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II. Scientific Evidence Regarding Rainforest Ecology and Protection

6. VICTORIAN RAINFOREST FIRE ECOLOGY

It has long been recognised by foresters, botanists and ecologists that under appropriate fire regimes in south eastern Australia, sclerophyll forests would replace rainforests and that, in the absence of fire, rainforests on many sites would replace adjacent sclerophyll vegetation (Casson 1952, Gilbert 1959, Cremer 1960, Cremer and Mount 1965, Webb 1968, Howard 1973a,b, Ashton and Frankenberg 1976, Ashton 1981a, Brown and Podger 1982, Smith and Guyer 1983, Hill and Read 1984, Ellis 1985, McMahon 1987, Ash 1988). Knowledge of eucalypt regeneration ecology has led to the extensive use of high intensity burning of logging residue and broadcast sowing of seed in logged areas with the purpose of encouraging the regeneration of eucalypts after clear felling (Frankcombe 1961, Ellis 1971, Gilbert and Cunningham 1972, Ellis and Pennington 1992, Attiwill 1994b). If the ecotone adjacent to clearfall areas burns, it will be detrimental to competing rainforest species. Although it is not the primary motivation, these fires may act to reduce the likelihood and severity of wildfire, but carry the cost of the potential for fire damage to rainforest species in the buffer from the site preparation burn itself (Horne and Hickey 1991). The moisture differential between eucalypt forest and rainforest has been used to limit the spread of fire (CFL 1987, 1989), usually in fuel reduction burns in damp forest containing warm temperate rainforest. If fire breaks are used instead, the risk to downhill rainforest vegetation in buffers is slight. Boundary tracks may have impacts of their own including increased soil disturbance and runoff, and elevated pioneer and weed invasion (see discussion below). However, it is not possible to be prescriptive about the use of boundary tracks or fire breaks without taking into account the character and local conditions of an area.

There is no doubt that southern Australian cool and warm temperate rainforests occasionally burn under natural disturbance regimes (Cameron 1992). For example, Neyland and Brown (1994) found charcoal in nearly all surface soils collected from 59 cool temperate rainforest stands in eastern Tasmania. The prevalence of fire in temperate rainforest communities is reflected in the ecological dynamics of these communities. Ellis (1985) described successional relationships among 12 vegetation types from the highlands of north-east Tasmania, including four postulated pathways by which rainforest may become established. The time taken for succession to proceed from eucalypt forest to rainforest depends on the fire history and topographically determined local climate. The long association between rainforest and fire may also be reflected in the possibility that fire may have shaped the evolution of the regeneration characteristics of many rainforest species. For example, A. smithii is resistant to irregular burning because it develops a lignotuber which confers a coppicing ability although the species does not coppice as successfully as adjacent eucalypts (Johnston and Lacey 1983). Most Victorian temperate rainforest species are not obligate reseeders and are able to resprout after fire (McMahon 1987, Chesterfield et al. 1990). McMahon (1992) reported that 39% of the perennial species of East Gippsland warm temperate rainforest are obligate reseeders and 41% have the ability to resprout. One important exception is Pittosporum undulatum, a primary warm temperate rainforest species and dry rainforest canopy species, which has no obvious means of resisting fire beyond seed stores and weak resprouting following crown damage.

The presence or absence of disturbance is not at issue in the management of Victorian rainforests. Rather, the frequency and intensity of fire and its spatial and temporal distribution are central to effective management. Frequent, hot fire will eliminate rainforest from a site. The importance of these factors is illustrated by the fact that A. smithii is eliminated by a single intense fire and by recurrent, frequent fire, even though the species is resistant to infrequent fire that involves canopy scorch only. Thus, Ashton and Frankenberg (1976) noted that A. smithii rainforest on Wilson's Promontory is a climax community only in fire-sheltered lowland gullies. The importance of fire in determining the distribution of temperate rainforest was emphasised by the results of distributional and bioclimatic studies of N. cunninghamii. Howard and Ashton (1973), Howard (1981), and Busby (1986) suggested that recurrent fire limits the distribution of N. cunninghamii, interrupting the expansion of the species to suitable habitat in the Central Highlands of Victoria. Busby (1986, 1987, 1992) concluded that Victorian primary cool temperate rainforests occupy only about 20% of their potential area, largely because of fire frequency. Thus, when considering rainforest protection, one must consider the impacts of human activities on fire frequency and intensity.

Cameron (1992) suggested that fires normally function to stabilise and maintain sharp boundaries between rainforest and sclerophyll vegetation, 'except under conditions of exceptional fire hazard such as prevailed in the 1982/83 fire season' (Cameron 1992, p. 20). In more usual conditions, the steep moisture gradients between the two communities ensure that the likelihood of spread of fire into rainforest is less than the likelihood of spread into adjacent sclerophyll communities. Similarly, Greenlef (1990) noted that moisture gradients are important determinants of vegetation patterns in each of five different historical fire regimes in Sequoia sempervirens communities in the western United States.

Within a fire interval, there is ample evidence that Victorian rainforest species will tend to spread into adjacent sclerophyllous vegetation, contracting abruptly with the ensuing fire event. Cameron suggested that if the fires are sufficiently intense and frequent, then sclerophyll vegetation will dominate and even permanently replace, rainforest (see also McMahon 1987). The long term prognosis for rainforest stands versus sclerophyll forest, depends on the frequency, intensity and variability of fire events. If the fire interval is short, or there is a high probability of successive fires with a relatively short fire interval, then it is likely that rainforest will be lost. The likelihood that rainforest will be lost after a fire is a function of the intensity of the fire, and of the post-fire regenerative success of eucalypts. Cameron (1992 p. 22) postulated 'moderately successful' initial recovery following a single fire catastrophe, and progressive structural collapse and ultimate elimination of rainforest following 'at most two further fire events'. That is, he suggested that secondary succession may be truncated by successive fire events if the fire interval is short. The distribution of rainforest and rainforest ecotones reflects both the detailed history of a site, in particular its fire history, and the underlying environmental gradients, especially on sites where gradients are gentle, and ecotones are broad and diffuse (Melick and Ashton 1991).

Fuel reduction burning and seed-bed preparation result in an element of risk of the burn escaping to adjacent rainforest. Current rotations at one coupe mean exposure to such risk every 60-100 years for some stands. Cameron (1992) speculated that, in addition, adjacent old sclerophyll forest affords more protection to rainforest from wildfire than does regrowth forest. This suggestion is based on three postulates. First, during the juvenile and mature growth phases, regrowth forest uses more water than old-growth forest and therefore the soil has lower moisture content. Second, more mesic species with broad-leaved canopies present in the old-growth forest will be replaced by more flammable, sclerophyll understorey species. Third, Cameron speculated that the greater height of the canopy fuel in old-growth forest contributes to a fire shadow in circumstances when crown fire may threaten rainforest.

Old forests occupy the least fire prone sites in the landscape. Topography, moisture levels, the composition of the understorey and the height of the canopy fuel may combine to reduce fire intensity and the severity of crown damage. In general, fire intensity is determined by the amount of fuel, its dryness and to some extent its vertical spatial distribution. Fuel loads in ash forests typically are very high in any forest type at the pole stage and beyond.

There is sufficient empirical evidence to suggest that some of Cameron's (1992) concerns have scientific justification. McMahon (1987, 1992) evaluated the impact of the 1982/83 wild fires on Victorian rainforest stands and summarised his qualitative observations as follows. Topographical features were one important factor determining fire intensity at a particular site. Warm temperate rainforest margins with mixed open canopies of A. smithii and eucalypt species invariably experienced a crown fire, whereas closed rainforest experienced surface fires or less intense crown fires. The fire effects on these two structural types were likely to be different because of moisture retention and flammability. The fires stimulated opportunistic species and fire pioneers usually rare in mature rainforest. Surface fires under a closed canopy had minimal structural or floristic effects. Narrow stands in close proximity to eucalypt forest were more susceptible to crown fire. In areas of complete rainforest canopy removal, post-fire opportunists dominated the stand. The distribution of sclerophyllous recruits was related to the degree of closure of the pre-fire canopy, the level of fire-induced canopy damage, and the availability of seed in the canopies of emergent eucalypts. Increasing severity of fire retards the rate of rainforest canopy recovery because Acmena smithii tends to regenerate from epicormic shoots and basal resprouts with increasing severity (see also, Chesterfield et al. 1990). It will only regenerate successfully from seed in undamaged stands. Abundant recruitment of sclerophyll species suggested that floristic and structural change following intense fire may be long-term (of the order of 300-500 years, the life expectancy of each eucalypt cohort), even in some areas that were previously closed mature rainforest (McMahon 1992; Woodgate et al. 1994).

Other studies provide confirmatory, if indirect, evidence that Cameron's (1992) speculations summarised above have substance. Brown and Podger (1982) found that the persistence of populations of Huon pine, King Billy pine, pencil pine (Athrotaxis cupressoides) and deciduous beech (Nothofagus gunnii) in cool temperate Tasmanian rainforest depend on the absence of fire for prolonged periods. Hill (1982) and Hill and Read (1984) found fire behaviour in cool temperate rainforest to be an important determinant of post-fire regeneration. Fire intensity and proximity of sclerophyll seed source may lead to long term structural and compositional change. Barker (1991) documented a population of the native conifer Podocarpus ssp. nov. (Ross 1993) in cool temperate rainforest at Goonmirk Rocks in Victoria. Barker suggested that its persistence in this unique habitat is facilitated by the broken canopy of the marginal habitat and the long-term absence of fire. Chesterfield et al. (1991) noted that populations of E. nitens which have since been described as the Victorian endemic E. denticulata, have a restricted distribution, even though the species is a relatively vigorously growing eucalypt. They suggested that it competes for habitat with rainforest understorey species in locations where fire is rare, and that it is relatively shade intolerant.

Overall, there are insufficient data available to judge conclusively the validity or otherwise of Cameron's (1992) arguments, and they remain speculative. The interaction between fire frequency, regeneration niche, and competitive tolerance is a pervasive feature of the ecology of Victorian forests. For example, it is responsible for structuring the boundaries between E. obliqua and E. regnans (Ashton 1981). Generally, the ignition probability and intensity of fires in these forests are closely related to fuel moisture content. Thus, our ability to predict accurately the impacts of management on the distribution of rainforest in the medium to long term will depend on our ability to understand the relationship between topography, climate, fuel loads, fuel moisture content, fire intensities, fire probabilities and other stochastic processes, and the competitive interactions among species. Such understanding is an essential component of strategic planning for rainforest protection.
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6.1 Characteristics of natural fires

Given that natural disturbance regimes are the appropriate model for forest management (Attiwill, 1994b) and that fire is the dominant ecological process driving disturbance dynamics in Australian forests (Attiwill 1994a), it would seem important to characterise the patterns of natural fire events in the past. Characterisation should include knowledge of fire probability and intensity in different forest types, and the spatial and temporal pattern of fire.

Perhaps surprisingly, there is very little known about historical, natural fire regimes in Victoria, or in other, analogous landscapes in Australia. King (1963, in Bowman and Jackson 1981) suggested that fires in high rainfall areas became less frequent but more intense as a result of colonisation of the Australian environment by Europeans and elimination of Aboriginal influence on the landscape. Cremer (1960) noted that a reduction in fire frequencies would have resulted in the expansion of rainforest species into adjacent sclerophyll forest. Jackson (1968) suggested that in Tasmanian forests, areas with fires at 150-200 year intervals carry mixed forest (rainforest with emergent eucalypts), and areas with lower frequency fires carry pure rainforest. All of these statements were essentially speculative, based on little or no direct field evidence.

Most fires in ash forests occur during relatively dry periods. Given the amount of fuel and the flammability of the vegetation, they tend to be major fires. Attiwill (1994b) noted six such fire events in the last 200 years in south eastern Australia, occurring in 1851, 1898, 1926, 1932, 1939 and 1983. Vines (1974) stated that 'very bad' fires occurred in Victoria in 1851, 1886, 1913, 1926, 1939, 1952 and 1965, and that minor forest-fires occurred in 1920, 1932, 1944 and 1960. Complete fire record dates are available only from 1910. Vines (1974) suggested that major wildfires occur about every 13 years in Victorian forests, based on an analysis of weather patterns and associated fire events. His prediction was not corroborated by subsequent fires. Such analyses treat State-wide or regional data and are somewhat irrelevant for ecological purposes unless quantified for different climate and landscape zones, and for a particular point in the landscape.

Woodgate et al. (1994) recorded that fire frequencies from eight trees ranging from regrowth to senescent in an E. sieberi stand in East Gippsland averaged about every 22 years between 1800 and 1992, and every 30 to 40 years prior to 1800. They concluded that fires at 20 to 50 year intervals are likely to represent the long-term average in this ridge community. In the absence of historical fire data such as these (scant though they are), any attempt to use natural disturbance regimes as a model for forest management will be hampered by a lack of necessary information. It is possible to suggest, for example, that fire frequencies and intensities in damp forest on ridge tops are very likely to be different to those in rainforest occurring at the bottom of gullies or adjacent to wet or montane forest. However, we do not know the expected frequency or the spatial or temporal variance of these fires. Such data are essential for specifying acceptable bounds on forest disturbance in managed landscapes.

Even if the characteristics of natural fire regimes were known, the silvicultural practices now in operation in the forests surrounding rainforest are unlikely to closely mimic natural disturbance. The effects of natural fires are unlikely to be the same as the effects of logging followed by site preparation burns because the spatial pattern, intensity, the effects on eucalypt, tree fern and understorey tree survival, and the distribution of surviving individuals as well as the distribution and type of fuels that remain after these disturbances will be different (eg., Mount 1979, Mueck and Peacock 1992, Ough and Ross 1992, Kutiel and Shaviv 1992; see the example in Figure 3). When State Forests available for timber production in Victoria burn, they are usually salvage logged. In contrast, natural fires leave behind much greater structural complexity in the form of dead stags, logs, recovering trees, resprouting shrubs and ferns than is normally found in areas that are clear felled, or areas that are salvage logged after natural fire (Franklin 1988, 1992, 1993). Furthermore, regeneration burns are likely to be smaller in area than other fires. For example, the fires of 1939 burned a larger area than will be logged and regenerated in the following 80 years. Wildfires may create more complex edges than are found in logged landscapes (Franklin 1993). They are likely to burn areas that would not normally be logged such as stream sides and forests on steep terrain. These differences are likely to influence many factors including wind dynamics and fuel loads, and hence future fire probabilities and intensities.

If we ignore, for the moment, the spatial extent of fires, we may examine the effect of random intervals between fires on the expected age structure of a forest when the extent of disturbance from logging and natural fire regimes is the same (Fig. 3). The deterministic regime is represented by a 200 year rotation, and one block is harvested each year. The fire regime is equivalent to a geometric random variable. Consider a forest as being composed of numerous areas. If each area is disturbed with an annual probability of p, then the expected proportion of areas that are n years old is equal to the proportion of areas disturbed n years ago (=p) multiplied by the probability that these areas are not disturbed subsequently (=[1-pn-1). Therefore, the age structure of the forest is described by the function which gives the proportion of forest that is expected to be n years old. The average time between disturbance events is equal to 1/p (after McCarthy and Burgman, in press).

McCarthy and Burgman (in press) suggested that the result in Figure 3 relates only to the "expected" age composition of the forest, representing the mean structure. There is no indication of fluctuations in age structure over time. If disturbance occurs at a small scale relative to the forest, such as may be due to individual treefall, or harvesting a small proportion of the forest each year, then the age structure of the entire forest will not fluctuate very much. At a finer scale (ie., at the scale of disturbance)
the age of the stands will fluctuate widely.
<p><div align='Left'><IMG SRC="/CA25677D007DC87D/LUbyDesc/0417.gif/$File/0417.gif" width='583' height='399' hspace='5' vspace='2' alt='Graph: Figure 3 The expected age structure of a forest under stochastic (fire) and deterministic (logging) disturbance regimes, for a forest divided into 200 blocks.'></div>

<b>Figure 3</b>. <font size=2 face="Arial">The expected age structure
of a forest under stochastic (fire) and deterministic (logging)
disturbance regimes, for a forest divided into 200 blocks.<br>
<br>
</font>
<p>
If disturbance occurs over wider spatial scales, then fluctuations
in the age structure of forests will be more important. This
may be illustrated by considering fluctuations in age structure
resulting from the simple stochastic disturbance regime described
above. We will define the spatial scale of disturbance as the
proportion of the forest that is affected by each disturbance
event. In this case, the standard deviation of the abundance
of a particular age class will be proportional to the square root
of the spatial scale of disturbance. If disturbance occurs over
moderately large areas, then we would expect the actual forest
age structure at any particular time to be different from the
expected forest age structure (McCarthy and Burgman, in press).
The model is much too simple to provide quantitative results
of any value, but the qualitative result has implications for
the management of forests that are subject to large scale disturbance
regimes such as the ash forests of Victoria. The age composition
that exists today (or at any other time) is unlikely to represent
an &quot;expected&quot; or &quot;average&quot; structure; the
natural regime is characterised by fluctuations in the age structure
of the forest. If the natural disturbance of forest ecosystems
is to be used as a model for management, then accounting for stochasticity
is likely to be important.<br>
<br>
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<p>
<a href="#top"><img src="http://exwb01.nre.vic.gov.au/images/c_top.gif" alt="Page Top" width=24 height=20 border=0></a><p>
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<font size=-1><b><a name=6.2>6.2 Ecotone disturbance</a><br>
</b></font>
<p>
An important issue in rainforest protection is concerned with
the interaction between sclerophyll forest and rainforest in the
ecotone between the two communities. In wood utilisation areas,
this ecotone is assumed to be largely within rainforest buffers.
This assumption is more likely to be true in steeper, more dissected
country, particularly for warm temperate rainforest stands. However,
the ecotone between cool temperate rainforest and sclerophyll<b>
</b>forest is typically wider, frequently exceeding 20m. Given
that, historically, the two vegetation types are in a state of
dynamic flux determined by topography and time since disturbance,
particularly fire intensity and frequency (Casson 1952, Webb 1968,
Ashton 1981a, Melick 1990a, Cameron 1992, McMahon 1992, Attiwill
1994b), the pattern of human disturbance of the ecotone and its
effect on rainforest is central to this review. <br>
<p>
Boundaries between rainforest and sclerophyll forest communities
have dynamics that span decades or centuries. Some eucalypt trees
that dominate the forests in which stands of rainforest are embedded
live for 400 years or more (see the section on 'Time Scales of
Rainforest Ecological Processes'). In areas where the two vegetation
types interact, there is direct competition for light, moisture
and space, and mature eucalypts may often be found in places where
there is currently no eucalypt recruitment. This region is the
edge of the competitive range of both sclerophyll community species
and rainforest species, and the predominance of one or the other
is largely a matter of local climatic conditions, topography,
and historical factors that have contributed to past fire frequency
and intensity. Thus, the conditions one observes at a point in
the present depend on the times since any previous disturbances,
the severity and kind of those disturbances, and the stochastic
processes that determine the ecological dynamics following each
disturbance event.<br>
<p>
There are many examples in which ecosystems that experience anthropogenic
disturbance appear to return to their predisturbance condition
(eg., Tranquillini 1979). However, there is clear evidence that
distinct vegetation boundaries can fail to recover from disturbance
(eg., Black and Bliss 1980), especially when plant communities
are at the edge of their range of tolerance of biotic and abiotic
conditions. Arguably, all three Victorian rainforest types fulfil
this edge of range scenario. There is little doubt that mature
rainforest can recover from intense wildfire in the absence of
eucalypt re-establishment, or that mature stands can be lost following
a single fire (eg., recovery of warm temperate rainforest at Jones
Creek following the 1982/83 fires). <b> </b>However, there is
no reason to believe that all Victorian rainforest stands will
necessarily recover after fire, or that they will be replaced
by sclerophyll forest after intense or frequent fires. Likewise,
one cannot be certain that changes in environmental conditions
will alter the ecology of rainforest, mediated through their effects
on fire risk (Cameron 1992) or otherwise. Different kinds of
rainforest are likely to respond differently to any disturbances
they experience, and the impact of climate and topography in ameliorating
the effects of fire are likely to have different impacts on different
rainforest types. Furthermore, stands of different kinds of rainforest
are embedded in different kinds of eucalypt forest. All of these
factors will interact to determine the probability of fire for
a rainforest stand, and the consequences of a fire, should it
occur. These factors suggest that the best way to evaluate such
impacts is to develop models to explain, predict and test fire
behaviour and fire ecology in relation to rainforest in each environment.
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