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MURDOCH RESEARCH REPOSITORY http://researchrepository.murdoch.edu.au

This is the author's final version of the work, as accepted for publication following peer review but without the publisher's layout or pagination.

Ruthrof, K.X. , Calver, M.C. , Dell, B. and Hardy, G.E.St.J. (2011) Look before planting: using smokewater as an inventory tool to predict the soil seed bank and inform ecological management and restoration.

Ecological Management & Restoration, 12 (2). pp. 154-157. http://researchrepository.murdoch.edu.au/5005

Copyright © 2011 Ecological Society of Australia.

It is posted here for your personal use. No further distribution is permitted.

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Look before planting: using smokewater as an inventory tool to predict the soil seedbank and

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inform ecological management and restoration. Katinka X. Ruthrof,1 Michael C. Calver,1 Bernard

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Dell1 and Giles E. St.J. Hardy1 (1 Centre of Excellence for Climate Change and Woodland and Forest

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Health, Murdoch University, South Street, Murdoch, 6150, Australia; Tel.: +61 (08) 9360 2605; Email:

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[email protected])

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Key words: smokewater, soil seedbank, land management, invasive species.

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Summary

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This study tested the efficacy of smokewater to determine the potential germination from soil seedbank in

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three management sites of the same National Park: a forest site prior to restoration, an ex-pine plantation

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site and an ex-mine site. This will provide further information to land managers so that more accurate

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planning can occur. Results showed that smokewater significantly increased the germination from the soil

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seedbank and significant differences in the level of germination of weed species from the soil seedbank

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were seen between the three management sites. This use of smokewater may be a useful tool to help

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predict differences in the soil seedbank compared with predicting soil seedbank based on land use history

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and recent condition.

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Introduction

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Successful land management requires accurate and reliable information on the characteristics of many

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plant species (Thomson, et al. 1993). Land managers are often faced with a high degree of uncertainty

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regarding the level of potential species establishment, particularly regarding weed species in the existing

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soil seedbank. This can make strategic planning difficult. However, it is sometimes possible to predict the

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response of vegetation to management activities from knowledge of the composition of the soil seedbank

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(Marks and Mohler 1985). If the soil seed bank could be analysed, it may be used as a predictive tool for

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land managers. However, collection of original seedbank data is often prohibitively labour intensive and

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expensive (Thomson, et al. 1993). This paper explores a glasshouse-based method that uses smokewater

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to help predict the potential weed soil seedbank and provide more information to land managers.

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Karrikinolide, a butenolide found in smoke (Flematti, et al. 2004), has been used in the form of aqueous

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smoke (smokewater) or aerosol smoke to enhance seed germination in the South African fynbos (De

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Lange and Boucher 1990), Western Australian bushland (Dixon, et al. 1995), post-mining situations

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(Roche, et al. 1997); un-mined woodland (Rokich, et al. 2002) and in old-fields (Standish, et al. 2007).

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Collectively, these results show firstly, the wide range of uses of smoke and secondly, that the use of

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smoke has practical application by providing more information about the soil seedbank. Smokewater may

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be a useful and comparatively inexpensive tool to conduct an inventory the soil seedbank prior to making

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management decisions.

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In this paper we aim to test the use of smokewater as an inventory tool at different management sites

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within a degraded Eucalyptus gomphocephala (Tuart) forest, the Ludlow Tuart Forest, in the south west

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of Western Australia. We focus on three management sites: a forest site pre-restoration, ex-pine plantation

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and ex-mine sites in which we aim to answer the following question: Can smokewater assist in predicting

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the weed soil seedbank of three different management sites within the same degraded forest and hence

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inform ecological management and restoration?

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Methods

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The Ludlow Tuart Forest (2049ha) is located 200 km south of Perth, Western Australia (33o35’08.72”S

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115o29’30.57”E). The average annual rainfall is 811.9 mm, 80% of which falls between May and

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September (BOM 2009a). The nearest weather station is located in Busselton Shire, 15 km south-west of

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the site. The soils are classified as part of the Spearwood Dune System, consisting of variable depths of

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siliceous brown and yellow leached sands.

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The three management areas in the Ludlow Tuart Forest are within 1 km of one another. The forest site

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prior to restoration is typical of degraded sites in the area, but has had some weed control over the past

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five years. It contains the dominant canopy species Tuart and low understorey diversity. The ex-mine site

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was in poor condition prior to mining and had the soil profile reestablished with original topsoil following

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mining (stored for 1-5 years) and was recently ripped. The ex-pine plantation site had been harvested for

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pine in the early 2000’s (after >30 years under Pinus radiata) and was fallow. It was noted in the previous

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wet season that the forest site and ex-pine sites seemed to have much higher weed loads than the ex-mine

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site. Each of the three management areas was paired with a site in the adjacent Tuart forest – referred to

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here as ‘off-site’ sites . These, ideally, would have been reference sites however; there are no nearby sites

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that have a low level of disturbance. Therefore, sites were typical of the rest of the forest, which is

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degraded with a long history of grazing, logging and weed invasion. Major weed species were: Annual

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Veldt Grass (Ehrharta longifolia) Dune Onion Weed (Trachyandra divaricata) and Arum Lily

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(Zantedeschia aethiopica) (DEC 2007). Thus, there were six plots in total: 1) the ex-mine and 2) the

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adjacent Tuart forest; 3) the State Forest and 4) the adjacent Tuart forest; and 5) the ex-pine plantation

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and 6) the adjacent Tuart forest.

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Soil samples were taken in May 2007 just prior to the start of the winter rains to allow

temperature-69

sensitive seeds to be responsive. At 25 sampling points located randomly within each of six plots, five

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sub-samples of 10 cm x 10 cm to 2 cm depth were collected at each sampling point, bulked and placed in

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calico bags. The majority of the soil seed bank is held within the top 2.5 cm of the surface (Bellairs and

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Bell 1993).

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The 150 (25 samples at 6 plots) soil samples were air-dried; each mixed thoroughly then split into two

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sub-samples. Sub-samples were spread to a depth of 2 cm over a layer of steam-sterilized sand in lined

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but free-draining plastic trays, which measured 11 cm (width) x 16.5 cm (length) x 5 cm (depth). A liquid

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soil wetting agent (Brunnings Easy Wetta ™) was applied to all trays to reduce hydrophobicity. Half of

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the trays were treated with diluted (10%v/v) smokewater at the equivalent of 50ml/m2. Smokewater was

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created by bubbling smoke from a drum of burning hay through a 20 L container of water for 1 hour

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(Dixon, et al. 1995). The seedling trays were arranged in an evaporative cooled glasshouse (Mean Max.

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28.5 oC Mean Min. 19.2 oC) in May 2007 and watered regularly. Trays were randomized fortnightly.

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Germinants were counted as they appeared after 3 and 10 months (data presented is that from the latter).

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Statistical analysis

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Data were analysed as a repeated measures ANOVA for each site separately. There were two repeated

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measures variables for each site: smoke water (present or absent) and location (on-site, offsite – i.e.

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different management sites and adjacent forest sites). There were only two levels of the repeated measures

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factors, so there was no need to incorporate corrections for possible violations of the sphericity

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assumption (von Ende 2001).

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Results

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Germination density rates were significantly higher in the smokewater treated plots relative to those not

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treated at all sites, and also higher in the ‘off-site’ sites. There was a difference in germination density

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between the three management areas, with the mine-site having the lowest germination, followed by the

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forest site, and the ex-pine site having the highest germination. At the Tuart forest site the interaction

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between smokewater treatment and ‘on-site’ and ‘off-site’ plots was also significant, with the effect of

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smokewater more strongly marked off-site (Table 1, Figure 1).

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Weed species dominated in the seedling trays at all sites, particularly Annual Veldt Grass (recorded in 54

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% of trays and from all sites). Other weeds noted were Dune Onion Weed (7 %), which occurred in all

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sites and Arum Lily (2 %) only in the ‘off-forest’ trays. The only recorded native species emerging was

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the mid-canopy dominant, Peppermint (Agonis flexuosa) (5 %) and it was recorded in all trays except for

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ex-pine and ex-mine sites.

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Implications for management

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The use of plant-derived smoke has been utilized as a means of determining the potential soil seed bank in

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old-fields (Standish, et al. 2007), and forest communities (Read, et al. 2000). However, the current study

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is the first to analyse different management areas within one forest with the aim of informing land

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managers. The differences in germination rates (the majority of which were weeds) between the three

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management areas could be a reflection of their previous land use. For example, the ex-mine site had a

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lower weed load than the other sites, perhaps because of topsoil storage and mixing during the ripping

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process. The fact that very little native germination and a high weed load were detected at all sites is

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probably due to the high level of disturbance at all sites in this forest. Nonetheless, the results depict the

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potential abundance of weeds and some of the species that may challenge land managers.

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There are clearly limitations to this type of study. Firstly, it only tested seed germination response to a

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particular smoke concentration; different concentrations or smoke in addition to heat should be studied.

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Secondly, restoration strategies should not only be based on this aspect of the soil seedbank but take into

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account other factors, including prior land use and management. However, this use of smokewater can

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provide land managers with a somewhat greater level of understanding of the composition of the soil

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seedbank rather than based on land use history and recent condition.

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Given this new information, land managers could make more informed decisions about prioritization of

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management options, including project type, site selection, timeline of activities and cost-effectiveness.

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They may wish to include such information in a decision support system. For example, managers may

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wish to postpone treatment until financial resources can match the amount of weed management input

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required, or budget and plan for a particular suite of weed control methods, depending on the invasive

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species found in the soil seedbank. Furthermore, managers may wish to select a particular native species

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mix and planting/seeding density as part of revegetation activities to ensure that species with particular

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functional traits are chosen that provide a high competitive ability in the face of particular invasive

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species composition and abundance.

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Acknowledgements

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The authors thank the Department of Environment and Conservation, Bemax Cable Sands and the

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Botanic Gardens and Parks Authority. This research was undertaken as part of the Tuart Health Research

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Group, which is funded through an Australian Research Council Linkage Grant (LP0668195), and the

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Western Australian State Centre of Excellence for Climate Change, Woodland and Forest Health.

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References

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Bellairs S. M. and Bell D. T. (1993) Seed stores for restoration of species-rich shrubland

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vegetation following mining in Western Australia. Restoration Ecology

1

, 231-240.

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De Lange J. H. and Boucher C. (1990) Autecological studies on Audounia capitata (Bruniaceae).

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I. Plant-derived smoke as a seed germination cue. South African Journal of Botany

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,

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700-703.

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DEC (2007) Ludlow Tuart Forest Bemax Titanium minerals mine (216.0ha) forest stratification

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report. In: Department of Environment and Conservation, Perth, Western Australia.

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Dixon K. W., Roche S. and Pate J. S. (1995) The Promotive Effect of Smoke Derived from

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Burnt Native Vegetation on Seed-Germination of Western-Australian Plants. Oecologia

149

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, 185-192.

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Flematti G. R., Ghisalberti E. L., Dixon K. W. and Trengove R. D. (2004) A compound from

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smoke that promotes seed germination. Science

305

, 977-977.

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Marks P. L. and Mohler C. L. (1985) Succession after elimination of buried seeds from a

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recently plowed field. . Bulletin of the Torrey Botanical Club 112, 376-382.

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Read T. R., Bellairs S. M., Mulligan D. R. and Lamb D. (2000) Smoke and heat effects on soil

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seed bank germination for the re-establishment of a native forest community in New South

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Wales. Austral Ecology

25

, 48-57.

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Roche S., Koch J. M. and Dixon K. W. (1997) Smoke enhanced seed germination for mine

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rehabilitation in the southwest of Western Australia. Restoration Ecology

5

, 191-203.

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Rokich D. P., Dixon K. W., Sivasithamparam K. and Meney K. A. (2002) Smoke, mulch, and

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seed broadcasting effects on woodland restoration in Western Australia. Restoration

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Ecology

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, 185-194.

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Standish R. J., Cramer V. A., Wild S. L. and Hobbs R. J. (2007) Seed dispersal and recruitment

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limitation are barriers to native recolonization of old-fields in western Australia. Journal of

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Applied Ecology

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, 435-445.

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Thomson K., Band S. R. and Hodgson J. G. (1993) Seed size and shape predict persistence in

166

soil. Functional Ecology

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, 236-241.

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von Ende, C. N., 2001. Repeated measures analysis: growth and other time-dependent measures. Pp.

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116-157 in Design and analysis of ecological experiments ed by S. M. Scheiner and J. Gurevitch.

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Oxford University Press, Oxford.

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Table 1. Results of repeated measures ANOVA for each site (forest, minesite, and pine), with smoke

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water (present or absent) and location (on-site and off-site) as the repeated measures variables.

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Forest Minesite Pine

Effect df (effect,

error)

F

p-level

F

p-level

F

p-level

Location

(on- or off-site)

1, 24

21.335

> 0.001 20.447

> 0.001 0.362

0.553

Smoke water treatment

(present or absent)

1, 24

35.541

> 0.001 10.784

0.003

6.736

0.016

Location, Smoke water

treatment

1, 24

7.339

0.012

3.766

0.064

0.035

0.854

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0 5 10 15 20 25 30 35

Forest Off Forest Mine Off Mine Pine Off Pine Management areas Tot a l ge rm ina nt s

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Figure 1. Mean number of germinants per plot for three management areas and paired sites (“Off”-) in

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the Ludlow Tuart Forest, with (black bars) and without (white bars) the addition of smoke water. Values

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are means (+ SE) of 25 samples.

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References

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