Journal of Arid Environments 92 (2013) 14e25

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Journal of Arid Environments

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Review

Soil salinity: A neglected factor in plant ecology and biogeography

E.N. Bui*
CSIRO Land and Water, GPO Box 1666, Canberra, ACT 2601, Australia

a r t i c l e i n f o a b s t r a c t

Article history: This paper argues that soil salinity needs to be more broadly acknowledged as a driving factor in plant
Received 11 June 2011 ecologydnot only in the ecology of halophytesdin order to understand and make more accurate pre-
Received in revised form dictions for the impact of environmental change on biodiversity and vegetation patterns throughout the
13 August 2012
semi-arid world. It summarizes recent research on soil salinity and plant distributions in semi-arid
Accepted 31 December 2012
Available online 11 February 2013
environments throughout the world: there is empirical as well as experimental evidence that soil sal-
inity, even at low levels, is an abiotic stress factor that influences vegetation patterns and diversification.
Lines of evidence demonstrating salinity’s potential influence as a selective agent in East Africa and North
Keywords:
Acacia
America are presented. The paper then synthesizes recent results from spatial ecology, plant and insect
Evolutionary ecology systematics and behavioral ecology, focusing on Australia, that support a role for salinity in evolutionary
Halophytes ecology of Acacia. On a shorter time scale, soil salinity may play a role in weed invasion and woody
Non-halophytes vegetation encroachment in Australia.
Speciation Crown Copyright Ó 2013 Published by Elsevier Ltd. All rights reserved.

1. Introduction
Recently there has been much activity related to predicting the
impact of climate change on biodiversity, nevertheless the question
of what determines the distribution of species, one of the most
basic in ecology, is still unanswered for most of Australia and much
of the rest of the world (Austin and Van Niel, 2011; Hughes, 2003;
Kier et al., 2005). Five categories of factors that determine distri-
bution and abundance of plant species are recognized (Mueller-
Dombois and Ellenberg, 1974): climatic; edaphic; geographic-
historic factors; species interactions; and perturbations. These
factors also govern the composition and structure of plant com-
munities. They also act as evolutionary stress factors.
That climatic and topographic data are not always sufficient to
explain vegetation patterns is evident (e.g., Austin and Van Niel,
2011; Bertrand et al., 2012; Bui and Henderson, 2003; Dirnbock
et al., 2002; Martin et al., 2006; Reed et al., 2009). Yet even when
edaphic factors are taken into consideration by plant ecologists and
biogeographic modelers, soil moisture/texture, pH, and nutrients
are generally the only variables considered. Although soil salinity is
recognized as a major limitation to cropping (e.g., Lauchli and
Luttge, 2002; US Salinity Laboratory, 1969; Zhu, 2001) and the
focus of many plant breeding efforts to produce salt-resistant crops
(e.g., Flowers et al., 2010; Lauchli and Luttge, 2002; Parida and Das,
2005), generally, it is neglected as a factor in plant ecology, except
* Tel.: þ61 2 6246 5935.
E-mail address: elisabeth.bui@csiro.au.
in the ecology of deserts (salt pans and playas) and wetlands (salt
marshes, mangroves) where its importance is obvious given the
prevalence of halophytes.
This paper first considers definitions of saline soils and halo-
phytes, and reviews the role of soil salinity in plant ecology and
geography around the world but then focuses on Australia. It
presents new evidence that salinity plays a role at the macro-level,
in the diversification of the genus Acacia, with more than 900
native Australian species. Australia is one of the most biologically
unique areas of the world, with one of the most extensive arid
zones; natural salinity is widespread, with extensive playa lake
systems and saline and sodic soils (Northcote and Skene, 1972;
Rengasamy, 2006).
1.1. What is soil salinity?
The definition of saline soil is confusing.1 Soils are considered
saline if they contain salt in a concentration sufficient to interfere
with the growth of most crop species. Saline soils have an electrical
conductivity >4 dS m
1 (w36 mM NaCl) measured on a saturated
soil paste extract at 25  C (US Salinity Laboratory, 1969). Electrical
1
Soil scientists distinguish saline soils with an electrical conductivity >4 dS m
1,
pH < 8.5, and Na < 15% of total exchangeable cations from sodic soils with
Na > 15%, and in older literature, alkali soils with pH > 8.5. Here I will be discussing
broadly salt-affected soil: soil that has been adversely modified for the growth of
most crop plants by the presence of soluble salts, with or without high amounts of
exchangeable Na (SSSA glossary, https://www.soils.org/publications/soils-
glossary#). Salt-affected soils can be saline and alkaline.

0140-1963/$ e see front matter Crown Copyright Ó 2013 Published by Elsevier Ltd. All rights reserved.
http://dx.doi.org/10.1016/j.jaridenv.2012.12.014
 E.N. Bui / Journal of Arid Environments 92 (2013) 14e25 15

conductivity is a measure of the ease with which electrical current

will pass through water: the greater the salinity, the greater the
conductivity; however, the relationship is a function of the specific
ions present in the solution and their concentrations. Depending on
the soil texture, the relative water and salt content, and osmotic
pressure of the saturation extract can be very different (Fig. 7, US
Salinity Laboratory, 1969). When soil water content less than satu-
ration, plants will experience salt concentration higher than that
measured in the test for soil salinity. Salinity level will change
down a soil profile with seasonal moisture fluctuations and relative
soil water content but salt is generally localized below the root
zone. Expressing salinity as a soil profile-integrated value (e.g. Bui
and Henderson, 2003) dilutes the presence of salt concentrated in
a single horizon only.
Salt accumulation is a phenomenon that is not unique to any
particular soil type in many soil classifications but it is typically
associated with “Soils influenced by water” especially Solonetz
(alkaline) and Solonchak (salt enrichment upon evaporation) Soil
Groups in the World Reference Base (IUSS, 2006). It occurs where
evaporation is high relative to precipitation (there is a seasonal
water deficit) and leaching is insufficient to move salts out of the
soil profile (Duchaufour, 1982; Schofield and Kirkby, 2003). Sec-
ondary salinization can arise when salts accumulate near the soil
surface as a result of rising water tables due to land management
practices such as irrigation or tree clearing (Cisneros et al., 1999;
Rengasamy, 2006; Runyon and D’Odorico, 2010; Silburn et al.,
2009; Williamson, 1986). Thus a large fraction of natural and sec-
ondary salinity occurs in desert and grassland biomes, in savanna
ecosystems, which are often used as rangelands, and are closely
associated with semi-arid, seasonally contrasted climates in the
tropics and sub-tropics. Nevertheless the occurrence of salt-
affected soils is not a perfect match with climate, even when also
taking lithology (a potential source of solutes released by weath-
ering) into account (Schofield and Kirkby, 2003). Saline and sodic Fig. 1. Well developed Solonetz at Keyneton, South Australia with a typical horizon
soils often occur in or near local topographic lows in the landscape sequence: a brown or black surface horizon with organic matter dispersed by Na
cations; pale (albic) eluviation horizon directly over a horizon with exchangeable Na
where soluble salts accumulate under endorheic drainage condi- >15% (natric), columnar structure, and pH of about 8.5 (indicative of the presence of
tions, thus they are often seasonally waterlogged and can be free sodium carbonate); a horizon with Ca-carbonate (calcic) is present below the
associated with wetlands. Approximately 10% of the Earth’s total natric horizon. The sequence of mineral precipitates in the profile follows the Hardie
land surface may be salt-affected (Schofield and Kirkby, 2003). and Eugster (1970) sequence for the evolution of closed basin brines thus indicating
the presence of a shallow (w2 m) water table with Ca < CO3 during the soil’s for-
Soil salinity can imply the presence of chlorides, sulfates, ni-
mation. (Photo courtesy of Rob Fitzpatrick). (For interpretation of the references to
trates, (and bicarbonates) of sodium (Na), calcium (Ca), magnesium color in this figure legend, the reader is referred to the web version of this article.)
(Mg), and potassium (K). High levels of carbonates are reflected by
soil pH >8.3. Saline soils often have a typical sequence of horizon
concentration that separates halophytes from glycophytes has been
chemistry that reflects the position and salt content and chemistry
set at different levels by different authors. Flowers et al. (2010) use
of the underlying water table, the dominant upward trend of water
a threshold of 200 mM NaCl which corresponds roughly to
movement through the profile, and the solubility of salts (e.g.,
20 dS m
1 but others have used 80 mM NaCl (w8 dS m
1) to dis-
Fig. 1). Microbial activity can also influence saline soil chemistry
tinguish between halophytes and non-halophytes. Obligate halo-
and vice versa (Miletto et al., 2008; Whittig and Janitsky, 1963;
phytes require saline conditions to grow whereas facultative
Wolicka and Jarzynowska, 2012). Vegetation zonation is demar-
halophytes are found in less saline habitats and are characterized
cated as a function of salts and their chemical composition (e.g.,
by broader physiological diversity that enables them to cope with
Cisneros et al., 1999; Richardson et al., 1994; Stewart and Kantrud,
saline and non-saline conditions2 (Parida and Das, 2005). Salt-
1972) and changes in vegetation cover lead to changes in the soil
tolerant glycophytes exhibit some physiological mechanisms to
hydrology and chemistry (e.g., Cisneros et al., 1999; Runyon and
reduce sodium toxicity although they are not halophytes (e.g.
D’Odorico, 2010; Silburn et al., 2009).
Flowers et al., 2010; Hamilton et al., 2001; Parida and Das, 2005).
Global patterns of salinization have changed over geologic time
Even low soil salinity levels (<4 dS m
1) can exert physiological
with climate (Kershaw et al., 2003). Importantly, as Schofield and
stress on plants by affecting osmotic potential, ion toxicity (Na, Cl,
Kirkby (2003) have shown, climate change will impact on the
and other ions could be toxic), photosynthesis (reduction in
spatial pattern of soil salinity by changing patterns of precipitation
and evapotranspiration, and landscape hydrology.

1.2. Plant response to soil salinity
Most plants are glycophytes that tolerate only low concentra-
tions of salt before they are adversely affected as evidenced by
decrease in productivity and/or death. The threshold salt
2
Thus they apparently show phenotypic plasticity (the ability of a single geno-
type to produce multiple phenotypes in response to variation in the environment)
as defined by Pfennig et al. (2010). Phenotypic plasticity can promote diversification
because the developmental pathways that underlie environmentally induced
phenotypes consist of many genetic components that can potentially respond to
selection.
16 E.N. Bui / Journal of Arid Environments 92 (2013) 14e25
efficiency of photosynthesis is linked to reduction in stomatal
conductance), nutrient absorption, and N-fixing symbioses
(Lauchli and Luttge, 2002; Parida and Das, 2005; US Salinity
Laboratory, 1969; Zhu, 2001). As a result, plants have developed
a plethora of biochemical and molecular mechanisms to cope with
salt stress including: (i) selective accumulation or exclusion of
ions; (ii) control of ion uptake by roots and transport into leaves;
(iii) compartmentalization of ions at the cellular and whole-plant
levels; (iv) synthesis of compatible solutes, e.g., proline; (v) change
in photosynthetic pathway, e.g., a shift from the C3 to the C4
pathway in response to salinity is observed in the halophyte
Atriplex lentiformis; (vi) alteration in membrane structure; (vii)
induction of antioxidative enzymes; and (viii) induction of plant
hormones, e.g., abscissic acid (Flowers et al., 2010; Lauchli and
Luttge, 2002; Parida and Das, 2005; Zhu, 2001). Response starts
at low levels of salt and the mechanisms involved are based on
regulatory processes for ions, water, and CO2 common to all plants
that also may control responses to other environmental stress
such as drought (Flowers et al., 2010; Hamilton et al., 2001; Parida
and Das, 2005; Zhu, 2001). Plant responses to salinity vary with
the degree and duration of stress and the developmental stage at
time of exposure.
Biochemical pathways leading to products and processes that
improve salt tolerance are likely to act additively and probably
synergistically, controlled by multiple genes (Flowers et al., 2010;
Javid et al., 2011; Parida and Das, 2005; Zhu, 2001). For example,
adaptive differentiation of the halophyte Helianthus paradoxus from
its glycophyte parental species has been related to transgressive
trait expression, in which trait values in hybrids are higher or lower
than those of both parents (Edelist et al., 2009).3 Transgressive
traits in H. paradoxus include more succulent leaves and higher leaf
contents of Na and sulfur than both parental species (Karrenberg
et al., 2006; Rieseberg et al., 2003). Previous genetic studies
imply that such transgressive traits result from complementary
gene action where alleles with opposing effects are recombined in
hybrids (Rieseberg et al., 2003).
Although the mechanisms for coping with alkalinity stress are
similar to those for salt stress, different genes may be involved
(Javid et al., 2011; Yang et al., 2012) and neither the mechanisms
nor the genes have been studied as much as those for salt stress
which have been investigated intensively for three decades at least
(Flowers et al., 2010; Parida and Das, 2005). Given the frequent
coincidence of alkalinity and sodicity in salt-affected soils, genes
that confer tolerance to alkalinity may play a role in salt tolerance
too.
Salt tolerance is widely distributed across the plant kingdom
and is thought to have evolved polyphyletically but exactly how
and how many times salt tolerance has evolved since the emer-
gence of land plants 450e470 Mya is unresolved (Flowers et al.,
2010). Given that Na is an essential element for many plants that
use the C4 photosynthetic pathway (Subbarao et al., 2003), is there
any relationship between salt-tolerance and the evolution of
a photosynthetic pathway adapted to low CO2, high irradiance, and
dry conditions? Is it only coincidental that of the order Car-
yophyllales, the Chenopodiaceae use C4 carbon fixation the most,
with 550 or 39% of species in this family using this pathway or was
salt-tolerance a pre-condition (Sage, 2001)? New phylogenetic
analyses suggest that salt-tolerance pre-dates evolution of the C4
pathway in Chenopodiaceae, in contrast to Poaceae where the
reverse appears true (Kadereit et al., 2012).
3
Under- or over-expressed genes related to potassium and calcium transport
were homologs of KT1, KT2, ECA1; homologs of the potassium transporter HAK8 and
of a transcriptional regulator were generally over-expressed in saline treatments.
2. A role for soil salinity in ecology over geologic time
Biogeographic patterns emerge over time scales of hundreds to
tens of thousands of years as organisms adjust to prevailing envi-
ronmental conditions. Evolutionary processes, such as selection,
adaptation, speciation, and extinction act over longer geologic time
scales (to 106 years). There is some evidence that salinity plays
a part in both and thus may be an agent an evolutionary ecology
(evolution of biodiversity and plant communities).
2.1. Phytogeography
Because grasslands and seasonally flooded savannas are some of
the most poorly studied biomes in the world (Kier et al., 2005) and
salinity is associated with them, there has been comparatively little
work to draw on to review the influence of soil salinity as a driver in
plant ecology and phytogeography. Most studies on factors
explaining vegetation distribution focus on the role of climate and
soil moisture or fire but there is evidence that salinity plays a role in
current patterns of plant distribution at a regional scale in Central
Europe, the Middle East, Africa, the Americas, and Australia. Except
for the Prairie Pothole Region in North America, there is little detail
on the chemistry of salts present.
2.1.1. Europe, the Mediterranean basin and Middle East
In seminal work published in the 1970s, Heinz Ellenberg
assigned ‘indicator values’ to vascular plants in Central Europe with
respect to climatic factors (light, temperature, continentality), soil
moisture, acidity, available nitrogen, salt, and to a lesser degree of
success, heavy metal content of soil (Ellenberg et al., 1991). The
occurrence of salinity and sodicity in the Danube-Tisza floodplain is
well documented (Toth and Rajkai, 1994) and Ellenberg’s proximity
to this region may have influenced his consideration of salinity as
a factor in plant distribution.
In coastal ecosystems in southern Europe (Molina et al., 2003;
Rogel et al., 2001), and around the Mediterranean, in north Africa
and the Middle East (El-Bana et al., 2002; Le Houerou, 2003;
Shaltout et al., 1997), plant distribution in dune, marshland, and
aquatic habitats can be related to soil electrical conductivity and
chemistry. Focusing principally on identifying floristic domains and
salt-tolerant fodder and timber species, a comprehensive review of
vegetation patterns in the Red Sea basin (810,000 km2) describes
vegetation zonation that reflects climate, topography, and soil sal-
inity and gives a useful classification of halophytes and salt-tolerant
glycophytes in terms of their ecology (Le Houerou, 2003). In the Red
Sea lowlands (210,000 km2), vegetation is characterized by desert
contracted vegetation with many halophyte communities to the
north, and tropical Acacia-Commiphora savanna and halophytic
communities to the south (Le Houerou, 2003). Many halophyte and
salt-tolerant glycophyte species (100 with 20 endemics) are pre-
sent in the coastal plains of Egypt, the Sudan, Eritrea, Djibouti, the
Gulf of Aden, and northern Somalia and their habitat varies in its
level of salinity. Because this review is trying to demonstrate the
role of soil salinity in ecology beyond that of halophytes, some of
the salt-tolerant glycophyte communities identified by Le Houerou
(2003) are worth noting: Sporobolus spicatus community, Acacia
tortilis s.l., Acacia ehrenbergiana, and Acacia oerfota. These occur on
soil where only the subsoil is saline.
2.1.2. Africa
In the savannas of east Africa, grasses respond to a climate,
topography, and soil chemistry gradient (Banyikwa, 1976, cited by
Belsky, 1986; Hamilton et al., 2001). Whereas Andropogon green-
wayi occurs on soils with low Na concentrations, Sporobolus iocla-
dos, Sporobolus kentrophyllus and S. spicatus reflect an increasing
 E.N. Bui / Journal of Arid Environments 92 (2013) 14e25
level of Na in the Serengeti plains (w30,000 km2). These individual
Sporobolus species have been shown to respond differently to
increasing Na concentrations in laboratory experiments, in a man-
ner that reflects their distribution in East Africa (Hamilton et al.,
2001). S. spicatus tolerates the highest levels of Na in the field
and experimentally (Fig. 2) and qualifies as a halophyte under the
Flowers et al. (2010) threshold. Sub-surface concentrations of Na
and presenceeabsence of mound-building termites accounted for
55% of the variability in vegetational heterogeneity in the Serengeti
(Belsky, 1988).
In her review of East African savanna ecology models, Belsky
(1990) explains that the vegetation pattern of the Serengeti is
complex, reflecting rainfall, grazing, and soil salinity and pH. She
continues to explain that these factors are not independent since
salts are leached out by the higher rainfall in the north, and that
grazing patterns reflect grass productivity that increases with
rainfall northward. She concludes “in the Serengeti Plains it is the
high salinity and shallow soils that exclude trees, not herbivores or
rainfall.” Recent modeling work by Reed et al. (2009) concludes that
both rainfall and topography are important contributors to the
distribution of woodlands and grasslands in the Serengeti but un-
fortunately does not consider soil chemistry.
Electrical conductivity and carbonate content are the main soil
attributes that account for the vegetation groups along the Turkwel
River floodplain in Kenya (Stave et al., 2003). The effect of electrical
conductivity is reflected at levels below those that would charac-
terize a saline soil. The A. tortilis/Cadaba rotundifolia and Hyphaene
compress/C. rotundifolia wooded shrubland communities reflect the
highest salt and carbonate levels.
In southern Africa, Ellery et al. (1997) concluded that ground-
water salinity and chemistry were one of the major determinants of
vegetation species distribution in the Okavango delta, where again
S. spicatus was associated with high Na and soil electrical conduc-
tivity levels. Ellery et al. (1997) explained the dynamic fluctuations
in plant zonation patterns in the Okavango as a feedback effect
between groundwater depth and chemistry, and evapotranspira-
tion. Similar observations were made on endorheic pans in South
Africa by Cilliers and Bredenkamp (2003). In quartz fields of
southern Africa, where 70% of the plant species are endemic,
Fig. 2. Mean  1 SE field leaf blade Na concentrations and mean  1 SE field soil Na
concentrations (mg/kg dry mass) for Andropogon greenwayi, Sporobolus ioclados,
S. kentrophyllus, and S. spicatus paired leaf and soil samples collected in the Serengeti
short-grass plains. Diamonds (A) represent experimental Na treatment concentrations
(from Hamilton et al., 2001, with permission). Except for S. spicatus, photosynthetic
activity and biomass production of all species were significantly reduced by the
addition of Na.
17
Schmiedel and Jurgens (1999) found two dominant patterns of
growth forms, one associated with high salt content, neutral to
slightly acid soil pH, and low stoniness, the other characterized by
low soil salinity, acid soil pH, and high stoniness. Further, Schmiedel
and Jurgens (1999) observed that diversification of Mesem-
bryanthemaceae reflects subtle edaphic gradients in the saline
quartz fields.
2.1.3. Americas
In North America, the Prairie Pothole Region, which occupies
900,000 km2 across north central USA and south central Canada, is
characterized by thousands of small wetlands whose chemistry and
vegetation patterns are highly variable over short distances. For
example, in surface water of wetlands, total soluble salts, domi-
nantly sulfates and carbonates, can vary from 150 mg kg
1 to
3000 mg kg
1 over a distance of 300 m (Goldhaber et al., 2011;
Heagle et al., 2013) and salinity level is associated with concentric
zonation of herbaceous plant communities (Richardson et al., 1994;
Stewart and Kantrud, 1972). In the Prairie Pothole wetlands, the
occurrence of different species of Typha and Scirpus across the
salinity gradient (Fig. 3) suggests that speciation may be associated
with salinity. Salinity is one of the drivers for the distribution of
Helianthus (sunflower) species in North America, H. paradoxus
having adapted to saline soil (Bush and Van Auken, 2004; Rieseberg
et al., 2003).
In South America, in the Chaco of Argentina, plant communities
are distributed along a soil moisture and salinity gradient (Hilgert
et al., 2003). In the flooding pampas of Argentina, where more
than 60% of the soils over the 90,000 km2 area are saline or sodic,
edaphic factors associated with subtle topographic features are the
principal environmental factors explaining vegetation structure
(Perelman et al., 2001). C4 grasses are more common than C3 in
more saline environments (Perelman et al., 2001); this is not sur-
prising given Na is an essential element for many C4 plants
(Subbarao et al., 2003).
2.1.4. Australia
In Australia, the local role of soil salinity (and alkalinity) in
structuring plant communities is recognized widely (Bell and
Williams, 1997; Fensham et al., 2007; Froend et al., 1987;
Keighery et al., 2004; Kreeb et al., 1995; Wardell-Johnson et al.,
1997). The potential regional role of environmental factors
including salinity in plant ecology was investigated in the Murray
basin of Australia by Noy Meir (1974) who found it significant but
concluded that the role of salinity reinforced that of soil texture and
moisture. Crowley (1994) suggested that salinity played a role in
determining vegetation patterns in the southeast over the Qua-
ternary. The low relief landscapes of southwest Australia where
saline ground- and surface water systems have persisted for mil-
lennia have promoted the evolution of halophyte diversity in many
phylogenetically independent lineages including Chenopodiaceae
and Eucalyptus (Hopper, 2009).
Other evidence that salinity has been a major driver in plant
ecology in Australia over geological time is from Queensland in
northern Australia. There, Bui and Henderson (2003) analyzed
modern geo-referenced vegetation data that cover three of Aus-
tralia’s biodiversity ‘hotspots’, the Einasleigh Uplands, Desert Up-
lands, and northern Brigalow Belt.4 Their results also extend over
the southern Brigalow Belt. Their work shows that profile-
integrated soil salinity and topsoil pH, respectively, are strongly
associated with the first two score axes after correspondence
4
The map of Australia’s bioregions is at http://www.environment.gov.au/parks/
nrs/science/bioregion-framework/ibra/index.html.
18 E.N. Bui / Journal of Arid Environments 92 (2013) 14e25
PLANT COMMUNITY
In order of increasing electrical
conductivity
Scirpus heterochaetus (8)
Scirpus fluviatilis (13)
Typha spp. (32)
Typha spp.-Scirpus acutus (33)
Scirpus acutus (42)
Scirpus acutus-Scirpus paludosus (22)
Scirpus paludosus-Scirpus acutus (16)
Scirpus paludosus (13)
0.1
Fig. 3. Relationship of dominant deep-marsh emergent communities to electrical conductivity of pond water (from Stewart and Kantrud (1972)). Number of ponds sampled for each
community is shown in parentheses. For details, see page D11 in Stewart and Kantrud (1972).
analysis of site by species data, an indirect environmental gradient
method (Fig. 4A). While three Acacia species (Acacia argyrodendron
Domin., Acacia harpophylla F. Muell., and Acacia cambagei R. T. Bak.)
are indicative of saline soil, the distribution of 20 other individual
species across many genera responds strongly to a salinity gradient,
starting at very low levels of salinitydbelow those that would
typify a saline soil (Fig. 4B). Until this work, climate, soil texture,
and soil moisture were believed to be the most important variables
in the ecology of A. harpophylla (brigalow) communities but adding
soil salinity to these other environmental variables in a Generalized
Additive Model to account for site scores improved the results
dramatically (Mean Squared Error decreased from 0.89 to 0.65 and
R2 improved from 0.41 to 0.57).
2.2. A role for soil salinity in evolutionary biology
Environment-dependent selection may be important in speci-
ation but the conceptual frameworks of speciation genetics and
environmental stress physiology have not been fully integrated
(Lexer and Fay, 2005). Perhaps the best illustration of soil salinity’s
role in evolutionary biology comes from Rieseberg and colleagues
(e.g. Edelist et al., 2006, 2009; Rieseberg et al., 2003) who dem-
onstrated experimentally how speciation in sunflowers (Helianthus
spp.) in North America has been a response to environmental
stress, including soil salinity, over the Quaternary. The role of
abiotic environmental stress factors on plant evolution is difficult to
study and Helianthus, because of its relatively recent (60,000e
200,000 years ago) speciation and niche adaptation, is one of
only a few genera that have been studied from the molecular,
physiological, and evolutionary angles. Unlike its progenitors, Hel-
ianthus annuus and Helianthus petiolaris, the homoploid hybrid
H. paradoxus grew better and had strongly reduced leaf potassium,
Ca, and Mg contents upon saline treatment while supporting more
than nine times higher leaf Na loads than its parental species,
suggesting that Na may replace other cations as vacuolar osmotica
(Karrenberg et al., 2006). Under- or over-expressed genes related to
EXPLANATION
Geometric mean
Range
Standard deviation
Standard error of the mean
0.316 1 3.16 10 31.6 100
Electrical conductivity at 25°C, dS m-1
K and Ca transport; thus, alterations in the expression or activity of
ion transport genes may have been important in the evolution of
salt tolerance in H. paradoxus (Edelist et al., 2009).
Also experimentally, Hamilton et al. (2001) showed that Na-
tolerance in C4 grasses (including three Sporobolus species) was
associated with heat-shock protein (Hsp) production and proline
accumulationdboth provide protection against salt stress by reg-
ulating osmotic potential. They demonstrated induction of Hsps in
response to Na levels similar to those in which each species grows
in the field. Photosynthetic tolerance correlated with the observed
levels of Na in field tissues, and with the molecular adaptation of
Na-induced Hsp production. Long-term biomass responses were
correlated with short- and long-term physiological responses in
a direct relationship with the field Na gradient. Their experimental
results supported the hypothesis that the individual species have
evolved species-specific tissue Na tolerances in relation to their
field soil Na concentrations. Hamilton et al. (2001) noted that S.
spicatus exudes Na salts from its leaves as do other halophytic
Sporobolus species.5
Further evidence from Australia supports a role for salinity as
a driver in speciation. Several independent lines of evidenced
phytogeography, plant and insect systematics, and insect behav-
ioral ecologydpoint to a potential role for soil salinity in macro-
evolutionary processes for the genus Acacia and, indirectly, for the
insect order Thysanoptera. Investigations of Acacia thrips system-
atics and their host-plant relationships (Crespi et al., 2004) show
that: 1) A. harpophylla and A. cambagei harbor two sister-species
pairs of elongate and round gall thrips (Kladothrips spp.); 2)
phyllode-glueing thrips also show host specificity; and 3) thrips on
A. harpophylla and A. cambagei display high species richness.
Chronological phylogenetic analyses indicate that the approximate
5
The existence of salt glands in species across several genera of the Poaceae
suggests that they had a common halophytic ancestor to Liphschitz and Waisel
(1974).
 E.N. Bui / Journal of Arid Environments 92 (2013) 14e25 19

A
EUNOR

4
TRMIT

site occurence profile (species scores 2) EUSET

ACSHI CYBOM
EUSIM
2
EUPE1
TRSP1 EUSHI
MESP1
ERSP4MENE1
HASP1
EUME1
PEPUB
GRSP1
ACSP2
ENSP1
EUPER GRPAREUPOL
ARSP1PLCAR
EUCA1 GRSTR
ERMIT ERSP1
ERAU1 EUSP1HETRI
SPCAR BUINC
ACAR2 EUBR1 CHFAL
SPSP1
0

PASP1
DISP2 EUPL1
EUTE2THTRI
log(tss) LYCA1 ACSA1 RHREP
EUPAP
HECO1
ACHAR CYSP1
ACCAM TEOB1 CHSP1
ATHEM BOSP1EUCR1
EUTES
ALBAS
BODE1
DISP1 BOEWA ACBID
EUAUR EUER1
DISE1
URMOS
EUCOO BOPER
-2

ACFAR
CECIL

pH OPEXA

EUORG

-4 -3 -2 -1 0
site occurrence profile (species scores 1)

B
0
-1
-2
log(TSS)
-3
-4
-5
-6

THTRI
TRMIT

BUINC
ERMIT

HETRI
EUSHI
ACSHI

ACBID
CECIL

EUCR1

EUSIM
RHREP
ACHAR

EUAUR

ACFAR

CHFAL
ATHEM
ACCAM

URMOS
EUNOR

GRSTR
DISP2
DISP1

EUER1
ACAR2
EUCA1

CHSP1

EUBR1

DISE1

PLCAR
ERAU1

EUTES

EUSET
EUTE2

HECO1
EUME1

MENE1
EUCOO
SPCAR

EUPER
TRSP1

EUORG

CYBOM
GRPAR

EUSP1
EUPE1

EUPL1

HASP1
LYCA1

CYSP1

ACSA1

ERSP4
ACSP2
ERSP1
ENSP1
ARSP1

EUPAP
PEPUB
BODE1
MESP1

EUPOL
TEOB1

GRSP1

BOPER
BOEWA

ALBAS
SPSP1

PASP1

BOSP1
OPEXA

Fig. 4. A) Results of correspondence analysis; B) Response of vegetation to salinity starts at very low %TSS (TSS¼Total Soluble Salts) (from Bui and Henderson (2003), with per-
mission). See Appendix for list of species names.

age of origin of gall thrips (Kladothrips spp.) is Miocene and that recent work on Acacia biogeography in western and central Aus-
their subsequent diversification is closely linked to host-plant tralia (Ladiges et al., 2006) suggests that speciation has been driven
evolution; host affiliation with Plurinerves is estimated to date by aridity, but it does not consider salinity specifically.
from 7.5 Mya (McLeish et al., 2007). Thus the congruence between Salinity in Queensland dates back to the mid-Tertiary (Gunn and
planteinsect phylogenies and apparent co-speciation may have Richardson, 1978; van Dijk, 1980). Salinity in the area studied by Bui
been driven by environmental factors including salinity. Other and Henderson (2003) is associated with the Campaspe Formation
20 E.N. Bui / Journal of Arid Environments 92 (2013) 14e25

which is dated at 3.5e1.3 Mya (Nind, 1988). Acacia appears to have

been established in eastern Australia by early Miocene and became
an important component of sclerophyll communities by late Neo-
gene (Macphail and Hill, 2001). That salt stress may have played
a role in evolutionary ecology by exerting selection pressure on
Australian vegetation is supported by taxonomic and recent
chloroplast-DNA evidence that A. harpophylla, A. cambagei and
A. argyrodendron are closely related species of microneurous Plu-
rinerves (Crespi et al., 2004; Maslin, 2004; confirmed by J.T. Miller,
CSIRO Plant Industry, pers. comm.). These are the three Acacia

Table 1
Acacia species, their reported habitat and associated species (http://www.
worldwidewattle.com).

Section Species Habitat and assoc species

Juliflorae A. catenulata invariably in pure stands or with emergent
eucalypts on shallow soils on scarps of
weathered sediments, sometimes adjoining,
but not mixing with, stands of A. shirleyi or
A. aneura
A. shirleyi Grows in shallow gravelly or skeletal sandy
soils on sandstone or laterite often in dense
stands, also in closed forests, low open forests
or mixed savannah woodlands
A. stowardii Common in shallow soils of stony or rocky
ridges and breakaways, also on sand dunes;
often in A. aneura communities on low hills
A. adsurgens Grows in reddish sandy and gravelly soils, on
flat plains and hillsides, commonly in spinifex
grassland communities
A. aneura var. usually on loamy or sandy soils in areas of low
aneura relief or on shallow rocky soils on hills
Plurinerves A. melanoxylon favors fertile soils in valleys and on flats in
mountainous areas, often growing in wet
sclerophyll forest and cooler rainforest.
A. oswaldii in arid, semi-arid and subtropical areas in
all mainland states S of 19 S; mainly in
calcareous sands or loam.
A. melvillei Grows in loam, clay and sand often in mixed
open woodland and woodland
A. papyrocarpa Grows in sandy loam, clay and calcareous
soil, in low open woodland and shrubland,
often with chenopods
A. pendula Grows mainly on floodplains in fertile alluvial
clay (and red earth soils in the South),
sometimes dominant in woodland and
open woodland
A. tephrina Grows mainly in heavier soils, including
alkaline and saline clays, in tall open
woodland and, in drier areas, low woodland
and shrubland
A. loderi Grows in solonized brown and red soils, in
low woodland and tall shrubland, sometimes
associated with chenopods, Casuarina cristata
or Acacia aneura
A. microsperma grows in shallow loam or clay over weathered
rock, often forming dense stands alone or
with A. cambagei or Eucalyptus thozetiana
A. cana Grows along watercourses on gibber plains
and on sandhills as pure stands of scattered
trees and in mixed woodland
A. maranoensis Grows in texture contrast soils, usually in
woodland of Eucalyptus populnea
A. harpophylla Grows on fertile clay and loamy clay
A. argyrodendron Grows mostly in dark cracking clay, either
alone or, on the southern and eastern edge
of its range, with Acacia harpophylla
A. cambagei Tolerates a wide range of soils but occurs
most commonly in dark cracking clay or
loam, as scattered individuals or in dense,
almost pure stands
A. georginae Usually grows on plains and along
watercourses in clay and loam, often
dominating in low woodland and low open
woodland
Fig. 5. Suggested environmental factors (see Table 1) associated with the splits of the
phylogram from Crespi et al. (2004) with additional species closely related botanically
in parentheses. Parkia timoriana is an outgroup. A. catenulata, A. stowardii, A. adsurgens,
and A. aneura are Juliflorae (boxed). All others are Plurinerves.
species that exhibit tolerance of the highest levels of salt in
Queensland (Bui and Henderson, 2003).
Using published data (Orchard and Wilson, 2001), Table 1 syn-
thesizes the ecology of Acacia species from sections Plurinerves and
Juliflorae of subgenus Phyllodineae (see Maslin, 2004) for which
phylogenetic relations have been advanced by Crespi et al. (2004).
The habitats described for these species suggest that environ-
mental factors could have played a role in the speciation of these
acacias as schematized in Fig. 5. Alkalinity (calcareous habitat) and
salinity are factors that appear to characterize the habitats of the
Plurinerves closely related to A. harpophylla. Although tolerance to
alkalinity seems to have appeared many times over the Acacia
phylogeny, not all Plurinerves are salt-tolerant, thus salinity ap-
pears to have played a role in the speciation within this clade.
The influence of environmental factors in biogeography and
evolutionary ecology are notoriously difficult to unravel and the
long record of landscape evolution in Australia suggests that cli-
mate and soil have interacted at length in driving the phylogeny of
Acacia (C. Gonzalez-Orozco, pers. comm.). The trend toward aridity
over the continent began in the mid-late Miocene (Martin, 2006)
and the current monsoonal pattern over Queensland became
established in the Holocene around 9000 years ago (Kershaw,
1978).
Acacia are legumes and in some species form symbioses with
rhizobia that are known to be tolerant of salt, drought, and high
temperatures (Thrall et al., 2009; Zahran, 2001; Zhang et al., 1991).
Fierer and Jackson (2006) found that edaphic variables and most
importantly pH control soil bacterial biogeography across the
Americas. Salinity may mediate subterranean processes that in-
fluence plantemicrobial interactions. Progress in the systematics of
bacterial N-fixing legume symbionts (Sawada et al., 2003; Zheng
et al., 2004) may shed further light on the role of salt-stress in
evolutionary ecology.
The convergence of three independent lines of evidence pro-
vides an exciting insight into the potential role of soil alkalinity and
salinity on macroevolutionary processes for the genus Acacia.
Acacia is a widespread genus (Maslin et al., 2003)6 and salinity may
play a role in its distribution and speciation outside of Australia.
6
See http://www.worldwidewattle.com/infogallery/distribution.php.
 E.N. Bui / Journal of Arid Environments 92 (2013) 14e25
3. A role for soil salinity over short time scales
Whereas the previous section has discussed the role of natural
variations in salinity over geologic time, this section addresses the
impact of changes in soil salinity brought about through anthro-
pogenic activity, that occur over short time frames (years to de-
cades). The threat of secondary dryland salinity to current patterns
of biodiversity in Australia has been dealt with by others (e.g.,
Briggs and Taws, 2003; Cramer and Hobbs, 2002 and other papers
in a special issue of Australian Journal of Botany; Keighery et al.,
2004; Seddon et al., 2007) and this paper will only consider two
aspects that have been overlooked: the possible role of salinity in
weed invasion and woody vegetation encroachment.
In the western USA, Scott et al. (2010) investigated possible
mechanisms driving the invasion of Bromus tectorum L. which was
introduced from Eurasia in the 1800s. B. tectorum is a selfing winter
annual grass that has achieved widespread dominance in semiarid
western North America since the 1930s and is actively invading salt
desert habitats. They found that seeds from saline playa pop-
ulations were able to establish better than those of most other
populations across all habitats, including two highly saline sites.
They concluded through molecular analysis that local population
differentiation in B. tectorum resulted from differential selection on
pre-adapted genotypes with characteristic micro-satellite marker
fingerprints and found little evidence for selection favoring novel
genotypes.
3.1. Invasive species and soil salinity in Australia
Weed invasions are currently a major concern for land managers
and conservation biologists in Australia. Current research has
focused on dispersal mechanisms, recruitment and survival, sub-
terranean root competition, functional traits, the role of grazing
animals, the impact of fire, and control strategies but the role of soil
salinity as a potential factor has been neglected. However, the data
in Bui and Henderson (2003) show that the spatial pattern of Buffel
grass (Cenchrus ciliaris L.), an introduced pasture grass now con-
sidered an invasive species (Fairfax and Fensham, 2000), is driven
largely by soil pH and salinity (CECIL in Fig. 4A), even though C.
ciliaris is only weakly salt-tolerant (Graham and Humphreys,
1970).7 Personal field observations in Queensland suggest that
the woody weed Parkinsonia aculeata L. appears to colonize saline
discharge areas where it forms monostands and that Parthenium
hysterophorus L. also seems to be widespread in saline areas. Others
have also reported a dominance of groundcover species by exotics,
including weeds, on salinized sites in Queensland and New South
Wales (Briggs and Taws, 2003). Tomar et al. (1998) have found P.
aculeata a suitable species for afforestation of saline soils in India.
Acacia nilotica (L.) Del. ssp. indica (Benth.) Brenan. is a woody
weed in Australia (Miller et al., 2011) that has received much
attention in the last decade (e.g. Kriticos et al., 2003). A. nilotica is
known to perform well under water-logged, saline conditions (e.g.
Marcar et al., 2003; Tomar et al., 1998). This raises the question of
invasion by A. niloticadis it related in part to soil salinity? A similar
question arises regarding Acacia farnesiana (L.) Willd., a woody
weed (Miller et al., 2011) that grows on calcareous cracking clay
soils and along saline artesian drains8 and is suitable for affor-
estation of saline soils (Tomar et al., 1998). The results of Bui and
Henderson (2003) support an important role for soil pH in its
7
According to the FAO grassland species database, it grows optimally on soils
with pH 7e8 and the Australian cultivar Biloela is more salt-tolerant than others
(http://www.fao.org/ag/AGP/AGPC/doc/Gbase/DATA/Pf000196.HTM).
8 9
See http://www.fao.org/ag/AGP/AGPC/doc/Gbase/DATA/Pf000113.HTM.
21
distribution in Queensland (ACFAR in Fig. 4A). Interestingly,
chloroplast-DNA analysis shows that A. nilotica and A. farnesiana
are sister species (Bukhari et al., 1999). Since A. nilotica originates
from Africa-Asia whereas A. farnesiana is from Central America, the
adaptation to similar edaphic conditions appears ancestral to both.
Prosopis L. is another woody weed in Australia with known salt-
tolerant species (e.g. Le Houerou, 2003) so the same question ap-
plies to its ability to invade. Tomar et al. (1998) found Prosopis
juliflora (Swartz) DC. to be suited for reclamation of saline soil in
India. Stave et al. (2003) report that Prosopis chilensis (Mol.) Stuntz,
introduced in Kenya around 1985, is invading the floodplain of the
Turkwel River where electrical conductivity and carbonate content
account for vegetation distribution. Thus the physical environment,
stressed by salinity, may play a key role in defining a niche op-
portunity for weed invasion for plants that tolerate salt to a degree,
especially at the seedling recruitment stage.
In 1999 the Australian Government and state and territory
governments endorsed a list of 20 Weeds of National Significance
(WONS).9 Of these, twelve show some salt-tolerance (Table 2).
Many WONS are phreatophytes or amphibious aquatic species that
invade seasonal wetlands. It appears that areas that are affected by
the interaction between shallow water tables and salt are partic-
ularly susceptible to weed invasions in Australia.
Tree clearing, which changes the hydrological balance of land-
scapes and can promote soil salinization (Runyan and D’Odorico,
2010; Williamson, 1986), may be a human activity that also pro-
vides an advantage to opportunistic invaders. Nevertheless salinity
has been neglected as a potential factor in invasion biology inland,
although its role in coastal ecosystems in America has been dem-
onstrated (e.g. Silliman and Bertness, 2004).
3.2. Woody vegetation encroachment into grassland and salinity
A. harpophylla, although a native Australian species endemic to
the Brigalow Belt, is considered a weed by rangeland managers
because of its suckering habit (Bortolussi et al., 2005; Nix, 1994). An
association between A. harpophylla and saline seeps is acknowl-
edged by Nix (1994) and empirically demonstrated again in Bui and
Henderson (2003) (ACHAR in Fig. 1). Emergent seedlings of
A. harpophylla can tolerate high salinity (Nix, 1994; Reichman et al.,
2006).
Thickening of brigalow communities and encroachment of
woody plants into grasslands has been a reported “nuisance” by
Regional Vegetation Management Planning Committees across the
Brigalow Belt, Desert Uplands, Mitchell Grass Downs, Gulf Plains
biogeographic regions in Queensland. Woodland thickening is
reportedly occurring on vast areas of rangelands, however, because
it is not possible to assess with the resolution of Landsat TM data
used to monitor statewide landcover change (Goulevitch et al.,
1999, 2002), it is difficult to assess the actual area affected. Ac-
cording to Goulevitch et al. (1999), 20 sites show evidence of
woodland encroachment into grassland from d13C which discrim-
inates between C produced by trees using the C3 photosynthetic
pathway from C produced by grasses using the C4 pathway. At one
such site in Queensland where A. cambagei R.T. Bak., a salt-tolerant
acacia species closely related to A. harpophylla (Bui and Henderson,
2003; Maslin, 2004), is encroaching on grassland, Krull et al. (2005)
found that there was a strong shift from C4- (grass) to C3-derived
(trees) soil C in the thickened woodland.
Woodland thickening in savannas is generally attributed to
changes in land management such as decreased fire incidence,
http://www.weeds.gov.au/weeds/lists/wons.html.
22 E.N. Bui / Journal of Arid Environments 92 (2013) 14e25

Table 2
Of the 20 WONS, twelve show some salt- or alkalinity-tolerance; seven are phreatophytes; five are nitrogen-fixing legumes. Clearly soil chemistry and moisture are key
variables in the ecology of weeds.

Weeds of National Significance (WONS) Salt-tolerant Phreatophyte Aquatic Leguminous Reference for salt-
(N2-fixer) tolerance
Alligator weed (Alternanthera philoxeroides) Y Y CRC Weed Managementa
Athel pine or tamarisk (Tamarix aphylla) Y Y, facultative Di Tomaso (1998)
Bitou bush (Chrysanthemoides monilifera ssp. rotundata) Y CRC Weed Management
Boneseed (Chrysanthemoides monilifera ssp. monilifera)
Blackberry (Rubus fruticosus aggregate) ? grows in gullies CRC Weed Management
Bridal creeper (Asparagus asparagoides) Alkalinity CRC Weed Management
Cabomba (Cabomba caroliniana) N Y CRC Weed Management
Chilean needle grass (Nassella neesiana) CRC Weed Management
Gorse (Ulex europaeus) Y CRC Weed Management
Hymenachne or Olive hymenachne N “not brackish water” Y CRC Weed Management
(Hymenachne amplexicaulis)
Lantana (Lantana camara) N CRC Weed Management
Mesquite (Prosopis species) Y Y Y FAOb
Mimosa (Mimosa pigra) Y Facultative? Y ABRS (2007)
Parkinsonia (Parkinsonia aculeata) Y Facultative? Y FAOb
Parthenium weed (Parthenium hysterophorus) Y? Personal observation
Pond apple (Annona glabra) Y Y CRC Weed Management
Prickly acacia (Acacia nilotica) Y Y Y FAOb
Rubber vine (Cryptostegia grandiflora) Facultative? CRC Weed Management
Salvinia (Salvinia molesta) Y Y CRC Weed Management
Serrated tussock (Nassella trichotoma) N CRC Weed Management
Willow (Salix spp.) Y Y Kennedy et al. (2003)
a
CRC Weed Management data sheets for WONS: http://www.weeds.gov.au/publications/guidelines/index.html.
b
FAO Grassland Index: http://www.fao.org/ag/AGP/AGPC/doc/Gbase/mainmenu.htm.

increasing grazing pressure, climate change and increasing at-
mospheric CO2 concentration (Archer, 1995; Bond and Midgley,
2012; Burrows, 2002; Fensham and Fairfax, 2005) but may be
due to or reinforced byein some places at leastethe interaction
between recent changes in landscape hydrology and soil salinity.
Given that most of the Brigalow Belt has been cleared, it is pos-
sible that more water is moving laterally through local recharge
areas into seepage areas, that salt is being transported, and that
Acacia remnants and re-growth are encroaching on grassland that
is becoming more saline close to the soil surface. Fairfax and
Fensham (2005) found that episodes of thickening and
encroachment of A. cambagei coincide with wet periods. Bui and
Moran (2000) show that soil salinity is greater downslope of sa-
line seepage areas under remnant brigalow stands in northern
Queensland; this is confirmed in Silburn et al. (2009). Woodland
encroachment, possibly responding to landscape salinization in
the Brigalow Belt of Australia, may be a good example of habitat
tracking in real time.
In East Africa however, the situation appears reversed: if trees
and bush thickets in low rainfall areas are left undisturbed long
enough, they develop into grasslands (Belsky, 1990). Belsky (1990)
concludes that trees do not grow in the Serengeti and Amboseli
plains, even when protected from herbivory and fire, and that it is
the high salinity and shallow soils that exclude trees from grass-
lands. There it seems that grasses are better adapted than trees to
prevailing environmental conditions including soil salinity. But this
situation may reflect in part the paleoenvironmental history of East
Africa and the relatively smaller number of Acacia species in Africa
(144) compared to Australian ones (w1000).
4. Implications for global change research
Evidence from Australia and elsewhere suggests a role for soil
salinity in biogeography and ecology (i.e., niche definition and se-
lection pressure), at a range of spatio-temporal scales. Over the
geologic time scale, it has played a part in driving speciation while
in more recent time, over decades, it is changing current patterns of
biodiversity.
Good predictive models of the distribution of biota and of their
responses to environmental stress are essential for evidence-based
management of biodiversity in Australia and throughout the
world. To construct such models, predictor variables are required
that can represent the underlying environmental gradient and
processes that control spatial distribution. Biogeographic models
throughout the world frequently rely on climatic and topographic
datasets for predictions of species distribution (e.g., Austin and
Meyers, 1996; Eeley et al., 1999; Reed et al., 2009). Perhaps this
worldwide tendency to focus on climate and topography data in
biogeographical studies also reflects the availability of those
datasets. The existence of computer programs such as ANUCLIM
makes the estimation of climatic variables as a function of mete-
orological and topographic data, and the definition of bioclimatic
zones, readily achievable (Houlder et al., 1999). Because plant
available water (and often other soil properties too) is a function of
the interaction between climate and topography, commonly
environmental gradient analyses focus on transects along climate
and topographic gradients. The recent focus on assessment of
climate change on ecosystems may also contribute to the relative
neglect of edaphic factors. Even so, soil data are often not available.
For Australia, that lacuna has been filled by the Australian Soil
Resources Information System (ASRIS) datasets which generated
digital maps of soil pH, % organic C, total P, total N, texture, % clay,
and thickness for surface and subsurface horizons at 1-km reso-
lution (Johnston et al., 2003). On a global scale, soil texture, pH,
and soil type maps are also relatively more accessible than geo-
chemical (salinity, major and trace element) maps but substrate
geochemistry may be critical to account for observed floristic
patterns (e.g. Martin et al., 2006; Reed et al., 2009). Better global
hydrology coverage, including depth-to-groundwater and geo-
chemistry, is also badly needed.
Access to a comprehensive range of digital environmental
datasets linked with biological datasets will give biogeographers
and ecological modelers an opportunity to determine which factors
are major drivers for the distribution of biota in any given region. It
will be critical to evidence-based management of biodiversity;
extinction risks could be higher than predicted if future locations of
 E.N. Bui / Journal of Arid Environments 92 (2013) 14e25 23

suitable climate do not coincide with other essential resources such (continued )
as suitable soils (Austin and Van Niel, 2011; Bertrand et al., 2012; Species name Species
Thomas et al., 2004). At the same time, it will open the door to an abbreviation
emerging science area of “phylo-environmental modelling”, inte- Heteropogon triticeus HETRI
grative research for reconstructing genetic and spatial patterns of Lysiphillum carronii LYCA1
change through geological time. Melaleuca nervosa MENE1
Melaleuca species MESP1
Ophiurous exaltatus OPEXA
Acknowledgments Panicum species PASP1
Petalostigma pubescens PEPUB
Planchonia careya PLCAR
I want to thank Laurence Mound and the Australian Biological
Rhynchelytrum repens RHREP
Resources Study for encouragement and permission to use Fig. 32 Sporobolus caroli SPCAR
from Crespi et al. (2004), and the two anonymous reviewers for Sporobolus species SPSP1
providing many useful comments and references for improving the Terminalia oblongata TEOB1
paper. Themeda triandra THTRI
Triodia mitchelii TRMIT
Triodia species TRSP1
Urochloa mosambicensis URMOS
Appendix

Species name Species

abbreviation References
Acacia argyrodendron ACAR2
A. bidwilii ACBID ABRS, 2007. Flora of Australia On-line. http://www.environment.gov.au/
A. cambagei ACCAM biodiversity/abrs/online-resources/flora/main/index.html (accessed 13.04.11.).
Archer, S., 1995. Harry Stobbs Memorial Lecture, 1993: herbivore mediation of
A. farnesiana ACFAR
grass-woody plant interactions. Tropical Grasslands 29, 218e235.
A. harpophylla ACHAR
Austin, M.P., Meyers, J.A., 1996. Current approaches to modelling the environmental
A. salicina ACSA1
niche of eucalypts: implication for management of forest biodiversity. Forest
A. shirleyi ACSHI Ecology and Management 85, 95e106.
A. species ACSP2 Austin, M.P., Van Niel, K.P., 2011. Impact of landscape predictors on climate change
Albizia basaltica ALBAS modelling of species distributions: a case study with Eucalyptus fastigata in
Aristida species ARSP1 southern New South Wales, Australia. Journal of Biogeography 38, 9e19.
Atalaya hemiglauca ATHEM Bell, D.T., Williams, J.E., 1997. Eucalypt ecophysiology. In: Williams, J.E.,
Bothriochloa decipiens BODE1 Woinarski, J.C.Z. (Eds.), Eucalypt Ecology: Individuals to Ecosystems. Cambridge
Bothriochloa ewartiana BOEWA University Press, Cambridge, pp. 168e196.
Bothriochloa pertusis BOPER Belsky, A.J., 1990. Tree/grass ratios in East African savannas: a comparison of
Bothriochloa species BOSP1 existing models. Journal of Biogeography 17, 483e489.
Bursaria incana BUINC Belsky, A.J., 1988. Regional influences on small scale vegetational heterogeneity
Cenchrus ciliaris CECIL within grasslands in the Serengeti National Park, Tanzania. Plant Ecology 74, 3e
10.
Chrysopogon fallax CHFAL
Belsky, A.J., 1986. Population and community processes in a mosaic grassland in the
Chloris species CHSP1
Serengeti, Tanzania. Journal of Ecology 74, 841e856.
Cymbopogon bombycinus CYBOM Bertrand, R., Perez, V., Gégout, J.C., 2012. Disregarding the edaphic dimension in
Cyperus species CYSP1 climate change: the case of Quercus pubescens in France. Global Change Biology
Dichanthium sericeum DISE1 18, 2648e2660.
Dichanthium species DISP1 Bond, W.J., Midgley, G.F., 2012. Carbon dioxide and the uneasy interactions of trees
Digitaria species DISP2 and savannah grasses. Philosophical Transactions of the Royal Society B 367
Enneapogon species ENSP1 (1588), 601e612. http://dx.doi.org/10.1098/rstb.2011.0182 1471e2970.
Erythroxylon australe ERAU1 Bortolussi, G., McIvor, J.G., Hodgkinson, J.J., Coffey, S.G., Holmes, C.R., 2005. The
Eremophila mitchelii ERMIT Australian beef industry: a snapshot. 4. Condition and management of natural
Eragrostis species ERSP1 resources. Australian Journal of Experimental Agriculture 45, 1109e1120.
Eriachne species ERSP4 Briggs, S.V., Taws, N., 2003. Impacts of salinity on biodiversitydclear understanding
Eulalia aurea EUAUR or muddy confusion? Australian Journal of Botany 51, 609e617.
Bui, E.N., Henderson, B.L., 2003. Vegetation indicators of soil salinity in north
Eucalyptus brownii EUBR1
Queensland. Austral Ecology 28, 539e552.
E. cambageana EUCA1
Bui, E.N., Moran, C.J., 2000. Regional-scale investigation of the spatial distribution
E. coolibah EUCOO
and origin of soluble salts in central north Queensland. Hydrological Processes
E. crebra EUCR1 14, 237e250.
E. erythrophloia EUER1 Bukhari, Y.M., Koivu, K., Tigerstedt, P.M.A., 1999. Phylogenetic analysis of Acacia
E. melanophloia EUME1 (Mimosaceae) as revealed from chloroplast RFLP data. Theoretical and Applied
E. normantonensis EUNOR Genetics 98, 291e298.
E. orgadophylla EUORG Burrows, W.H., 2002. Seeing the wood(land) for the trees e an individual per-
E. papuana EUPAP spective of Queensland woodland studies (1965e2005). Tropical Grasslands 36,
E. peltata EUPE1 202e217.
E. persistens EUPER Bush, J.K., Van Auken, O.W., 2004. Relative competitive ability of Helianthus para-
E. platyphylla EUPL1 doxus and its progenitors, H. annuus and H. petiolaris (Asteraceae), in varying
E. polycarpa EUPOL soil salinities. International Journal of Plant Sciences 165, 303e310.
Cilliers, S.S., Bredenkamp, G.J., 2003. Vegetation of inland endorheic pans in the
E. setosa EUSET
North-West Province, South Africa. Phytocoenologia 33, 289e308.
E. shirleyii EUSHI
Cisneros, J.M., Cantero, J.J., Cantero, A., 1999. Vegetation, soil hydrophysical prop-
E. similis EUSIM
erties, and grazing relationships in saline-sodic soils of Central Argentina.
E. species EUSP1 Canadian Journal of Soil Science 79, 399e409.
E. tereticornis EUTE2 Cramer, V.K., Hobbs, R.J., 2002. Ecological consequences of altered hydrological
E. tessellaris EUTES regimes in fragmented ecosystems in southern Australia: impacts and possible
Grevillea parallela GRPAR management responses. Austral Ecology 27, 546e564.
G. species GRSP1 Crespi, B.J., Morris, D.C., Mound, L.A., 2004. Evolution of Ecological and Behavioural
G. striata GRSTR Diversity: Australian Acacia Thrips as Model Organisms. Australian Biological
Hakea species HASP1 Resources Study and Australian National Insect Collection, CSIRO, Canberra.
Heteropogon contortus HECO1 Crowley, G.M., 1994. Quaternary soil-salinity events and Australian vegetation
history. Quaternary Science Review 13, 15e22.
24 E.N. Bui / Journal of Arid Environments 92 (2013) 14e25
Dirnbock, T., Hobbs, R.J., Lambeck, R.J., Caccetta, P.A., 2002. Vegetation distribution
in relation to topographically driven processes in southwestern Australia.
Applied Vegetation Science 5, 147e158.
Di Tomaso, J.M., 1998. Impact, biology, and ecology of saltcedar (Tamarix spp.) in the
southwestern United States. Weed Technology 12, 326e336.
Duchaufour, P., 1982. Pedology. George Allen and Unwin.
Edelist, C., Lexer, C., Dillmann, C., Sicard, D., Rieseberg, L.H., 2006. Microsatellite
signature of ecological selection for salt tolerance in a wild sunflower hybrid
species, Helianthus paradoxus. Molecular Ecology 15, 4623e4634.
Edelist, C., Raffoux, X., Falque, M., Dillmann, C., Sicard, D., Rieseberg, L.H.,
Karrenberg, S., 2009. Differential expression of candidate salt-tolerance genes
in the halophyte Helianthus paradoxus and its glycophyte progenitors
H. annuus and H. petiolaris (Asteraceae). American Journal of Botany 96 (10),
1830e1838.
Eeley, H.A.C., Lawes, M.J., Piper, S.E., 1999. The influence of climate change on the
distribution of indigenous forest in KwaZulu-Natal, South Africa. Journal of
Biogeography 26, 595e617. http://dx.doi.org/10.1046/j.1365-2699.1999.00307.x.
El-Bana, M., Khedr, A.H., Van Hecke, P., Bogaert, J., 2002. Vegetation composition of
a threatened hypersaline lake (Lake Bardawil), North Sinai. Plant Ecology 163,
63e75.
Ellenberg, H., Weber, H.E., Dull, R., Wirth, V., Werner, W., Paulisen, D., 1991. Scripta
Geobotanica, vol. 18. Verlag Erich Goltze KG, Göttingen.
Ellery, W.N., Ellery, K., McCarthy, T.S., 1997. Plant-distribution in islands of the
Okavango delta, Botswanaddeterminants and feedback interactions. African
Journal of Ecology 31, 119e134.
Fairfax, R.J., Fensham, R.J., 2000. The effect of exotic pasture development on floristic
diversity in central Queensland, Australia. Biological Conservation 94, 11e21.
Fairfax, R.J., Fensham, R.J., 2005. Preliminary assessment of gidgee (Acacia cambagei)
woodland thickening in the Longreach district, Queensland. Rangeland Journal
27, 159e168.
Fensham, R.J., Silcock, J., Biggs, A., 2007. Vegetation-soil relations in a highly sodic
landscape, Yelarbon, southern Queensland. Cunninghamia 10 (2), 273e284.
Fierer, N., Jackson, R.B., 2006. The diversity and biogeography of soil bacterial
communities. Proceedings of the National Academy of Sciences of the United
States of America 103, 626e631.
Flowers, T.J., Galal, H.K., Bromham, L., 2010. Evolution of halophytes: multiple ori-
gins of salt tolerance in land plants. Functional Plant Biology 37, 604e612.
Froend, R.H., Heddle, E.M., Bell, D.T., McComb, A.J., 1987. Effects of salinity and
waterlogging on the vegetation of Lake Toolibin, Western Australia. Australian
Journal of Ecology 12, 281e298.
Goldhaber, M.B., Mills, C., Stricker, C.A., Morrison, J.M., 2011. The role of critical zone
processes in the evolution of the Prairie Pothole Region wetlands. Applied
Geochemistry 26, S32eS35.
Goulevitch, B.M., Danaher, T.J., Stewart, A.J., Harris, D.P., Lawrence, L.J., 2002. Map-
ping woody vegetation cover over the state of Queensland using Landsat TM
and ETMþ imagery. In: Proceedings of the 11th Australasian Remote Sensing
and Photogrammetry Conference, Brisbane, Australia, September 2002. http://
www.nrme.qld.gov.au/slats/pdf/0127anav.pdf.
Goulevitch, B.M., Danaher, T.J., Walls, J.W., 1999. The Statewide Landcover and Trees
Study (SLATS) e monitoring land cover change and greenhouse gas emissions
in Queensland. In: Proceedings of IEEE International Geoscience and Remote
Sensing Symposium, Hamburg, June 1999. http://www.nrme.qld.gov.au/slats/
pdf/igarss99overview.PDF.
Graham, T.W.G., Humphreys, L.R., 1970. Salinity response of cultivars of buffel grass
(Cenchrus ciliaris). Australian Journal of Experimental Agriculture and Animal
Husbandry 10, 725e728.
Gunn, R.H., Richardson, D.P., 1978. The nature and possible origins of soluble salts in
deeply weathered landscapes of eastern Australia. Australian Journal of Soil
Research 17, 197e215.
Hamilton, E.W., McNaughton, S.J., Coleman, S.J., 2001. Molecular, physiological, and
growth responses to sodium stress in C4 grasses from a soil salinity gradient in
the Serengeti ecosystem. American Journal of Botany 88, 1258e1265.
Hardie, L.A., Eugster, H.P., 1970. The evolution of closed-basin brines. Mineralogical
Society of America Special Publication 3, 273e290.
Heagle, D., Hayashi, M., van der Kamp, G., 2013. Surfaceesubsurface salinity dis-
tribution and exchange in a closed-basin prairie wetland. Journal of Hydrology
478, 1e14.
Hilgert, N.I., Pensiero, J.F., Marino, G., Lewis, J.P., D’Angelo, C., 2003. Vegetation of
the Saladillo area (province of Santa Fe) in the south Chaco, Argentina. Inter-
ciencia 28, 512e520.
Hopper, S.D., 2009. OCBIL theory: towards an integrated understanding of the
evolution, ecology and conservation of biodiversity on old, climatically buf-
fered, infertile landscapes. Plant and Soil 322 (1), 49e86.
Houlder, D., Hutchinson, M., Nix, H., McMahon, J., 1999. ANUCLIM Version 5.0. User
Guide. Centre for Resource and Environmental Studies, The Australian National
University, Canberra.
Hughes, L., 2003. Climate change and Australia: trends, projections and impacts.
Austral Ecology 28, 423e443.
IUSS Working Group WRB, 2006. World Reference Base for Soil Resources 2006.
World Soil Resources Reports No. 103. FAO, Rome.
Javid, M., Nicolas, M., Ford, R., 2011. Current knowledge in physiological and genetic
mechanisms underpinning tolerances to alkaline and saline subsoil constraints
of broad acre cropping in dryland regions. In: Shanker, Arun, Venkateswarlu, B.
(Eds.), Abiotic Stress in Plants e Mechanisms and Adaptations. InTech, pp. 193e
214. Available from: http://www.intechopen.com/books/abiotic-stress-in-
plants-mechanisms-and-adaptations/current-knowledge-in-physiological-and-
genetic-mechanisms-underpinning-tolerances-to-alkaline-and-sa.
Johnston, R.J., Barry, S.J., Bleys, E., Bui, E.N., Moran, C.J., Simon, D.A.P., Carlile, P.,
McKenzie, N.J., Henderson, B., Chapman, G., Imhoff, M., Maschmedt, D.,
Howe, D., Grose, C., Schokneckt, N., Powell, B., Grundy, M., 2003. ASRIS: the
database. Australian Journal of Soil Research 41, 1021e1036.
Kadereit, G., Ackerly, D., Pirie, M.D., 2012. A broader model for C4 photosynthesis
evolution in plants inferred from the goosefoot family (Chenopodiaceae s.s.).
Proceedings of the Royal Society B 279 (1741), 3304e3311. http://dx.doi.org/
10.1098/rspb.2012.0440 1471e2954.
Karrenberg, S., Edelist, C., Lexer, C., Rieseberg, L., 2006. Response to salinity in the
homoploid hybrid species Helianthus paradoxus and its progenitors H. annuus
and H. petiolaris. New Phytologist 170 (3), 615e629.
Keighery, G.J., Halse, S.A., McKenzie, N.L., Harvey, M.S. (Eds.), 2004. A Biodiversity
Survey of the Western Australian Agricultural Zone. Western Australian
Museum, Perth, Records of the Western Australian Museum. Supplement 67,
384 p.
Kennedy, S.A., Ganf, G.G., Walker, K.F., 2003. Does salinity influence the distribution
of exotic willows (Salix spp.) along the Lower River Murray? Marine Freshwater
Research 54, 825e831.
Kershaw, A.P., 1978. Record of last interglacial cycle from north-eastern Queensland.
Nature 272, 159e162.
Kershaw, A.P., Moss, P., Van Der Kaars, S., 2003. Causes and consequences of long-
term climatic variability on the Australian continent. Freshwater Biology 48,
1274e1283.
Kier, G., Mutke, J., Dinerstein, E., Ricketts, T.H., Kuper, W., Kreft, H., Barthlott, W.,
2005. Global patterns of plant diversity and floristic knowledge. Journal of
Biogeography 32, 1107e1116.
Kreeb, K.H., Whalley, R.D.B., Charley, J.L., 1995. Some investigations into soil and
vegetation relationships associated with alkaline-saline soil surfaces in the
Walcha area, Northern Tablelands, New South Wales. Australian Journal of
Agricultural Research 46, 209e224.
Kriticos, D.J., Sutherst, R.W., Brown, J.R., Adkins, S.W., Maywald, G.F., 2003. Climate
change and the potential distribution of an invasive alien plant: Acacia nilotica
ssp. indica in Australia. Journal of Applied Ecology 40, 111e124.
Krull, E.S., Skjemstad, J.O., Burrows, W.H., Bray, S.G., Wynn, J.G., Bol, R., Spounder, L.,
Harms, B., 2005. Recent vegetation changes in Australia: evidence from d13C
and 14C analyses of soil organic matter. Geoderma 126, 241e259.
Ladiges, P.Y., Ariati, S.R., Murphy, D.J., 2006. Biogeography of the Acacia victoriae,
pyrifolia and murrayana species groups in arid Australia. Journal of Arid Envi-
ronments 66, 462e476.
Lauchli, A., Luttge, U., 2002. Salinity: EnvironmentePlantseMolecules. Kluwer Ac-
ademic Publishers.
Le Houerou, H.N., 2003. Bioclimatology and phytogeography of the Red Sea and
Aden Gulf Basins: a monograph (with a particular reference to the highland
evergreen sclerophylls and lowland halophytes). Arid Land Research and
Management 17, 177e256.
Lexer, C., Fay, M.F., 2005. Adaptation to environmental stress: a rare or frequent
driver of speciation? Journal of Evolutionary Biology 18, 893e900. http://
dx.doi.org/10.1111/j.1420-9101.2005.00901.x.
Liphschitz, N., Waisel, Y., 1974. Existence of salt glands in various genera of the
Gramineae. New Phytologist 73, 507e513.
Macphail, M.K., Hill, R.S., 2001. Fossil record of Acacia in Australia: Eocene to pre-
sent. In: Orchard, A.E., Wilson, A.J.G. (Eds.), Flora of Australia. Mimosaceae,
Acacia Part 1, vol. 11A. CSIRO Publishing, Melbourne, pp. 13e29.
Marcar, N.E., Ansari, R., Khanzada, A.N., Khan, M.A., Crawford, D.F., 2003. Perfor-
mance of several tree species on a saline site in southern Pakistan. Journal of
Tropical Forest Science 15, 457e468.
Martin, C. San, Subiabre, M., Ramirez, C., 2006. A floristic and vegetational study of
a latitudinal gradient of salt marshes in South-Central Chile. Ciencia e Inves-
tigacion Agraria 33 (1), 33e40.
Martin, H.A., 2006. Cenozoic climatic change and the development of the arid
vegetation in Australia. Journal of Arid Environments 66, 533e563.
Maslin, B.R., 2004. Botanical Annexe. In: Crespi, B.J., Morris, D.C., Mound, L.A. (Eds.),
Evolution of Ecological and Behavioural Diversity: Australian Acacia Thrips as
Model Organisms. Australian Biological Resources Study and Australian Na-
tional Insect Collection. CSIRO, Canberra, pp. 97e112.
Maslin, B.R., Miller, J.T., Seigler, D.S., 2003. Overview of the generic status
of Acacia (Leguminosae: Mimosoideae). Australian Systematic Botany 16,
1e18.
McLeish, M.J., Chapman, T.W., Schwarz, M.P., 2007. Host-driven diversification of
gall-inducing Acacia thrips and the aridification of Australia. BMC Biology 5, 3.
http://dx.doi.org/10.1186/1741-7007-5-3.
Miletto, M., Loy, A., Antheunisse, A.M., Loeb, R., Bodelier, P.L.E., Laanbroek, H.J., 2008.
Biogeography of sulfate-reducing prokaryotes in river floodplains. FEMS
Microbiology Ecology 64 (3), 395e406.
Miller, J.T., Murphy, D.J., Brown, G.K., Richardson, D.M., González-Orozco, C.E., 2011.
The evolution and phylogenetic placement of invasive Australian Acacia species.
Diversity and Distributions 17, 848e860. http://dx.doi.org/10.1111/j.1472-
4642.2011.00780.x.
Molina, J.A., Casermeiro, M.A., Moreno, P.S., 2003. Vegetation composition and soil
salinity in a Spanish Mediterranean coastal ecosystem. Phytocoenologia 33,
475e494.
Mueller-Dombois, D., Ellenberg, H., 1974. Aims and Methods of Vegetation Ecology.
John Wiley and Sons.
 E.N. Bui / Journal of Arid Environments 92 (2013) 14e25
Nind, M.A.P., 1988. Age of Campaspe Formation, north Queensland. Search 19 (1),
30e32.
Nix, H., 1994. Brigalow. In: Dovers, S. (Ed.), Australian Environmental History: Essays
and Cases. Oxford University Press, pp. 198e233.
Northcote, K.H., Skene, J.K.M., 1972. Australian Soils with Saline and Sodic Prop-
erties. CSIRO Soil Publ. No. 27. CSIRO, Melbourne.
Noy-Meir, I., 1974. Multivariate analysis of the semi-arid vegetation in south-eastern
Australia. II. Vegetation catenae and environmental gradients. Australian Jour-
nal of Botany 22, 115e140.
Orchard, A.E., Wilson, A.J.G. (Eds.), 2001. Flora of Australia. Mimosaceae, Acacia, vol.
11B. CSIRO Publishing, Melbourne.
Parida, A.K., Das, A.B., 2005. Salt tolerance and salinity effects on plants: a review.
Ecotoxicology and Environmental Safety 60, 324e349.
Perelman, S.B., Leon, R.J.C., Oesterheld, M., 2001. Cross-scale vegetation patterns of
Flooding Pampa grasslands. Journal of Ecology 89, 562e577.
Pfennig, D.W., Wund, M.A., Snell-Rood, E.C., Cruickshank, T., Schlichting, C.D.,
Moczek, A.P., 2010. Phenotypic plasticity’s impacts on diversification and spe-
ciation. Trends in Ecology & Evolution 25, 459e467.
Reed, D.N., Anderson, T.M., Dempewolf, J., Metzger, K., Serneels, S., 2009. The spatial
distribution of vegetation types in the Serengeti ecosystem: the influence of
rainfall and topographic relief on vegetation patch characteristics. Journal of
Biogeography 36, 770e782.
Reichman, S.M., Bellairs, S.M., Mulligan, D.R., 2006. The effects of temperature and
salinity on Acacia harpophylla (brigalow) (Mimosaceae) germination. Rangeland
Journal 28, 175e178. http://dx.doi.org/10.1071/RJ06027.
Rengasamy, P., 2006. World salinization with emphasis on Australia. Journal of
Experimental Botany 57, 1017e1023.
Richardson, J.L., Arndt, J.L., Freeland, J., 1994. Wetland soils of the Prairie Potholes.
Advances in Agronomy 52, 121e171.
Rieseberg, L.H., Raymond, O., Rosenthal, D.M., Lai, Z., Livingstone, K., Nakazato, T.,
Durphy, J.L., Schwarzbach, A., Donovan, L.A., Lexer, C., 2003. Major ecological
transitions in wild sunflowers facilitated by hybridization. Science 301, 1211.
Rogel, J.A., Silla, R.O., Ariza, F.A., 2001. Edaphic characterization and soil ionic
composition influencing plant zonation in a semi-arid Mediterranean salt
marsh. Geoderma 99, 81e98.
Runyon, C.W., D’Odorico, P., 2010. Ecohydrological feedbacks between salt accu-
mulation and vegetation dynamics: role of vegetation-groundwater in-
teractions. Water Resources Research 46 (W11561), 11.
Sage, R.F., 2001. Environmental and evolutionary preconditions for the origin and
diversification of the C4 photosynthetic syndrome. Plant Biology 3, 202e213.
http://dx.doi.org/10.1055/s-2001-15206.
Sawada, H., Kuykendall, L.D., Young, J.M., 2003. Changing concepts in the system-
atics of bacterial nitrogen-fixing legume symbionts. Journal of General and
Applied Microbiology 49, 155e179.
Schmiedel, U., Jurgens, N., 1999. Community structure on unusual habitat islands:
quartz-fields in the succulent Karoo, South Africa. Plant Ecology 142, 57e69.
Schofield, R.V., Kirkby, M.J., 2003. Application of salinization indicators and initial
development of global soil salinization scenario under climatic change. Global
Biogeochemical Cycles 17, 1078. http://dx.doi.org/10.1029/2002GB001935.
Seddon, J.A., Zerger, A., Doyle, S.J., Doyle, S.J., Briggs, S.V., 2007. The extent of dryland
salinity in remnant woodland and forest within an agricultural landscape.
Australian Journal of Botany 55, 533e540.
Scott, J.W., Meyer, S.E., Merrill, K.R., Anderson, V.J., 2010. Local population differ-
entiation in Bromus tectorum L. in relation to habitat-specific selection regimes.
Evolutionary Ecology 24, 1061e1080.
Shaltout, K.H., El-Halawany, E.F., El-Garawany, M.M., 1997. Coastal lowland vege-
tation of eastern Saudi Arabia. Biodiversity and Conservation 6, 1027e1044.
25
Silburn, D.M., Cowie, B.A., Thornton, C.M., 2009. The Brigalow Catchment Study
revisited: effects of land development on deep drainage determined from non-
steady chloride profiles. Journal of Hydrology 373, 487e498.
Silliman, B.R., Bertness, M.D., 2004. Shoreline development drives invasion of
Phragmites australis and the loss of plant diversity on New England salt
marshes. Conservation Biology 18, 1424e1434.
Stave, J., Oba, G., Bjora, C.S., Mengistu, Z., Nordal, I., Stenseth, N.C., 2003. Spatial and
temporal woodland patterns along the lower Turkwel River, Kenya. African
Journal of Ecology 41, 224e236.
Stewart, R.E., Kantrud, H.A., 1972. Vegetation of Prairie Potholes, North Dakota, in
Relation to Quality of Water and Other Environmental Factors. Geological Sur-
vey Professional Paper 585-D. U.S. Geological Survey, Denver.
Subbarao, G.V., Ito, O., Berry, W.L., Wheeler, R.M., 2003. Sodiumda functional plant
nutrient. Critical Reviews in Plant Sciences 22, 391e416.
Thrall, P.H., Broadhurst, L.M., Hoque, M.S., Bagnall, D.J., 2009. Diversity and salt
tolerance of native Acacia rhizobia isolated from saline and non-saline soils.
Austral Ecology 34, 950e963.
Thomas, C.D., Cameron, A., Green, R.E., Bakkenes, M., Beaumont, L.J.,
Collingham, Y.C., Erasmus, B.F.N., Ferreira de Siqueira, M., Grainger, A.,
Hannah, L., Hughes, L., Huntley, B., van Jaarsveld, A.S., Midgley, G.F., Miles, L.,
Ortegua-Huerta, M.A., Townsend Peterson, A., Phillips, O.L., Williams, S.E., 2004.
Extinction risk from climate change. Nature 427, 145e148.
Tomar, O.S., Gupta, R.K., Dagar, J.C., 1998. Afforestation techniques and evaluation of
different tree species for waterlogged saline soils in semiarid tropics. Arid Soil
Research and Rehabilitation 12, 301e316.
Toth, T., Rajkai, K., 1994. Soil and plant correlations in a solonetzic grassland. Soil
Science 157, 253e262.
United States Salinity Laboratory Staff, 1969. Diagnosis and Improvement of Saline
and Alkali Soils. Agriculture Handbook No. 60, Rev. ed. United States Depart-
ment of Agriculture, Washington, DC.
van Dijk, D.C., 1980. Salt and Soil Reaction Patterns in the Tara Brigalow Lands,
South-east Queensland. Division of Soils Divisional Report No. 47. CSIRO,
Melbourne.
Wardell-Johnson, G.W., Williams, J.E., Hill, K.D., Cumming, R., 1997. Evolutionary
biogeography and contemporary distribution of eucalypts. In: Williams, J.E.,
Woinarski, J.C.Z. (Eds.), Eucalypt Ecology: Individuals to Ecosystems. Cambridge
University Press, Cambridge, pp. 92e128.
Whittig, L.D., Janitsky, P., 1963. Mechanisms of formation of sodium carbonate in
soils. 1. Manifestations of biological conversions. Journal of Soil Science 14 (2),
322e333.
Williamson, D.R., 1986. The hydrology of salt affected soils in Australia. Reclamation
and Revegetation Research 5, 181e196.
Wolicka, D., Jarzynowska, L., 2012. Microbiological reduction of sulphates in salty
environments and mineralogical characterization of the transformation prod-
ucts. Geomicrobiology Journal 29 (6), 528e536.
Yang, C.W., Zhao, N., Xu, C.M., Liu, B., Shi, D.C., 2012. Regulation of ion homeostasis
in rice subjected to salt and alkali stresses. Australian Journal of Crop Sciences 6,
724e731.
Zahran, H.H., 2001. Rhizobia from wild legumes: diversity, taxonomy, ecology, ni-
trogen fixation and biotechnology. Journal of Biotechnology 91, 143e153.
Zhang, X.P., Harper, R., Karsisto, M., Lindstrom, K., 1991. Diversity of Rhizobium
bacteria isolated from the root nodules of leguminous trees. International
Journal of Systematic Bacteriology 41, 104e113.
Zheng, J.F., Liu, G.R., Zhu, W.F., Zhou, Y.G., Liu, S.L., 2004. Phylogenetic clusters of
rhizobia revealed by genome structures. Science in China Series C e Life Sci-
ences 47, 268e278.
Zhu, J.K., 2001. Plant salt tolerance. Trends in Plant Science 6, 66e71.