Journal of the Royal Society of Western Australia, 91; 163-174, 2008 Post-fire juvenile period of plants in south-west Australia forests and implications for fire management N D Burrows’, G Wardell-Johnson^ & B Ward^ ‘ Department of Environment & Conservation, Locked Bag 104, Bentley Delivery Centre, WA 6983 ^Natural & Rural Systems Management, The University of Queensland, Gatton, Qld 4343 ^Department of Environment & Conservation, Locked Bag 2, Manjimup, WA 6258 Corresponding author; ^ neilb@calm.wa.gov.au Manuscript received November 2007; accepted April 2008 Abstract Regular prescribed burning to manage the accumulation of flammable live and dead vegetation (fuel) is a strategy for ameliorating wildfire impacts in fire-prone environments. The interval between prescribed fires needs to be sufficient to manage fuel accumulation but it should also be ecologically acceptable. Time to first flowering after fire (juvenile period) is a biological indicator that can be used to guide minimum intervals between fires to conserve plant diversity. A survey of 639 plant species in forests and associated ecosystems of south-west Western Australian revealed that 97% of understorey species readied flowering age within 3 years of fire and all species reached flowering age within 5 years of fire. Within species variation was evident, with plants at the drier end of their range taking longer to reach flowering age. Fire sensitive plants, being obligate seeder species with longer juvenile periods (> 3 years), mostly occurred in low rainfall zones so took longer to mature, or in habitats that were less prone to fire because they remained moist for a longer period or because surface fuels were inherently sparse and discontinuous. Due to uncertainty about the reproductive biology and seed bank dynamics of most of the flora, we recommend that the conservative minimum interval between fires that are lethal to fire sensitive plants is about twice the juvenile period of the slowest maturing species in the community. Occasional landscape fires at shorter intervals would be ecologically acceptable only if these fires were of a sufficiently low intensity as to not kill plants with long juvenile periods, or were patchy and did not burn the habitats in which they occur. Keywords: fire, juvenile period, seeders, resprouters, prescribed burning Introduction timber and amenity. Large, intense wildfires can also be environmentally damaging and present a threat to Plant communities in the south-west forest region of ecosystem processes and conservation values including Western Australia display a remarkable array of physical old growth forests, communities and species sensitive to and biological traits that enable them to persist in a fire high fire intensities, and extant populations of rare flora prone environment, and in some cases depend upon fire and fauna (see Abbott and Burrows 2003). Fire intensity, for structural and floristic diversity (e.g., Gardner 1957, or the rate of heat energy release, is a measure of the Christensen and Kimber 1975, Christensen and Abbott severity of the fire; its damage potential, suppression 1989 and Burrows and Wardell-Johnson 2003). Fire difficulty and killing power. Wildfires derive their regime is the history of fire frequency, season and energy from the quantity' and arrangement of live and intensity (Gill 1981) with fire frequency affecting both dead vegetation (fuel) that bums. Fire intensity is further the structure of the vegetation and its floristic influenced by fuel dryness, weather conditions and composition (Muir 1987, Burrows and Wardell-Johnson topography. Of the factors that determine potential fire 2003). Fire frequency also directly influences fire intensity, fuel quantity and weather conditions are the intensity because the quantity of live and dead most influential, but fuel quantity is the only factor that vegetation, which becomes fuel for a bushfire, can be effectively managed. Fuel reduction burning, the accumulates rapidly in the first 10 years after fire, after practise of purposefully setting low intensity fires under which the rate of fuel accumulation slows (Burrows 1994, defined coneJitions of fêl, weather and topography, is a McCaw etal. 1996). fire management technique to minimise the impact of Today, intense summer wildfires threaten human life, wildfires by reducing the potential size and intensity of a destroy property and damage forest values such as wildfire, thereby reducing its damage potential and enhancing suppression opportunities. Prescribed burning does not prevent wildfires from occurring but where a © Royal Society of Western Australia 2008 significant proportion of the landscape is managed this 163 Journal of the Royal Society of Western Australia, 91(2), June 2008 way, wildfire impact can be considerably reduced. While measure but flowering is a more practical measure. Gill there are few published cases where this practice has and Nicholls (1989) suggest that doubling of the juvenile effectively reduced the impact of wildfires and period of a species is a useful guide to when the species contributed to suppression success (e.g. Rodger 1961, is likely to be able to replenish its seed bank following Underwood et al. 1985, Grant and Wouters 1993), there fire. This 'rule of thumb' is consistent with a number of are numerous unpublished accounts by firefighters and individual species studies of seed production with time land managers where prescribed burning has (see review by Burrows and Wardell-Johnson 2003). ameliorated wildfires (Cheney 1994). However, the However, there is a paucity of information about the practice of prescribed burning for fuel reduction is reproductive biology and seed bank dynamics of plant controversial with concerns that it is ecologically species in Australian ecosystems (Whelan 1995, Burrows damaging in the long term. and Wardell-Johnson 2003). Until more is known, knowing the juvenile period of species in an ecosystem There have been many studies and observations of the subjected to regular fire is a useful criterion for ecological effects of a single fire on various plant species determining the minimum fire interval between lethal and communities in south-west ecosystem.s, however fires and for assisting with defining ecologically there are few published data on the long term ecological sustainable fire regimes (Tolhurst 1999, Tolhurst 2004). If effects of repeated fuel reduction burning in south-west the interval between fires that are lethal to the parent Australian forests (Christensen and Abbott 1989, plants is shorter than the time to first flowering (juvenile Burrows and Wardell-Johnson 2003). Burrows and period) and seed set, then it is reasonable to assume that Wardell-Johnson (2003) reported that 30 years of very obligate seeder species especially, could be at risk of frequent burning (3-4 year intervals) of an experimental decline. However, if the intensity or patchiness of fires at site in a southern jarrah {Eucalyptus marginata) forest short intervals is such that parent plants are not killed by did not result in any species losses, but caused changes fire, then populations may be able to persist and thrive in the relative abundance of species over time including under a regime of frequent but very low intensity, patchy a decline in the abundance of two obligate seeder species fires. and an increase in abundance of several other species. They also reported that none of the experimental fire This paper reports on post-fire regeneration strategies, regimes favoured all species, including long periods of post-fire juvenile period and flowering phenology of 639 fire exclusion. There was evidence that burning at 5-7 species that occur predominantly in the southern forests year intervals in spring and autumn caused no and associated ecosystems of south-west Western significant changes to species richness or abundance at Australia. We also discuss how other factors such as the study site. climate (rainfall), regeneration strategy and life form influence age to first flowering. In areas zoned for fuel reduction by prescribed burning, rrtanagers aim to maintain fine surface fuel quantity (dead leaves, twigs, bark and floral parts <6 mm in diameter) below about 8 t ha ’ and 19 t ha ’ for jarrah Methods and karri (E. diversicolor) forests respectively. The higher fuel loading for karri reflects its more mesic, hence less Study sites flammable condition for a longer period throughout the Information about post-fire regeneration strategies, year (McCaw and Hanstrum 2003). The time taken for flowering time (month) and juvenile period (months) fuels to reach these levels, hence the interval between was gathered from study sites in the southern forest prescribed fires, depends on the rate of fuel region of the south-west of Western Australia (Fig. 1). A accumulation. This varies across the forest region from brief description of each site is given in Table 1. The about 6 to 10 years, depending on site productivity and McCorkhill and Yendicup forest sites each comprised 10 rainfall (Sneeuwjagt and Peet 1985, Burrows 1994). For small (each 4 ha) adjacent plots that were experimentally example, in most jarrah forests, fine surface fuel quantity burnt at various times as part of a broader investigation reaches quasf-equilibrium in 15-17 years (Burrows 1994). into the long term effects of fire (see Burrows and While about 70% of jarrah forest plant species have Wardell-Johnson 2003). The Lindsay forest site (Table 1) the capacity to resprout following fire, the remainder was similar except the plots were smaller (each about 0.5 depend on seed (obligate seeders), stored either in the ha). Data from the Walpole study site (Table 1) were soil (soil-stored seeders) or in woody capsules in the accumulated from some 400 vegetation sample sites canopy (canopy-stored seeders), for regeneration located throughout a broad range of vegetation types (Christensen and Kimber 1975, Burrows and Wardell- represented in the Walpole-Nomalup National Park (see Johnson 2003). An adequate viable seed store and Wardell-Johnson et al. 1989). Post-fire regeneration conditions favourable for seed germination and plant strategy and juvenile period were assessed at each site growth are essential for flowering plants to persist in an by regular (3-4 weekly) Inspections until all species at ecosystem that is regularly burnt. For some species, fruit the site had reached flowering age. The juvenile period is production and seed set may not occur in the first quite variable amongst individuals of the same species in flowering year (Benson 1985, Wark et al. 1987, Burrows the same population, so we define juvenile period as the and Wardell-Johnson 2003) and seed production time taken for at least 50% of individuals in a population sufficient to provide adequate seed reserves to restore to reach flowering age after fire. This was assessed the population to pref-fire abundance levels may take visually. Visual inspection also ascertained the post-fire many years (Gill and McMahon 1986, Gill and Nicholls regeneration strategy, and plants were classified as 1989, Wooller et al. 2002). As observed by Gill and shown in Table 2, which has been adapted from Gill Nicholls (1989), seed production is the most relevant (1981). Some species regenerated by more than one of the 164 Burrows et ah: Post-fire juvenile period of plants in SW Australia Figure 1. Location of survey sites at McCorkhill forest, Lindsay forest, Yendicup forest and the Walpole-Nornalup National Park in south-west Western Australia from which information about the post-fire juvenile period, regeneration strategy and time of flowering of 639 plant species was gathered. Table 1 Location and general description of vegetation, soils, landform and climate of sites in south-west Australia forest and associated ecosystems surveyed for post-fire regeneration and flowering responses. Location General description of vegetation Soils and landform Climate McCorkhill forest: Havel and Mattiske (2000) vegetation complex Yellow brown gravelly Mediterranean-type 20 km west of the type 'Kingia'. Open forest, overstorey dominated sands on ridges and climate with cool wet town of Nannup. by Eucalyptus marginata and Corymbia calophylla undulating slopes. winters and warm Latitude 33° 57.4' S, 20-25 m tall. Second storey of Aliocasuarina dry summers. Mean Longitude 115° 32.5' E fraseriana, Banksia grandis and Persoonia annual rainfall longifolia to 6 m tall. Low woody shrub and herb 1 000 mm. understorey to 1 m tall. Lindsay forest: Havel and Mattiske (2000) vegetation complex Yellow duplex soils with Mediterranean-type 12 km west of the town type 'Crowea'. Tall open forest of C calophylla pale brown sandy loam with cool wet winters of Manjimup. and E. marginata with some E. diversicolor topsoil on ridges and and warm dry Latitude 34° 14.6' S, 30-35 m tall. Second storey of Xylomelum midslopes. summers. Mean longitude 115° 58.0' E. occidentals, B. grandis and P. longifolia to 6m tall. annual rainfall Woody shrub and herb understorey to 1.5 m tall. 1 150 mm. Yendicup forest: Havel and Mattiske (2000) vegetation complex Yellow brown gravely Mediterranean-type 45 km west of the town type 'Frankland Hills'. Woodland of E. marginata duplex soils with some with coot wet winters of Manjimup. and C. calophylla 15-20 m tall. No significant laterite boulders. and warm dry summers. Latitude 34° 11.6' S, second storey. Open woody shrub and herb Mean annual rainfall longitude 116° 37.0' E. understorey to 0.5 m tall. 750 mm. Walpole-Nornalup Wardell-Johson et al. (1989) described 15 Diversity of soils and Mediterranean type National Park: vegetation communities including a variety of landforms. See Wardell- climate with cool wet 18 390 ha on the south heathlands, woodlands, low open eucalypt Johnson et al. (1989) and winters and mild coast of Western forests and tall eucalypt forests. CALM (1990). summers. Mean Australia. Surrounds annual rainfall varies the town of Walpole across the Park from 1 200-1 300 mm. 165 Journal of the Royal Society of Western Australia, 91(2), June 2008 Table 2 Classification of post-fire regeneration strategies of vascular plants in south-west Australia ecosystems (adapted from Gill 1981). Seeders Resprouters (1) Stem girdling or 100% scorch kills, depends on canopy (4) Survives stem girdling or 100% scorch, soil suckers (rhizome, stored seed corm, bulb, tuber) (2) Stem girdling or 100 % scorch kills, depends on soil (5) Survives stem girdling or 100% scorch, basal sprouts stored seed (lignotuber) (3) Stem girdling or 100% scorch kills, no stored seed (6) Survives 100% scorch, epicormic shoots (8) Stem girdling or 100% scorch kills, any of 1,2,3 above (7) Survives 100% scorch, large apical bud (10) Ferns and Allies (spores) (9) Survives 100% scorch, any of 4,5,6,7 above categories shown above. For example some lower tree documented for 639 plant species representing 249 and overstorey tree species in particular, including genera and 72 families (Table 3). The most commonly Banksia grandis and Eucalyptus marginata, were capable observed families were Orchidaceae, Proteaceae, of resprouting from either a subterranean lignotuber or Papilionaceae, Myrtaceae, Epacridaceae, Asteraceae and epicormic buds, or of regenerating from seed, depending Mimosaceae. The least commonly observed families on the fire intensity' and the physiological age of the plant included Portulaceae, Cephalotaceae, Lindsaeaceae, (Burrows 1985). In these cases, species were deemed to Zamiaceae, Loranthaceae, Podocarpaceae and be resprouters and allocated to one of the resprouter Dennstacdtiaceae. At the intermediate and high rainfall categories shown in Table 2. Some thin barked tall shrub sites (McCorkhill forest near Nannup, Lindsay forest species that are typically characterised as obligate near Manjimup and Walpole-Nornalup National Park), seeders, such as Melaleuca viminea, have occasionally all upland forest understorey species reached flowering been observed to resprout following very low intensity age within in 3 years of fire, with most (90%) flowering fire which scorched the canopy but did not girdle the within 2 years (Fig. 2). At the lower rainfall site main stem. Such species were classified as obligate (Yendicup forest near Perup) all upland forest seeders in this study. understorey species reached flowering age within 4 years of fire. Of the 639 species observed, 17 (3%) species had juvenile periods >3 years and three species were Results observed to take longer than 4 years to reach flowering after fire. These were Banksia brownii (5 yrs), B. Information on post-fire regeneration strategy, seminuda (5 yrs) and Melaleuca viminea (5 yrs) (Table juvenile period and peak flowering month was 4). Species with longer juvenile periods were mostly Perup ----(cid:9632)(cid:9632)---Nannup ----A----Manjimup - - x- -Walpole Time since fire (months) Figure 2. Cumulative proportion of species to have reached flowering age with time since fire for 639 species from four locations in the south-west forest region of Western Australia. A species was deemed to have reached flowering age when at least 50% of the population had flowered. 166 Burrows et al: Post-fire juvenile period of plants in SW Australia Table 3 canopy-stored seeders that occurred in less flammable Plant families surveyed ranked by number of species in each habitats such as moist parts of the landscape {e.g., family riparian zones, some swamps) or where surface fuels were discontinuous or sparse (e.^., rock outcrops, some Family No. Genera No. Species swamps) (Table 4). Orchidaceae 20 76 Intra-species variation in juvenile period was evident Protcaceae 19 64 locally and regionally with plants growing in the lower Papiliopaceae 18 59 rainfall areas often taking considerably longer to flower Myrtaceae 17 46 than the same species at higher rainfall sites. Some Epacridaceae 9 28 Asteraceae 18 28 examples are shown in Table 5. Mimosaceae 2 27 Mean juvenile period, or secondary juvenile period in Poaceae 15 23 Stylidiaceae 2 23 the case of resprouters (Gill 1975), for various plant life Goodeniaceae 6 20 forms is shown in Table 6. As to be expected, trees, lower Cyperaceae 9 16 trees and woody shrubs took longer to flower after fire Apiaceae 9 16 than geophytes, grasses and annual herbs. All overstorey Dilleniaceae 1 16 Dasypogonaceae 5 15 tree species {Eucalyptus spp.) and many lower tree Antnericaceae 8 15 species (e.^. some Banksia spp.) were capable of both re¬ Rutaceae 5 13 sprouting and regenerating from seed after fire; the Thymelaeccae 1 9 juvenile period reported here is for mature plants that Haemodoraceae 3 9 Droseraceae 1 8 have re-sprouted. The juvenile period of plants Restionaceae 6 8 regenerating from seed would be considerably longer Sterculiaceae 3 6 (probably 10-20 years for eucalypts). There was a Loboliaceae 1 6 statistically significant difference (a = 0.05) in juvenile Tremandraceae 3 6 Iridaceae 2 6 period between seeders and resprouters in almost all life Colchicaceae 2 6 form categories with resprouters flowering sooner than Polygalaceae 2 5 obligate seeders, the exception being grasses, where Lamiaceae 2 5 seeders flowered sooner. Mean juvenile period for plants Santalaceae 3 4 Haemodoraceae 1 4 in each of the nine post-fire regeneration categories is Euphorbiaceae 4 4 shown in Figure 3. Overall, canopy-stored seeders had Loganiaceae 2 4 the longest juvenile period (mean about 36 months) and Rubiaceae 1 3 geophytes (especially Orchidaceae, Droseraceae) and Pittosporaceae 2 3 Rhamnaceae 2 3 plants with large apical buds (Xanthorroeaceae and Casuarinaceae 1 3 Dasypogonaceae) took the shortest time to flower (mean Lentibulariaceae 1 3 about 10 months). Kingia australis (Dasypogonaceae) had Haloragaccae 2 2 the shortest juvenile period with most of the population Stackliousiaceae 2 2 Xanthorroeaceae 2 2 flowering within 2 months of fire; plants with stems Lauraceae 1 2 shorter than about 100 cm had few or no flowers, Ranunculaceae 2 2 suggesting they had not reached reproductive maturity. Phormiaceae 2 2 Geraniaceae 2 2 While flowering occurred throughout the year, there Scraphylariaceae 2 2 is a distinctive flowering peak during spring when about Solanaceae 1 2 60% of plants flower. Few plants flowered during the Campanulaceae 1 2 Hypoxidaceae 1 2 Onagaceae 1 2 Amarathaceae 1 2 Caesalpiniaceae 1 1 Gyrostemonaceae 1 1 Violaceae 1 1 Phormiaceae 1 1 Centrolepidaceae 1 1 Aspleniaceae 1 1 Linaceae 1 1 Malvaceae 1 1 Brassicaceae 1 1 Primulaceae 1 1 Olacaceae 1 1 Saxifragaceae 1 1 Byblidaceae 1 1 Maryanthaceae 1 1 Sapindaceae 1 1 Xyridaceae 1 1 Portulaceae 1 1 Cephalotaceae 1 1 Lindsaeaceae 1 1 Zamiaceae 1 1 Loranthaceae 1 1 Podocarpaceae 1 1 Dennstaedtiaceae 1 1 Figure 3. The mean juvenile period for plant species categorised according to post-fire regeneration response strategy. 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Post-fire juvenile period of plants in south-west Australia forests and implications for fire management

Similar Post-fire juvenile period of plants in south-west Australia forests and implications for fire management