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Clarke, Peter J
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Given Name
Peter J
Peter
Surname
Clarke
UNE Researcher ID
une-id:pclarke1
Email
pclarke1@une.edu.au
Preferred Given Name
Peter
School/Department
School of Environmental and Rural Science
14 results
Now showing 1 - 10 of 14
- PublicationHow do drought and fire influence the patterns of resprouting in Australian deserts?Rainfall is the key driver of woody cover and life-history attributes in arid grassy biomes where disturbance is mostly rare and of low intensity. However, relatively little is known about the causes of woody community assembly in arid systems that are subject to periodic intense fire disturbance. In the central Australian desert region, grassland and shrubland fire can occur following above average rainfall. Patterns of species regeneration response (resprouting vs. reseeding) are poorly documented in this region. We tested the effects of rainfall and fire on species' resprouting response across the latitudinal rainfall-fire gradient using constrained ordination of 385 sites and general linear models. A resprouting response was significantly greater in grassland habitat as well as at the high end of the rainfall-fire gradient. The frequency of epicormic stem resprouting also increased along the rainfall-fire gradient. We attribute this pattern to the combined effects of frequent fire and rapid gap closure on seedlings of slow-growing, fire-killed woody species in higher rainfall grasslands. In addition, we also demonstrated that rapidly maturing fire-recruiting species are similarly favoured by high fire disturbance. In arid grassy ecosystems, unlike in mesic savanna, flammable grassland supports a mix of resprouting and recruitment functional types, and habitat membership cannot be predicted by resprouting capacity. Regions, such as central Australia, that are characterised by grassland-shrubland mosaics of high and low fuel biomass, respectively, pose specific challenges to fire ecology research that are possibly best dealt with by focussing modelling at the habitat scale.
- PublicationSeed traits and seed bank longevity of wet sclerophyll forest shrubsIn wet sclerophyll forests seedling recruitment either occurs after intermittent fire events or continuously during intervals between fires in gaps created by small-scale disturbances. The dormancy and dispersal characteristics of seeds will influence how plant species exploit these contrasting recruitment opportunities. For example, long-lived seed banks may be crucial for persistence of species that are unable to recruit during intervals between fires if the length of fire intervals exceeds the life span of standing plants (senescence risk). To better understand mechanisms of population persistence during prolonged absence of fire in montane wet sclerophyll forests, we studied seed bank dynamics in four understorey species. We chose two species thought to have fire event-driven recruitment, 'Banksia integrifolia' subsp. monticola (Proteaceae) and 'Goodia lotifolia' (Fabaceae), and two species that are thought to have canopy gap-phase recruitment, 'Trochocarpa laurina' (Ericaceae) and 'Tasmannia stipitata' (Winteraceae). We measured seed rain, seed bank density and used seeds buried in nylon mesh bags to estimate rates of seed decay in the soil over time. All species produced a substantial seed crop on an annual basis. The annual seed crop in three species ('G. lotifolia', 'T. stipitata' and 'T. laurina') was released in a dormant state and developed a persistent seed bank, while one species ('B. integrifolia') lacked dormancy and rapidly germinated under laboratory and field conditions. Seed bank characteristics of 'G. lotifolia' appear to promote episodic recruitment after large landscape-scale fires, those of 'B. integrifolia' appear to promote more continuous recruitment in response to smaller fires and other disturbances that avoid widespread mortality of established plants, while seed bank characteristics of 'T. stipitata' and 'T. laurina' may facilitate both episodic and continuous recruitment under respective types of disturbance. The four species appeared to have varied vulnerabilities and mechanisms for reducing immaturity risk and senescence risk to persistence of their populations under recurrent disturbance. Dormancy, seed bank longevity and seed rain are likely to be useful syndromes for predicting the response of wet sclerophyll forest understorey species to changed disturbance regimes.
- PublicationResprouting as a key functional trait in woody plants: challenges to developing new organizing principlesMost global biomes are shaped by disturbances, such as fire or herbivory, that damage or kill the aboveground biomass of plants. In many of these biomes, however, the damaged plants do not die; rather, they persist through sprouting. In disturbance-prone environments, resprouting from meristems stabilizes plant populations where disturbance may cause demographic bottlenecks. The advantage of resprouting is that it confers persistence under disturbance; however, sprouting has disadvantages such as potentially reducing sexual reproduction and limiting gene flow (Bond & Midgley, 2001; Lamont & Wiens, 2003). Understanding resprouting is critically important for understanding long-term vegetation dynamics, extinction risks, carbon balance and woody plant management. A Vegetation Function Working Group (Working Group 67; http://www.vegfunction.net/wg/67/67_Sprouting.htm) was established in 2009 to identify challenges encountered in developing coherent models of the functional role of resprouting in fire-prone environments, with an emphasis on savanna and Mediterranean biomes. Fire is the most pervasive disturbance and is integral to the evolutionary ecology of these biomes. This broad disciplinary group met at the International Ecological Conference (INTECOL) in Brisbane in September 2009 and reconvened in July 2010 at The University of New England, Armidale, Australia, to review progress. In this report we highlight major challenges encountered in developing unifying models of the functional role of resprouting.
- PublicationFire severity, feedback effects and resilience to alternative community states in forest assemblagesAlternative stable state theory is often used to explain the occurrence of flammable vegetation adjacent to less flammable vegetation where fire regimes mediate the shift between states. In 2002 an extreme landscape scale fire burnt extensive areas of forests in eastern Australia, including rainforests that are rarely severely burnt. This unique event allowed us to test long-held assumptions that predict landscape scale change after major disturbance. We tested three assumptions for detecting alternative community states; (1) that the scale of the event was large enough to remove canopy dominants, (2) fire feedbacks, both positive and negative are present, and (3) shifts in the floristic composition of communities are detected. We also examined whether high severity fires resulted in a community shift from less flammable to more flammable vegetation (e.g. from rainforest to wet sclerophyll vegetation), by examining floristic composition of vegetation communities (rainforest, wet sclerophyll forest, and dry sclerophyll forest) when burnt at different fire severities (high and low). Conversely, we tested whether there was a state shift from flammable to less flammable vegetation communities in sclerophyll forests long unburnt. In addition, we determined if there was any evidence that antecedent fire regimes and fire severity influenced flammability feedbacks. Severe fire caused significant ongoing disruption to forest canopies and fire effects were still detectable some 7 years after fire. Whilst some pyrogenic environmental feedbacks were detected from historic fire regimes, we found no shifts in the floristic composition or pyrogenic traits of forests burned at high severity. Medium-term (30 year) fire exclusion did not result in the sclerophyll forest becoming more pyrophobic although some fire-cued species senesced in the absence of fire. Contrary to expectation, we found no evidence that the floristic composition of less flammable vegetation burnt at high severity became more similar to flammable vegetation burnt at low severity or that were unburnt. Conversely, with more than 30 years of fire exclusion there was no evidence that the sclerophyllous communities became more floristically similar to rainforest. We have shown that species assemblages in warm-temperate rainforest were resilient to a catastrophic fire event and propose that these forests are unlikely to represent alternative community states driven by fire alone.
- PublicationResilience of a high-conservation-value, semi-arid grassland on fertile clay soils to burning, mowing and ploughing(Wiley-Blackwell Publishing Asia, 2010)
;Lewis, Tom; ; In grassland reserves, managed disturbance is often necessary to maintain plant species diversity. We carried out experiments to determine the impact of fire, kangaroo grazing, mowing and disc ploughing on grassland species richness and composition in a nature reserve in semi-arid eastern Australia. Vegetation response was influenced by winter–spring drought after establishment of the experiments, but moderate rainfall followed in late summer–autumn. Species composition varied greatly between sampling times, and the variability due to rainfall differences between seasons and years was greater than the effects of fire, kangaroo grazing, mowing or disc ploughing. In the fire experiment, species richness and composition recovered more rapidly after spring than autumn burning. Species richness and composition were similar to control sites within 12 months of burning and mowing, suggesting that removal of the dominant grass canopy is unnecessary to enhance plant diversity. Two fires (separated by 3 years) and post-fire kangaroo grazing had only minor influence on species richness and composition. Even disc ploughing caused only a small reduction in native richness. The minor impact of ploughing was explained by the small areas that were ploughed, the once-off nature of the treatment, and the high degree of natural movement and cracking in these shrink-swell soils. Recovery of the composition and richness of these grasslands was rapid because of the high proportion of perennial species that resprout vegetatively after fire and mowing. There appears to be little conservation benefit from fire, mowing or ploughing ungrazed areas, as we could identify no native plant species dependent on frequent disturbance for persistence in this grassland community. However, the ability of the 'Astrebla'- and 'Dichanthium'-dominated grasslands to recover quickly after disturbance, given favourable seasonal conditions, suggests that they are well adapted to natural disturbances (e.g. droughts, fire, flooding and native grazing). - PublicationResprouting as a key functional trait: how buds, protection and resources drive persistence after fire(Wiley-Blackwell Publishing Ltd, 2013)
; ;Lawes, MJ ;Midgley, JJ ;Lamont, BB ;Ojeda, F ;Burrows, GE ;Enright, NJResprouting as a response to disturbance is now widely recognized as a key functional trait among woody plants and as the basis for the persistence niche. However, the underlying mechanisms that define resprouting responses to disturbance are poorly conceptualized. Resprouting ability is constrained by the interaction of the disturbance regime that depletes the buds and resources needed to fund resprouting, and the environment that drives growth and resource allocation. We develop a buds-protection-resources (BPR) framework for understanding resprouting in fire-prone ecosystems, based on bud bank location, bud protection, and how buds are resourced. Using this framework we go beyond earlier emphases on basal resprouting and highlight the importance of apical, epicormic and below-ground resprouting to the persistence niche. The BPR framework provides insights into: resprouting typologies that include both fire resisters (i.e. survive fire but do not resprout) and fire resprouters; the methods by which buds escape fire effects, such as thick bark; and the predictability of community assembly of resprouting types in relation to site productivity, disturbance regime and competition. Furthermore, predicting the consequences of global change is enhanced by the BPR framework because it potentially forecasts the retention or loss of above-ground biomass. - PublicationFire severity and nutrient availability do not constrain resprouting in forest shrubsPlants often survive disturbances such as fire by resprouting which involves having both protection traits and carbohydrate storage capacity. Protection traits not only act directly to insulate meristems but also prevent combustion of carbohydrate stores. Rapid stem growth also allows replenishment of carbohydrate stores ensuring persistence through another event. Resource availability may, however, constrain the ability to develop resilience to high-severity fires through either nutrient limitation or light limitation. We tested whether fire severity influenced resprouting ability of woody plants in two contrasting environments, low nutrient dry sclerophyll forest and more fertile wet sclerophyll forest. We tested which fire protection and growth traits were associated with resprouting ability (27 species) and resprouting vigour (16 species). Fire severity did not limit the ability of most species to resprout in either forest type. There was no generalized protection syndrome for surviving top kill, but combinations of bud protection and growth together with storage capacity appear to drive resprouting ability. In nutrient-limited forests, low specific leaf area (SLA) may reduce stem growth in resprouters, causing more reliance on bud protection through bark thickness. Conversely, in the more fertile forests, where light becomes limiting with time-since-fire, high SLA appears to increase the capacity for rapid stem growth with less emphasis on developing thicker bark. These different syndromes appear to be adaptive as fire severity did not influence survival in either forest type.
- PublicationCosts and benefits of relative bark thickness in relation to fire damage: a savanna/forest contrast1. In fire-prone ecosystems, bark protects the stem bud bank from fire. Absolute bark thickness is a good indicator of this protective function, but it depends on stem size as well as inherent differences between species. Relative bark thickness (i.e. relative to stem diameter) takes the latter into account. We argue that relative bark thickness is an important functional trait offering insights to the evolution of species persistence in fire-prone habitats. 2. During growth ontogeny different species can acquire absolutely thick bark through having: (i) relatively thick bark (i.e. an early commitment to thick bark) or (ii) relatively thin bark but fast stem diameter growth rates. We test the hypothesis that the most effective way of protecting tree stems from frequent fire is by having relatively thick-barked small stems. We predict that species with higher relative bark thickness are more common in fire-prone habitats. In habitats with long fire-free intervals such as rainforest, delayed investment in bark thickness results in thin bark. 3. We examined the relative bark thickness of woody congeners from Australian non-fire-prone forest and fire-prone savanna and in other tree-dominated systems world-wide. We determined the relative cost of acquiring absolute bark thickness of 0.5 cm for different rates of bark allocation. The insulating benefits of bark were considered a linear function of bark thickness. 4. Synthesis. We suggest that relatively thick bark minimizes the costs of acquiring absolutely thick bark, and it confers greater protection to smaller stems. The cost of acquiring thick bark prevents small trees from merely accumulating bark as a consequence of fast height or stem diameter growth. Accordingly, our field survey indicated that forest species had relatively thin bark and acquired thick bark only as a consequence of very large size, while fire-prone savanna species had relatively thick-barked small stems. Based on this, relative bark thickness appears to be a good predictor of local fire regimes and is a useful plant functional trait.
- PublicationEcology of plant resprouting: populations to community responses in fire-prone ecosystemsResprouting has emerged as a key functional trait in plant ecology over the past decade with more than 400 papers published since 2000 (Web of Science). This special issue of Plant Ecology brings together a set of papers that advance our understanding of this functional trait, in the quest for developing a better conceptual framework for predicting community response to disturbance. This special issue highlights current research on all aspects of the effects of fire and other disturbances on plant resprouting behaviour and the importance of the persistence niche in structuring plant communities.
- PublicationFire intensity, serotiny and seed release in 19 woody species: evidence for risk spreading among wind-dispersed and resprouting syndromesSeed storage in woody fruits on plants has been much studied, whereas trait variation in seed release has been given scant attention. In non-Mediterranean climates, some species release seeds immediately after fire, whereas others retain seeds in open fruits/cones for longer. We expected that species with wind-dispersed seeds and those killed by fire would spread their recruitment risks by having stronger cues for fruit opening and slower seed release once fruits were open. We therefore tested whether fire intensity (heat) affected fruit opening and seed release in 19 species. We then contrasted fruit opening and seed release among (1) serotiny levels (weak, moderate, strong), (2) dispersal (wind v. unassisted) and (3) resprouting ability (killed v. resprout) traits. Only three species required heat for fruit opening. Most species, however, retained varying proportions of seeds in open fruits. Strongly and moderately serotinous species retained seeds in open fruits longer than did weakly serotinous species. Both species with wind-dispersed seeds and fire-killed species required stronger heat effects for fruits to open but retained seeds in open fruits longer than did species with alternative traits. By delaying seed release after fruits have opened, species with wind-dispersed seeds, and those that are killed by fire, maximise the ability of seeds to arrive at safe sites after fire.