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The ecological consequences of Buffel Grass

(Cenchrus ciliaris) establishment within
remnant vegetation of...

Article in Pacific Conservation Biology · January 2002

DOI: 10.1071/PC020099

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 The ecological consequences of BufIel Grass
Cenchrus ciliaris establishment within remnant
vegetation of Queensland
ANDREW J. FRANKS 1,2·

The effects of Buffel Grass Cenchrus ciliaris presence and dominance in 78 Poplar Box Eucalyptus populnea
woodland remnants of south central Queensland are documented, Buffel Grass was recorded from the majority of sites
sampled, As the relative cover of Buffel Grass increased at both the quadrat and site level, the number of native ground
cover species declined significantly. Most commonly recorded ground cover species displayed significant changes in
frequency as Buffel Grass cover increased, Only two species increased with increased Buffel Grass cover: Desert
Goosefoot Chenopodium desertorum subsp. anidiophyllum and Galvanised Burr Sclerolaena birchii.
The average cover of Buffel Grass decreased significantly from the edge of remnants towards the core areas. The
foliage projected cover of the sub-canopy layers appeared to be an important factor affecting the relative cover of
Buffel Grass at anyone site indicating that shading and competition with the woody layers are important determinants
in the structure and composition of the ground cover layer. A number of sub-canopy species were found to produce
leachates that were capable of significantly reducing the germination or growth of Buffel Grass seeds, alluding that
allelopathy may play a minor role in determining ground cover species assemblages. However, this result can only be
interpreted conservatively in relation to its ecological relevance with effects related to competition with overstorey woody
species being of greater importance.
This study quantifies the long observed effect that the exotic perennial Buffel Grass excludes other ground cover
species, which may have detrimental ramifications on the functioning of remnant native vegetation over much of the
state. With the recent introduction of regulations controlling broad scale tree clearing on both freehold and leasehold
lands in Queensland, it is now important to identify a range of issues pertaining to the effective on ground management
of remnant vegetation located outside the existing nature refuge network,
Key words: Exotic species; Invasive species; Mulga lands; Native vegetation management; Pasture grass; Poplar Box
woodlands.

INTRODUCTION suitability, it is estimated that approximately 4

ALTHOUGH thought to have been
accidentally introduced into north-western
Australia sometime during the 1870s (Marriot
1955; Humphreys 1967; Paull and Lee 1978;
Cavaye 1991; Low 1999), Buffel Grass Cenchrus
ciliaris L., including its many cultivars, has since
been actively promoted as a pasture species.
This promotion has facilitated its establishment
and spread in pasture lands across much of
northern and inland Australia (Griffin 1993).
Buffel's ability to establish and spread under
highly variable climatic conditions has converted
many "low producing scrub lands to highly
productive grazing lands" (Paull and Lee 1978).
Native to southern and eastern Africa and
parts of Asia, Buffel Grass has also been utilized
in erosion mitigation (Bryant 1961; Maass et al.
1988; Smith et al. 1992), and rehabilitation of
degraded and mined land, both in Australia and
overseas (Lawrie and Gunness 1985; Warren and
Aschmann 1993; Harwood et al. 1999). It is an
extremely drought tolerant introduced pasture
grass, making it a critical component of property
development through much of the semi-arid
region of Australia. Based on environmental
'Queensland Herbarium (Townsville Office), Environmental and Technical Services, Environmental Protection Agency, P.O. Box 5391, Townsville MC, Queensland,
Australia 4810.
'Department of Botany, The University of Queensland, St Lucia, Queensland, Australia 4072.
'Present address: School of Environmental Science, Murdoch University, Murdoch, Western Australia, Australia 6150.
PACIFIC CONSERVATION BIOLOGY Vol. 8: 99-107. Surrey Beatty & Sons, Sydney. 2002.
486 000 km 2 of Australia (Lawson et al.,
submitted) and 220 000 km 2 of Queensland
(Walker and Weston 1990; Cavaye 1991) are
highly to very highly suited for its establishment.
Buffel's success as a pasture species is mainly
due to its ease of establishment, rapid growth
rate, fast maturation, prolonged flowering/
fruiting periods, prolific seed production, and
high seed dispersal ability coupled with
relatively long seed dormancy (Fenton and
Campbell 1981; Franks et al. 2000). It is easily
naturalized in most climates and on a range of
soil types and quickly forms self-sustaining
populations under a range of disturbance
regimes. Buffel Grass is also encouraged by
burning (Low 1997; Butler and Fairfax,
submitted) and is tolerant of persistent grazing
(Hodgkinson et al. 1989). Its deep root system,
the accumulation of carbohydrates in the swollen
stem base, and horizontally orientated nodal
tillers assist in its tolerance of drought, fire and
grazing and its ability to sprout rapidly after
such disturbance (Fenton and Campbell 1981;
Hodgkinson et al. 1989). Buffel Grass is also
agamospermic, being able to produce viable
seed without fertilization (De Lisle 1963).
100 PACIFIC CONSERVATION BIOLOGY
The life-history traits that make BufId Grass
such a successful pasture species are the same
traits that have led to its unsolicited
establishment in other parts of the landscape,
including remnant vegetation adjacent to
pastures. By the year 2000 it was estimated that
Buffel Grass had naturalized between 30 and 50
million hectares of Queensland (Hannah and
Thurgate 2001). Outside of Queensland it now
covers large areas of Western Australia, Northern
Territory, South Australia and New South Wales
(Piggott 1995; Low 1999; Pitts and Albrecht
2000). It has been recorded as establishing
"without seedbed preparation" under Mulga
Acacia aneura (Paull and Lee 1978). In
Queensland it is rapidly encroaching upon a
number of conservation areas including several
key national parks (Low 1997; Butler and
Fairfax, submitted). It is also proving problematic
in Uluru-Kata Tjuta National Park and the
MacDonnell Ranges in the Northern Territory
(Pitts and Albrecht 2000), and has been recorded
in many conservation reserves in Western
Australia (Pigott 1995). It has been declared a
noxious weed in Arizona (USA), where it has
naturalized along waterways and invaded nearby
areas (Piggott 1995), and is problematic in
Hawaii (Daehler and Carino 1998).
Over the past decade it has become
increasingly evident that Buffel Grass has spread
from pasture areas into non-target habitats
throughout Queensland, particularly remnant
vegetation in semi-arid areas. However, there
has been little in the way of documenting the
implications of this spread, particularly the
effects upon native ground cover species within
remnant vegetation. Whereas Fairfax and
Fensham (2000) documented changes in floristic
diversity associated with clearing of remnant
woodlands for conversion to pastures (including
Buffel Grass pastures), this study aims to
highlight the impacts associated with Buffel
Grass establishment on the species composition
and floristics of native ground cover within
remnant vegetation of south central Queensland.
Some of the possible factors controlling the
degree of Buffel Grass establishment and
dominance, including the physical structure of
a remnant or possible allelopathic effects are
also investigated. The ecological consequences
and management implications that may
accompany this establishment are also described.
MATERIAL AND METHODS
Study area
The study sites were located within remnant
Poplar Box Eucalyptus populnea woodlands of the
West Balonne Plains subregion of the Mulga
Lands bioregion (Sattler and Williams 1999),
south-central Queensland (Fig. 1). In total, 78
remnants were sampled across 11 000 km 2
between St George and Bollon, bordered by
latitudes 27°30'S and 28°30'S and longitudes
147°30'E and 148°30'E (Fig. 1). Falling between
the 400 and 600 mm isohyets, the climate of the
study area can be broadly described as
mesothermic semi-arid (Bureau of Meteorology
1986). The study area occurs within a broad
transitional zone between the predominantly
winter rainfall regime of southern Australia and
the summer rainfall regime of northern
Australia.
The most widespread soils in the study area
are fine sandy clay loam to light clay massive
earths, which grade into limited areas of
gilgaied grey clays/red duplexes (Manning, D.
2000, pers. comm. 31 January). The massive
earths usually support woodlands of Poplar Box,
Yellow Jacket (Forest ·Gum) Eucalyptus intertexta
and Mulga with a moderately dense shrub layer
of Wilga Geijera parviflora and False Sandalwood
Eremophila mitchellii.
Between 1995 and 1997 approximately one-
half of the annual clearing of native vegetation
for the Mulga Lands bioregion occurred in the
West Balonne Plains subregion (DNR 1999). The
Balonne Shire, which is located within this
subregion, recorded the highest rate of tree
clearing for any Local Government Area in
Queensland for the combined period between
1995 and 1999 (DNR 1999; DNR 2000).
Clearing of eucalypt woodlands and Acacia
communities within this subregion is often
followed with pasture improvement by sowing of
exotic species, particularly Buffel Grass.
Sampling methods
All sites were located in remnant Poplar Box
woodlands with remnants varying in their size
(5.6 to 25 620 ha), shape (linear to elliptic) and
landscape configuration. Climate, geology and
soils were relatively uniform across all sites.
Potential study sites were identified using aerial
photographs and Landsat TM imagery. Final site
selection was based on ground truthing of these
potential sites. All remnants have been grazed
to varying degrees by domestic stock (both sheep
and cattle), introduced vertebrates (feral goats)
and native wildlife.
A series of thirty 1 X 1 m quadrats was used
to assess the composition and frequency of
ground cover species within the 78 remnants
(n = 2 340). The quadrats were located along
two sampling lines parallel to the edge of the
remnant. The first line of fifteen quadrats were
located approximately 2.5 m from the pasture/
remnant interface. These were defined as "edge"
quadrats, which may be greatly influenced by
adjacent pasture processes. The second series
of fifteen quadrats were located approximately
 FRANKS: BUFFELL GRASS IN REMNANT VEGETATION
148
Bollon +
148
Legend 50 o 50
•Towns/cities ----
CJ Study area
I 1Bioregions
Fig. 1. Location of the study area in south central Queensland indicating bioregional boundaries and major population
centres.
47.5 m from the remnant edge. These were
defined as "core" quadrats. It was anticipated
that the positioning of these two quadrat lines
would indicate a gradation from edge area to
core within the remnant thereby highlighting
any edge preferences displayed by Buffel Grass.
For each quadrat, ground cover species
presence and their life form were recorded. The
cover/abundance of Buffel Grass was also
recorded. The cover/abundance rating used was
a modified Domin-Kranjina cover-abundance
scale (Mueller-Dombois and Ellenberg 1974) and
was later converted into a per cent cover scale.
Ground cover species were defined as those non-
woody species less than 0.5 m in height.
Furthermore, a rating of the amount of stock
dung present within a quadrat was given with
the relative frequency of dung at a site used as
101
150 152
150 152
Bioregion data supplied courtesy of
100 Kilometers
the Environmental Protection Agency.
a surrogate for grazing intensity. These relative
frequency values were then assigned to an
appropriate grazing intensity class; very light,
light, moderate, and high.
The physical structure of each remnant
sampled was assessed by three randomly located
50 m line transects, running from the edge and
perpendicular to the remnant/pasture interface.
Commencing at 2.5 m along the transect and
spaced at 5 m intervals, the foliage projected
cover (FPC) of the canopy, understorey (tall
shrubs and short trees between 1.5 and 10 m
in height) and low shrub « 1.5 m in height)
layer was determined for each point (ten points
per transect, 30 points per site).
All field work was carried out between July
1999 and June 2000. Rainfall for the study area
was above average in both 1998 and 1999
102 PACIFIC CONSERVATION BIOLOGY
making conditions amenable for plant growth
and optimal for identification. Nomenclature
used in this study follows Henderson (2002) and
botanical binomials presently accepted by the
Queensland Herbarium. Bryant (1961) reported
that by the beginning of the 1960s over 80
cultivars of Buffel Grass had been introduced
into Australia. It is likely that many more
cultivars have been introduced since this time.
This paper includes all cultivars of Buffel Grass
under a unified "Cenchrus ciliaris" complex.
A bioassay experiment was undertaken to
determine the possible chemical inhibition of
germination and growth of Buffel Grass by
commonly occurring canopy and sub-canopy
species. Methodologies follow those described by
Hobbs and Atkins (1991). Fresh foliage was
collected from the study area of six commonly
encountered canopy and sub-canopy species
present at the majority of sites sampled;
Eucalyptus populnea, Acacia aneura, Callitris
glaucophylla, Eremophila mitchellii, Geijera parviflora
and Senna artemisioides. Fifty grams of leaf
material of each species was soaked in 250 ml
of distilled water for 24 hours. Ten Buffel Grass
seeds (cv. Gayndah) were removed from their
surrounding bristles and were placed between
filter paper in a Petri dish with five dishes given
5 ml of stirred and filtered leachate (n = 50 for
each species sampled). An additional five dishes
received an equivalent amount of distilled water
as a control. Dishes were placed within a
germination cabinet and maintained in the dark
with temperatures ranging from 18°C to 34°C on
a 24-hour cycle. The dishes were removed after
ten days, at which time the number of germinated
seeds were counted and the length of the radicle
of each germinated seed was measured.
Analyses
Quadrat data were combined according to the
converted per cent cover classes of Buffel Grass
in each. Hence, six per cent cover classes for
Buffel Grass were defined; 0%, <5%, 5-25%,
25-50%, 50-75%, and >75% cover. The average
number per square metre of ground cover
species other than Buffel Grass was calculated
for each of these per cent cover ranges. One-
way ANOVA was used to test for significant
differences in the average number of ground
cover species for each Buffel Grass per cent
cover class.
The average Buffel Grass cover and ground
cover species richness was calculated for each
remnant sampled with the relationship between
the two examined with linear regression.
Multiple regression was used to investigate the
influence of the average FPC of the canopy and
sub-canopy layers on Buffel Grass cover for each
site. A paired t-test was used to examine
differences in the germination and radicle length
of Buffel Grass seed with each bioassay trial.
The quadrat data were divided into edge and
core quadrats, with the average number per
square metre of ground cover species other than
Buffel Grass calculated for each of the six per
cent cover classes defined above. A one-way
ANOVA was used to test for significant
differences in the average number of ground
cover species along the edge and within the core
habitat areas for each Buffel Grass per cent
cover class. The average cover of Buffel Grass
along the edge and within the core was
calculated for each of the remnants sampled and
compared using a z-test.
The relative frequency of the most commonly
encountered native ground cover species (i.e.,
those recorded in more than 10% of quadrats)
was examined between those quadrats where
Buffel Grass was absent compared to those with
any Buffel cover. This was performed to
determine which ground cover species were
affected by or resilient to Buffel Grass presence
and cover. The relative cover of Buffel per site
for each grazing intensity class determined the
response of Buffel Grass to grazing intensity.
RESULTS
Of the 2 340 quadrats sampled, Buffel Grass
was recorded in 51.6%, with 57.4% of edge and
45.8% of core quadrats containing Buffel Grass.
The native ground cover species richness
decreased significantly as Buffel Grass cover
increased (F5.2334 = 16.956, P < 0.001). The
average number of native ground cover species
per square metre in quadrats without Buffe! Grass
(n = 1 132) was 4.3 (Fig. 2). This dropped to 1.6
species per square metre when Buffe! Grass cover
was greater than 75% (n = 15) (Fig. 2).
The average cover of Buffel Grass per site
varied significantly between the edge (13.9%)
and core (8.7%) areas (z = 3.23, P < 0.01).
6
:l 5
l4
..
>
~ 3
I:
g2
.I:
:; 1
0
0% <5% 5·25% 25·50% 50-75% >75%
Buffel cover
Fig. 2. The mean number (and standard error) of native
ground cover species/m2 in relation to Buffel Grass
cover from all quadrats sampled (n = 2 340).
 FRANKS: BUFFELL GRASS IN REMNANT VEGETATION 103

The quadrats located along the edge of the low shrub layer (multiple regression, F3•74 =
remnants also recorded a significant reduction 5.22, P = 0.0025, r2 = 0.14) but the FPC of the
in species richness (F5 • 1164 = 8.82, P < 0.001), tree canopy layer appeared to have little or no
with the average number of species per square effect.
metre decreasing from 4.17 (with Buffel Grass
Table 1 indicates that foliar leachate from
absent, n = 498) to 2.3 (with Buffel Grass cover
Eremophila mitchellii significantly reduced the
greater than 75%, n = 10). A very strong
germination of Buffel Grass seed. Leachate from
negative correlation exists between Buffel Grass
Acacia aneura, Eremophila mitchellii, Senna
cover and ground cover species richness
artemisioides and Eucalyptus populnea significantly
(Spearman's rs = -0.9425 P < 0.05 df = 6). A
reduced the growth of the radicle. Foliar
similar response was recorded for the core
1eachates from Callitris glaucophylla and Geijera
quadrats with a significant reduction (F5•1164 =
parviflora did not appear to adversely affect
8.202, P < 0.001) in ground cover species
either the germination or radicle growth of
richness as Buffel Grass cover increased. Again
Buffel Grass seeds.
the average number of species decreased from
4.35 with Buffel Grass absent (n = 634) to 0.2 In all, 130 taxa of ground cover species were
with Buffel cover greater than 75% (n = 5). recorded. The response of the most commonly
recorded native ground cover species with Buffel
The relationship between Buffel Grass cover
Grass was variable (Table 2). All native ground
per site and ground species richness is
cover species that were recorded in more than
illustrated in Figure 3. There was a strong
10% of quadrats had a significant change
negative relationship between Buffel Grass cover
in frequency when Buffel Grass was present
at a site and ground cover species richness
(Table 2; T = 20, P < 0.05, Wilcoxon's test for
(Fig. 3; F 1•76 = 10.31, P = 0.0019, r2 = 0.119).
matched pairs). Compared with other native
Buffel Grass cover at a site was negatively ground cover species, Mulga Mitchell Thyridolepis
correlated with the FPC of the understorey and mitchelliana was recorded most often in those
quadrats with Buffel Grass present. However, its
45.-----------------------------------, relative frequency dropped from 44.4% with
40 •
• • Buffel Grass absent to 25.3% when Buffel was
• y; -0.2678x + 29.362 present (Table 2). Only two species appeared
en 35
",
~ 30"
••
. . ...
• •
••
•• •
..
•• • opportunistic to Buffel Grass presence, these
.g~ ~~.~J~~~~~ .:.~.-~--~----
being Desert Goosefoot Chenopodium desertorum
25 •• •• __ • •
en • • ••• • • • subsp. anidiophyllum and Galvanised Burr
.. 20 • • •••
115 •• • • Sclerolaena birchii .
If)
• • •
10 Although the average relative cover of Buffel
5
• Grass increased with grazing intensity (Fig. 4),
O+---~---,----r_--~--_,--_.~--,_--~ this response was not statistically significant
o 5 10 15 20 25 30 35 40 (F3.74 = 1.8525, P = 0.145). However, it is difficult
Buffel cover (%) to determine whether this response was due to
Fig. 3. The relationship between species richness of native grazing or whether the grazing intensity had
ground cover species at a site and Buffel Grass cover increased in response to increased Buffel Grass
from all remnants sampled (n = 78). cover.

Table 1. The effect of foliar leachate on the germination of Buffel Grass seeds (mean
peF Petri dish, five dishes per treatment, ten seeds per dish) and radicle length
(mean over all germinated seeds per treatment ± 1 S.E.) ten days after
germination. Significant differences from distilled water control, as determined by
t-test, are indicated (* = p < 0.05; ** = P < 0.01; *** = P < 0.001).

Number germinated
Treatment (out of 10) Radicle length (mm)
Control (distilled water) 10 ± 0.0 31.48 ± 1.92 (n = 50)
Acacia ancura 9.6 ± 0.2 16.42 ± 1.77 (n = 48)***
CallitTis glaucophylla 9.6 ± 0.2 29.75 ± 1.46 (n = 48)
Eremophila mitchellii 7.8 ± 0.4** 6.83 ± 1.07 (n = 39)***

Eucalyptus populnea 9.4 ± 0.4 25.15 ± 1.89 (n = 47)*

Geijera parviflora 9.6 ± 0.2 31.66 ± 1.34 (n = 48)
Senna artemisioides 9.4 ± 0.4 25.87 ± 1.59 (n = 47)**
 104 PACIFIC CONSERVATION BIOLOGY

Table 2. The response of the most commonly recorded ground cover species from remnant Poplar Box woodlands in quadrats
with and without Buffel Grass.

Species % Frequency % Frequency Change in

without Buffel (n) with Buffel (n) % frequency
Increase in frequency (>5% increase)
Sclerolaena birchii 3.89 (44) 12.58 (152) 8.69
Chenopodium desertorum subsp. anidiophyllum 11.93 (135) 19.12 (231) 7.19
No change in frequency «5% change)
Paspalidium wnstrictum 17.32 (196) 12.83 (155) -4.49
Einadia nutans subsp. nutans 15.02 (170) 11.18 (135) -3.84
Scleroiaena convexula 24.65 (279) 21.94 (265) -2.71
Calotis cuneifolia 14.40 (163) 12.58 (152) -1.82
Boerhavia dominii 14.40 (163) 15.48 (187) 1.08
Enchylaena tomentosa 11.40 (129) 10.43 (126) -0.97
Enteropogon acicularis 14.00 (158) 14.82 (179) 0.82
Sclerolaena diacantha 16.52 (187) 17.30 (209) 0.78
Decrease in frequency (>5% decrease)
Thyridolepis mitchelliana 44.43 (503) 25.29 (305) -19.14
Aristida echinata 10.60 (120) 3.39 (41) -7.21
Chamaesyce drummondii 16.78 (190) 9.93 (120) -6.85
Tripogon loliiformis 10.10 (114) 4.14 (50) -5.96
Hibiscus sturtii var. grandiflorus 12.19 (138) 6.63 (80) -5.56
Aristida jerichoensis var. subspinulifera 12.10 (137) 7.04 (85) -5.06

30~----------------------------------~
C 25
~
8 20
:l
15
:t!; 10 1 disrupts the natural ecological functioning of
"
.c
: 5
~ 0~__n_:_5_______n_:_2_5______n_:_3_2__~__n_:_1_6__~
Very light Light Moderate
Grazing intensity
Fig. 4. The relationship between Buffel Grass cover and
grazing intensity per site indicating a general
increase of cover as grazing intensity increased.
DISCUSSION
Buffel Grass was recorded at varying degrees
of cover from all sites sampled indicating the
susceptibility of Poplar Box woodland remnants
to invasion. In fact, soil seed bank analysis of
eight of the largest remnants of the study area
revealed that Buffel Grass had a mean density
of 25 germinable seeds/m2 recorded from the
central 10% or "core" areas (Moore 2000).
Although relatively under represented in the
germinable soil seed banks of the core areas of
remnants (mean density of germinable Buffel
Grass seed from the surrounding pasture matrix
was calculated at 861 seeds/m2), it must be
pointed out that seeds are present and may
germinate if the conditions are amenable. Ad hoc
observations suggest that Buffel Grass seeds are
triggered to germinate by even minor forms of
soil disturbance including breaking of the soil
surface by stock movement. Hence populations
of Buffel Grass may establish relatively easily in
the interior "core" habitat of Poplar Box
woodlands with little to no disturbance and
further disrupt the functioning of these systems.
Once established within remnant native
vegetation, Buffel Grass alters the character and
these systems in several key ways. As is clearly
evident in this study, Buffel Grass, like any other
competitively dominant species, has the capacity
Heavy
of forming dense swards that displace native
plant species leading to a loss of heterogeneity
and floristic diversity. Fairfax and Fensham
(2000) recorded a similar response in a study
comparing exotic and native pastures to
uncleared woodlands of central Queensland.
They found substantial declines in native species
richness and diversity associated with Buffel
Grass pasture development following tree
clearing for both brigalow and eucalypt
woodlands (Fairfax and Fensham 2000). Table 2
also indicates that many native ground cover
species decline with increased Buffel Grass cover
with some showing a substantial reduction in
relative frequency. Not only does Buffel Grass
physically displace and exclude species but
studies have also indicated that it may
chemically suppress the germination of some
plant species seed (Cheam 1984). Buffel Grass,
like many other exotic species, may also
effectively prevent regeneration of canopy and
sub-canopy species (e.g., Panetta and Hopkins
1991) through competition and changes in the
microclimate of the near ground environment.
The relative homogenous nature of dense stands
of Buffel Grass also impacts on native fauna,
encouraging some species to the detriment of
others (Ludwig et at. 2000; Hannah and
Thurgate 2001).
 FRANKS: BUFFELL GRASS IN REMNANT VEGETATION
The cover of Buffel Grass within remnants
appears to be strongly affected by the degree of
closure of the understorey and low shrub layers,
with those sites with more open sub-canopy
layers having a greater amount of Buffel Grass
cover. This may indicate sub-canopy woody
species may effectively exclude Buffel Grass
possibly through shading and competition for
water and nutrients. Other studies in both
Mulga (Beale 1973) and Eucalyptus woodlands
(Scanlan and Burrows 1990) have indicated that
an increase in density of the canopy and sub-
canopy layers (and the related increase in basal
area) leads to a decrease in herbaceous yields of
ground cover species. However, there may also
be some minor chemical inhibition occurring
between canopy and/or sub-canopy species and
Buffel Grass seeds as indicated by the crude
bioassay experiment performed here. It must be
stressed that the use of a bioassay using foliar
leachates is of limited value (see Webb et al.
1961) but these preliminary results indicate that
some inhibition of Buffel Grass germination may
be occurring. However, Webb et al. (1961) found
that while fresh leaf material of Eremophila
mitchellii inhibited the germination of a number
of herbaceous species, the leaf litter of the same
species had no effect. Hence the type of
allelopathic response documented here could be
ecologically irrelevant with the primary factor
determining the cover and dominance of Buffel
Grass in remnant woodlands being the density
of the sub-canopy and, to a lesser extent, canopy
layers.
As is evident by the significant reduction of
Buffel Grass cover from edge to core areas of
each remnant, some edge effects may be
operating. Despite this decline in Buffel Grass
cover, ground cover species were still being
excluded in core areas where Buffel had formed
dense swards. Stock movement into these core
areas may aid the incursion of Buffel Grass
further into remnants from the pasture/remnant
interface. Other studies have indicated that stock
grazing significantly increases the level of weedy
invasion into remnant vegetation (e.g.,
Abensperg-Traun et al. 1998). The interaction
between remnant degradation, Buffel Grass
cover and grazing may be closely tied, with
fragments providing shelter to stock, stands of
Buffel Grass attracting grazing further into the
remnant, and disturbance associated with stock
movement allowing Buffel to further expand.
Not unlike other exotic grass species, Buffel
Grass may significantly alter the natural fire
regime, through increasing fuel loadings causing
extensive, intense and more frequent fires than
those carried by native grasses, thereby
destroying seedlings and detrimentally altering
soil seed banks (Humphries et al. 1991;
D'Antonio and Vitousek 1992; Daehler and
105
Carino 1998; Mack and D'Antonio 1998; Butler
and Fairfax, submitted). Buffel Grass produces
around two to three times the flammable
biomass of displaced native grasses resulting in
hotter and more intense fires particularly later
in the dry season (Humphries 1993). Fire
sensitive communities such as Brigalow Acacia
harpophylla, Gidgee A. cambagei and softwood
scrubs that are heavily infested with Buffel Grass
may be at greatest risk (Butler and Fairfax,
submitted). For example, a recent fire fuelled by
Buffel Grass within Mazeppa National Park,
central Queensland, has either killed overstorey
trees or significantly reduced their crown cover
thereby facilitating the further incursion and
thickening of existing Buffel stands (Butler and
Fairfax, submitted). It is these cumulative effects
that have contributed to Buffel Grass being
identified as one of Queensland's sixteen and
Australia's eighteen worst environmental weeds
(Humphries et al. 1991; Environmental
Protection Agency 1999).
Buffel Grass provides a paradox in the
Queensland context: it is a valuable pasture
species forming a critical component of the
grazing industry, but it has also demonstrated
an ability to detrimentally impact upon and alter
the natural ecology of remnant vegetation. With
the recent proclamation of the U!getation
Management Act 1999 (Queensland) to regulate
tree clearing in rural areas, it is now important
to identify key issues in relation to the effective
on-ground management of remnant vegetation
particularly those areas outside the established
conservation reserve network. If remnants are to
be managed to maintain and promote their
regenerative capacity and to safe guard against
long-term landscape degradation then it is
imperative to document the effects that Buffel
Grass and other problematic plants are having
on the composition, structure and functioning of
these systems. There is a need for realistic
strategies to be implemented to control Buffel
Grass expansion, protect remnant native
vegetation and to assist in the off-reserve
management of endangered flora and fauna
across Queensland. As the distribution of Buffel
Grass across northern Australia continues to
expand, it is essential for state, territory and
federal governments to decide whether the
economic gains associated with this species
outweigh the loss of native species and the
degradation of natural systems.
ACKNOWLEDGEMENTS
This study forms part of the larger Remnant
Vegetation-Condition and Function project and
was financially supported by the Forest
Ecosystem Research and Assessment (FERA) unit
of the former Department of Natural Resources.
Don Butler, Russell Fairfax, Renee Moore,
 106 PACIFIC CONSERVATION BIOLOGY
Lachlan Pegler, Rod Fensham and John N eldner
are thanked for reviewing an early version of the
manuscript. Renee Moore (NR&M) kindly
supplied soil seed bank data. Terri Sutcliffe
(Forestry & Wildlife, QPWS) prepared Figure 1.
Jodi Rees, Jodi Neal, Renee Moore and Annie
Kelly provided thorough and at times
entertaining field assistance. Thanks are also
extended to the many landholders of the
Balonne Shire who provided unrestricted access
to their properties and gave keen insights into
their local environment.
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