CSIRO PUBLISHING

www.publish.csiro.au/journals/ajb Australian Journal of Botany, 2009, 57, 163170

Multi-decadal increases in shrub abundance in non-riverine

red gum (Eucalyptus camaldulensis) woodlands occur
during a period of complex land-use history

Jodi N. Price A,B,C and John W. Morgan A

A
Department of Botany, La Trobe University, Bundoora, Vic. 3086, Australia.
B
Present address: Ecosystem Management, University of New England, Armidale, NSW 2351, Australia.
C
Corresponding author. Email: jprice20@une.edu.au

Abstract. The history of land-use was examined in Eucalyptus camaldulensis Dehnh. woodland in the Victoria Valley of
the Grampians National Park, south-eastern Australia, to help interpret changes in vegetation there during the last 50 years.
We used aerial photography and dendrochronological data to quantify the amount of, and the rate of change in, the woody-
vegetation cover between 1948 and 1997, and historical data to document land-use changes during this time. Aerial
photographs indicated that in 1948, 56% of study area had <50% cover of woody plants. By 1997, 90% of the study area had
>50% woody-plant cover. The native shrub Leptospermum scoparium J.R. Forst & G. Forst (Myrtaceae) was predominantly
responsible for the increases in cover. Demographic analyses indicated that recruitment has been ongoing rather than
episodic; large numbers of shrubs, however, have recruited since 1994. We hypothesise that the vegetation changes observed
are likely a response to changes in land-use that have occurred since European occupation. Increased woody-plant cover
followed the removal of sheep grazing in the long-term absence of re. It is very likely that the long history of stock grazing,
coupled with selective logging and associated soil disturbance, initiated a change in understorey vegetation by reducing
competitive native tussock grasses and fuel loads to carry res and this reduction was initially responsible for the
encroachment of shrubs into the woodland. Recruitment has been ongoing in the absence of any recent land-use
changes (most utilisation ceasing after the declaration of National Park status in 1984) and hence, this transformation
from species-rich herbaceous woodland to shrubby woodland is expected to continue in the future.

Introduction historical links between disturbance and differing vegetation

Woody plants have been increasing in savanna and woodland
ecosystems in recent decades on a global scale, a process often
referred to as encroachment (Archer et al. 1995; Lunt 1998a;
Roques et al. 2001; Bartolome et al. 2005; Briggs et al. 2005).
Shifts in vegetation structure are attributed to changes in climate
and CO2 concentration as well as modications to land-use
practices such as grazing and re regimes (Archer 1989;
Bahre and Shelton 1993; Polley et al. 1994; Van Auken 2000;
Heisler et al. 2003; Fensham et al. 2005). Little attention has
been given to woody-plant encroachment in land managed
primarily for conservation. Many conservation reserves have
traditionally had a non-interventionist management
philosophy and changes in vegetation condition are often
attributed to successional processes. However, many studies
attribute land-use change as an important driver of vegetation
dynamics as well as reporting negative consequences for plant
biodiversity (Withers and Ashton 1977; Gleadow and Ashton
1981; Kirkpatrick 1986; McMahon et al. 1996; Rose and
Fairweather 1997; Lunt 1998a, 1998b; Costello et al. 2000).
Temporal data are required to place vegetation change in the
context of disturbance history to (a) determine the relative
inuences of anthropogenic and successional processes on
vegetation dynamics and (b) enable managers to incorporate
states into future management strategies.
Increases in woody-plant cover have been documented in
south-eastern Australia (e.g. in Leptospermum laevigatum,
Kunzea ericoides, Pittosporum undulatum, Allocasuarina
littoralis, Acacia sophorae, A. paradoxa) in a variety of
ecosystems such as grasslands, heathlands, woodlands and
open forest (Withers and Ashton 1977; Gleadow and Ashton
1981; Bennett 1994; Lunt 1998a; Singer and Burgman 1999;
Costello et al. 2000; Franco and Morgan 2007). Changes have
mostly occurred in the last 50 years although may be a response
to longer-term changes in land-use. Encroachment has often
been due to reductions (or alterations) in re frequencies
(Withers and Ashton 1977; Gleadow and Ashton 1981;
Kirkpatrick 1986; Molnar et al. 1989; Singer and Burgman
1999) and modications to grazing regimes (Bennett 1994;
Singer and Burgman 1999; Costello et al. 2000; Franco and
Morgan 2007). Other factors such as increased dispersal
capabilities (Gleadow and Ashton 1981; McMahon et al.
1996; Rose and Fairweather 1997), changes in hydrology
(Bren 1992) and phenotypic plasticity (Price and Morgan
2006) have also been implicated in encroachment.
There is increasing concern about potential (negative) impacts
of shrub encroachment in grassy woodlands of south-eastern

 CSIRO 2009 10.1071/BT07079 0067-1924/09/030163

164 Australian Journal of Botany
Australia and the role of land-use and/or disturbance in this
expansion (Bennett 1994; Lunt 1998b; Franco and Morgan
2007). The native shrub L. scoparium has increased in
abundance in herb-rich E. camaldulensis (red gum) woodlands
in the Grampians National Park, southern Australia (National
Parks Service Victoria 1985; Calder 1987; Price and Morgan
2006). Aerial photographs of the Victoria Valley in the central
Grampians suggested an open woodland in the 1940s that has
since been thickening; concern has been raised by the
management agency (Parks Victoria) and local botanists about
the encroachment of shrubs into the area. Reductions in species
richness have been found with increased cover of L. scoparium
(Price and Morgan 2008) in communities that, at small scales, are
some of the most species-rich in temperate regions, with up to
45 species per m2 (Lunt 1990).
Leptospermum scoparium occurs naturally in New Zealand,
south-eastern Australia and Tasmania (Thompson 1989). In
New Zealand, L. scoparium occurs over a wide geographic
range and is commonly an early successional species in
previously forested regions as well as occupying a more
permanent role at both extremes of the rainfall gradient
(Burrell 1965; Yin et al. 1984; Stephens et al. 2005). Its role
as a pioneer in succession has seen an expansion of its range
with increased human disturbance in New Zealand (Stephens
et al. 2005). The species has invaded into tussock grassland in the
absence of re (Calder et al. 1992) and into cushion bog
communities in response to frequent burning (Johnson 2005).
L. scoparium is believed to be strongly serotinous in Australia,
with variable serotiny found in New Zealand in relation to re
history (Bond et al. 2004). The species has been observed to
reproduce vegetatively by root suckers (Agnew et al. 1993;
Price and Morgan 2006) and can resprout after re, depending
on re severity (J. N. Price and J. W. Morgan, pers. obs.
2006). Establishment of L. scoparium in the Victoria Valley
region is favoured by greater summer soil moisture than in
other regions within the Grampians National Park, where
recruitment appears limited by summer drought (Price and
Morgan 2006).
Fig. 1. Aerial photographs (1 : 20 000) of the study area in the Victoria Valley of the central Grampians, demonstrating dramatic woody thickening between 1948
(left panel) and 1997 (right panel).
J. N. Price and J. W. Morgan
The aim of the present study was to quantify the changes in
woody-plant cover that have occurred in the last 50 years in herb-
rich woodlands in the Victoria Valley of the Grampians National
Park, as well as to document the land-use changes which may be
implicated in this process.
Materials and methods
Study area
The study was conducted in non-riverine, herb-rich
E. camaldulensis woodland in the Victoria Valley of the
Grampians National Park, south-eastern Australia (37
170 S,
142
240 E). The climate is temperate, with cool winters (mean
minimum temperature of the coldest month (July) is 4
C) and
warm summers (mean maximum temperature of the
warmest month (JanuaryFebruary) is 26.6
C) (Bureau of
Meteorology, Hamilton recording station, ~35 km to the south-
west of the study site). The average annual rainfall is 687 mm,
with a mean monthly maximum of 77 mm in August and a
minimum of 32 mm in February. We selected an area of
~370 ha in the Victoria Valley where aerial photographs
suggested a very open woodland (Fig. 1). The site was located
on an alluvial plain; the soils are duplex, consisting of a loamy
sand, 0.5 m in depth, above a heavy mottled clay. Soils are
typically waterlogged in winter and spring. Lunt (1990)
described the herb-rich woodlands of western Victoria as
being of national signicance because of the high small-scale
species richness of the ground ora. The ora is primarily
herbaceous, perennial (~65%) and native (~87%) (Lunt 1990).
The native vegetation of the Grampians National Park has been
modied by stock grazing, timber and wattle-bark harvesting and
soil disturbance since European occupation (~170 years ago).
Increases in woody-plant cover in the study area occurred during
a long re-free period (>50 years).
Data collection
A variety of methodologies was utilised to assess land-use and
vegetation changes in herb-rich woodlands during the last
Long-term changes in woody-plant cover in woodlands Australian Journal of Botany 165

50200 years. We utilised a series of aerial photographs taken in
1948 (black and white: 1 : 15 000), 1971 (colour: 1 : 20 000) and
1997 (colour: 1 : 20 000) to determine vegetation changes.
Woody-plant cover was quantied by overlaying a transparent
grid over a 370-ha region and determining in each grid square
(900 m2) the percentage cover occupied by woody vegetation.
The following three shrub-cover classes were identied for
analysis: 025% cover, 2650% cover, >50% cover of woody
species. The total area (ha) occupied by each cover class was
determined within the 370-ha study region.
On the basis of the most recent aerial photos (1997), four
areas were located for ground-truthing, two in dense cover and
two in relatively open areas. At each location, a 100-m transect
was established with 5, 10  10 m quadrats placed at regular
intervals. The identity and cover of shrubs and other woody
plants were estimated in each quadrat in August 2004. A
dendrochronological study was also undertaken to determine
the age-structure of shrubs. All basal diameters of the dominant
shrub (L. scoparium) were recorded in each quadrat. A previous
study found a strong relationship between basal diameter and
annual growth rings in this species (Price and Morgan 2006);
hence, basal diameter can be used as a surrogate for age.
We searched for historical information on vegetation and
land use in the accounts of early settlers and explorers, local
histories and land-use reports. Small areas of red gum woodland
occur in the Grampians National Park, with the largest area being
in the Victoria Valley. Hence, we are certain any land-use records
that refer to red gum woodland in the Victoria Valley occurred in
the study area. Information on early pastoral runs was sourced
from manuscript collections held at the Royal History Society,
Victoria, and the State Library Victoria. Information on land
use during the State Forest era is contained at the Public
Records Ofce Victoria (PROV), in the form of the Forests
Commissions ofcial correspondence rather than detailed
management records.
Rainfall history of the study site was examined by using long-
term data from the Hamilton Rainfall Station (18701983) and the
Hamilton Airport (19842005). These stations are located
~35 km from the study area and best represent the climatic
conditions of the study area. A drought index was calculated
by the methodology outlined in Fensham et al. (2005), i.e. it was
calculated as the actual annual rainfall less mean annual rainfall
divided by the mean annual rainfall for each year. This index
identies sustained dry and wet periods and was calculated
for 6-year periods (i.e. the 6-year drought index for 1877 is the
running total of the drought index from 1871 to 1877).
Plant nomenclature follows Ross and Walsh (2003).
Results
Vegetation change
There was a large increase in woody-plant cover in the Victoria
Valley during the 49-year period of our aerial photographs
(Table 1, Figs 1, 2). In 1948, 163 ha (44% of the study area)
had >50% cover of woody plants and this had increased to 333 ha
(90% of the study area) by 1997. The area of mostly herbaceous
cover (025% shrub cover) was reduced between 1948 and 1997
from 118 ha to 7.4 ha. Increases in woody-plant cover were
recorded in both time sequences (19481971, 19711997),
with a faster rate of change recorded in the rst time period
(Table 1). Ground-truthing conrmed that 90% of the quadrats
identied as relatively open (<50% cover) and 80% of the
quadrats identied as dense (>50% cover) on the aerial photos
were correctly allocated, and that L. scoparium was implicated in
these changes. Leptospermum continentale was also common in
the study area, although with a low cover, and appeared to be
limited to wetter, low-lying areas. The size-class distribution of
shrubs (Fig. 3) was skewed to the left, with the majority of
individuals having a basal diameter of <30 mm, (~10 years
old). The oldest individuals recorded were ~30 years old.
Rainfall history
The drought index (Fig. 4) identied a series of droughts in the
late 1800s, early 1900s and in the 1990s. Rainfall in the period
Table 1. Changes in the area occupied by woody vegetation from 1948 to
1997 in the Victoria Valley, by using interpretation of aerial photographs
Woody-cover Area covered in ha (% of total area)
class (%) 1948 1971 1997
025 118 (32) 0 (0) 7 (2)
2650 89 (24) 104 (28) 30 (8)
>50 163 (44) 266 (72) 333 (90)

National Park
100 025
Timber harvesting
2650
75 >50
Total area (%)

Wattle barking
50
Grazing
25

0
40

50

60

70

80

90

00

10

20

30

40

50

60

70

80

90

00
18

18

18

18

18

18

19

19

19

19

19

19

19

19

19

19

20

Year

Fig. 2. Percentage changes in the woody-plant cover in the three cover classes, derived from the aerial-photograph analysis with the major land-use phases.
166 Australian Journal of Botany J. N. Price and J. W. Morgan

250
19941997 in the north and the Victoria Valley in the south (Halls Gap and
Grampians Historical Society 2006). These were predominantly
200 extensive unfenced sheep runs of between 24 000 and 60 000 ha,
with low stocking rates (~1 animal to 2.5 ha) (Sibley 1967).
Frequency

150 The rst known occupant of the Moora Moora run was
R. H Bunbury in late 1844, stocking it with heifers and
100 bullock (Halls Gap and Grampians Historical Society 2006).
1975 The records then indicate a shift to sheep grazing as the run went
50
through a series of different owners. Pastoral records indicate the
0 area occupied by each run, numbers of sheep grazed and
<5 515 1630 3150 51100 101150 >150 ownership. For example, in 1862, the Robertson run in Moora
Size-class Moora consisted of 46 000 ha, with an estimated capacity of
20 000 sheep (Chappel 1977). The Victoria Valley run also
Fig. 3. Total number of individuals of Leptospermum scoparium, by stem- had grazing leases from 1852. In the 1860s, various Land Acts
class diameter (mm), for all individuals observed in 20 quadrats sampled were passed which gave applicants the right to select smaller
during 2004 in the study area. Approximate year of establishment is indicated, areas of land (Sibley 1967) and several large squatted runs were
based on the following equation: number of growth rings = 3.7 + 0.18 basal
subdivided around this time.
diameter (mm) + 0.7 (Price and Morgan 2006).
Agricultural settlement in the Victoria Valley was deemed to
be unsuccessful by the early 1870s: northern areas rst became
before the rst aerial photograph (1948) was mostly average or grazing country of the Moora Moora station but in a decade or
above average. Rainfall in the rst time sequence of photos so it was recognised as unsuitable for that purpose (Halls Gap
(19481971) had periods of excess rainfall, followed by and Grampians Historical Society 2006, p. 141). The causes
drought in the last 10 years of this time period. Drought were attributed to poor country, grievous losses of sheep by
conditions were then experienced for most of the second time rot and uke, increasing number of rabbits in the early 70s,
sequence (19711997), during which increases in woody-plant unsuitable selection applicants (Chappel 1977). Bushres were
cover were reported. From 1987 to 2005, drought conditions also blamed, with res recorded in 1874, when 18 selectors
were experienced. lost fencing and grass, and in 1875, 1600 ha of grass was
destroyed (Chappel 1977). All richly grassed country known
Land-use history as Moora Moora has been burnt (Halls Gap and Grampians
1. Early European occupation phase Historical Society 2006, p. 133).

Indigenous people inhabited the area covering the Grampians

National Park (Gariwerd) for at least 22 000 years before
European occupation (Bird and Frankel 2005). Early
observations report on extensive Aboriginal burning in parts of
Victoria (Gott 2005) and it has been suggested that this promoted
the openness of the vegetation in the Victoria Valley (Bird and
Frankel 2005). The rst European to visit the Grampians was
Major Thomas Mitchell in 1836. His favourable reports of
good grazing land led to extensive land settlement (Calder
1987). In the 1840s, all of the Victoria Valley was covered by
grazing leases; initially, there were two main runs in the Victoria
Valley that covered the area of the present study Moora Moora
2. Forest Reserve phase (18721984)
Most of the Victoria Valley was set aside as Forest Reserve
in 1872 and the remaining squatters were forced out of the
valley. Before the State Forest Reservations were completed,
ve grazing leases were issued and when they expired, this
land also became part of the Grampians State Forest. Initially,
the Forest Authority (from 1907) managed the Grampians State
Forest, and in 1918, it was replaced by the Forests Commission
of Victoria. During this phase, main land uses were timber and
wattle-bark harvesting, with apiculture and forest grazing as
important secondary uses (Fig. 2).

1.0

0.5

0
Drought index

1875 1881 1887 1893 1899 1905 1911 1917 1923 1929 1935 1941 1947 1953 1959 1965 1971 1977 1987 1993 1999 2005

0.5

1.0

1.5

2.0

2.5
Year

Fig. 4. Drought index values for 6-year periods at Hamilton weather station. Negative numbers represent rainfall decit and positive numbers rainfall excess.
Long-term changes in woody-plant cover in woodlands
Large areas of black wattle (Acacia mearnsii) in the Victoria
Valley were utilised for the tannin extracted from the bark.
Most of the harvested bark from Victoria came from the
Victoria Valley and this was the main industry there in the
late 1800s and throughout the 1900s, with up to 600 t of bark
harvested per annum (Calder 1987; Searle 1991). Management
of these areas included treatment and improvement to produce
a larger and sustained yield (Searle 1991, p. 16) as well
as burning to encourage wattle regeneration. I, Thomas
McGinnisken, employed by the Forests Commission as
foreman in charge of the gang working in the State Forest in
the Parish of Jalur. . . was on 22-3-33, burning off under control
for the purposes of both re protection and the creation of a
natural regeneration of wattle. . . . The burning was carried out
in red gum and wattle which would not take re and throw out
sparks (PROV 1933). However, no record could be found
of the regularity or intensity of the burns, or the extent of the
area managed this way. Cultivation of wattles was attempted
in several areas in Victoria following an enquiry into wattle
barking in 1878 and was attempted in the Victoria Valley in
1901 (Searle 1991). Forestry Commission maps indicate the
location of a wattle plantation in the Victoria Valley in the
vicinity of our study site (PROV 1933). The wattle-bark
industry became uneconomical in the early 1900s (Searle
1991) and the Victoria Valley was one of the last areas where
bark was harvested, up to the 1960s (Calder 1987).
Sherry (1971) documents two typical areas in the
Grampians where wattle bark was harvested, one of these
being Eucalyptus melliodora woodland with a shrubby
understorey of Leptospermum myrsinoides and L. scoparium,
and a dense understorey of A. mearnsii. The second area was
described as mixed forest community with large eucalypts of
E. camaldulensis and E. aromaphloia and only a few scattered
wattles (~15 per ha). There was no mention of L. scoparium in
the understorey of this community; instead, there was a light
ground cover of grass and bracken with scattered Emu bush
[Astroloma conostephioides] and Hibbertia fasciculata (Sherry
1971, p. 37).
Timber harvesting was one of the earliest industries in the
Victoria Valley, controlled by the Forests Commission by a
system of licenses. Records of logging allocations are held
at the PROV. These records document concern about the
availability of Red Gum in the Victoria Valley in 1959 when
logging allocations were reduced. This allocation is a 50%
reduction on the quota granted last year. The reason for this
is the increased shortage of mature Red Gum available
(PROV VPRS 11563/P0001. Forests Commission, General
correspondence les, Unit 3, report re. logging allocations
(Melbourne)). It appears logging continued as licenses to
harvest red gum continued to be allocated in extended areas
of the Victoria Valley until no area was left untouched. Logging
was phased out in 1987 after the declaration of the National Park
(Fig. 2).
Stock grazing continued in the Victoria Valley until the
1930s, when all grazing licenses were abolished (Fig. 2). The
Commission is now considering the necessity of abolishing all
grazing occupation licenses in Victoria Valley and admitting
during part of the year limited numbers of sheep and cattle for
temporary agistment only (PROV 1933). Reasons cited were
Australian Journal of Botany 167
because of re danger and protection of the wattle industry.
You are informed that the Commission now considers this
area should not be let for the present at any rate as there is
too much danger from re and you consider it a valuable wattle
area (PROV 1933).
3. National Park phase (1984present)
The Grampians was declared a National Park in 1984, with
conservation one of its major priorities, and most utilisation
ceased at this stage. A draft management plan for the National
Park mentions L. scoparium encroachment because of concern
for Diuris punctata (a rare orchid) that was known to occur in the
study area: closure of Moora track will protect D. punctata and
any form of soil disturbance could cause disappearance of the
plants. If tea-trees [Leptospermum scoparium] on the stand
perimeter prove invasive, they must be removed by autumn
burning and further where shrub growth becomes signicant
in red gum forests burning at a low frequency (every 710 years)
may be undertaken to reduce re hazard and open the canopy to
allow light to reach the ground layer plants (National Parks
Service Victoria 1985, p. 25). Despite this, no burning was
attempted and some tea-trees were removed by hand.
Discussion
There was a substantial increase in woody-plant cover in the
woodlands of the Victoria Valley between 1948 and 1997.
Such vegetation change occurred at a faster rate between 1948
and 1971 than between 1971 and 1997. Encroachment did
not occur in response to the declaration of a National Park
(i.e. 1984); rather, it was likely a response to a complex land-
use history that has characterised the vegetation since European
occupation. Size-class analysis indicates that recruitment is
ongoing (at least since approximately 1975), with large
numbers of individuals establishing since 1994, rather than
event-driven in response to disturbance or favourable
wet years for establishment. Long-term rainfall data did not
show any clear trends associated with increased L. scoparium
cover. Rainfall may interact with disturbance regime to inuence
recruitment processes; however, it does not appear to be the key
driver inuencing shrub encroachment, particularly the recent
encroachment, which has occurred during the past 30 years,
despite long periods of drought.
Changes in atmospheric CO2 have also been hypothesised to
be partly responsible for transitions from grasslands to shrublands
during the past 200 years (Bond et al. 2003), although this has
rarely been tested. Recently, Morgan et al. (2007) found evidence
that rising CO2 may be partly responsible for woody-plant
encroachment. This is due to the increased responsiveness to
atmospheric CO2 by C3 woody plants, compared with the C4
grasses. In contrast, many studies have found C4 grasses to be
more responsive to CO2 than previously thought (see Archer
et al. 1995, for a detailed discussion). Increased atmospheric
CO2 may be an explanation for woody-plant encroachment
in the present study; however, this does not explain the
replacement of the mostly C3 grasses in the study by a C3
shrub (Archer et al. 1995), and this requires further exploration.
The oldest individuals recorded were ~30 years old, indicating
population turnover since 1948. Seedling recruitment at these
168 Australian Journal of Botany
sites is driven by the natural cycle of mortality and subsequent
release of canopy-stored seeds into woodlands with low-biomass
ground-layer vegetation. The results we report here concur with
vegetation changes that have been alluded to in the literature for
some time. Calder (1987, p. 4) referred to vegetation change in
the Victoria Valley: no longer is it possible to gallop a horse
through unroaded parts of the Victoria Valley as could be done at
the turn of the century. Further in a number of places tall tea-
trees appear to be invading red gum forest and establishing a
dense shrubby understorey (Day et al. 1984, p. 41).
Our study provides evidence of ongoing recruitment of
L. scoparium in the long-term absence of re, despite the
species being serotinous in Australia. Encroachment is not
related to a recent change in burning practises, as has been
reported for other range-expanding species (Withers and
Ashton 1977). This does not rule out re as a driver of change
in the early 1900s, at a time when burning was used to promote
Acacia for the wattle-bark industry. However, re no longer plays
a role in the ongoing expansion of this species in the Grampians
National Park. A re-return interval that occurs before
L. scoparium reaches reproductive maturity, however, may
maintain an open community as the species does not retain a
soil seed bank in New Zealand (Mohan et al. 1984) and few
germinable seeds were found under mature L. scoparium patches
at the study site (J. N. Price and J. W. Morgan, unpubl. data).
Historical documents refer to res occurring in the Victoria
Valley in the early settlement years and exclosure experiments
(to exclude kangaroos) indicate the accumulation of grassy fuel
can be substantial (J. N. Price and J. W. Morgan, pers. obs. 2006).
More recently, we observed that despite landscape-scale re
affecting the Grampians National Park in 2006 (burning ~60%
of the park, and the area surrounding the study site), woodlands
are now largely non-ammable because of the lack of ne ground
fuels. Reductions in re frequencies because of overgrazing can
favour establishment of woody plants and have been frequently
reported in semiarid and savanna regions (Harrington et al. 1976).
Sharp and Whittaker (2003) suggested that encroachment in
northern Australian savanna habitats was a direct consequence
of extreme overgrazing by cattle reducing the ammability of the
herbaceous vegetation and, in the absence of regular re, woody
vegetation increased rapidly. Roques et al. (2001) also suggested
that the critical mechanism by which grazing inuences
encroachment is through its effect on re, whereas Van Auken
(2000) argued the woody-plant encroachment is largely driven by
high levels of herbivory which reduce biomass and consequently,
re frequencies, hence allowing re-sensitive shrubs to recruit.
Stock grazing may have initially changed the grassy-
woodland vegetation and facilitated incursion of woody plants.
Indeed, early reports of the Victoria Valley describe a land that
is no longer observable today, even within remaining open
patches. References in historical documents to the area being
good grass country, implies a dense layer of grasses that may
have competitively excluded shrubs and the danger from re
suggests that the understorey vegetation was ammable, which
may have minimised shrub establishment. Competition from
pasture species has been found to decrease germination of
L. scoparium in New Zealand (Ledgard and Davis 2004) and,
hence, a mechanism that prevented widespread establishment
of shrubs can be readily hypothesised, i.e. competition from
J. N. Price and J. W. Morgan
perennial tussock grasses inhibits growth and survival of
woody invaders, and regular re kills re-sensitive seedlings.
Woody-plant expansion appears to have followed the
removal of stock grazing from the Victoria Valley in the
1930s, consistent with the results of other studies in south-
eastern Australia (Bennett 1994; Costello et al. 2000; Franco
and Morgan 2007). A shift in grazing pressure from stock and
macropod herbivory to mostly macropods can have an impact
on vegetation structure and composition because of selective
grazing and palatability (Gardiner 1986a, 1986b; Bennett 1994).
Kangaroos are generally considered grass eaters and will seldom
browse shrubs, whereas sheep are more of a generalist herbivore
(Dawson 1989; Edwards 1989). In semiarid regions, kangaroo
grazing, by suppressing the recruitment and survival of perennial
grasses, has promoted the spread of woody weeds (Gardiner
1986a). Expansion of Leptospermum laevigatum in southern
Australia was associated with the removal of cattle grazing as
well as increased densities of kangaroos providing gaps for shrub
recruitment (Bennett 1994).
Despite the obvious convenience of suggesting that grazing
and re suppression play a (the) key role in vegetation changes at
the Victoria Valley, other disturbances were also common. The
area has been subject to a long history of forestry activities
(c. 100 years), including wattle-bark stripping, and it is unclear
what impact this may have had on vegetation patterns and
dynamics. The removal of trees can inuence understorey
patterns by modifying resource availability and by altering
competitive interactions (Walker et al. 1986; Harrington and
Johns 1990). Large-scale removal of wattles, which were the main
midstorey species, may also have had a dramatic impact on co-
occurring species and, in particular, may have created canopy
gaps for L. scoparium establishment (although this would also
depend on the response of herbaceous species to canopy
removal). Experiments are needed to manipulate key resources
(light, bare ground, re) to further rene our understanding of the
factors that promote shrub establishment in woodlands.
Conclusions
Land-use changes appear to be a common factor implicated in the
establishment and encroachment of woody plants at broad scales
in southern Australia. This seems to be particularly true where
relative abundances of indigenous shrubs have increased in situ,
rather than considering the case of the range expansion of native
shrubs into new habitats, where other factors such as increased
dispersal may be partly responsible (Gleadow and Ashton 1981).
On the basis of these studies in southern Australia, we propose a
generalised model, relating land-use history to vegetation change.
It is generally accepted that indigenous re management strongly
affected vegetation patterns (Bowman 1998; Bradstock et al.
2002) and that re frequencies have been reduced since 1750.
Prior to the introduction of stock grazing, it has been suggested
that regular burning because of the abundant fuel build up resulted
in high mortality of woody plants (Noble 1997). Stock grazing
may have played the same role when at high density and in areas
where re frequencies were reduced. Several studies have
reported increases in cover of re-sensitive woody plants in
the long-term absence of re (Withers and Ashton 1977;
Gleadow and Ashton 1981; Molnar et al. 1989), as well as the
Long-term changes in woody-plant cover in woodlands
encroachment of vegetation, coinciding with the removal of
stock grazing (Bennett 1994; Singer and Burgman 1999;
Costello et al. 2000). Stock grazing may have reduced dense
native (competitive) vegetation and this, combined with lack of
mortality from re, enabled woody plants to recruit. In some
cases, this process may have been amplied by increased
densities of native herbivores, which selectively graze
competitive native grasses and avoid shrubs.
By incorporating an historical perspective, it becomes
apparent that areas that are now considered shrubby
woodland are a recent transition in response to a particular
land-use history, rather than natural processes in a classic
successional sense. If current conservation management in
Victoria is shifting open ecosystems to shrub thickets on a
broad scale, and further studies are required to determine this,
then it may be necessary to incorporate the overabundance of
native species into the conservation-management framework.
This may require making value judgements about what
communities or states we want to conserve. Open woodlands
with grassy understoreys were more prevalent, in this region, at
the time of European occupation and have been dramatically
reduced in extent and condition (Lunt and Bennett 1999).
Increased shrub cover has been associated with reductions in
species richness in the community studied here (Price and Morgan
2008) and in other communities (Mullett and Simmons 1995;
Costello et al. 2000). Loss of biodiversity resulting from
woody-plant expansion therefore poses signicant threats to
conservation of herb-rich woodlands in Victoria. In the
absence of any major land-use change to combat thickening,
it is predicted that encroachment will continue to occur in many
open ecosystems in southern Australia, with possible negative
implications for conservation. Further information is required
to determine whether recovery of shrub-encroached sites is
possible. However, the ability of the herbaceous vegetation to
recover will be constrained by (a) the lack of soil seed banks
for many of the open-woodland species (Price and Morgan 2008)
and (b) ongoing heavy grazing by native herbivores.
Acknowledgements
Logistical and nancial support for this research was provided by the
Department of Botany, La Trobe University. Marty Gent assisted with
aerial photograph analysis and Ian Lunt provided us with valuable insights
into the wattle-bark industry. The manuscript was greatly improved by helpful
comments from Wal Whalley, Bob Parsons and four anonymous reviewers.
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Manuscript received 26 April 2007, accepted 31 March 2009