Freshwater Biology (2006) 51, 206–223 doi:10.1111/j.1365-2427.2005.01484.

Flooding, soil seed bank dynamics and vegetation

resilience of a hydrologically variable desert floodplain
S A M A N T H A J . C A P O N * , † A N D M A R G A R E T A . B R O C K †, ‡
*Centre for Riverine Landscapes, Griffith University, Nathan, Qld, Australia
†
Co-operative Research Centre for Freshwater Ecology, Canberra, ACT, Australia
‡
NSW Department of Natural Resources, University of New England, Armidale, NSW, Australia

SU M M A R Y
1. This paper explores soil seed bank composition and its contribution to the vegetation
dynamics of a hydrologically variable desert floodplain in central Australia: the Cooper
Creek floodplain. We investigated patterns in soil seed bank composition both temporally,
in response to flooding (and drying), and spatially, with relation to flood frequency.
Correlations between extant vegetation and soil seed bank composition are explored with
respect to flooding.
2. A large and diverse germinable soil seed bank was detected comprising predominantly
annual monocot and annual forb species. Soil seed bank composition did not change
significantly in response to a major flood event but some spatial patterns were detected
along a broad flood frequency gradient. Soil seed bank samples from frequently flooded
sites had higher total germinable seed abundance and a greater abundance of annual
monocots than less frequently flooded sites. In contrast, germinable seeds of perennial
species belonging to the Poaceae family were most abundant in soil seed bank samples
from rarely flooded sites.
3. Similarity between the composition of the soil seed bank and extant vegetation increased
following flooding and was greatest in more frequently flooded areas of the floodplain,
reflecting the establishment of annual species. The results indicate that persistent soil seed
banks enable vegetation in this arid floodplain to respond to unpredictable patterns of
flooding and drying.

Keywords: arid zone, Australia, drought, germination, wetland plants

shifts in composition in response to fluctuating

Introduction
The salient feature of freshwater ecosystems in desert
regions is hydrological variability (Davies et al., 1994;
Williams, 1995; Puckridge et al., 1998). As a result of
variable flooding and drying regimes, desert rivers
and their associated floodplains and wetlands are
characterised by high levels of temporal and spatial
heterogeneity (Stromberg, 2001). Vegetation in these
environments is usually dynamic, exhibiting dramatic
Correspondence: Samantha J. Capon, School of Biological
Sciences, Monash University, Clayton Campus, Clayton,
Vic. 3800, Australia.
Email: samantha.capon@sci.monash.edu.au
206
hydrological conditions (Capon, 2003) and often
displays clear spatial patterns related to flood history
(Carr, 1998; Stromberg, 2001; Capon, 2005). Persistent
soil seed banks can facilitate such plant community
dynamics by allowing species to persist in a dormant
state until suitable conditions for their germination
and establishment occur (Parker, Simpson & Leck,
1989; Bonis, Lepart & Grillas, 1995; Baskin & Baskin,
1998) thus affording vegetation in these variable
environments some degree of resilience (Brock et al.,
2003).
Numerous recent studies have demonstrated the
critical role of flooding and drying in the development
of plant communities germinating from soil seed

2005 Blackwell Publishing Ltd

banks of variable wetland habitats (Seabloom, van Der
Valk & Moloney, 1998; Casanova & Brock, 2000;
Crosslé & Brock, 2002; Nicol, Ganf & Pelton, 2003;
Warwick & Brock, 2003). As germination in wetland
plants often occurs in response to specific hydro-
logical conditions, such as flood timing or depth
(Leck, 1989; Leck & Brock, 2000), flooding can be
perceived as an ‘environmental sieve’ through which
vegetation composition is determined as a function of
differences in germination cues between species resi-
ding in the soil seed bank (van der Valk, 1981).
Flooding also affects other life history stages of
wetland plants by influencing growth rates or repro-
ductive vigour (Gerritsen & Greening, 1989; Coops &
van der Velde, 1995; Coops, van den Brink & van der
Velde, 1996; Warwick & Brock, 2003) or through
hydrochoric (i.e. via floodwaters) propagule dispersal
(Blom & Voesenek, 1996; Cronk & Fennessy, 2000).
Each of these life history stages can in turn influence
patterns of soil seed bank depletion and replenish-
ment. Consequently, flooding has the potential to
structure wetland soil seed banks both temporally and
spatially as well as determine their contribution,
through germination, to the extant vegetation.
Despite the attention given to successional fluctua-
tions in extant wetland vegetation (Keddy, 2000),
remarkably few studies have investigated temporal
dynamics of wetland soil seed banks, particularly
with respect to the effects of flooding. Bonis et al.
(1995) observed marked interannual fluctuations in
seed densities in two Mediterranean temporary mar-
shes over a period of 4 years and, in studies of North
American prairie marshes, sharp increases in seed
densities of some opportunistic species have been
observed between years in association with periods of
hydrological drawdown (van der Valk & Davis, 1979;
Smith & Kadlec, 1983; Poiani & Johnson, 1989).
However, Poiani & Johnson (1989) found that the
composition of the bulk of a prairie marsh soil seed
bank remained fairly constant between years despite
the occurrence of a drawdown event. Some evidence
of reduced seed densities following flood-induced
recruitment has been reported though for salt marsh
species (Egan & Ungar, 1999). In contrast, studies of
spatial patterns in soil seed bank composition have
been conducted in wetlands from a wide range of
climatic regions and hydrological regimes (Leck,
1989). Within wetland systems, lower seed densities
tend to occur in areas which are most permanently

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Soil seed banks of an arid floodplain 207
inundated (Keddy & Reznicek, 1982; Leck, 1989) and
several cases of declining seed density with increasing
water depth have also been recorded (Poiani &
Johnson, 1989; Wilson, Moore & Keddy, 1993).
However, Bonis et al. (1995) reported higher seed
densities in the centre of Mediterranean marshes and
decreasing densities towards their edges. Seed density
was not found to vary significantly along elevational
gradients in playa wetlands (Haukos & Smith, 1994).
Similarly, Brock & Rogers (1998) did not detect any
differences in seed densities or species richness
between the seed banks of wetlands with varying
flow regimes on a South African ephemeral flood-
plain.
The contribution of soil seed banks to extant
vegetation dynamics also varies considerably between
wetland systems. In freshwater tidal wetlands of
North America, for example, there is a high concor-
dance between the composition of extant plant com-
munities and their soil seed banks (Leck & Simpson,
1987, 1995) and the soil seed banks of prairie marshes
contain suites of species which appear in the extant
vegetation during different hydrological phases (van
der Valk & Davis, 1978). In contrast, the composition
of soil seed banks in some North American lake
shores and inland marshes resembles that of draw-
down vegetation rather than that which is prevalent
under submerged conditions (Leck, 1989). In many
other cases, little similarity exists between soil seed
banks and extant plant communities, particularly
where seed banks are overwhelmingly dominated
by one or two species which are absent from the
vegetation or vice versa (Thompson & Grime, 1979;
McGraw, 1987; Hughes & Cass, 1997).
Australia’s large desert rivers are amongst the most
hydrologically variable river systems in the world
(Puckridge et al., 1998) and many are associated with
extensive floodplains which experience both long
periods of drought and huge flood pulses that
transform vast areas into wetlands, often for months
at a time. Vegetation in these heterogeneous habitats
exhibits significant temporal shifts in composition and
structure in response to flooding and drying (Capon,
2003) and varies spatially with relation to flood
history (Capon, 2005). It might be expected, therefore,
that many plant species inhabiting these desert
floodplains maintain large and long-lived soil seed
banks that enable persistence through such variable
hydrological conditions. Although deserts have been

208 S.J. Capon and M.A. Brock
the subject of many soil seed bank investigations (e.g.
Freas & Kemp, 1983; Henderson, Petersen & Redak,
1988; Pake & Venable, 1996), studies of large wetland
systems within Australia’s desert regions are rare and,
amongst the wetland soil seed bank literature, studies
of riverine and floodplain systems are also lacking
(Goodson et al., 2001). To address this knowledge gap,
this study examines the soil seed bank composition of
the Cooper Creek floodplain in the arid Lake Eyre
Basin of central Australia. In particular, this paper
investigates temporal and spatial patterns in soil seed
bank composition with relation to flooding and
explores the contribution of the soil seed bank to
flow-related vegetation dynamics. Results are presen-
ted from germination studies of soil seed bank
collections made along a broad flood frequency
gradient before and after a major flood event and
again following a period of drought. The specific
questions addressed in this paper are:
1 What is the composition of the germinable soil
seed bank in this desert floodplain and does this vary
temporally in response to flooding (or drying)?
2 Does the composition of the germinable soil seed
bank vary spatially in relation to flood frequency? and
3 How does soil seed bank composition compare
with that of the extant vegetation and does this vary
temporally or spatially as a result of flooding?
Methods
Study area
The Cooper Creek catchment is located in the Lake
Eyre Basin of central Australia and is approximately
296 000 km2 in area (Fig. 1). The majority of the
catchment is hot, with summer temperatures ranging
between 25 and 40 C and winter temperatures
between 5 and 25 C, and arid, with mean annual
rainfall c. <400 mm. Surface water discharge in the
Cooper Creek is extremely variable and historically
includes many periods of low or no flow (Fig. 2),
during which time the river exists as a string of
disconnected waterholes. Flood pulses in the system
are driven predominantly by northern cyclone activity
and are variable in their timing, magnitude and
duration. A complex channel network combined with
the catchment’s low topographic gradient facilitate
the inundation of vast areas of floodplain following
high flow events. In April 1990, for example, almost
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26 000 km2 were inundated south of Windorah as a
result of a relatively rare, large flood pulse with a peak
discharge of 1 460 000 ML day)1 and a peak height of
7.95 m (Queensland Department of Natural Re-
sources, 1998). By way of comparison with the
historical flood record (Fig. 2), the 1974 flood peak,
for which the greatest ever instantaneous flow in the
catchment was recorded, had a peak height of 8.48 m
at Windorah (Queensland Department of Natural
Resources, 1998). In contrast, the comparatively small
flood pulse which occurred in December 1984 had a
peak height of 3.9 m, a peak discharge of
26 000 ML day)1 at Currareva and inundated an area
of approximately 3400 km2 (Queensland Department
of Natural Resources, 1998). According to anecdotal
evidence, such events generally inundate areas close
to channels near Windorah about once every 2 years.
Conversely, areas on the periphery of the floodplain
may be inundated less than once every 10 years and
only then by events of extreme magnitude such as
those which occurred in 1974 or 1990.
Vegetation on the floodplain is mainly short grass
and forb associations which fluctuate in composition
in response to hydrological conditions (Capon, 2003).
Open succulent shrubland dominated by Chenopodium
auricomum Lindl. (Queensland bluebush) and
Muehlenbeckia florulenta Meisn. (tangled lignum) also
occurs in more frequently flooded areas. Trees, e.g.
Eucalyptus camaldulensis Dehnh. (river red gum),
Eucalyptus coolabah Blakely & Jacobs (coolibah) and
Acacia cambagei R.T. Baker (gidgee), are mostly
restricted to the margins of channels.
Data collection
The study was conducted on the Cooper Creek
floodplain in the vicinity of Windorah (Fig. 1). Due
to the lack of detailed hydrological data for the
catchment, local landholders were consulted to iden-
tify floodplain areas possessing broadly different
flood frequencies. These included a near-channel
zone, flooded approximately once every 2 years, a
mid-floodplain zone, comprising areas flooded
approximately once every 5 years and a far-floodplain
zone which consisted of areas only flooded once every
10 years or less frequently. Twelve sites, each com-
prising a 50 m2 quadrat, were then selected with four
sites haphazardly located in each of the three broad
flood frequency zones. Sampling was conducted on

2005 Blackwell Publishing Ltd, Freshwater Biology, 51, 206–223
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Soil seed banks of an arid floodplain 209

er
riv Longreach
n
so
m
Tho
Currareva gauging er
station o riv
r co
Ba

Windorah

South Australia

Cooper Creek

Queensland
Lake
Eyre

0 100 200 km

Fig. 1 The Cooper Creek catchment. Inset shows location in central Australia. Latitude and longitude for Windorah are )25.4228S and
142.6564E respectively.

three occasions, February, May and October, during
the year 2000. A major flood event, comparable in size
to the April 1990 event, inundated all sites between
the first and second sampling dates and a period of
drought preceded the third sampling date (Fig. 3). It
is also worth noting that conditions prior to the
commencement of the survey were relatively moist,
particularly in the near-channel zone, as both rainfall
and a small flood event had occurred in late 1999.
At each survey time, five soil cores of 0.055 m
diameter and 10 cm depth were collected from each
site and aggregated in a single plastic bag to form a
sample. Individual soil cores were collected from
random locations within the 50 m2 quadrat to increase

2005 Blackwell Publishing Ltd, Freshwater Biology, 51, 206–223
the likelihood of detecting species with patchy seed
distributions (Bigwood & Inouye, 1988; Blanch &
Brock, 1994; Brock, Theodore & O’Donnell, 1994). A
list of species occurring in the extant vegetation
within each 50 m2 quadrat was also compiled on each
sampling occasion. Soil samples were then transpor-
ted to Griffith University and dried and stored at
room temperature until germination trials com-
menced.
Soil seed bank composition was examined using the
seedling emergence method. Unlike manual sifting
and counting of seeds, this method does not allow for
a complete assessment of seed abundance (Hutchings
& Russell, 1989). However, total seed count methods
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210 S.J. Capon and M.A. Brock
25 1998). In this case, the seedling emergence method
was considered appropriate for determining the
20
Discharge (mL × 106)

composition of the germinable soil seed bank and

15 detecting temporal and spatial trends within this. For
the germination trials, each aggregated sample was
10 divided evenly between four plastic trays,
17 cm · 12 cm, with drainage holes, providing a soil
5
depth of approximately 2 cm and thereby increasing
0 the surface area available for germination. All 144
1958

1961

1964

1967

1970

1973

1976

1979

1982

1985

1988
trays were then randomly arranged on tables in a
glasshouse at Griffith University. A pilot study indi-
Fig. 2 Annual discharge of Cooper Creek between 1958 and cated that fewer species germinated under flooded
1988 recorded at Currareva gauging stating near Windorah (see
conditions than moist, so a single watering treatment
Fig. 1). As this gauging station was closed following this period,
no more recent discharge data is available. (Data obtained from was selected where trays were kept moist by means of
the Queensland Department of Natural Resources). an automatic watering system operating four times
daily. Species which require complete submergence
for germination could be missed under a moist
(a) 160
watering treatment; however, these were not observed
in the pilot study which examined germination from
Rainfall (mm)

120
the soil seed bank under a range of watering
80
treatments including submergence for up to 3 months
40 duration. ter Heerdt, Schutter & Bakker (1999) and
Boedeltje, ter Heerdt & Bakker (2002) support this use
0
of a single watering treatment determined through
(b) pilot studies when experiments are constrained by
6 time and space. Trays were rotated at regular intervals
Discharge (mL × 106)

5
4
3
2
1
0
O D F A
1999 2000
Fig. 3 (a) Total monthly rainfall (mm) at Windorah (source:
Bureau of Meteorology) and (b) total monthly discharge (mil-
lions of ML) at Longreach from October 1999 to December 2000
(source: Queensland Department of Natural Resources). Arrows
indicate sampling times. The Currareva gauging station (see
Fig. 1) was closed in 1988, hence the flow data from upstream on
the Thomson River are the best available for the study period.
Discharge in the Cooper Creek is determined principally by
precipitation associated with northern monsoons and local
rainfall rarely results in floodplain inundation. Rainfall is pre-
sented here to illustrate the period of local drought between the
second and third sampling dates rather than to demonstrate any
relationship with discharge.
are time consuming, have the potential to overlook
species with very small seeds and provide no
information on seed viability (Baskin & Baskin,
to ensure an even distribution of watering and were
kept in the glasshouse for 1 year to allow exposure to
a wide range of temperatures. Germination trials for
samples from the first and second sample times
commenced in August 2000, following completion of
J A O D
the pilot study, and those from the third survey in late
October 2000. Temperatures in the glasshouse over
this period ranged between 20 and 45 C during
summer and 10 and 35 C in the winter months. As
seedlings germinated from the trays, they were
recorded and removed once their identification was
possible.
Data analysis
Seedling count data for the four trays from each
sample were pooled to give an abundance value for
each species at each site for each survey time. Species
richness was considered to be the total number of
species occurring within these pooled samples and
total abundance was calculated by summing all
species abundances from each pooled sample. Species
were also assigned to groups on the basis of life form

2005 Blackwell Publishing Ltd, Freshwater Biology, 51, 206–223

(i.e. shrubs, sub-shrubs, forbs and monocots) and life
span (i.e. annual or perennial) using descriptions
contained in Cunningham et al. (1992), resulting in a
total of seven plant groups; shrubs, annual sub-
shrubs, perennial sub-shrubs, annual forbs, perennial
forbs, annual monocots and perennial monocots. The
use of plant functional groups defined on the basis of
how plants germinate, establish and reproduce in
relation to surface water (i.e. submerged, amphibious
and terrestrial) has provided a fruitful approach for
considering the spatial and temporal dynamics of
temporary wetland plant communities (Brock &
Casanova, 1997). However, the autecological informa-
tion necessary to assign many species to such
previously defined functional groups is lacking for
many plants of the Cooper Creek floodplain. There-
fore, following Higgins, Rogers & Kemper (1997),
groups based on life form and life span were used
under the assumption that plants within such groups
are likely to utilise resources and respond to stressors
in a comparable manner. This approach has also been
successfully employed in a previous study of spatial
patterns in Cooper Creek floodplain vegetation
(Capon, 2005). The abundance of seeds belonging to
each of these groups was then calculated for each
sample. This was also done at the family level within
each of these major groups.
Abundance values were transformed, [log10(x + 1)],
to meet the requirements of analysis of variance
(A N O V A ) (Zar, 1999). Untransformed species richness
and transformed abundance variables were then
analysed by two-way A N O V A to determine the effects
of sample time (February, May or October) and flood
frequency zone (near-channel, mid-floodplain and far-
floodplain). Significantly different means were deter-
mined using Tukey’s b-test. Univariate analyses were
all conducted in SPSS (SPSS, 2001).
Further exploration of temporal and spatial patterns
in soil seed bank composition was achieved by
performing a semi-strong hybrid (SSH) ordination
on the untransformed species abundance data using
Bray–Curtis dissimilarity measures. These multivari-
ate analyses were performed in PATN (Belbin, 1995).
Three dimensions were used for the ordination and
default options were selected throughout. The ‘good-
ness of fit’ of SSH ordinations can be assessed via a
stress value computed by PATN in conjunction with a
solution. Stress for the ordinations presented here
were below the recommended upper value of 0.2

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Soil seed banks of an arid floodplain 211
(Belbin, 1995). Principal axis rotation and correlation
and Monte-Carlo randomisation procedures were also
conducted to determine species vectors correlating
significantly with the ordination space.
Comparison of soil seed bank and extant vegetation
composition were initially made by means of a further
SSH ordination based on a matrix of species presence
and absence data. Additionally, Sorenson’s SI were
calculated for each site and survey time using the
formula SI ¼ 2c/a + b, where c was the number of
species common between the soil seed bank and
extant vegetation and a and b were the total numbers
of species in the soil seed bank and extant vegetation
respectively (Sorenson, 1948). Similarity indices were
converted into percentages by multiplying by 100 and
effects of flood frequency zone and sample time on
similarity indices were examined using two-way
ANOVA.
Results
Soil seed bank composition
Over the duration of the germination trials, a total of
5483 seedlings germinated from the soil seed bank
samples with 2011, 2064 and 1408 germinating from
samples from the first, second and third sample times
respectively. Of these, 56 species from at least 22
families were present (four forb species and one
annual grass species could not be identified) (Appen-
dix). Annual monocots were the most abundant group
and represented over half of the total seedlings
recorded. Annual forbs were the next most abundant
group (35%), followed by perennial monocots (11%)
and perennial forbs (4%). No shrub or sub-shrub
seedlings germinated during the trials. At the family
level, seedlings belonging to the Poaceae family in the
annual monocot group were the most abundant and
the annual grass, Eragrostis tenellula, was the most
common species, germinating in samples from all
flood frequency zones and sample times (Table 1).
Asteraceae seedlings in the annual forb group, e.g.
Calotis hispidula, Centipeda minima and Pseudognapha-
lium luteoalbum, were the second most abundant
family germinating during the trials. The annual
monocot Cyperus difformis and the perennial monocot
Cyperus bifax, both belonging to the Cyperaceae
family, were also frequently recorded from most
samples as were the annual forbs Ammania multiflora
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212 S.J. Capon and M.A. Brock
Table 1 Common species germinating from soil seed bank samples in each flood frequency zone at each sample time

February May October

Species NC MF FF NC MF FF NC MF FF

Alternanthera nodiflora d d d d
Ammania multiflora d d d d d d d
Arabidella eremingena d
Aristida contorta d d d d d d
Calotis hispidula d d d d d
Centaurium spicatum d d d
Centipeda minima d d d
Chloris pectinata d d
Cullen cinereum d d d
Cyperus bifax d d d d d d d d d
Cyperus difformis d d d d d d
Daucus glochidiatus d d d d d d
Echinochloa turneriana d
Eragrostis tenellula d d d d d d d d d
Eriochloa crebra d
Haloragis aspera
Iseilema sp. d
Mimulus gracilis d d d d d d d d
Panicum decompositum d d
Phyllanthus virgatus d
Plantago cunninghamii d
Pseudognaphalium luteoalbum d d d d
Spergularia rubra d
Trigonella suavissima d
Wahlenbergia gracilis d d d

Symbols indicate that species was amongst the 10 most abundant germinating from these samples (families for each species are listed
in the Appendix).
NC, near-channel zone; MF, mid-floodplain zone; FF, far-floodplain zone.

(Lythraceae), Alternanthera nodiflora (Amaranthaceae) near-channel zone. Furthermore, the abundance of

and Daucus glochidiatus (Apiaceae).
Temporal and spatial patterns in soil seed bank
composition
Sample time did not have a significant effect on either
species richness or total seed abundance of the
germinable soil seed bank, although seed abundance
did tend to decline in samples from the near-channel
zone at the third sample time (Fig. 4, Table 2).
Samples from the near-channel zone also had signi-
ficantly higher total seed abundance than those from
the mid-floodplain or far-floodplain zones (Fig. 4,
Table 2). However, species richness did not differ
significantly with flood frequency zone (Fig. 4, Ta-
ble 2). None of the major plant groups exhibited
significant changes in seed abundance between sam-
ple times, but a reduction in annual monocots and
forbs was apparent in the third sample time for the
both monocot groups differed significantly between
flood frequency zones with annuals occurring most
frequently and perennials least frequently in samples
from the near-channel zone (Fig. 5, Table 2).
At the family level, sample time had a significant
effect on seed abundance of three annual forb families
(Table 3). The abundance of Fabaceae seedlings was
significantly lower in samples from the second survey
time and Asteraceae seedlings germinated least fre-
quently and Portulacaceae seedlings most frequently
in samples from the third sample time (Fig. 6,
Table 3). In the perennial monocot group, seedlings
belonging to the Poaceae family were also signifi-
cantly more abundant in samples from the third
sample time and were present only in samples from
the far-floodplain zone during the first and second
sample times (Fig. 6, Table 3). Seed abundances of
nine other families differed significantly between
flood frequency zones including the annual forb

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(a) 20
Species richness ± SE
15
10
5 zone and the grasses Aristida contorta, Chloris pectinata
0
(b)
Near channel
300 Mid floodplain
Seed abundance ± SE
Far floodplain
150
0 Comparison of the soil seed bank and extant vegetation
Feb May Oct
Sample time
Fig. 4 (a) Mean species richness and (b) mean total seed abun-
dance of soil seed bank samples from each flood frequency zone
at each sample time. Error bars indicate standard error.
families Fabaceae, Campanulaceae and Caryophylla-
ceae which all exhibited significantly higher abun-
dances in samples from the near-channel zone (Fig. 6,
Table 3). Conversely, Amaranthaceae seedlings were
most abundant in samples from the far-floodplain
zone and Lythraceae seedlings increased in abun-
dance with decreasing flood frequency (Fig. 6, Ta-
ble 3). Seedlings belonging to the Haloragaceae family
in the perennial monocot group were significantly
more abundant in the mid-floodplain samples (Fig. 6,
Table 3). Amongst the monocots, both annual and
perennial species belonging to the Cyperaceae family
germinated least frequently in samples from the near-
channel zone while those belonging to the Poaceae
family in the annual monocot group were significantly
more abundant in these samples (Fig. 6, Table 3).
Clear temporal changes in overall soil seed bank
composition were not evident from the ordination but
samples did separate in the ordination space with
relation to flood frequency zone (Fig. 7). The compo-
sition of the germinable soil seed bank from the far-
floodplain zone appears to be particularly distinctive
whilst considerable overlap is evident in the compo-
sition of samples from the near-channel and mid-
floodplain zones. This spatial differentiation of soil

2005 Blackwell Publishing Ltd, Freshwater Biology, 51, 206–223
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Soil seed banks of an arid floodplain 213
seed bank composition correlated strongly with the
abundance of common annual monocot species with
high abundances of the grasses E. tenellula and Iseilema
sp. correlating with the position in the ordination
space of samples from the near-channel zone, the
sedge C. difformis with those from the mid-floodplain
and Eragrostis parviflora with far-floodplain samples
(Fig. 7, Table 4). Amongst the common annual forb
species, Calotis hispidula (Asteraceae) correlated with
samples from the near-channel zones, Trigonella suav-
issima (Fabaceae) and D. glochidiatus (Apiaceae) with
the mid-floodplain zone and Alternanthera nodiflora
and Centipeda minima (Asteraceae) with the far-flood-
plain zone (Fig. 7, Table 4).
A total of 99 species were recorded in both the soil
seed bank germination trials and the extant vegetation
surveys across all three sample times. Of these, 39
species were common to both (Appendix). Seventeen
species were observed only from the soil seed bank,
including the five unidentified species which germi-
nated in very low numbers. The remainder included
five annual monocot species [Cyperus pygmaeus and
an unknown Cyperus spp. (Cyperaceae) and Aristida
contorta, Elytrophorus spicatus and Eragrostis basedowii
(Poaceae)], five annual forb species [C. minima and
Pterocaulon sphacelatum (Asteraceae), Eryngium plant-
agineum (Apiaceae), Arabidella eremigena (Brassicaceae)
and Mollugo cerviana (Molluginaceae)], one perennial
monocot [Paspalidium jubiflorum (Poaceae)] and one
perennial forb species [Minuria denticulata (Astera-
ceae)]. A much higher number of species (43) was
observed only in the extant vegetation. Almost two
thirds of these were perennial species belonging to the
perennial monocot, perennial forb, perennial sub-
shrub and shrub groups. In particular, sub-shrubs
(e.g. Sclerolaena spp. and Atriplex spp.) and shrubs,
including the dominant species Chenopodium aurico-
mum and M. florulenta, were recorded only from the
extant vegetation (Appendix).
Soil seed bank samples tended to resemble each
other in composition more than they did the extant
plant communities at corresponding locations or
sample times. The ordination of species presence
and absence data indicates that the overall composi-
tion of soil seed bank samples was quite distinct from
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214 S.J. Capon and M.A. Brock
Table 2 Summary of two-way A N O V A s
Homogeneous investigating effects of sample time and
subsets for flood flood frequency zone on species richness,
frequency zone† total seed abundance and the abundance
of seeds in each major plant group
Source d.f.* F P NC MF FF

Species richness
Time 2 0.034 0.966
Flood frequency zone 2 0.114 0.893 a a a
Time · flood frequency zone 4 0.233 0.917
Total seed abundance
Time 2 0.930 0.407
Flood frequency zone 2 4.108 0.028 a b ab
Time · flood frequency zone 4 1.519 0.225
Annual forbs
Time 2 1.189 0.320
Flood frequency zone 2 2.450 0.105 a a a
Time · flood frequency zone 4 2.269 0.088
Annual monocots
Time 2 1.044 0.366
Flood frequency zone 2 5.518 0.010 a b b
Time · flood frequency zone 4 0.599 0.667
Perennial forbs
Time 2 1.519 0.237
Flood frequency zone 2 0.815 0.453 a a a
Time · flood frequency zone 4 0.485 0.746
Perennial monocots
Time 2 2.565 0.095
Flood frequency zone 2 6.296 0.006 a b b
Time · flood frequency zone 4 0.0.75 0.989

*Numerator only is reported here as in all tests the denominator ¼ 27.

†
Determined by Tukey’s b test: NC, near-channel; MF, mid-floodplain; FF, far-flood-
plain.

that of extant vegetation samples (Fig. 8). The com-
position of extant plant communities shifted substan-
tially following flooding and subsequent drying, as
illustrated by the trajectories of samples within the
ordination space. Extant vegetation composition
appeared to become more similar to that of the soil
seed bank over this time which is particularly appar-
ent in the position within the ordination space of
extant vegetation samples from the mid-floodplain
and far-floodplain zones at the third sample time
(Fig. 8). The similarity of soil seed bank samples and
their corresponding extant plant communities, as
indicated by Sorenson’s similarity index, was signifi-
cantly influenced by both sample time (F2,27 ¼ 10.046,
P ¼ 0.001) and flood history zone (F2,27 ¼ 4.182, P ¼
0.026) (Fig. 9). Although significant interaction be-
tween these factors was found (F2,27 ¼ 2.960, P ¼
0.038), similarity clearly increased following flooding
in samples from all flood frequency zones and, in
the far-floodplain samples, increased again at the
third sample time after drying. At all times, simi-
larity between the soil seed bank and extant veget-
ation tended to be higher in samples from the
near-channel zone and lowest in those from the far-
floodplain zone.
Discussion
Soil seed bank composition
The results indicate that the Cooper Creek floodplain
has a large and diverse soil seed bank composed
predominantly of seeds belonging to annual monocot
and forb species. The dominance of graminoid species
observed here is typical of soil seed banks in tempor-
ary wetlands as is the absence of woody species (Leck,
1989; LaDeau & Ellison, 1999; Rossell & Wells, 1999).
It is possible that viable seeds of some species not
observed as seedlings during this study were present
in the soil seed bank but their germination require-

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Soil seed banks of an arid floodplain 215
(a) 250 Annual forbs
Annual monocots
Temporal and spatial patterns in soil seed bank
Seed abundance ± SE

200
Perennial forbs
composition
150 Perennial monocots
The major flood event during this study had little
100 effect on the composition of the germinable soil seed
50 bank despite evidence in the extant vegetation of
extensive germination following inundation (Capon,
0
2003). Annual species belonging to the Fabaceae
(b) 250 family were the sole group for which a significant
decline in seed abundance was observed directly after
Seed abundance ± SE

200
flooding and, following the subsequent drying period,
150 seed numbers amongst these species had returned to
100 their previous levels. Wetland plants often have fast
50
growth rates and rapid life cycles, especially in
response to flooding (Blom & Voesenek, 1996).
0
Consequently, local replenishment of the soil seed
(c) 250 bank may have occurred in other species prior to the
second sample time. Further additions to the soil seed
Seed abundance ± SE

200
150
100
50
0 (Brock & Rogers, 1998; Brock, 1998; Leck & Brock,
Feb May Oct
Sample time
Fig. 5 Mean seed abundance in major plant groups for each
sample time in (a) near-channel zone sites, (b) mid-floodplain
zone sites and (c) far-floodplain zone sites. Error bars indicate
standard error.
ments were not met under glasshouse conditions.
However, due to the long incubation period of
sediments through a range of temperatures and the
total absence of shrub or sub-shrub seedlings in the
germination trials, it seems unlikely that species in
these groups could rely on persistent soil seed banks
for recruitment in the Cooper Creek floodplain.
Conversely, the persistence of many short-lived
annual monocot and forb species in this unpredictable
environment is probably facilitated largely by abun-
dant soil seed banks. The actual size of the viable soil
seed bank is likely to have been underestimated by
the seedling emergence method employed here as, in
many wetland plants, only a fraction of viable seeds
germinate in response to any one germination event
leaving substantial stores of seed from year to year
despite depletion by germination (Brock, 1998; Brock
& Rogers, 1998; Leck & Brock, 2000).

2005 Blackwell Publishing Ltd, Freshwater Biology, 51, 206–223
bank before this sample time are likely to have
resulted from widespread seed dispersal by flood-
waters. The lack of temporal change in response to
flooding may also reflect the low proportion of seeds
expected to germinate in a single germination event
2000; Haukos & Smith, 2001).
However, some differences in soil seed bank com-
position were apparent between the second and third
sample times, following the period of drying. These
included a decline in the abundance of annual forbs
and monocots germinating from near-channel zone
samples, although propagules of these groups were
probably present in higher numbers in samples from
the earlier survey times as a result of germination and
seed production following the small flood which
occurred prior to the survey period. It is likely that the
subsequent reduction occurred as a result of scouring
which could be expected to be greater in this zone.
Perennial grasses and annual forbs belonging to the
Portulacaceae family also increased in abundance in
samples from the third sample time from all flood
frequency zones. Such increases may have resulted
from wind dispersal during the intervening period
and could reflect the timing of seed fall amongst adult
plants in these species. However, the absence of
germinable perennial grass seeds in samples from the
near-channel and mid-floodplain zones during the
first and second sample times suggests that soil seed
banks in these species may be transient rather than
long lived and persistent (Thompson & Grime, 1979).
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216 S.J. Capon and M.A. Brock
Table 3 Summary of two-way A N O V A s
Homogeneous investigating effects of sample time and
subsets for flood
flood frequency zone on seed abundance
frequency zone†
of forb and monocot families. Only those
Source d.f.* F P NC MF FF forb families where a significant effect
(P < 0.05) was found are shown.
Annual forbs
Amaranthaceae
Time 2 0.156 0.857
Flood frequency zone 2 14.218 <0.001 b a c
Time · flood frequency zone 4 0.501 0.735
Asteraceae
Time 2 6.632 0.005
Flood frequency zone 2 1.397 0.265 a a a
Time · flood frequency zone 4 1.467 0.240
Campanulaceae
Time 2 2.493 0.102
Flood frequency zone 2 31.069 <0.001 a b b
Time · flood frequency zone 4 1.280 0.302
Caryophyllaceae
Time 2 0.771 0.473
Flood frequency zone 2 5.565 0.009 a b b
Time · flood frequency zone 4 0.320 0.862
Fabaceae
Time 2 7.005 0.004
Flood frequency zone 2 14.596 <0.001 a b b
Time · flood frequency zone 4 0.829 0.518
Lythraceae
Time 2 1.463 0.249
Flood frequency zone 2 5.439 0.010 a ab b
Time · flood frequency zone 4 1.546 0.217
Portulacaceae
Time 2 4.821 0.016
Flood frequency zone 2 0.916 0.412 a a a
Time · flood frequency zone 4 1.141 0.358
Perennial forbs
Haloragaceae
Time 2 1.632 2.14
Flood frequency zone 2 4.710 0.018 ab b b
Time · flood frequency zone 4 2.078 0.111
Annual monocots
Cyperaceae
Time 2 0.951 0.399
Flood frequency zone 2 11.527 <0.001 a b b
Time · flood frequency zone 4 0.492 0.742
Poaceae
Time 2 0.355 0.704
Flood frequency zone 2 12.721 <0.001 a b b
Time · flood frequency zone 4 0.744 0.571
Perennial monocots
Cyperaceae
Time 2 1.627 0.215
Flood frequency zone 2 8.666 <0.001 a b b
Time · flood frequency zone 4 0.261 0.834
Poaceae
Time 2 14.231 <0.001
Flood frequency zone 2 1.385 0.268 a a a
Time · flood frequency zone 4 0.174 0.950

*Numerator only is reported here as in all tests the denominator ¼ 27.

†
Determined by Tukey’s b test: NC, near-channel; MF, mid-floodplain; FF, far-floodplain.

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Soil seed banks of an arid floodplain 217

(a) (b)
60 60 Near channel

Seed abundance
Mid floodplain
40 40 Far floodplain

± SE 20 20

0 0

(c) (d)
60 60
Seed abundance

40 40
± SE

20 20

0 0

(e) (f)
Seed abundance

200 200
± SE

100 100

0 0

(g) (h)
200
Seed abundance

200
± SE

100 100

0 0
Feb May Oct Feb May Oct

Sample time

Fig. 6 Mean seed abundance in samples from each flood frequency zone and sample time for annual forb families: (a) Amaranthaceae,
(b) Asteraceae, (c) Fabaceae and (d) Lythraceae; annual monocot families; (e) Cyperaceae and (f) Poaceae; and perennial monocot
families; (g) Cyperaceae and (h) Poaceae. Error bars indicate standard error. Data for the annual forb families Campanulaceae,
Caryophyllaceae and Portulacaceae and the perennial forb family Haloragaceae have been omitted from this figure due to small
sample sizes.

Finally, the significant reduction of seeds belonging to
the Asteraceae family in samples from the third
sample time was probably due to germination as
many of these species are known to germinate in
response to winter temperatures (Cunningham et al.,
1992).
Although flooding did not alter soil seed bank
composition substantially during the sample period,
some spatial differentiation was evident with relation
to flood frequency. This can be attributed partially to
variations in the extant vegetation which have devel-
oped as a result of differences in flood frequency and
duration. Perennial grasses, for example, were most
abundant in the soil seed bank of the far-floodplain
zone where flooding is rare and these species tend to

2005 Blackwell Publishing Ltd, Freshwater Biology, 51, 206–223
dominate the extant vegetation (Capon, 2003, 2005).
Annual grasses which are able to complete their life
cycles rapidly in periods intervening flood events are
generally most common in more frequently flooded
areas of floodplains (Menges, 1986; Capon, 2003, 2005)
and this was reflected by their high seed abundance in
the soil seed bank of the near-channel zone. Soil seed
bank samples from the near-channel zone also had
significantly higher total seed abundance and greater
abundance of several annual forb families than those
from the less frequently flooded areas. It is likely that
more frequent and prolonged inundation in this zone
leads to more reproductively successful germination
events, providing greater local additions to the soil
seed bank. More frequent additions of seeds dis-
 13652427, 2006, 2, Downloaded from https://onlinelibrary.wiley.com/doi/10.1111/j.1365-2427.2005.01484.x by Supreme Court Library, Wiley Online Library on [19/06/2023]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License
218 S.J. Capon and M.A. Brock
(a) (b)

Axis 2

Axis 1 Axis 1

Fig. 7 Ordination of sites by seed abundance of species and species vectors significantly correlating with the ordination space. (a) SSH
ordination (stress ¼ 0.157). Symbol shape indicates flood frequency zone; ¼ near-channel, d ¼ mid-floodplain and ¼ far-
floodplain. Shading signifies sample time; black ¼ February, grey ¼ May and open symbols ¼ October. (b) Species vectors. Species
indicated by each number are provided in Table 4.

Table 4 Species correlating significantly to the ordination

(Fig. 7) as indicated by the Monte-Carlo randomisation proce-
dure

ID Species Plant
(Fig. 7b) name group Family

1 Calotis hispidula** af Asteraceae

2 Wahlenbergia gracilis*** af Campanulaceae
3 Eragrostis tenellula** am Poaceae
Axis 2

4 Spergularia rubra** af Caryophyllaceae

5 Cullen cinereum*** af Fabaceae
6 Iseilema sp.** am Poaceae
7 Cyperus bifax*** pm Cyperaceae
8 Centipeda minima* af Asteraceae
9 Calotis porphyroglossa* af Asteraceae
10 Alternanthera nodiflora*** af Amaranthaceae
11 Chloris pectinata* am Poaceae
12 Eragrostis parviflora* am Poaceae
13 Ammania multiflora*** af Lythraceae
14 Aristida contorta*** am Poaceae
15 Dicanthium sericeum* pm Poaceae Axis 1
16 Sida rohlenae* pf Malvaceae
Fig. 8 Ordination of species presence and absence (stress ¼
17 Leiocarpa brevicompta* af Asteraceae
0.18) in the extant vegetation (open symbols) and soil seed bank
18 Senecio lautus*** af Asteraceae
(closed symbols). Samples from each flood frequency zone are
19 Trigonella suavissima** af Fabaceae
indicated by symbols: ¼ near-channel, d ¼ mid-floodplain
20 Daucus glochidiatus* af Apiaceae
and ¼ far-floodplain. Numbers indicate sample times (i.e.
21 Elytrophorus spicatus* am Poaceae
1 ¼ February, 2 ¼ May and 3 ¼ October).
22 Cyperus difformis*** am Cyperaceae
23 Centaurium spicatum* af Gentianaceae
24 Calotis multicaulus* af Asteraceae persed by floodwaters could also contribute to the
higher seed abundance of the soil seed bank in the
Plant group (af, annual forb; pf, perennial forb; am, annual
monocot; pm, perennial monocot) and family are also given. near-channel zone, although this could be offset by
*P < 0.05, **P < 0.01, ***P < 0.001. losses due to scouring. Many of the most common

2005 Blackwell Publishing Ltd, Freshwater Biology, 51, 206–223

50
40
Similarity index (%)
30
20
Near-channel
Mid-floodplain
10
Far-floodplain
0
February May October
Sample time
Fig. 9 Mean similarity index (%) of soil seed bank samples and
extant vegetation in each flood frequency zone at each sample
time. Error bars indicate standard error.
species germinating from the soil seed bank in this
study were distributed throughout samples from all
flood frequency zones (Table 1) and the absence of
further spatial patterns in soil seed bank composition
may be attributed to widespread dispersal of seeds by
wind or floodwaters during large events (Haukos &
Smith, 1994; Brock, 1998).
Contribution of the soil seed bank to vegetation
dynamics
Desert floodplains are characterised by highly vari-
able flooding regimes (Walker, Sheldon & Puckridge,
1995; Puckridge et al., 1998) and plants inhabiting
these environments must persist through periods of
both drought and inundation. Soil seed banks enable
plants to escape from the stresses associated with such
disturbances and contribute to the extant vegetation
when conditions are suitable for their germination
and growth (Bonis et al., 1995; Blom & Voesenek,
1996). Germination in many wetland plants occurs
when sediments are waterlogged rather than sub-
merged (van der Valk, 1981; Welling, Pederson & van
der Valk, 1988; Baskin & Baskin, 1998) and it is likely,
therefore, that many of the annual species residing in
the soil seed bank of the Cooper Creek floodplain
contribute to the extant vegetation predominantly
during the moist conditions following floodwater
recession before drought or further inundation occur.
Greater concordance between the soil seed bank and
the extant vegetation which developed after flooding
was apparent in this study both temporally and
spatially as similarity tended to decline with decreas-

2005 Blackwell Publishing Ltd, Freshwater Biology, 51, 206–223
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Soil seed banks of an arid floodplain 219
ing flood frequency of sites. However, the widespread
distribution of common species throughout the soil
seed bank of all flood frequency zones implies that
plant communities across the flood frequency gradi-
ent have the potential to respond to flooding with
similar shifts in composition. The soil seed bank of the
Cooper Creek floodplain also contains seeds of species
more akin to desert ephemerals which are likely to
contribute to the extant vegetation following rainfall
during drier periods (Freas & Kemp, 1983; Henderson
et al., 1988). Members of the Asteraceae family, in
particular, germinate in winter months when floods
are rare and these probably account for the higher
similarity observed between the soil seed bank and
extant vegetation of the far-floodplain zone at the
October sample time.
In contrast with annual species, perennial plants of
the Cooper Creek floodplain do not appear to depend
on soil seed banks for persistence. Perennial species
were poorly represented in soil seed bank samples
and shrub and sub-shrub species were completely
absent despite their prevalence in the extant veget-
ation. Long-lived species may allocate more resources
to tolerance rather than avoidance of disturbances
such as droughts and floods via physiological or
morphological traits, e.g. adventitious roots (Blom &
Voesenek, 1996). Reproduction in many perennial
plants of the Cooper Creek floodplain may be
predominantly vegetative and propagules resulting
from sexual reproduction might be short-lived or
stored in aerial seed banks. In some desert plants,
seeds are held in the canopies of dead plants and are
released in response to rainfall (Baskin & Baskin, 1998;
Narita & Wada, 1998). The timing of sexual reproduc-
tion in some species may coincide with conditions
favourable for germination and establishment, e.g.
drawdown (Blom et al., 1990). The dominant shrub,
M. florulenta, for example, persists through drought in
a vegetatively dormant state and produces leaves and
flowers in response to rainfall or flooding (Roberts &
Marston, 2000). Dispersal of seeds from neighbouring
land systems such as sand dunes may also contribute
to the recruitment of perennial plant species.
In summary, soil seed banks appear likely to
contribute substantially to flow-related vegetation
dynamics of the Cooper Creek floodplain, partic-
ularly with regards to short-term shifts in composi-
tion of the annual grass and forb communities which
dominate this landscape. The lack of temporal

220 S.J. Capon and M.A. Brock
change in soil seed bank composition observed here
in response to flooding implies a high degree of
resilience amongst these plant communities in the
short term. The results indicate that, at the flood
pulse scale, it would be unlikely for soil seed bank
species to be lost from the system should conditions
following flood-induced germination not lead to
successful reproduction. However, the abundance
of seeds in the soil seed bank was related to flood
frequency suggesting that alterations to the flooding
regime over the longer term, through water resource
development or climate change, could lead to chan-
ges in soil seed bank composition, perhaps as a
result of fewer additions through local seed fall or
reductions in seed dispersal by floodwaters. Without
flood-related replenishment, soil seed banks of rarely
inundated peripheral floodplain areas could decline
in abundance (and ultimately diversity) simply as a
result of seed mortality. The potential of floodplain
plant communities to respond to future variability
would be reduced as a consequence. A greater
understanding of processes of soil seed bank deple-
tion (e.g. germination, mortality and granivory) and
replenishment (primary and secondary dispersal),
with a view towards constructing quantitative mod-
els of seed bank dynamics, might therefore be a
target for future research in this area.
Acknowledgements
This research was completed as part of a doctoral
project supervised by Professor Stuart Bunn, Dr Fran
Sheldon and Dr Margaret Brock. Financial support for
the work was received from Griffith University, the
CRC for Freshwater Ecology and Land and Water
Australia (GRU26). Much appreciated hospitality was
received from the people of Windorah, particularly the
Kidd family, and thanks is given to Robert and Connie
Capon for their valuable assistance in the field. The
manuscript also benefited from comments provided
by Dr Paul Reich and two anonymous referees.
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Appendix List of species recorded from the soil seed bank of the Cooper Creek floodplain

Plant group Family Species name Species author*

Annual forbs Amaranthaceae Alternanthera nodiflora R. Br.

Apiaceae Daucus glochidiatus
Eryngium plantagineum†
Asteraceae Calotis hispidula
Calotis multicaulis
Calotis porphyroglossa
Centipeda minima†
(Labill.) Fisch., C.A. Mey & Ave-Lall.
F. Muell.
(F. Muell.) F. Meull.
F. Muell. Ex Benth.
(L.) A. Braun & Asch. var. minima

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Soil seed banks of an arid floodplain 223
Appendix (Continued)

Plant group Family Species name Species author*

Leiocarpa brevicompta (F. Muell.) Paul G. Wilson

Pseudognaphalium luteoalbum (L.) Hilliard & B.L. Burtt
Pterocaulon sphacelatum† (Labill.) F. Muell.
Senecio lautus G. Forst. Ex Willd.
Brassicaceae Arabidella eremigena† (F. Muell) E.A. Shaw
Lepidium sagittulatum Thell.
Campanulaceae Wahlenbergia gracilis (G. Forst.) A.DC.
Caryophyllaceae Spergularia rubra (L.) J. Presl & C. Presl
Convolvulaceae Ipomea diamantinensis J.M. Black ex Eardley
Euphorbiaceae Phyllanthus virgatus G. Forst.
Fabaceae Aeschynomene indica L.
Cullen cinereum (Lindl.) J.W. Grimes
Trigonella suavissima Lindl.
Gentianaceae Centaurium spicatum (L.) Fritsch ex Janch.
Lythraceae Ammania multiflora Roxb.
Molluginaceae Mollugo cerviana† (L.) Ser.
Plantaginaceae Plantago cunninghamii Decne.
Portulacaceae Portulaca oleracea L.
Annual monocots Cyperaceae Cyperus difformis L.
Cyperus pygmaeus† Rottb.
Poaceae Aristida contorta† F. Muell.
Chloris pectinata Benth.
Dactyloctenium radulans (R. Br.) P. Beauv.
Echinocholoa turneriana (Domin) J.M. Black
Elytrophorus spicatus† (Willd.) A. Camus
Eragrostis basedowii† Jedwabn.
Eragrostis parviflora (R. Br.) Trin.
Eragrostis tenellula (Kunth) Steud.
Iseilema sp.
Perennial forbs Asteraceae Minuria denticulata† (DC.) Benth.
Goodeniaceae Goodenia fascicularis F. Muell & Tate
Haloragaceae Haloragis aspera Lindl.
Malvaceae Sida rohlenae Domin subsp. Rohlenae
Marsileaceae Marsilea drummondii A. Braun
Scrophulariaceae Mimulus gracilis R. Br.
Verbenaceae Verbena officinalis L.
Perennial monocots Cyperaceae Cyperus bifax C.B. Clarke
Eleocharis sp.
Poaceae Dicanthium sericeum (R. Br.) A. Camus subsp. Sericeum
Eragrostis eriopoda Benth.
Eriochloa crebra S.T. Blake
Panicum decompositum R. Br. var. decompositum
Paspalidium jubiflorum† (Trin.) Hughes

*Nomenclature follows Henderson (2002).

†
Indicates species present in soil seed bank only.

2005 Blackwell Publishing Ltd, Freshwater Biology, 51, 206–223