Fire Regimes in the Spinifex Landscapes of Australia. in 'Flammable Australia: Fire Regimes and Biodiversity Ofa Continent'

References

Allan GE, Southgate R (2001) Fire regimes in the spinifex landscapes of Australia. In 'Flammable Australia: Fire Regimes and Biodiversity ofa Continent'. (Eds RA Bradstock, JE Williams and AM Gill) pp. 145-176. (Cambridge University Press: Cambridge)

Bellingham PJ, Sparrow AD (2000) Resprouting as a life history strategy in woody plant communities. Oikos 89, 409-406.

Bond W, Midgley 11 (2001) Ecology of sprouting in woody plants: the persistence niche. TRENDS in Ecology and Evolution 16, 45-53.

Bond W, Midgley JJ (2003) The evolutionary ecology of sprouting in woody plants. International Journal ofPlant Science 164, 103-1 14.

Bond WJ, Van Wilgen BW (1996) 'Fire and Plants.' (Chapman & Hall: London)

Bowen BJ, Pate JS (1993) The significance of root starch in post-fire shoot recovery of the resprouter Stirlingia latifolia R.Br. (Proteaceae). Annals ofBotany 72 7-16.

Bradstock RA, Auld TD, Ellis ME, Cohn 1S (1992) Soil temperatures during bushfires in semi-arid, mallee shrublands. Australian Journal ofEcology 17, 433-440.

Burbidge NT (1943) Ecological succession observed during regeneration of Triodia pungens after burning. Journal ofthe Royal Society ofWestern Australia 28, 149-156.

Canadell J, Lloret F, Lopez-Soria L (1991) Resprouting vigor oftwo Mediterranean shrub species after experimental fire treatments. Vegetatio 95, 119-126.

Cary GJ, Morrison DA (1995) Effects offire frequency on plant species composition of sandstone communities in the Sydney region: Combinations of inter-fire intervals. Australian Journal ofEcology 20, 418-426.

Clarke PJ, Knox KJE (2002) Post-fire response of shrubs in the tablelands ofeastern Australia: do existing models explain habitat differences? Australian Journal ofBotany 50, 53-62.

Clarke PJ, Knox KJE, Wills KE, Campbell M (2005) Landscape patterns ofwoody plant response to crown fire: disturbance and productivity influence sprouting ability. Journal ofEcology 93, 544-555.

Cruz A, Beatriz P, Moreno 1M (2003) Resprouting ofthe Mediterranean-type shrub Erica australis with modified lignotuber carbohydrate content. Journal ofEcology 91, 348-356.

144 Drewa PB, Platt WJ, Moser EB (2002) Fire effects on resprouting of shrubs in headwaters ofsoutheastern longleaf pine savannahs. Ecology 83, 755-767.

FI inn MA, Wein RW (1977) Depth of underground organs and theoretical survival during fire. Canadian Journal ofBotany 55, 2550-2554.

Grant CD, Lonegaran WA, Koch JM, Bell DT (1997) Fuel characteristics, vegetation structure and fire behavior of 11-15 year old rehabilitated bauxite mines in Western Australia. Australian Forestry 60, 147-157.

Hodgkinson KC (1982) Adaptation of shrub species to fire in the arid zone. In 'Evolution ofthe flora and fauna ofarid Australia'. (Eds WR Barker and PJM Greenslade). (Peacock Publications: Frewville)

Hodgkinson KC (1998) Sprouting success ofshrubs after fire: height-dependent relationships for different strategies. Decologia 115, 64-72.

Lloret F, Siscart 0, Dalmases C (2004) Canopy recovery after drought dieback in holm oak mediterranean forests ofCatalonia (NE Spain). Global Change Biology 10, 2092 2099.

Malanson GP, Trabaud L (1988) Vigor of post-fire resprouting by Quercus coccifera L. Journal ofEcology 76,351-365.

Matlack GR, Gibson OJ, Good RE (1993) Regeneration ofthe shrub Gaylussacia baccata and associated species after low intensity fire in an Atlantic coastal plain forest. American Journal ofBotany 80, 119-126.

Moreno JM, Oechel WC (1991) Fire intensity and herbivory effects on post-fire resprouting ofAdenostomafasciculatum in southern Californian chaparral. Decologia 85, 429-433.

Morgan JW (2001) Seedling recruitment patterns over 4 years in an Australian perennial grassland community with different fire histories. Journal ofEcology 89, 908-919.

N ieuwenhuis A (1987) The effect offire frequency on the sclerophyll vegetation ofthe West Head, New South Wales. Australian Journal ofEcology 12, 373-385.

Noble JC (1989) Fire regimes and their influence on herbage and mallee coppice dynamics. In 'Mediterranean landscapes in Australia: mallee ecosystems and their management'. (Eds JC Noble and RA Bradstock) pp. 168-181. (CSIRO: East Melbourne)

Odion DC, Davis FW (2000) Fire, soil heating, and the formation ofvegetation patterns in chaparral. Ecological Monographs 70, 149-169.

145 Olson MS, Platt WJ (1995) Effects ofhabitat and growing season fires on resprouting of shrubs in longleaf pine savannahs. Vegetatio 119, 101-118.

Rice SK (1993) vegetation establishment in post-fire Adenostoma chaparral in relation to fine-scale pattern in fire intensity and soil nutrients. Journal ofVegetation Science 4.

Schimmel J, Granstrom A (1996) Fire severity and vegetation response in the Boreal Swedish forest. Ecology 77, 1436-1450.

Scotter DR (1970) Soil temperatures under grass fires. Australian Journal ofSoil Research 8,273-9.

Verdaguer 0, Ojeda F (2005) Evolutionary transition from resprouter to seeder life histories in two Erica (Ericaceae) species: insight from seedling axillary buds. Annals of Botany 95,593-599.

Vesk PA, Warton 01, Westoby M (2004) Sprouting by semi-arid plants: testing a dichotomy and predictive traits. Gikos 107, 72-89.

Wade DO, Johansen RW (1986) 'Effects offire on southern pine: observations, and recommendations.' USDA Forest Service General Technical Report SE-41.

Watson P, Wardell-Johnson G (2004) Fire frequency and time-since-fire effects on the open-forest and woodland flora of Girraween National Park, south-east Queensland, Australia. Austral Ecology 29, 225-236.

Williams PR, Congdon RA, Grice AC, Clarke PJ (2004) Soil temperature and depth of legume germination during early and late dry season fires in a tropical eucalypt savannah of north-eastern Australia. Austral Ecology 29, 258-263.

146 Chapter 8

General conclusions

Introduction This thesis sought to examine the effects of fire regime on the vegetation dynamics in spinifex grasslands in central Australia. This chapter synthesizes the results of this research and puts forward the hypothesis that plants within spinifex grasslands possess adaptations that have been selected for under a fire regime characterised by periodic, summer fire. It also summarizes the management implications of these findings, and points towards potential future research directions.

Spinifex fire regimes Chapter 2 showed that contemporary fire regimes in spinifex grasslands are characterised by extensive wildfire events that occur in response to fuel accumulations after extreme rain. During these events, the scale of fires can be enormous, with one fire during the 1980s burning over 6000 km2 of sand dune and range country. The almost complete absence of fires during inter-event periods was also striking, with less than 1% (~ 80 2 km ) of the study area burning during the entire period from 1985-2000. The main season of fire occurrence in spinifex grasslands is over the late spring and summer months, when air temperatures are highest and humidity levels are low. The potential for fires during these months is enhanced by the increased likelihood of summer lightning strikes (Griffin, Price et al. 1983). Chapter 2 also showed that spinifex grasslands are occasionally subjected to short fire intervals, and that these fires occur when sufficient rains fall soon after initial fires. It is suggested that these short interval fires are fuelled by inter-hummock grass species, such as Aristida holathera and A. inaequiglumis, as these species were common in the seed bank (see Chapter 5) and Triodia fuels are virtually absent during early stages of post-fire succession (see Chapter 4). Chapter 3 showed that these short interval fires are of lower intensity and severity than initial Triodia-fuelled fires (see Chapter 6), but as was demonstrated in Chapters 4 and 5, their ecological impact can still be significant.

147 While the data from this thesis only pertain to the very recent history of fires, it was suggested in Chapter 2 that the contemporary relationship between rain and widespread fire in spinifex grasslands may also have prevailed prior to the arrival of humans in Australia. Such a hypothesis may be plausible, given that fires in arid environments across the globe are constrained by low biomass levels, and that opportunities for fire only ever occur when high rainfalls promote substantial fuel accumulation (Frost 1985; Kitzberger, Swetnam et al. 2001; Veblin, Kitzberger et al. 1999). At this stage no data are available to directly validate this hypothesis, although it could be tested in the future using research techniques similar to those of Atahan et al. (2004) and Dodson et al. (2005), who used Pliocene charcoal records from palaeo-lakes to show that pre-human fire regimes in south-western Australia were characterised by fires that were highly seasonal and frequent. In Chapter 2 it was also suggested that the seasonality of fires in spinifex grasslands may not have changed considerably since pre-human times, as fuel combustibility during these times would still have been highest in summer. This hypothesis has previously been put forward by Suijendorp (1981), who also postulated that pre-human fire regimes may have been even more seasonal than contemporary ones, owing to the absence of anthropogenic ignition sources and the dependence on summer lightning strikes to initiate fires. If these hypotheses hold, then it can be presumed that the spinifex flora has spent a considerable portion of its evolutionary history adapting to fire regimes characterised by infrequent summer fires that occur in response to extreme rainfall events. In addition, the results of Chapters 4 and 6 suggest that two other selective forces have played major roles in the evolution of the contemporary flora: 1) high levels of post-dispersal seed predation; and 2) a climatically unpredictable arid environment in which highly competitive perennial grasses (Triodia spp.) and shrubs restrict recruitment to early stages of pyric succession (during which time these species lose competitive dominance). The following section describes some of the life history groups that I believe have evolved under this 'triad' ofselective forces.

148 Plant life histories

Fire-cued ephemerals Chapter 4 showed that the spinifex grasslands are home to a highly diverse ephemeral flora that increases dramatically in abundance during periods shortly after fire. It is suggested that the evolution of this life history group has been evoked by the competitive dominance of the established Triodia/woody component in long unburned vegetation with the ephemeral species 'escaping' these competitive effects by restricting establishment to periods shortly after fire. As evidenced in Chapter 5, the seed banks of these ephemeral species are persistent, with high numbers of ephemeral seeds in both recently burned and long unburned vegetation. Their seeds are also normally dormant, but are released from dormancy by fire-related cues such as heat shock or smoke (as in the case of Brachyscome ciliaris and Aristida holathera - see Chapter 5). The adult plants of this group reach reproductive maturity quickly (- 1-2 years), and from the seed bank work in this thesis it has been shown that there are large inputs of seeds during periods shortly after fire (see Fig. 8.1).

establi~hment Fire ___.... Rain -. Vegetation + seed production

r Vegetation

D D D 0 Seed bank

Time

Fig. 8.1 Seed bank and vegetation dynamics of ephemeral fire-cued plant species. Squares represent seed abundances, while shading in squares indicate that seeds have been released from dormancy by fire. Germination, predation and decay account for the decline in seed abundances after seed rain. The large increase in seed abundance shortly after fire is accounted for by the input of seed from the recently established standing vegetation.

149 These seeds enter the soil as a dormant seed bank, and are gradually reduced over time, probably through a cOlnbination of predation, decay and intermittent germination, until replenishment occurs after the next fire. The seeds of these species are usually small, which encourages the formation of a persistent seed bank by facilitating seed entry into small fissures in the soil surface and increases the likelihood ofescape from predators. Chapter 4 revealed that fire-cued ephemeral species may be negatively affected by short-fire intervals. This finding refuted predictions from the seed bank literature, that ephemeral species should be resilient to frequent disturbances owing to the shortness of their life history. The precise mechanisms behind the observed negative effects of short intervals on fire-cued ephemerals are uncertain. However, it is postulated that our results reflect establishment failure (rather than exhausted seed banks), perhaps caused by insufficient post-fire rain after follow-up fires. It was, however, shown in Chapter 4 that these negative effects are normally transient, and that populations appear to recover over time.

Herbaceous clonal plants Herbaceous clonal plants, such as Leptosema chambersii and Bonamia erecta, are commonly referred to as 'underground trees' as they have long-lived life histories that are largely analogous to the woody component of the spinifex grasslands. Rather than 'escaping' the dominance of the other perennial flora, these species have evolved a life history that maintains strong competitive viability in both recently burned and long unburned landscapes by regenerating asexually via vegetative expansion. Thus they maintain high levels of cover regardless of pyric intrusion (see Chapter 4). Chapter 5 also found that these plants maintain a persistent seed bank in both burned and unburned habitats (see Fig. 8.2), although it is unclear to what extent these plants rely on seedling recruitment during post-fire or post-drought regeneration. It was also shown in Chapter 4 that the vegetation of these species is susceptible to short interval fires; possibly indicating that depletion of carbohydrate reserves occurs as the result of repeated disturbances. However, as with the ephemeral species, the negative effect of short interval fires on these species disappears over time.

150 Fire ---...- Rain

r Q) Vegetation u c en "'0 c ~ «.0 D o D D D D Seed bank

Time

Fig. 8.2 Seed bank and vegetation dynamics of clonal plant species. Squares represent seed abundances, while line represents vegetation cover (neither of which change) over time.

Triodia species Adult Triodia schinzii plants in the Haast Bluff study are killed by fire and population are entirely dependent on seedling regeneration for persistence after fire (see Chapter 4). This seemingly perilou reliance on only one means of post-fire regeneration appears paradoxical, given that this species of Triodia is the dominant perennial component and drives many of the competitive interactions between species in these highly flammable ecosystems. However, the findings of this thesis indicate that the recruitment syndromes of T. schinzii have a highly robust ability to tolerate the range of fire regimes that they are likely to experience. Triodia pecies are opportunistic seeders and produce bumper crops in 'mast' years following extreme rains (see 'opportunistic fruiters' - Fig. 8.3). Having a phenological cycle that maximizes seed output after extreme rains ensures that a supply of seed is always available at a time when fuel levels are high and fire events are likely to occur. Increa es in seed output satiate seed predators and maximize the likelihood that seedlings will establish in the post-fire environment.

151 Extreme rain ------.... Fire ...Post.-fire ~ Recruitment ram

I,ll IIII ---. 00 D Pi DD D I I I I Opportunistic seeders

IIII ---. 0 D 0 D D D I II I Seasonal seeders IIII IIII 1111 IIII 'III II II II I' Serotinous D tJ DDD D D I I

spring winter spring

Time

Fig. 8.3 Seed bank syndromes of woody species in relation to the rain-driven fire cycle that characterises the contemporary fire regime. Squares represent seed abundances, while shading in squares indicates that seeds have been released from dormancy by fire. Seed of opportunistic seeders has a 'pulse' approximately 6-12 months after extreme rain; seed of seasonal seeders has a 'pulse' each spring; serotinous species release seed after fire. Germination, predation and decay account for the decline in seed abundances after seed rain.

Chapter 5 revealed that during inter-event periods, Triodia species maintain only a sparse seed bank, as their large seeds are rapidly removed after seed fall by seed predators. Thus, it seems likely that recruitment potentials for these species become greatly reduced in the event of post-fire drought or fires that occur in non- 'mast' years (as seed supplies are low during these periods). Further research may demonstrate that rodent seed 'caches' or ant seed 'mounds' (areas close to nest entrances where seeds are abandoned by ants once food bodies are removed from seeds) are important re-colonization sites under these situations (Berg] 975; Hughes and Westoby 1990).

]52 Chapter 4 indicated that Triodia schinzii is also resilient to short fire intervals (over the long term). This is surprising given that seedlings are fire-killed and that initial seed crops are rapidly removed by seed predators. It is known that under favourable conditions the primary juvenile period for these plants can be as short as 2 yrs (pers. obs.). So once again, it may be high rainfall that stimulates opportunistic production of a seed crop (albeit a much smaller one than would be produced in unburnt spinifex), thereby ensuring that a seed bank is present at the time when follow-up fire events occur. It is also possible that rodent or ant seed 'caches' and wind dispersal of seed from unburned areas play important roles in re-colonization following these events.

Woody species This thesis demonstrated that woody species In the spinifex grasslands have 'dual regeneration strategies', possessing complex 'predator satiating' recruitment strategies (see Chapter 6) and strong resprouting responses (see Chapter 7). The three main recruitment strategies expressed among these species are; 1. Plants with regular, spring-time production of fire-cued seed (seasonal seeders see Fig 8.3); 2. Plants with opportunistic production offire-cued seed (opportunistic seeders - see Fig 8.3); and 3. Plants with canopy held seed banks (serotinous - see Fig 8.3). The seasonal seeding syndrome appears to have evolved as an adaptation to the interaction between seed predation and the seasonality of fire in spinifex grasslands. This syndrome synchronises seed production with the main season of fire occurrence (summer), thus maximizing the likelihood that some seeds will escape predation, become cued by fire and germinate in the post-fire environment. The opportunistic syndrome is largely analogous to the recruitment syndrome displayed by Triodia species and appears to have evolved as an adaptation to the interaction between predation and the sequence of events that precede widespread fire occurrence in spinifex grasslands: extreme rain --+ fuel accumulation --+ fire. These species produce bumper seed crops in 'mast' years following periods of high rainfall, thus maximizing the likelihood that a proportion ofthe seed crop will escape predation at a time when fire occurrence is most likely (1-2 years

153 after extreme rain events). Finally, serotiny represents an adaptation to fire per se, not necessarily to a particular season or regime of fire. These species have canopy-held seed banks with seeds that are non-dormant and are released to the soil surface via the heat of fire. This mass release of seed after fire maximizes the likelihood that some seeds will escape predation and establish in the post-fire environment. While all of these syndromes increase the potential resilience of woody populations to pyric disturbance, there are still very high risks associated with post-fire seedling recruitment in spinifex grasslands. The climatic unpredictability of these arid environments means that there is always the possibility that rains will be delayed for considerable periods after fire. Under such circumstances, seeds that would normally be abundant in the post-fire environment are rapidly removed by predators (see Fig 8.3). As a result, seed banks may be completely exhausted by the time rains finally occur. Recruitment failures can also transpire following 'aseasonal' winter fires (see Chapter 4 and 6). Such fires considerably reduce the recruitment potentials of both seasonal fruiters and opportunistic fruiters, primarily due to the sparseness of seed during this season (owing to depletion by seed predators), but also due to the lower soil temperatures that are experienced (which induce lower rates of dormancy release for fire-cued seeds - see Chapters 3 and 6). The high risk associated with post-fire seedling recruitment is likely to have been a strong selective force in the evolution of resprouting systems among woody species in spinifex habitats. Having dual regeneration strategies (both resprouting and seedling recruitment) means that plants have a buffer against the possibility of recruitment failure, and ensures that at least some progeny will persist after fire. In Chapter 7 we observed variability in the overall strength of resprouting among different Acacia species, although there were consistent trends concerning the effects of the different components of fire regime on plant survival. High severity burns produced higher mortality than low severity burns (among all Acacia species), and it is presumed that this was due to the increased penetration of heat in soils during high severity burns. Also, both spring and summer burns produced higher mortality than winter burns (only among A. maitlandii and A. melleodora). It is presumed that this was due to a combination of the increased penetration of heat in soils during spring and summer, as well as the reduced

154 physiological resilience of species to severe perturbation in these seasons (perhaps driven by water availability and/or carbohydrate accumulations). Further experimental research is required here to determine the precise impact that water availability and carbohydrate reserves have on seasonal variation in resprouting among woody species. This thesis also demonstrated that many woody species are able to cope with short-interval Aristida-fuelled fires. Not only can adult plants resprout from underground organs following short-interval fires (provided the severity of burn to a given plant is low-see Chapter 7), but they also possess very short secondary juvenile periods (2-3 yrs). Having short secondary juvenile periods means that even if most of the seed produced prior to an initial fire has recruited or been removed by predators, the resprouts of burned plants will already be producing seed by the time follow-up fires occur. Thus, the synchrony of a follow-up fire with seed produced by a secondary seed-rain event may have an additive effect on populations, provided a large proportion ofadults and juveniles resprouted following the second fire (see A. melleodora in Chapter 7). Those seedlings recruited after a short interval fire will have the additional advantage of establishing in an environment in which the highly competitive Triodia species have been greatly reduced (see Chapter 3).

Conclusion This thesis has shown that fire is a dynamic driver of ecological processes in spinifex grasslands, and is capable of causing dramatic compositional changes among a range of plant life-history groups. It is a crucial factor in releasing the competitive grip of the established perennial component and in allowing the ephemeral flora to establish. This release from competition is also crucial in allowing the seedlings of the long-lived Triodia and woody species to establish. Fire and competition also interact strongly with seed predation, and over evolutionary time, it is suggested that these interactions have played strong roles in evoking the various complex recruitment syndromes that are now in evidence among the spinifex flora. At this stage it is unclear whether spinifex grasslands in the Haasts Bluff region would benefit significantly from management attempts to alter the seasonality, frequency or extent of fires. The recruitment syndromes ofthe flora appear to be inextricably linked to the summer-dominated rain driven fire cycles that currently prevail (see Chapter 6).

155 Also, under most situations, the resprouting systems of the perennial component are largely resilient to fire - as was shown by the high survival rates of resprouters after the 2000-02 fires (see Chapter 4). It is also uncertain whether the extent of fires in these grasslands is manageable at landscape scales. Previous attempts at reducing fire size in these systems have largely failed, owing to the overwhelming bulk and flammability of cured fuels that accumulate following extreme rains (Allan, Phillips et al. 2003, Mathews D., Pers. comm., Finke Gorge National Park, 2002, Yates P., Pers.comm., Anangu Pitjantjatjarra Yankuntjatjarra lands, 2004). Finally, any attempt at managing vegetation in these fire-prone grasslands will always be largely reliant on chance. This is because it is the completely unpredictable influence of post-fire rainfall that is the ultimate driver of both seedling recruitment and resprouting regeneration in these systems.

156 References

Alexander ME (1982) Calculating and interpreting forest fire intensities. Canadian Journal ofBotany 60,349-357.

Allan GE (1993) 'The fire history of central Australia. CSIRO/CCNT Bushfire Research Project, volA. CSIRO/CCNT Technical Report.' CSIRO, Alice Springs.

Allan GE, Griffin GF (1986) Fire ecology ofthe hummock grasslands ofcentral Australia. In 'Proceedings ofthe 4th Australian Rangeland Society Conference'. Armidale, NSW pp. 126-129

Allan GE, Phillips NR, Hookey P (2003) Learning lessons from an exceptional period of fires in central Australia: 1999 to 2002. In 'Proceedings ofthe 3rd International Wildland Fire Conference'. Sydney, Australia

Ashton DH (1976) The development ofeven aged stands ofEucalyptus regnans F.Muell. in central Victoria Australian Journal ofBotany 24, 397-414.

Aston AR, Gill AM (1976) Coupled soil moisture, heat and water vapour transfers under simulated fire conditions. Australian Journal ofSoil Research 14, 55-66.

Atahan P, Dodson JR, Itzstein-Davey F (2004) A fine resolution Pliocene pollen and charcoal record from Yallalie, Southwestern Australia. Biogeography 31, 199-205.

Auld TD (1990) The survival ofjuvenile plants ofthe resprouting shrub Angophora hispida (Myrtaceae) after a simulated low-intensity fire. Australian Journal ofBotany 38, 255-260.

Auld TD (1995) Soil seed bank patterns of four trees and shrubs from arid Australia. Journal ofArid Environments 29,33-45.

Auld TD, Bradstock RA (1996) Soil temperatures after the passage offire: do they influence the germination of buried seeds. Australian Journal ofEcology 21, 106-109.

Auld TD, O'Connell MA (1991) Predicting patterns of post-fire germination in 35 eastern Australian Fabaceae. Australianjournal ofEcology 16, 53-70.

157 Ball DA, Miller SO (1989) A comparison oftechniques for estimation ofarable soil seed banks and their relationship to weed flora. Weed Research 29, 365-373.

Baver LD (1940) 'Soil Physics.' (Chapman and Hall: London)

Beadle NCW (1940) Soil temperatures during forest fires and their effect on the survival ofvegetation. Journal ofEcology 28, 180-192.

Beadle NCW (1981) 'The Vegetation of Australia.' (Gustav Fischer Verlag: Stuttgart)

Bell DT (1999) Turner Review No.1: The process of germination in Australian species. Australian Journal ofBotany 47,475-517.

Bell DT, Plummer JA, Taylor SK (1993) Seed germination ecology in south western Western Australia. The Botanical Review 59, 24-59.

Berg RY (1975) Myrmecochorous plants in Australia and their dispersal by ants. Australian Journal ofBotany 23, 475-508.

Bird OW, Bird RB, Parker CH (2005) Aboriginal burning regimes and hunting strategies in Australia's western desert. Human Ecology 33, 443-464.

Bolton BL, Latz PK (1978) The western hare-wallaby, Lagorchestes hirsutus (Gould) (Macropodidae), in the Tanami Desert. Australian Wildlife Research 5,285-293.

Bond WJ (1984) Fire survival ofCape Proteaceae - influence offire season and seed predators. Vegetatio 56, 65-74.

Bond WJ, Van Wilgen BW (1996) 'Fire and Plants.' (Chapman & Hall: London)

Bowen BJ, Pate JS (1993) The significance of root starch in post-fire shoot recovery of the resprouter Stirlingia latifolia R.Br. (Proteaceae). Annals ofBotany 72 7-16.

Bowen BJ, Pate JS (2004) Effect ofseason of bum on shoot recovery and post-fire flowering performance in the resprouter Stirlingia latifolia R.Br. (Proteaceae). Austral Ecology 29, 145-155.

Bowman DMJS (1998) Transley Review No. 101: The impact ofAboriginal landscape burning on the Australian biota. New Phytologist 140, 385-410.

Bowman DMJS, Latz PK (1993) Ecology of Callitris glaucophylla (Cupressaceae) on the Macdonnell Ranges, Central Australia. Australian Journal ofBotany 41,217-225.

Bowman DMJS, Latz PK, Panton WJ (1994) Pattern and change in an Acacia aneura shrub land and Triodia hummock grassland mosaic on rolling hills in central Australia. Australian Journal ofBotany 43,25-37.

158 Bradstock RA (1989) Dynamics ofa perennial understorey. In 'Mediterranean landscapes in Australia: Mallee ecosystems and their management'. (Eds JC Noble and RA Bradstock) pp. 141-154. (CSIRO: Melbourne)

Bradstock RA, Auld TD (1995) Soil temperatures during experimental bushfires in relation to fire intensity: consequences for legume germination and fire management in south-eastern Australia. Journal ofApplied Ecology 32, 76-84.

Bradstock RA, Auld TD, Ellis ME, Cohn JS (1992) Soil temperatures during bushfires in semi-arid, mallee shrublands. Australian Journal ofEcology 17, 433-440.

Bradstock RA, Myerscough PJ (1988) The survival and population response to frequent fires oftwo woody resprouters Banksia serrata and Isopogun anemonifolius. Australian Journal ofBotany 36, 415-431.

Burbidge NT (1943) Ecological succession observed during regeneration of Triodia pungens after burning. Journal ofthe Royal Society ofWestern Australia 28, 149-156.

Burbidge NT (1953) The genus Triodia. Australian Journal ofBotany 1, 121-184.

Burrows ND, Christensen PES (1990) 'A survey of Aboriginal fire patterns in the Western Desert of Australia.' U.S. Department of Agriculture Forest Service, General Technical Report SE-69. (Southeastern Forest Experimental Station: Ashville, NC.).

Burrows ND, Ward B, Robinson A (1991) Fire behaviour in spinifex fuels on the Gibson Desert ature Reserve, Western Australia. Journal ofArid Environments 20, 189-204.

Byram GM (1959) Combustion offorest fuels. In 'Forest fire: control and use'. (Ed. KP Davis) pp. 61-89. (McGraw-Hill: New York)

Canadell J, Lloret F, Lopez-Soria L (1991) Resprouting vigor oftwo Mediterranean shrub species after experimental fire treatments. Vegetatio 95, 119-126.

Casson NE (1994) The biology and ecology of Triodia basedowii and T wiseana associations in relation to aridity and fire. Unpublished PhD thesis, Curtin University.

Cheney P (1981) Fire behaviour. In 'Fire and the Australian Biota'. (Eds AM Gill, RH Groves and IR Noble). (Australian Academy of Science: Canberra)

Cheney NP, Sullivan A (1997) 'Grassfires: fuel, weather and fire behavior.' (CSIRO Publishing: Melbourne)

159 Cohn JS, Bradstock RA (2000) Factors affecting post-fire seedling establishment of selected mallee understorey species. Australian Journal ofBotany 48, 59-70.

Cook L (1939) A contribution to our information on grass burning. South African Journal o/Science 36, 270-282.

Craig AB (1992) The post-fire regeneration of plant communities dominated by Triodia pungens in the vicinity ofNewman, W.A. Unpublished MSc thesis, Curtin University of Technology.

Craig AB (1999) Fire management of rangelands in the Kimberly low-rainfall zone: a review. Rangelands Journal 21, 39-70.

Cruz A, Beatriz P, Moreno JM (2003) Resprouting ofthe Mediterranean-type shrub Erica austrlis with modified lignotuber carbohydrate content. Journal ofEcology 91, 348-356.

Davidson DW, Morton SR (1984) Dispersal adaptations of some Acacia species in the Australian arid zone. Ecology 65, 1038-1051.

Davison EA (1982) Seed utilization by harvester ants. In 'Ant-plant interactions in Australia'. (Ed. RC Buckley) pp. 1-6. (Junk Publishers: The Hague) de Groot WJ, Wein RW (1999) Betula glandulosa Michx. Response to burning and post fire growth temperature and implications ofclimate change. International Journal of Wildland Fire 9,51-64. de Groot WJ, Wein RW (2004) Effects offire severity and season of burn on Betula glandulosa growth dynamics. International Journal ofWildland Fire 13,287-295.

Dodson JR, Robinson M, Tardy C (2005) Two fine resolution Pliocene charcoal records and their bearing on pre-human fire frequency in south-western Australia. Austral Ecology 30, 592-599.

Drewa PB, Platt WJ, Moser EB (2002) Fire effects on resprouting of shrubs in headwaters of southeastern longleaf pine savannahs. Ecology 83, 755-767.

Enright NJ, GoldblulTI D (1999) Demography ofa non-sprouting and resprouting Hakea species (Proteaceae) in fire prone Eucalyptus woodlands of southeastern Australia in relation to stand age, drought and disease. Plant Ecology 144, 71-82.

Enright NJ, Kintrup A (2001) Effects ofsmoke, heat and charred wood on the germination ofdormant soil-stored seeds from a Eucalyptus baxteri heathy-woodland in Victoria, SE Australia. Austral Ecology 26,132-141.

160 Envans EC, Long JPM (1965) The Aborigines of western central Australia. The Geographical Journal 131, 318-329.

Flinn MA, Wein RW (1977) Depth of underground organs and theoretical survival during fire. Canadian Journal ofBotany 55, 2550-2554.

Floyd AG (1966) Effect offire upon weed seeds in the wet sclerophyll forests ofnorthern New South Wales. Australian Journal ofBotany 14, 243-256.

Friedel MH, Nelson D, Sparrow AD, Kinloch JE, Maconochie JR (1994) Flowering and fruiting of arid zone species ofAcacia in central Australia. Journal ofArid Environments 27, 221-239.

Frost PGH (1985) The responses of savanna organisms to fire. In 'Ecology and management of the world's savannas'. (Eds JC Tothill and JJ Mott) pp. 232-237. (Australian Academy of Science: Canberra)

Geiger R (1957) 'The climate near the ground.' (Harvard University Press: Cambridge)

Gibbs WJ (1969) Meteorology and climatology. In 'Arid Lands ofAustralia'. (Eds RO Slatyer and RA Perry) pp. 33-45. (Australian National University Press: Canberra)

Giles WEP (1979) 'Australia Twice Traversed.' (Doubleday: Sydney)

Gill AM (1981) Adaptive responses of Australian vascular plants to fire. In 'Fire and the Australian biota'. (Eds AM Gill, RH Groves and IR Nobel) pp. 243-272. (Australian Academy of Science: Canberra)

Gill AM (2000) Fire pulses in the heart of Australia: fire regimes and fire management in Central Australia.

Gould RA (1971) Uses and effects offire among the Western Desert Aborigines of Australia. Mankind 8, 14-24.

Griffin GF (1990) Characteristics ofthree spinifex alliances in Central Australia. Journal ofVegetation Science 1, 445-452.

Griffin GF (1992) Will it burn-should it burn?: management ofthe spinifex grasslands of inland Australia. In 'Desertified grasslands: their biology and management'. (Ed. GP Chapman) pp. 63-76. (Academic Press: London)

Griffin GF, Allan GE (1984) Fire behavior. In 'Anticipating the inevitable: a patch-burn strategy for fire management at Uluru (Ayers Rock-Mt Olga) National Park'. (Ed. EC Saxon) pp. 55-68. (CSIRO Australia: Melbourne)

161 Griffin GF, Allan GE (1985) Fire and the management of Aboriginal owned land In 'Proceedings ofthe Application of Science and Technology to Aboriginal Development in Central Australia. Workshop.' Alice Springs pp. 46-52

Griffin GF, Friedel MH (1984) Effects offire on central Australian rangelands. II: Changes in tree and shrub populations. Australian Journal ofEcology 9, 395-403.

Griffin GF, Friedel MH (1985) Discontinuous change in central Australia: some implications for major ecological events for land management. Journal ofArid Environments 9, 63-80.

Griffin GF, Morton SR, Allan GE (1990) Fire created patch dynamics for conservation management in the hummock grasslands of central Australia. In 'Proceedings ofthe International Symposium on Grassland Vegetation'. Beijing, China. (Science Press)

Griffin GF, Price NF, Portlock HF (1983) Wildfires in the Central Australian Rangelands, 1970-1980. Journal ofEnvironmental Management 17, 311-323.

Gunawardene NR, Majer JD (2005) The effect offire on ant assemblages in the Gibson Desert Nature Reserve, Western Australia. Journal ofArid Environments 63,725-739.

Halcombe S (2004) Traditional owners and 'community-country' anangu: distinctions and dilemmas. Australian Aboriginal Studies 2, 64-71.

Harper JL (1957) The ecological significance ofdormancy and its importance in weed control. International Congress ofPlant Protection 4, 415-420.

Harrington GN, Wilson AD, Young MD (1984) Management of Rangeland Ecosystems. In 'Management ofAustralia's Rangelands'. (Eds GN Harrington, AD Wilson and MD Young) pp. 3-13. (CSIRO Division of Wildlife Research: CSIRO, Australia)

Hartford RA, Frandson WH (1992) When its hot, its hot. .. or maybe its not! (surface flaming may not portend extensive soil heating). International Journal ofWildland Fire 2,139-144.

Haydon DT, Friar JK, Pianka ER (2000) Fire-driven dynamic mosaics in the Great Victoria Desert, Australia - 1. Fire geometry. Landscape Ecology 15, 373-381.

Hodgkinson KC (1982) Adaptation of shrub species to fire in the arid zone. In 'Evolution ofthe flora and fauna of arid Australia'. (Eds WR Barker and PJM Greenslade). (Peacock Publications: Frewville)

Hodgkinson KC (1986) Response ofrangeland plants to fire in water-limited environments. In 'Rangelands: A resource under siege. Proceedings ofthe Second International Rangeland Conference'. (Eds PJ Joss, PW Lynch and OB Williams). (Australian Acdemy of Science: Canberra)

162 Hodgkinson KC (1991) Shrub recruitment response to intensity and season offire in a semi-arid woodland. Journal ofApplied Ecology 28, 60-70.

Hodgkinson KC (1998) Sprouting success ofshrubs after fire: height-dependent relationships for different strategies. Oecologia 115, 64-72.

Hodgkinson KC, Harrington GN, Miles GE (1980) Composition, spatial and temporal variability ofthe soil seed pool in a Eucalyptus populnea shrub woodland in central New South Wales. Australian Journal ofEcology 5,23-29.

Hodgkinson KC, Oxley RE (1990) Influence offire and edaphic factors on germination ofthe arid zone shrubs Acacia aneura, Cassia nemophila and Dodonea viscosa. Australian Journal ofBotany 38,269-279.

Holland PG (1967) Biomass and pattern in the ecology of mallee vegetation. PhD thesis thesis, Australian National University.

Hughes L, Westoby M (1990) Removal rates ofseeds adapted for dispersal by ants. Ecology 71, 138-148.

Hughes L, Westoby M (1992) Fate ofseeds adapted for dispersal by ants in Australian sclerophyll vegetation. Ecology 73, 1285-1299.

Ireland C, Andrew MH (1995) Ants remove virtually all western myall (Acacia papyrocarpa Benth.) seeds at Middleback, South Australia. Australian Journal of Ecology 20, 565-570.

Jacobs SWL (1982) Relationships, distribution and evolution of Triodia and Plectrachne (Gramineae). In 'Evolution ofthe flora and fauna ofarid Australia'. (Eds WR Barker and PJM Greenslade) pp. 287-290. (Peacock Publications: Frewville)

Jacobs SWL (1992) Spinifex (Triodia, Plectrachne, Symplectrodia and Monodia: Poaceae) in Australia. In 'Desertified grasslands: Their Biology and Management'. (Ed. GP Chapman) pp. 47-62. (Academic Press: London)

James CD, Landsberg J, Morton SR (1995) Ecological functioning in arid Australia and research to assist conservation of biodiversity. Pacific Conservation Biology 2, 126-142.

Janzen DH (1971) Seed predation by animals. Annual Review ofEcological Systematics 2,465-492.

Johnson EA (1992) 'Fire and vegetation dynamics.' (Cambridge University Press: New York)

163 Keeley JE (1977) Seed production, seed populations in soil, and seedling production after fire for two congeneric pairs of sprouting and non-sprouting Chaparral shrubs. Ecology 58, 820-829.

Keith DA (1996) Fire-driven extinctions ofplant populations: a synthesis oftheory and review ofevidence from Australian vegetation. Proceedings ofthe Linnean Society of New South Wales 116, 37-77.

Kemp PR (1989) Seed banks and vegetation processes in deserts. In 'Ecology of soil seed banks'. (Eds MA Leck, TV Parker and RL Simpson) pp. 257-281. (Academic Press: San Diego)

Kershaw AP, Clark JS, Gill AM, D'Costa DM (2002) A history offire in Australia. Fire regimes in the spinifex landscapes of Australia. In 'Flammable Australia: Fire regimes and biodiversity of a continent'. (Eds RA Bradstock, JE Williams and AM Gill) pp. 3-12. (Cambridge University Press: Cambridge)

Kimber R (1983) Black Lightning: aborigines and fire in central Australia and the Western Desert. Archaeology in Oceania 18, 38-44.

Kinloch JE, Freidel MH (2005) Soil seed reserves in arid grazing lands ofcentral Australia. Part 1: seed bank and vegetation dynamics. Journal ofArid Environments 60, 133-161.

Kinloch JE, Friedel MH (2005) Soil seed reserves in arid grazing lands ofcentral Australia. Part 1: seed bank and vegetation dynamics. Journal ofArid Environments 60, 133-161.

Kitzberger T, Swetnam TW, T.T. V (2001) Inter-hemispheric synchrony offorest fires and the El Nino-Southern Oscillation. Global Ecology and Biogeography 10, 315-326.

Lakon G (1949) The topographic tetrazolium method for determining the germination capacity of seeds. Plant Physiology 24, 389-394.

Lamont BB, Swanborough PW, Ward D (2000) Plant size and season of burn affect flowering and fruiting ofthe grasstree Xanthorrhea preissii. Austral Ecology 25, 268 272.

Letnic M, Dickman CR, McNaught G (2000) Bet-hedging and germination in the Australian arid zone shrub Acacia ligulata. Austral Ecology 25, 368-374.

Lloyd MV, Dixon KW, Sivasithamparam K (2000) Comparative effects ofdifferent smoke treatments on germination of Australian native plants. Austral Ecology 25, 610 615.

164 Long J (1989) Leaving the desert: actors and sufferers in the Aboriginal exodus from the Western Desert. Aboriginal History 13, 9-43.

Lunt ID, Morgan JW (1999) Effect of fire frequency on plant composition at the Laverton North grassland reserve, Victoria. The Victorian Naturalist 116, 84-89.

Maconochie JR (1982) Regeneration ofarid zone plants: a floristic survey. In 'Evolution ofthe Flora and Fauna of Arid Australia'. (Eds WR Barker and PJM Greenslade) pp. 141 144. (Peacock Publications: Hong Kong)

Malanson GP, Trabaud L (1988) Vigour of post-fire resprouting by Quercus coccifera L. Journal ofEcology 76, 351-365.

Malone CR (1967) A rapid method for enumeration ofviable seeds in soil. Weeds 15, 381-382.

Masters P, Nano T, Southgate R, Allan GE, Reid J (1997) 'The Mulgara: its distribution in relation to landscape type, fire age, predators, and geology in the Tanami Desert.' Parks and Wildlife Commission NT: Alice Springs.

McArthur AG (1972) Fire control in the arid and semi-arid lands ofAustralia. In 'The use oftrees and shrubs in the dry country of Australia'. (Ed. N Hall) pp. 488-515. (Australian Government Publishing Service: Canberra)

Meney KA, Nielssen GM, Dixon KW (1994) Seed bank patterns in Restionaceae and Epacridaceae after wildfire in Kwongan in southwestern Australia. Journal ofVegetation Science 5, 5-12.

Minnich RA (1983) Fire mosaics in southern California and northern Baja California. Science 219, 1287-1294.

Molina MJ, Llinares JV (200 1) Temperature-time curves at the soil surface in marquis summer fires. International Journal ofWildland Fire 10, 45-52.

Moreno JM, Oechel WC (1991) Fire intensity and herbivory effects on post-fire resprouting ofAdenostomafasciculatum in southern Californian chaparral. Oecologia 85, 429-433.

Morgan JW (1998) Composition and seasonal flux ofthe soil seed bank of species-rich Themeda triandra grasslands in relation to burning history. Journal ofVegetation Science 9, 145-156.

165 Morgan JW (1999) Defining grassland fire events and the response of perennial plants to annual fire in temperate grasslands ofsouth-eastern Australia. Plant Ecology 144, 127 144.

Morrison DA (2002) Effects offire intensity on plant species composition of sandstone communities in the Sydney region. Austral Ecology 27, 433-441.

Morrison DA, Cary GJ, Pengelly SM, Ross DG, Mullins BJ, Thomas CR, Anthony TS (1995) Effects offire frequency on plant species composition of sandstone communities in the Sydney region: Inter-fire interval and time since fire. Australian Journal of Ecology 20,239-247.

Morrison DA, McClay K, Prorter C, Rish S (1998) The role ofthe lens in controlling heat-induced breakdown oftesta-imposed dormancy in native Australian legumes. Annals ofBotany 82, 35-40

Morton SR (1990) The impact of European settlement on the vertebrate animals ofarid Australia: a conceptual model. Proceedings ofthe Ecological Society ofAustralia 16, 201-213.

Morton SR, Stafford-Smith OM, Friedel MH, Griffin GF, Pickup G (1995) The stewardship ofarid Australia: ecology and landscape management. Journal ofArid Environments 43, 195-217.

Nano CEM (2005) 'Within- and between-habitat coexistence in mulga (Acacia aneura) shrubland-spinifex (Triodia spp.) hummock grassland mosaics in central Australia.' (University ofNew England: Phd thesis)

Navie SC, Cowley RA, Rogers RW (1996) The relationship between distance from water and the soil seed bank in a grazed, semi-arid subtropical rangeland. Australian Journal of Botany 44, 421-431.

Ne'eman G, Izhaki I (1999) The effect of stand age and microhabitat on soil seed banks in Mediterranean Aleppo pine forests after fire. Plant Ecology 144, 115-125.

Nieuwenhuis A (1987) The effect offire frequency on the sclerophyll vegetation ofthe West Head, New South Wales. Australian Journal ofEcology 12, 373-385.

Noble JC (1982) The significance offire in the biology and evolutionary ecology of mallee Eucalyptus populations. In 'Evolution ofthe flora and fauna ofarid Australia'. (Eds WR Barker and PJM Greenslade) pp. 153-159. (Peacock Publications: Adelaide)

Noble JC (1989a) Fire regimes and their influence on herbage and mallee coppice dynamics. In 'Mediterranean landscapes in Australia: mallee ecosystems and their management'. (Eds JC Noble and RA Bradstock) pp. 168-181. (CSIRO: East Melbourne)

166 Noble JC (1989b) Fire studies in mallee (Eucalyptus spp.) communities of western New South Wales: The effects offire applied in different seasons on herbage productivity and their implications for management. Australian Journal ofEcology 14, 169-187.

Noble JC (1993) Fire studies in mallee (Eucalyptus spp.) communities ofwestem New South Wales: grass fuel dynamics and associated weather patterns. RangelandJournal 15, 270-297.

Noble JC, Smith A W, Leslie HW (1980) Fire in the mallee shrublands of western New South Wales. Australian Rangelands Journal 2, 104-114.

O'Connell JF, Allen FJ (1998) When did humans first arrive in greater Australia, and why is it important to know? Evolutionary Anthropology 6, 132-146.

O'Donoghue JG (1916) Rambles in Raak. Victorian Naturalist 33, 7-15.

O'Dowd OJ, Gill AM (1984) Predator satiation and site alteration following fire: mass reproduction of Alpine Ash (Eucalyptus delagatensis) in southeastern Australia. Ecology 65, 1052-1066.

Odion DC, Davis FW (2000) Fire, soil heating, and the formation ofvegetation patterns in chaparral. Ecological Monographs 70, 149-169.

Olano JM, Menges ES, Martinez E (2006) Carbohydrate storage in five resprouting Florida shrub plants across a fire chronosequence. New Phytologist 170, 99-106.

Olson MS, Platt WJ (1995) Effects ofhabitat and growing season fires on resprouting of shrubs in longleaf pine savannahs. Vegetatio 119, 101-118.

Pannel JR, Myerscough PJ (1993) Canopy-stored seed banks ofAllocasuarina distyla and A.nana in relation to time since fire Australian Journal ofBotany 41, 1-9.

Perry RA, Litchfield WH, Quinlan T (1962) 'Land systems ofthe Alice Springs area.' CSIRO, Melbourne.

Perry RA, Mabbutt JA, et al. (1962) 'General Report on Lands ofthe Alice Springs Area, Northern Territory, 1956-57: Land Research Series no. 6.' CSIRO, Melbourne.

Planisek SL, Pippen RW (1984) Do sand dunes have seed banks? The Michigan Botanist 23,169-177.

Preece PB (1971) Contributions to the biology of mulga II. *Germination. Australian Journal ofBotany 19, 39-49.

167 Preisler HK, Haase SM, Sackett SS (2000) Modeling and risk assessment for soil temperatures beneath prescribed forest fires. Environmental and Ecological Statistics 7, 239-254.

Prescott EE (1941) Germination ofthe seed of mal lee eucalypts. Victorian Naturalist 58, 8.

Prescott 1A (1952) 'The soils ofAustralia in relation to vegetation and climate.' (Commonwealth Scientific and Industrial Research Organization: Melbourne)

Quinn GP, Keough M1 (2002) 'Experimental Design and Data Analysis for Biologists.' (University Press: Cambridge)

Raison R1 (1979) Modification ofthe soil environment by vegetation fires, with particular reference to nitrogen transformations: a review. Plant Soil 51, 73-108.

Read TR, Bellairs SM, Mull igan DR, Lamb D (2000) Smoke and heat effects on soil seed bank germination for the re-establishment of a native forest community in New South Wales. Austral Ecology 25,48-57.

Riba M (1997) Effects of cutting and rainfall pattern on resprouting vigor and growth of Erica arborea L. Journal ofVegetation Science 8, 401-404.

Rice B, Westoby M (1999) Regeneration after fire in Triodia R. Br. Australian Journal of Ecology 24, 563-572.

Rundel PW (1981) Fire as an ecological factor. In 'Physiological plant ecology. 1 Responses to the physical environment.' (Eds OL Lange, PS Nobel, CB Osmond and H Zeigler) pp. 50 1-538. (Springer-Verlag: Berlin)

Russel-Smith 1, Ryan PG, Cheal DC (2002) Fire regimes and the conservation of sandstone heath in monsoonal northern Australia: frequency, interval, patchiness. Biological Conservation 104, 91-106.

Russel-Smith J, Ryan PG, Durieu R (1997) A LANDSAT MSS- derived fire history of Kakadu National Park, monsoonal northern Australia, 1980-94: seasonal extent, frequency and patchiness. Journal ofApplied Ecology 34, 748-766.

Schimmel J, Granstrom A (1996) Fire severity and vegetation response in the Boreal Swedish forest. Ecology 77, 1436-1450.

Scotter DR (1970) Soil temperatures under grass fires. Australian Journal ofSoil Research 8, 273-9.

168 Noble JC (1989b) Fire studies in mallee (Eucalyptus spp.) communities of western New South Wales: The effects offire applied in different seasons on herbage productivity and their implications for management. Australian Journal ofEcology 14, 169-187.

Noble JC (1993) Fire studies in mallee (Eucalyptus spp.) communities ofwestern New South Wales: grass fuel dynamics and associated weather patterns. RangelandJournal 15,270-297.

Noble JC, Smith A W, Leslie HW (1980) Fire in the mallee shrublands ofwestern New South Wales. Australian Rangelands Journal 2, 104-114.

O'Connell JF, Allen FJ (1998) When did humans first arrive in greater Australia, and why is it important to know? Evolutionary Anthropology 6, 132-146.

O'Donoghue JG (1916) Rambles in Raak. Victorian Naturalist 33, 7-15.

O'Dowd OJ, Gill AM (1984) Predator satiation and site alteration following fire: mass reproduction of Alpine Ash (Eucalyptus delagatensis) in southeastern Australia. Ecology 65, 1052-1066.

Odion DC, Davis FW (2000) Fire, soil heating, and the formation ofvegetation patterns in chaparral. Ecological Monographs 70, 149-169.

Olano JM, Menges ES, Martinez E (2006) Carbohydrate storage in five resprouting Florida shrub plants across a fire chronosequence. New Phytologist 170, 99-106.

Olson MS, Platt WJ (1995) Effects of habitat and growing season fires on resprouting of shrubs in longleaf pine savannahs. Vegetatio 119, 101-118.

Pannel JR, Myerscough PJ (1993) Canopy-stored seed banks ofAllocasuarina distyla and A.nana in relation to time since fire Australian Journal ofBotany 41, 1-9.

Perry RA, Litchfield WH, Quinlan T (1962) 'Land systems ofthe Alice Springs area.' CSIRO, Melbourne.

Perry RA, Mabbutt JA, et of. (1962) 'General Report on Lands ofthe Alice Springs Area, Northern Territory, 1956-57: Land Research Series no. 6.' CSIRO, Melbourne.

Planisek SL, Pippen RW (1984) Do sand dunes have seed banks? The Michigan Botanist 23, 169-177.

Preece PB (1971) Contributions to the biology of mulga II.*Germination. Australian Journal ofBotany 19, 39-49.

167 Preisler HK, Haase SM, Sackett SS (2000) Modeling and risk assessment for soil temperatures beneath prescribed forest fires. Environmental and Ecological Statistics 7, 239-254.

Prescott EE (1941) Germination of the seed of mallee eucalypts. Victorian Naturalist 58, 8.

Prescott JA (1952) 'The soils of Australia in relation to vegetation and climate.' (Commonwealth Scientific and Industrial Research Organization: Melbourne)

Quinn GP, Keough MJ (2002) 'Experimental Design and Data Analysis for Biologists.' (University Press: Cambridge)

Raison RJ (1979) Modification ofthe soil environment by vegetation fires, with particular reference to nitrogen transformations: a review. Plant Soil 51, 73-108.

Read TR, Bellairs SM, Mulligan DR, Lamb 0 (2000) Smoke and heat effects on soil seed bank germination for the re-establishment of a native forest community in New South Wales. Austral Ecology 25, 48-57.

Riba M (1997) Effects ofcutting and rainfall pattern on resprouting vigor and growth of Erica arborea L. Journal ofVegetation Science 8, 401-404.

Rice B, Westoby M (1999) Regeneration after fire in Triodia R. Br. Australian Journal of Ecology 24, 563-572.

Rundel PW (1981) Fire as an ecological factor. In 'Physiological plant ecology. 1 Responses to the physical envirorunent.' (Eds OL Lange, PS Nobel, CB Osmond and H Zeigler) pp. 50 1-538. (Springer-Verlag: Berlin)

Russel-Smith J, Ryan PG, Cheal DC (2002) Fire regimes and the conservation of sandstone heath in monsoonal northern Australia: frequency, interval, patchiness. Biological Conservation 104, 91-106.

Schimmel J, Granstrom A (1996) Fire severity and vegetation response in the Boreal Swedish forest. Ecology 77, 1436-1450.

Scotter DR (1970) Soil temperatures under grass fires. Australian Journal ofSoil Research 8, 273-9.

168 Setterfield SA (1997) The impact ofexperimental fire regimes on seed production in two tropical eucalypt species in northern Australia. Australian Journal ofEcology 22, 279 287.

Shea SR, McCormick J, Portlock CC (1979) The effect offires on the regeneration of leguminous species in the northern jarrah (Eucalyptus marginata Sm) forest of Western Australia. Australian Journal ofEcology 4, 195-205.

Simpson B (1999) 'Rehabilitation at the Granites gold mine: a study of unmined vegetation and seed banks.' (Flinders University: Honours thesis)

Smith MA (1989) The case for a resident human population in the Central Australian Ranges during full glacial aridity. Archaeology in Oceania 24, 93-105.

Smith MA, Loneragan WA, Grant CD, Koch JM (2000) Effect oftop-soil seed banks of rehabilitated bauxite mine sites in the jarrah forest of western Australia. Ecological Management and Restoration 1, 50-60.

Snyman HA (2005) The effect of fire on the soil seed bank ofa semi-arid grassland in South Africa. South African Journal ofBotany 71, 53-60.

Steward FR, Peter S, Richon JB (1990) A method for predicting the depth of lethal heat penetration into mineral soils exposed to fires of various intensities. Canadian journal of Forest Research 20, 919-926.

Stretch J (1996) Fire management ofspinifex pastures in the coastal and west Pilbara. Miscellaneous publication 23/96. Agriculture Western Australia, Perth.

Stuart JM (1975) 'The journals ofJohn McDouall Stuart during the years 1858, 1859, 1860, 1861, and 1862. .' (Libraries Board of South Australia: Adelaide)

Suijdendorp H (1967) The influence of management practices on 'spinifex' (Triodia pungens) grazing. Unpublished M.Sc. (Agric.) thesis, University of Western Australia.

Suijdendorp H (1981) Responses ofthe hummock grasslands of northwestern Australia to fire. In 'Fire and the Australian Biota'. (Eds AM Gill, RH Groves and IR Nobel) pp. 417 426. (Australian Academy of Science: Canberra)

Tang Y, Boulter SL, Kitching RL (2003) Heat and smoke effects on the germination of seeds from soil seed banks across forest edges between subtropical rainforest and eucalypt forest at Lamington National Park, south-eastern Queensland, Australia. Australian Journal ofBotany 51,227-237.

Taylor JE, Monamy V, Fox BJ (1998) Flowering ofXanthorrhoeafulva: the effect of fire and clipping. Australian Journal ofBotany 46, 241-251.

169 Thomas PB, Charles Morris E, Auld TO (2003) Interactive effects of heat shock and smoke on germination of nine species forming soil seed banks within the Sydney region. Austral Ecology 28, 674-683.

Tieu A, Dixon KW, Meney KA, Sivasithamparam K (2001) The interaction ofheat and smoke in the release of seed dormancy in seven species from southwestern Western Australia. Annals ofBotany 88,259-265.

Tothill JC, Shaw NH (1968) Temperatures under fires in bunch spear grass pastures of south-east Queensland. The Journal ofthe Australian Institute ofAgricultural Science, 94-97.

Turner EJ (1979) 'Report on management ofthe eucalypt, spinifex woodlands ofcentral western Queensland.' Report 79/2. Division of Land Utilization, Queensland Department of Primary Industries.

Valette J, Gomendy V, Marechal J, Houssard C, Gillon D (1994) Heat transfer in the soil during very low intensity experimental fires: the role ofduff and soil moisture content. International Journal ofWildland Fire 4, 225-237. van Leeuwen SJ, Start AN, Bromilow RB, Fuller PJ (1995) Fire and the floristics of mulga woodlands in the Hammersley Ranges, Western Australia. In 'Ecological Research and Management in the Mulga lands'. (Eds MJ Page and TS Beutel) pp. 169-175. (University ofQueensland, Gatton College) van Wagner CE (1973) Height of scorch in forest fires. Canadian Journal ofForest Research 3, 373-378.

Veblin TT, Kitzberger T, Villalba R, Donnegan J (1999) Fire history in northern Patagonia: The roles ofhumans and climatic variation. Ecological Monographs 69, 47 67.

Vesk PA, Warton 01, Westoby M (2004) Sprouting by semi-arid plants: testing a dichotomy and predictive traits. Gikos 107, 72-89.

Vlok JH, Yeaton RI (2000) The effect of short fire cycles on the cover and density of understorey sprouting species in South African mountain fynbos. Diversity and Distributions 6, 233-242.

Wade DO, Johansen RW (1986) 'Effects offire on southern pine: observations, and recommendations.' USDA Forest Service General Technical Report SE-41.

Warburton PE (1875) 'Journey across the Western Interior ofAustralia.' (Sampson, Low, Marston, Low, and Searle: London)

170 Wellington AB (1984) Leaf water potentials, fire and the regeneration of mallee eucalypts in semi-arid, south-eastern Australia. Gecologia 64, 360-362.

Wellington AB (1989) Seedling regeneration and the population dynamics ofeucalypts. In 'Mediterranean Landscapes in Australia: mallee ecosystems and their management'. (Eds JC Noble and RA Bradstock). (CSIRO: East Melbourne)

Wellington AB, Noble IR (1985) Seed dynamics and factors limiting recruitment ofthe mallee Eucalyptus incrassata in semi-arid, south-eastern Australia. Australian Journal of Ecology 73, 657-666.

Westoby M, Rice B, Griffin GF, Friedel MH (1988) The soil seed bank of Triodia basedowii in relation to time since fire. Australian Journal ofEcology 13, 161-169.

Whelan RJ (1995) 'The Ecology of Fire.' (Cambridge University Press: Cambridge)

Williams PR, Congdon RA, Grice AC, Clarke Pl (2004) Soil temperature and depth of legume germination during early and late dry season fires in a tropical eucalypt savannah of north-eastern Australia. Austral Ecology 29, 258-263.

Williams PR, Congdon RA, Grice AC, Clarke Pl (2005) Germinable soil seed banks in a tropical savannah: seasonal dynamics and effects offire. Austral Ecology 30, 79-90.

Wi lis TJ, Read J (2002) Effects of heat and smoke on germination of soi I stored seed in a south-eastern Australian sand heathland. Australian Journal ofBotany 50, 197-206.

Winkworth RE (1967) The composition of several arid spinifex grasslands ofcentral Australia in relation to rainfall, soil water relations, and nutrients. Australian Journal of Botany 15, 107-130.

Wright HA, Bunting SC, Neuenschwander LF (1976) Effect of fire on honey mesquite. Journal ofRange Management 29,467-471.

Zammit CA, Zedler PH (1988) The influence ofdominant shrubs, fire, and time since fire on soil seed banks in mixed chaparral. Vegetatio 75, 175-187.

Zedler PH, Gautier CR, Mcmaster GS (1983) Vegetation change in response to extreme events: the effect ofa short interval between fires in Californian Chaparral and coastal scrub. Ecology 64, 809-818.

Zimmer WJ (1940) Plant invasions in the Mallee. Victorian Naturalist 56, 143-147.

171 1CXXl 13853

800 48944

700 I 10893 600 ' 3252

500 '

400

300

200 100 ~~i~~~~~~~~~~~~~~~~~~~~~~~~~UI~~~~~~~~~~~~~~~~II~~~~I~~ij~~~~~~~~ ~~~~~~~~W~~I~ o ~ L{) co ,... ~ ~ ~ ~ ~ ffi ~ ffi § § § § ~ ~ ~ ~ ~ ~ ~ ffi ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ § § ~ !J; ffi ,... ~ ~ ~ ~ ~ ~ ~ ~ b> r r r r r r r r r r , , , , , , , , , , , , , , , ,... ,... ,... ffi ~ ~ ~ ~ ~ ~

Appendix 2.1 Annual rainfall records for Alice Springs region. Data are the averaged totals from Alice Springs Post Office, Alice Springs Air Port, and Derwent, Bond Springs and New Haven stations. Shaded bars indicate times when widespread fire events are known to have occurred across central Australia (Kimber 1983; Ralph 1984; Griffin and Friedel 1985; Gill 2000). Numbers above shaded bars indicate cumulative rainfall (mm) since last known fire event.

172 Appendix 4.1 Species groups used in supplementary matrices in RDA analyses in Chapters 4 and 5. Families follow Angiosperm Phylogeny Group (http://www.mobot.org; 5/1/2007). Names of other taxa follow Australian Plant Name Index (http://www.anbg.gov.au; 5/1/2007)). Abbreviations are as follows: R-resprouters; OS-obligate-seeders; FE-fire ephemerals; S serotinous woody; NS-non-serotinous woody; T-Triodia; C-clonal. Fire response and life history classifications were taken from NT herbarium 'Southern NT species fire response database', as well as from consultations with NT herbarium botanists and from my own field observations.

R OS FE S NS T C Apiaceae Trachymene glaucifolia (F. Muell.) Benth. * Apocynaceae subfam. Asclepiadoideae Marsdenia australis (R. Br.) Druce * Asteraceae Brachyscome ciliaris (Labill.) Less. * Minuria tridens (D.A Cooke) Lander * Boraginaceae Halgania cyanea Lindl. * Halgania erecta Ewart & B. Rees * Trichodesma zeylanicum (Burm. f.) R Br. (Boraginaceae) * Casuarinaceae Allocasuarina decaisneana (F. Muell.) L.AS. Johnson * * Cleomaceae Cleome viscosa L. * Convolvulaceae Bonamia erecta RW. Johnson * Evolvulus alsinoides L. * Euphorbiaceae Euphorbia drummondii Boiss. * Euphorbia tannensis Spreng. var. eremophila (A Cunn) Hassal * Euphorbia wheeleri Baill. * Fabaceae subfam. Faboideae Crata/aria eremaea F. Muell. * Erythrina verspertilio Benth. * /ndigofera psammophila P.G. Wilson * Leptosema chambersii F. Muell *

173 R OS FE S NS T C Fabaceae subfam. Mimosoideae Acacia adsurgens Maiden & Blakely * * Acacia ancistrocarpa Maiden & Blakely * * Acacia aneura F. Muell. ex Benth. ** Acacia bivenosa (DC.) subsp. wayi (Maiden) Pedley * * Acacia coriacea DC. * * Acacia cuthbertsonii J.G. Luehmann * * Acacia estrophia/ata F. Muell. * * Acacia inaequilatera Domin. * * Acacia kempeana F. Muell. * * Acacia ligulata A. Cunn. ex Benth * * Acacia maitlandii F. Muell. * * Acacia melleodora Pedley * * Acacia murrayana F. Muell. ex Benth. * * Acacia pruinocarpa Tindale * * Acacia ramulosa W.V. Fitzg. ** Acacia sessiliceps F. Muell. * * Acacia tennuissima F. Muell. * * Acacia tetragonophylla F. Muell. * * Acacia victorae Benth. * * Fabaceae subfam. Caesalpinoideae Senna form taxon 'helmsii' (Symon) Randell * Senna form taxon 'petio/aris'Randell * Senna form taxon 'sturtii' (R.Br.) Randell * Senna form taxon 'filifolia' Randell * Goodeniaceae Brunonia australis Sm. Goodenia larapinta Tate * Goodenia triodiophila Carolin * Gyrostemonaceae Codonocarpus cotinifolius F. Muell. * Gyrostemon tepperi (F. Muell. ex H. Walter) A.S. George * * Lamiaceae subfam. Prostantheroideae Oicrastylis gilesii F. Muell * Malvaceae subfam. Byttnerioideae Keraudrenia integrifolia Steud *

174