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Effects of Fire Frequency on Plant Species Composition of Sandstone Communities in the Sydney Region: Inter-Fire Interval and Time-Since-Fire

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Effects of Fire Frequency on Plant Species Composition of Sandstone Communities in the Sydney Region: Inter-Fire Interval and Time-Since-Fire Australian Joumal of Ecology (1995) 20, 239-247 Effects of fire frequency on plant species composition of sandstone communities in the Sydney region: Inter-fire interval and time-since-fire DAVID A. MORRISON, GEOFFREY J. CARY,* STUART M. PENGELLY, DAVID G. ROSS, BRUCE J. MULLINS, COLLETTE R. THOMAS AND TIMOTHY S. ANDERSON Department of Applied Biology, University of Technology Sydney, PO Box 123, Broadway, NSW 2007, Australia Abstract Eire frequency is the number of fires experienced by a particular community within a given time period. This concept can potentially be resolved into a number of interacting variables, including: time since the most recent fire, the length of the inter-fire intervals, and the variability of the length of the inter-fire intervals. We estimated the effects of these three variables on the floristic composition of 65 samples from dry sclerophyll vegetation with different fire histories in Brisbane Water, Ku-ring-gai Chase and Royal National Parks near Sydney. Our analyses suggest that fire frequency may account for about 60% of the floristic variation among our samples. They confirm the hypothesis that the recent (<30 years) fire frequency produces effects on floristic composition of fire-prone communities that can recognizably be attributed both to the time since the most recent fire and to the length of the intervals between fires. These effects are equal in magnitude but are different in the nature of the floristic variation they are associated with. Increasing time-since-fire is associated with a decline in the evenness of fire-tolerant species, indicating that fewer of these species come to dominate the community in the prolonged absence of fire. Herbs and small shrubs decrease in abundance, while larger shrubs increase in abundance. Inter-fire intervals of decreasing length are associated with a decrease in the evenness of the fire-sensitive species, particularly those large Proteaceae shrubs that often dominate the community biomass in dry sclerophyll shrublands of southeastern Australia. Furthermore, the variation associated with inter-fire intervals is not necessarily solely related to the shortest inter-fire interval, but is related to combinations of inter-fire intervals through time. Thus, increasing variability of the length of the inter-fire intervals is associated with an increase in the species richness of both fire-sensitive and fire-tolerant species, implying that it may be variation of the inter-fire intervals through time that is primarily responsible for maintaining the presence of a wide variety of plant species in a particular community. Our results also suggest that the floristic variation associated with different inter-fire intervals decreases with increasing time-since-fire. Key words: fire, frequency, inter-fire interval, species composition. INTRODUCTION Australia, and has been since the mid-Quaternary (Gill et al. 1981). Fire regimes are usually considered to have Fire is a common source of change in the plant species ^j^^^^ interrelated components that can affect plant composition of the dry sclerophyll communities of community composition: fire frequency, season of occur- rence and fire intensity (Gill 1975). Effects of fire intensity .^ ,, _ _ . „ u c u 1 r on the floristic composition of dry sclerophyll com- *Present address: Ecosystetti Dynatnics, Research School of . • , , , • , n j- j • A I- Biological Sciences, Attstralian Natiotial University, ACT 2601, ^"1^'"^^ ^ave been relatively well Studied in Australia Australia. (c-g- Gill & Groves 1980; Christensen et al. 1981; Accepted for publication May 1994. Bradstock & Myerscough 1988; Auld & O'Connell 1991; 240 D. A. MORRISON ET AL. Hamilton et al. 1991), and fire intensity influences seed period; e.g. Bradstock & O'Connell 1988). Similarly, new release, breaking of seed dormancy and the suitability of individuals of fire-tolerant species will not be recruited the seed-bed for germination. On the other hand, the to the population if an inter-fire interval is shorter than effects of fire season are relatively poorly understood the time taken for the plants to reach first post-fire (e.g. Bradstock & O'Connell 1988; Enright & Lamont reproduction (the secondary juvenile period) and the 1989; Cowling et al. 1990; Lamont et al. 1991), but time taken by juvenile plants to reach fire-tolerant size appear to be important for the establishment of seedlings (e.g. Bradstock & Myerscough 1988). Furthermore, and the survival of resprouting individuals. species may disappear from the above-ground community However, most Australian work on plant-fire inter- during long fire-free periods, persisting as seeds or actions has concentrated on the effects of fire frequency. below-ground storage organs, but eventually becoming Fire frequency as originally described by Gill (1975) is a locally extinct if fire is withheld (e.g. Specht et al. 1958). function of the number of fires experienced by a particular It is thus apparent that time-since-fire and inter-fire community within a given time period. The effects of fire interval may have distinctly different effects on plant frequency on plant communities can therefore be resolved species composition (via effects on different demographic into at least two variables (Nieuwenhuis 1987): (i) the characteristics of the plants), and that these different shortest time interval between fires; and (ii) the time effects have not always been clearly distinguished in since the most recent fire. These variables are interrelated, previous studies of fire frequency. Consequently, the because as the number of fires per unit time changes so nature of the relationship between the components of fire does the average length of the inter-fire intervals in any frequency remain largely unknown, as is their relative one period as well as the time-since-fire. However, the importance. Therefore, we wished to investigate the majority of studies concerning fire frequency effects on interactive effects of the components of fire frequency in Australian plant communities have focused on the more detail. In this paper we report a study ofthe effects successional effects of time-since-fire (e.g. Jarrett & Petrie of fire frequency on the floristic composition of common 1929; Specht et al. 1958; Cochrane 1963; Purdie & plant communities of the Sydney region. In particular, Slatyer 1976; Siddiqi et al. 1976; Purdie 1977; Russell & we estimated the effects of: time since the most recent Parsons 1978; Bell & Koch 1980; Clemens & Franklin fire, the average length of the inter-fire intervals and the 1980; Posamentier et al. 1981; Brown & Podger 1982; variability of the length of the inter-fire intervals. Brown & Hopkins 1983; Bell et al. 1984; Clark 1988; Fox Our study was a purely descriptive (i.e. non-manipu- 1988; McFarland 1988) while the effects of inter-fire lative) one, relying entirely on the sampling of fortuitously interval have been less intensively studied (e.g. Fox & available areas in the field with different recent histories Fox 1986a; Bowman et al. 1988; Fensham 1990). Only of fire frequency. Such an experimental design is not Bradfield (1981) and Nieuwenhuis (1987) have explicitly ideal (e.g. see Gill 1977), but, given the practical compared the relative effects of these two variables on a difficulties of manipulating fire regimes over long time single plant community. periods (particularly near urban areas), large-scale manipu- The floristic composition of a community after a fire is lative field experiments for the study of fire frequencies dependent on the pre-bum composition, but it can vary may be logistically impossible. Furthermore, our study significantly away from it. In particular, species richness was only concerned with the above-ground component and diversity tend to increase immediately after the fire of plant species composition, as we did not quantify the and then to gradually decline (e.g. Specht 1981; Fox & soil-stored seed bank for any species (see Leek et al. Fox 1986b). Initial regrowth is usually by those species 1989). whose adult plants can survive fire (fire-tolerant species), while those species whose adult plants do not normally survive fire (fire-sensitive species) tend to contribute more to the above-ground community several years later. METHODS Furthermore, the abundance of fire-sensitive species Our work was carried out in dry sclerophyll communities tends to decrease as the number of fires increases, while in three of the coastal national parks of the Sydney fire-tolerant species show more variable responses (Fox region. Samples were collected from the area between & Fox 1986a; Nieuwenhuis 1987). Staples Lookout and Kariong in Brisbane Water National The demographic effects of fire frequency therefore Park in winter 1992 (33°27'S, 151°17'W: 22 samples), on appear to be primarily concerned with the time taken for the Lambert Peninsula in Ku-ring-gai Chase National the plants to reach a life-history stage that enables the Park in spring 1991 (33°37'S, 151°16'W: 24 samples), population to survive a subsequent fire. For example, the and between Loftus Heights and WattamoUa in Royal local extinction of fire-sensitive species will occur if an National Park in spring 1992 (34°07'S, 151°05'W: 19 inter-fire interval is shorter than the time taken for the samples; Table 1). All samples were taken from Hawkes- plants to reach first reproduction (the primary juvenile bury Sandstone plateaus and ridgetops {ca 150 m a.s.l.). FIRE FREQUENCY AND PLANT SPECIES COMPOSITION 241 in closed scrub and low to low-open woodland or forest all fires since 1964. Suitable replicate samphng areas (Benson & Fallding 1981; Thomas & Benson 1985). were then chosen with respect to vegetation type and These are the most widespread vegetation types of the ease of access. dominant sandstone areas in the Sydney region (Benson Our sampling design assumed that the fire history of & Howell 1990). each of the areas within each park was similar (or at least Sample areas were chosen to represent the range of fire randomized) before detailed records were started in the frequencies present in each national park in terms of early 1960s.
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