Mechanisms behind persistence of a fire-sensitive alternative stable state system in the Gibson Desert, Western Australia.

Alternative vegetation types that switch from one to another under contrasting fire regimes are termed fire-mediated alternative stable states (FMASS). Typically, pyrophylic communities (i.e., vegetation assemblages favored by burning) dominate under high frequencies or intensities of fire. Conversely, fire-sensitive (pyrophobic) vegetation types persist under long fire-free conditions. As the persistence traits of plants of FMASS systems are generally poorly researched, threshold levels of pyric disturbance that trigger 'state-switching' are often unknown. Dense thickets of the obligate-seeder shrub waputi (Aluta maisonneuvei ssp. maisonneuvei [Myrtaceae]) form fire-retarding woody islands within highly flammable spinifex (Triodia spp.) grasslands in arid Australia. To examine the tolerance of Aluta thickets to burning, we investigated: (1) the influence of post-fire rainfall and fire severity on recruitment (a field study); (2) soil seedbank densities (a field study); and (3) fire-related dormancy cues in seeds (a germination trial). We found a positive relationship between recruitment and post-fire rainfall volume, and much higher mean recruitment at sites with high- (5.9 seedlings/m2) than low-severity-burnt (2.2 seedlings/m2) and unburnt shrubs (0.03 seedlings/m2). Post-fire regeneration was mediated by dense soil-borne seedbanks, and the germination trial indicated that smoke promoted germination. Although Aluta shrubs are invariably fire-killed, high-severity fires are unlikely to lead to state shifts from shrubland to grassland because of the ability of mature stands to regenerate from dense, fire-cued seedbanks. Nevertheless, given that Aluta seedlings are exceptionally slow-growing, post-fire droughts combined with fire-return intervals less than the Aluta primary juvenile period of c. 5 years could drive conversion from Aluta- to Triodia-dominated vegetation.

Evidence that shrublands and hummock grasslands are fire-mediated alternative stable states in the Australian Gibson Desert.

Alternative stable state theory predicts that different disturbance regimes may support contrasting ecosystem states under otherwise analogous environmental conditions. In fire-prone systems, this theory is often invoked to explain abrupt ecotones, especially when adjacent vegetation types have contrasting flammabilities and differing tolerances to pyric perturbation. Despite being well-documented in forest-savanna transitions, unambiguous examples of fire-driven alternate stable states (FDASS) in arid systems are rare. The current study examined whether flammable spinifex (Triodia spp.) grasslands and fire-sensitive waputi (Aluta maisonneuvei subsp. maisonneuvei) shrublands in Australia's Gibson Desert represent FDASS. Specifically, analyses of soil and topographic variables assessed whether environmental differences explain habitat zonation. To determine whether different flammabilities of Aluta and Triodia systems may perpetuate alternative states via vegetation-fire feedback processes, community-level fuelloads were quantified to provide an indirect measure of flammability. To determine the propensity for fire to trigger 'state-shifting', community responses to a single high-severity fire were evaluated. Habitat segregation did not relate to between-site environmental differences, and the fuel-load study indicated that the more pyrophytic community (Triodia grassland) is more flammable, and hence more likely to experience higher frequency fire-cycles. Fire was identified as a potential vector of 'state-change', because although both systems regenerated well after fire, Triodia reproduced more prolifically at a younger age than Aluta, and hence should tolerate shorter fire-return intervals. In the absence of between-community topographic and edaphic differences, or herbivores that consume either plant, it is likely that Aluta shrublands and Triodia grasslands represent fire-mediated alternative equilibrium states.

Fire timing in relation to masting: an important determinant of post-fire recruitment success for the obligate-seeding arid zone soft spinifex (Triodia pungens).

Background and Aims: Plant species with fire-triggered germination are common in many fire-prone ecosystems. For such plants, fire timing in relation to the timing of reproduction may strongly influence post-fire population regeneration if: (a) flowering occurs infrequently (e.g. plants are mast seeders); and (b) seed survival rates are low and input from the current year's flowering therefore contributes a large proportion of the viable dormant seedbank. The role of fire timing in relation to masting as a driver of post-fire recruitment has rarely been examined directly, so this study tested the hypothesis that fires shortly after masting trigger increased recruitment of the obligate-seeding arid zone spinifex, Triodia pungens R. Br., an iteroparous masting grass with smoke-cued germination. Methods: Phenological monitoring of T. pungens was conducted over 5 years, while a longitudinal seedbank study assessed the influence of seeding events on soil-stored seedbank dynamics. Concurrently, a fire experiment with randomized blocking was undertaken to test whether T. pungens hummocks burnt shortly after masting have greater post-fire recruitment than hummocks burnt when there has not been recent input of seeds. Key Results: Triodia pungens flowered in all years, though most flowerings were characterized by high rates of flower abortion. A mast flowering with high seed set in 2012 triggered approx. 200-fold increases in seedbank densities, and seedbank densities remained elevated for 24 months after this event. The fire experiment showed significantly higher recruitment around hummocks burnt 6 months after the 2012 mast event than around hummocks that were burnt but prevented from masting by having inflorescences clipped. Conclusions: Fires shortly after masting trigger mass recruitment in T. pungens because such fires synchronize an appropriate germination cue (smoke) with periods when seedbank densities are elevated. Interactions between natural fire regimes, seedbank dynamics and fire management prescriptions must be considered carefully when managing fire-sensitive masting plants such as T. pungens.

Fire after a mast year triggers mass recruitment of slender mulga (Acacia aptaneura), a desert shrub with heat-stimulated germination.

PREMISE OF THE STUDY: Fire typically triggers extensive regeneration of plants with heat-stimulated germination by causing short periods of intense soil heating. If plants with heat-stimulated germination are also subject to seed predation and display mast-seeding cycles, postfire recruitment may be contingent on the seedfall density of prefire masts, and on whether granivores are satiated at the time of fire. METHODS: We conducted a longitudinal seedbank study and a mensurative field experiment in central Australia to examine whether fire and the variation in seedfall density across sites in a mast year interact to influence recruitment of slender mulga (Acacia aptaneura), an iteroparous masting shrub with heat-stimulated germination. KEY RESULT: The seedbank study showed seedbank pulsing after masting, with mean seed counts in the upper 4-cm soil layer being 132.8 seeds/m2 12-mo after a dense seedfall, but only 3.8 seeds/m2 following a year with no seed production. Consistent with this, recruitment increased postfire at sites where denser seedfall had occurred during the preburn mast year. Conversely, little recruitment occurred at unburnt populations, irrespective of prefire seedfall density. CONCLUSIONS: We attribute our findings to: (1) elevated soil temperatures during fires stimulating germination of heat-cued seeds; and (2) granivore satiation following masting facilitating assimilation of seeds into the soil seedbank. These results highlight the importance of rare seed-input events for regeneration in fire-prone systems dominated by masting plants, and provide the first example from an arid biome of fire interacting with masting to influence recruitment.