. Controlled mangrove growth experiments under elevated CO2

have found little growth enhancement in high-salinity conditions but more growth

enhancement in low-salinity conditions in a less salt-tolerant species (Ball et al.

1997). This effect could lead to expansion into brackish or freshwater areas, as we

have observed in this system. However, controlled environment experiments growing

mangroves and saltmarsh species alone and in competition under elevated CO2 found

that competition reduced the impact of elevated CO2 on mangrove growth suggesting

other factors may be required to explain mangrove expansion (McKee and Rooth,

2008). Without further experimental data we can’t reject the CO2 fertilizer effects as a

plausible contributor to of the mangrove expansion. However, experiments

comparing the growth of the two invading mangrove species at the study site

(Avicennia marina and Sonneratia lanceolata) with and without the presence of

dominant sedge (Schoenoplectus littoralis) under elevated CO2 are required to more

fully evaluate the importance of the putative fertilizer effect in this system. A further

question arises as to the time scale over which any CO2 fertilization effect will be

apparent. Simulations of forest dynamics in response to elevated CO2 in temperate

forests have found significant increases in basal area over a time span of 50-150 years

(Bolker et al. 1995). Free-air carbon enrichment (FACE) experiments have

demonstrated increased carbon flux and photosynthesis in forests in response to CO2

enrichment, an effect that appears generally stronger in trees and C3 species, but

structural changes in woody ecosystems have generally not been observed over the

period of time these studies have been running (Ainsworth and Long 2005). It is

possible rapid-growing, disturbance tolerant mangrove communities may be capable

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of showing a rapid response to CO2 enrichment, and the application of FACE

experiments to these systems would be informative.

The discontinuous landward expansion and seaward contraction is broadly correlated

with corresponding fluctuations of sea level over the study period. Therefore this

study shows general agreement with previous studies qualitatively linking mangrove

distribution changes with trends in sea-level rise (McTainsh et al. 1986; Ross et al.

2000) or that have been based on analysis of aerial photographs (Alongi 2008;

Dowling 1978; Gilman et al. 2007). Like our study, these latter authors found

mangrove invasion of brackish swamps at the limit of the inland tidal extent, and a

coincident loss of seaward mangroves. An obvious explanation for the landward

expansion of mangroves is the increased landward penetration of seawater (Lacerda et

al. 2007). Simulation modelling of mangrove habitat based on projected sea-level in

Florida over the next 100 years showed significant replacement of freshwater marsh

and swamp environments by mangroves (Doyle et al. 2003), and a similar outcome

was predicted for Homebush Bay, New South Wales based on fuzzy set modelling

(Zeng et al. 2007). In our study, direct time-series modelling of mangrove

distribution against sea level or rainfall trends was not possible due to the low number

of available repeat images and fluctuating tidal gauge data over the last decades

(Church et al. 2006). Such variability combined with the sporadic sequence of

imagery makes it difficult to directly link mangrove dynamics to sea level rise,

although saline intrusion, possibly as the result of sea-level rise, is a clear driver of

inland mangrove expansion in nearby areas (Woodroffe and Mulrennan 1993; Bell et

al. 2001; Winn et al. 2006).

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Northern Australia is showing a trend in increasing rainfall, particularly during the

wet season (Smith 2004, Taschetto and England 2009), although this trend is not

evident in the mean annual rainfall data obtained for the years of this study (Figure 4).

Increased rainfall is expected to lead to increased growth rates in mangroves (Ellison

2000). Periods of high rainfall are also expected to contribute to mangrove

establishment in the hypersaline flats due to dilution of the salts in the soil (Field

1995; Duke 1997; Saintilan and Williams 1999; Harty 2004). However, expansion

into hypersaline salt flats is limited at our study site (Table 3). Indeed the major area

of mangrove expansion was into brackish swamps that are inundated by the highest

spring tides in October (at the end of the dry season) before being filled by freshwater

in the wet season. This suggests mangrove expansion into these brackish areas is

driven by increasing rather than decreasing salinity.

The expansion of mangroves observed in these coastal swamps is temporally and

spatially consistent with the woody densification and expansion observed in nearby

but ecologically distinct savannah and rainforest ecosystems. Densification in these

systems has often been attributed to changes in fire regime or feral animal populations

(Banfai et al. 2007, Bowman et al. 2008), factors that should not affect the mangroves.

Conversely, mangrove expansion is often attributed to sea-level rise, a factor unable

to influence inland forests. Little is known about the two potential effects of those

environmental drivers shared by both coastal mangrove forests and inland forests:

atmospheric CO2 concentration and changing rainfall patterns. Studies examining

mangrove tree growth across a wider area of northern Australia, so local differences

in substrate types, disturbance history and rainfall can be statistically evaluated would

be a useful future direction for research, as would paired comparison of mangrove and

rainforest tree growth within local climatic zones.

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Conclusion 339

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Despite severe damage from a tropical cyclone the coastal swamps examined in this

study all show landward mangrove expansion over a 30-year period, primarily

replacing brackish reed swamps in the upper tidal zone. Expansion rate is particularly

high in the swamp that has undergone hydrological changes as the result of human

engineering works, but given the expansion is also occurring in the other swamps,

direct human interference alone cannot be established as the cause. While the

observed changes are similar to those expected to be seen with sea-level rise, this

cannot be confirmed as the primary driver of change given the fragmentary aerial

photographic record. The mangrove expansion is consistent with densification trends

in other ecologically distinct ecosystems in northern Australia, including savannas and

rainforests, suggesting regional-scale factors are driving woody expansion. Plausible

candidates for this change include changed local hydrology, changes in sea level and

elevated atmospheric CO2 concentrations.

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ACKNOWLEDGEMENTS

Funding for this project was provided by the Australian Research Council (ARC)

Linkage Grant (No. LP0667619), Northern Territory Department of Health and

Community Services, Australian Bureau of Meteorology, Northern Territory

Research and Innovation Fund, Australian Department of Defence and Charles

Darwin University. The authors would like to thank the Medical Entomology Branch

of the Northern Territory Health Department, especially Peter Whelan for expertise

and assistance, the Northern Territory Department of Planning and Infrastructure for

assistance with spatial data and equipment, and Lubomir Bisevac and Dimity Boggs

for assistance in the field.

REFERENCES

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Table 1. Mean and standard deviation of environmental variables for three landscape zones in the three swamps. Elevation is based upon Australian Height Datum, creek distance is the distance from nearest tidal creek, shore distance is the nearest distance to the low tide point.

Swamp Coastal Intertidal Upper-tidal Elevation Leanyer 6.95 ± 0.47 6.92 ± 0.48 7.46 ± 0.15(meters) Holmes Jungle 6.51 ± 0.31 6.92 ± 0.21 7.05 ± 0.29 Micket 6.62 ± 0.36 6.92 ± 0.70 7.40 ± 0.43Creek Distance Leanyer 0.32 ± 0.22 0.27 ± 0.25 0.91 ± 0.51(kilometers) Holmes Jungle 0.19 ± 0.14 0.43 ± 0.33 0.66 ± 0.49 Micket 0.18 ± 0.14 0.07 ± 0.06 0.53 ± 0.40Shore Distance Leanyer 0.65 ± 0.37 1.78 ± 0.39 3.58 ± 0.59(kilometers) Holmes Jungle 0.57 ± 0.34 2.04 ± 0.67 4.04 ± 0.74 Micket 0.62 ± 0.37 2.19 ± 0.50 3.48 ± 0.43

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Table 2. Change in the percentage of all vegetation/landscape units for the three swamps broken down by coastal, intertidal and upper-tidal zones between 1974 and 2004. Salt flat includes all vegetation-free areas inundated by the highest tides. Typha, Eleocharis, and Schoenoplectus are graminoid dominated vegetation types that fall along a gradient of increasing salinity, while the grass/sedge category comprises the remaining treeless vegetation. Woodland comprises all non-mangrove woody vegetation, including savanna and coastal thicket.

Swamp Position Salt Flat % Water % Grass/Sedge % Eleocharis % Typha % Schoenoplectus % Mangrove % Woodland % Holmes Jungle Coast 20.8 → 28.3 0.5 → 0.5 3.7 → 2.7 0.0 → 0.0 0.0 → 0.0 0.0 → 0.0 72.2 → 64.2 2.7 → 4.3 1974 → 2004 Inter 58.4 → 71.2 1.5 → 0.3 14.8 → 6.2 0.0 → 0.0 0.0 → 0.0 0.0 → 0.0 24.9 → 21.4 0.3 → 0.9 Upper 21.2 → 18.0 0.0 → 0.0 41.7 → 39.0 9.9 → 8.7 0.9 → 1.7 5.2 → 2.6 20.3 → 28.2 0.9 → 1.7 Leanyer Coast 32.7 → 28.4 0.5 → 1.1 5.2 → 6.1 0.0 → 0.0 0.0 → 0.0 0.0 → 0.0 54.7 → 58.4 6.9 → 6.1 1974 → 2004 Inter 44.4 → 36.2 0.3 → 0.6 14.3 → 7.7 0.0 → 0.0 0.0 → 0.0 0.0 → 0.0 35.2 → 51.0 5.7 → 4.6 Upper 13.8 → 27.1 0.0 → 0.0 26.9 → 44.8 17.8 → 0.0 0.0 → 0.0 28.4 → 0.0 5.5 → 22.7 7.6 → 5.4 Micket Coast 8.2 → 10.6 8.8 → 15.3 0.0 → 0.0 0.0 → 0.0 0.0 → 0.0 0.0 → 0.0 82.9 → 74.1 0.0 → 0.0 1974 → 2004 Inter 37.5 → 32.9 3.9 → 5.4 3.9 → 4.9 0.0 → 0.0 0.0 → 0.0 0.0 → 0.0 51.9 → 54.2 2.8 → 2.6 Upper 19.8 → 16.7 0.0 → 0.0 30.6 → 30.6 14.4 → 14.9 0.0 → 0.0 14.0 → 6.8 11.7 → 24.8 9.5 → 6.3

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Table 3. Vegetation transition matrix, showing proportion of points in each category converting to other categories between 1974 and 2004. Vegetation/landscape units are as in Table 2.

2004 Grass/Sedge Mangrove Salt Flat Schoenoplectus Woodland Eleocharis Water Urban Typha

Grass/Sedge 71.0% 4.2% 20.6% 0.0% 3.5% 0.5% 0.0% 0.2% 0.0%Mangrove 1.0% 87.7% 7.2% 0.2% 0.5% 0.0% 3.2% 0.1% 0.0%Salt Flat 5.0% 18.7% 75.4% 0.1% 0.5% 0.0% 0.1% 0.2% 0.0%Schoenoplectus 9.4% 44.1% 28.3% 12.6% 0.8% 4.7% 0.0% 0.0% 0.0%Woodland 21.5% 10.3% 4.7% 0.0% 57.0% 5.6% 0.0% 0.9% 0.0%Eleocharis 27.0% 7.0% 13.9% 4.3% 1.7% 42.6% 0.0% 0.9% 2.6%Water 0.0% 33.3% 10.3% 0.0% 0.0% 0.0% 56.4% 0.0% 0.0%Urban 0.0% 10.0% 0.0% 0.0% 30.0% 0.0% 0.0% 60.0% 0.0%

1974

Typha 0.0% 0.0% 0.0% 0.0% 0.0% 0.0% 0.0% 0.0% 100.0%

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130°58'0"E130°57'30"E130°57'0"E130°56'30"E130°56'0"E130°55'30"E130°55'0"E130°54'30"E

12°20'30"S

12°21'0"S

12°21'30"S

12°22'0"S

12°22'30"S

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VegetationSalt Pan / Bare

WaterWoodland

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Brackish SedgeFreshwater Reed

Leanyer

HolmesJungle

Micket

0 1 20.5 km

LandscapePosition

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Intertidal

Upper-tidal

Darwin

Figure 1. Map of 2004 vegetation communities in the study. The coastal, inter-tidal and upper-tidal zones used in the analysis are shown. The zones were defined in respect of tidal regimes and vegetation types (Table 2).

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0.0 0.2 0.4 0.6 0.8

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1975 1980 1985 1990 1995 2000 2005

0.0 0.2 0.4 0.6 0.8

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Leanyer Holmes Jungle Micket

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grov

e P

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rtion

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Figure 2. Proportion of sample points with mangroves over time for the coastal, intertidal and upper-tidal zones across the three swamps for all intervals between 1974 and 2004.

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0

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LeanyerHJMicketTotal

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0

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LeanyerHJMicketTotal

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Figure 3. Mean proportion of points showing mangrove increase (a) and decrease (b) between 1974 and 2004. Error bars indicate standard deviation reported by multinomial logistic regression.

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Figure 4. Preceding 5-year mean sea level and mean annual rainfall for study intervals in Darwin.

3900

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1974 1976 1979 1983 1985 1990 1997 2004

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n Se

a Le

vel (

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)

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1974 1976 1979 1983 1985 1990 1997 2004

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ainf

all (

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Figure 5. Change in extent of mangrove vegetation in the upper-tidal areas of the three swamps between 1974 and 2004.

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