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Is Kanuka and Manuka Establishment in Grassland Constrained by Mycorrhizal Abundance?

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Is Kanuka and Manuka Establishment in Grassland Constrained by Mycorrhizal Abundance? 172 AvailableNew on-lineZealand at: Journal http://www.newzealandecology.org/nzje/ of Ecology, Vol. 37, No. 2, 2013 Is kanuka and manuka establishment in grassland constrained by mycorrhizal abundance? Murray Davis1*, Ian A. Dickie2, Thomas Paul3 and Fiona Carswell2 1Scion, PO Box 29237, Christchurch 8540, New Zealand 2Landcare Research, PO Box 69040, Lincoln 7640, New Zealand 3Scion, Private Bag 3020, Rotorua 3046, New Zealand *Author for correspondence ([email protected]) Published online: 14 May 2013 Abstract: Two indigenous small tree and shrub species, kanuka (Kunzea ericoides) and manuka (Leptospermum scoparium), have potential as reforestation species in New Zealand as they are forest pioneer species that can invade grassland naturally from present seed sources. The aim of this study was to determine if establishment of kanuka and manuka from seed in grassland distant from stands of these species might be constrained by lack of appropriate mycorrhizal fungi. Both species were grown in an unsterilised grassland soil from a low- productivity montane site assumed to be devoid of appropriate mycorrhizal fungi and inoculated with sterilised or unsterilised O-horizon or mineral soil from beneath three kanuka and three manuka communities expected to contain such fungi. Inoculation with unsterilised O-horizon soil improved kanuka biomass by 36–92%, depending on the source of the inoculant. Inoculation did not improve manuka biomass. No ectomycorrhizal infection was observed on either kanuka or manuka in samples examined under binocular microscope. The biomass response by kanuka to inoculation may be due to introduction of more effective arbuscular mycorrhizal fungi from kanuka communities or possibly to the introduction of soil microorganisms. Testing of inoculation under field conditions will be essential to determine whether establishment of either species in grassland soil by seeding is seriously constrained by lack of appropriate mycorrhizal fungi or soil microorganisms. Keywords: arbuscular mycorrhiza; ectomycorrhiza; Leptospermum; reforestation; seedling establishment; soil inoculation Introduction – arbuscular mycorrhizal fungi (AMF) or ectomycorrhizal fungi (EMF). Arbuscular mycorrhizal fungi produce arbuscles, Indigenous forest once covered about 75% of New Zealand hyphae and vesicles within root cortex cells whereas EMF (Newsome 1987), but much of the forest has been removed form a mantle around roots and a Hartig net between root cells and the land converted to agricultural use, particularly to (Brundrett et al. 1996). The mycorrhizas also differ functionally, grassland for pastoral farming. Increasingly, there is interest with EMF generally having a greater ability to obtain nitrogen in re-establishing indigenous forests for carbon sequestration and phosphorus from organic polymers than AMF (Read & or other environmental benefit on land that is marginal for Perez-Moreno 2003). Kānuka and mānuka are unusual among pastoral use because of isolation, steepness or low productivity. New Zealand’s flora in that they are colonised by both AMF Two indigenous small tree and shrub species, kānuka (Kunzea and EMF (Orlovich & Cairney 2004; McKenzie et al. 2006). ericoides (A.Rich.) Joy Thomps.) and mānuka (Leptospermum In agricultural and forestry systems, plant performance scoparium J.R.Forst. & G. Forst.), have potential as is frequently constrained by lack of appropriate mycorrhizal reforestation species as they are forest pioneer species that can fungi (Smith & Read 1997; Lekberg & Koide 2005). The extent invade grassland naturally from present seed sources (Wardle to which the invasion of kānuka and mānuka into grassland 1991). For land areas that lack adjacent seed sources, these communities is constrained by abundance of appropriate species can be established by planting and possibly by seeding mycorrhizal fungi is unknown. Grassland communities are (Stevenson & Smale 2005; Dodd & Power 2007; Ledgard et al. dominated by species infected by AMF and it is possible 2008), which is potentially a lower cost option than planting. that some of these fungi will also infect kānuka and mānuka Once established, kānuka or mānuka should facilitate forest and facilitate the establishment of these species. Growth and succession by providing ectomycorrhizal inoculum (Dickie competitiveness of kānuka and mānuka may be improved, et al. 2012) or ameliorating microsite conditions (Wardle however, by introduction of AMF from kānuka or mānuka 1970; Davis et al. 2013). communities as has been shown for Podocarpus cunninghamii Successful restoration of indigenous vegetation may in New Zealand (Williams et al. 2011). This would also be the require restoration of components of the microbial community case for EMF, which are unlikely to be present in grassland (Williams et al. 2011). The root systems of all indigenous communities (Dickie et al. 2012). To test whether kānuka and forest tree and shrub species are infected by mycorrhizal mānuka establishment in grassland soil may be constrained fungi that form symbioses with the host plant and play an by lack of appropriate mycorrhizal fungi, both species were essential role in the nutrition and water uptake of the host. grown in kānuka and mānuka free grassland soil inoculated Many of New Zealand’s native woody flora form mycorrhizas or not inoculated with O-horizon and mineral soil collected exclusively with one or other of two types of mycorrhizal fungi from beneath extant stands of the respective species. We New Zealand Journal of Ecology (2013) 37(2): 172-177 © New Zealand Ecological Society. Davis et al.: Establishment of kānuka and mānuka 173 hypothesised that, if establishment of kānuka and mānuka treatments, namely site of inoculant collection, inoculant type in grassland soil was constrained by lack of appropriate (O-horizon or mineral soil), and sterilisation (unsterilised mycorrhizal fungi, inoculation with soil from extant stands or sterilised), being applied in factorial combination. An would provide functionally appropriate fungal species and uninoculated control treatment, which had 5 mm of grassland improve growth of the host species. soil in place of an inoculant layer, was included. The two trials were conducted simultaneously but separately in the same glasshouse on adjacent benches. Methods Seeds of each species were germinated on sterilised seed testing paper in Petri dishes and seedlings were transplanted Two soil inoculation trials, one with kānuka and the other with into pots (four per pot) when they were large enough (c. 5 mm) mānuka, were conducted simultaneously in a greenhouse. to handle. Kānuka seedlings were thinned to two per pot after A soil mapped as Acheron stony loam (Soil Bureau 1968), 8 weeks’ growth. The trials were planted in mid-winter (June) collected from kānuka- and mānuka-free grassland near Mt and harvested after 5 months. Watering was by an automated Barker (Rakaia Valley, Canterbury) was used as the growth overhead misting system. The pots were re-randomised twice medium for both trials. No attempt was made to remove during the trial period. Prior to harvest the pots were soaked in the existing bacteria and fungi by sterilisation. The nearest water for 1–2 h and soil was gently freed from the root systems kānuka and mānuka stands were 300 m and more than 3000 m under water. After cleaning, root systems were removed from distant respectively from the site where the soil was collected. shoots and stored moist to allow examination for mycorrhizas. o Vegetation at the site was of low productivity and dominated Roots and shoots were oven-dried at 65 C and weighed. by the native tussock-grass Festuca novae-zelandiae (Hack.) All roots of the first replicate pot of each treatment Cockayne and the adventive herb Pilosella officinarum Vaill. harvested were examined for presence of EMF infection Analysis showed the soil to be moderately acidic (pH 5.3), have by counting a minimum of 200 fine root tips and recording medium carbon (6.4%) and nitrogen (0.47%) levels and C:N whether or not each tip was ectomycorrhizal. For most ratio (13.6), and a low level of bicarbonate (Olsen) extractable seedlings, 50% of the root system was examined by displaying phosphorus (6.9 mg kg–1). Exchangeable cation levels were 1.9, roots over a clear plastic grid and examining alternate rows 6.4 and 0.48 cmol+ kg–1 for potassium, calcium and magnesium under a stereo microscope (4× to 50×). Where there was any respectively. The top 5 cm of mineral soil was removed and doubt, root tips were mounted on glass slides and examined with discarded before the underlying soil was collected, to reduce compound microscopy to confirm mycorrhizal assessments the potential of inclusion of ectomycorrhizal fungal spores that (presence of mantle, Hartig net, and/or extraradical hyphae; 40× might infect kānuka or mānuka. The soil was potted to a depth to 1000×). It was assumed the plants would develop arbuscular of 75 mm, in pots 80 × 80 × 100 mm deep, and covered by a mycorrhizas from propagules present in the grassland soil, 5-mm-deep layer of inoculum (see below), which was in turn hence no attempt was made to quantify AMF infection. After covered by a further 10-mm layer of grassland soil to reduce examination, roots were oven-dried and weighed as above. the potential of cross contamination of inocula between pots. The biomass response of kānuka and mānuka plants was Two types of soil inoculum were collected from beneath analysed using SAS Version 9.2. Normality and homogeneity of
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