19 Extension Note. Introduction

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1 19 Extension Note FEBRUARY 1998 Ectomycorrhizal Diversity of Paper Birch and Douglasfir Seedlings Grown in Single-species and Mixed Plots in the ICH Zone of Southern British Columbia Melanie D. Jones Daniel M. Durall S. Michelle Harniman Debbie C. Classen Department of Biology Okanagan University College 3333 College Way Kelowna BC V1V 1V7 and Suzanne W. Simard British Columbia Ministry of Forests Kamloops Forest Region 515 Columbia Street Kamloops BC V2C 2T7 Introduction There are a number of incentives to retain broadleaf tree species such as paper birch (Betula papyrifera Marsh.) following logging in the Interior Cedar Hemlock biogeoclimatic zone. For example the roots of paper birch are intimately associated with numerous species of ectomycorrhizal fungi. The retention in plantations of paper birch may therefore contribute to belowground biodiversity by hosting a variety of mycorrhizal fungi. Furthermore the retention of paper birch in young mixed plantations appears to reduce the incidence of Armillaria ostoyae among Douglasfir (Pseudotsuga menziesii (Mirb.) Franco) (Morrison et al. 1988; Simard 1997) and improve both the nitrogen and ph status of soils. Finally the presence of paper birch increases the diversity of tree species and contributes to the structural diversity of a stand (Simard and Vyse 1994) (Figure 1). Ectomycorrhizae are the nutrientand water-absorbing organs of most woody plants and as such mycorrhization of planted seedlings is thought to be critical for adequate growth and survival. However ectomycorrhizal fungi do not persist in the absence of a plant host and therefore the quantity and diversity of fungal inoculum often decreases after logging. Ectomycorrhizal species such as paper birch may act as reservoirs (refuge plants) for ectomycorrhizal fungi on a site thereby maintaining a diversity of fungi that may eventually associate with planted conifers. Interior Cedar Hemlock forests are often considered rich in tree species; however as with most temperate forests the diversity of plants FIGURE 1 Ectomycorrhizal diversity increases in mixed stands of paper birch and Douglas-fir. For full details see Jones et al Ministry of Forests Research Program

2 is small compared to the enormous diversity of soil organisms. For example there are an estimated 5 species of fungi capable of forming ectomycorrhizae. Douglasfir alone can associate with an estimated 2 species of ectomycorrhizal fungi and there can be as many as 1 different species in a typical Douglas-fir stand. Among the many species of ectomycorrhizal fungi considerable differences exist with respect to their physiology and ecology. Ectomycorrhizal fungi differ in their ability to take up various forms and types of nutrients in their tolerance to water stress and temperature extremes and in their resistance to pathogens. Therefore it is hypothesized that seedlings associated with a diverse array of ectomycorrhizal fungi will have an increased capability to fully exploit the heterogeneous soil through which the roots are growing. The objectives of this study were to investigate the effects of planting mixtures of paper birch and Douglas-fir at various densities and proportions on the diversity of ectomycorrhizal fungi. This study was carried out at three different sites over three years ( ) in the Interior Cedar Hemlock biogeoclimatic zone (ICH) of the southern Interior of British Columbia. Two of the three sites Adams Lake and Malakwa Lake are located in the ICHmw3 variant whereas the site at Hidden Lake is located in the ICHmw2 variant. All sites were harvested prior to 1988 and planted with Douglas-fir. In the fall of 1991 the sites were de-stumped to reduce the inoculum load of A. ostoyae and as a result the previously planted seedlings were also removed. Methods In June containerized paper birch and Douglas-fir were planted in single-species and mixed plots at various densities and proportions: Ep/16Fd 8Ep/8Fd 16Ep/Fd Ep/32Fd 32Ep/Fd 8Ep/4Fd and 32Ep/16Fd (numbers represent stems per hectare of paper birch [Ep] and Douglas-fir [Fd]). A 32Ep/16Fd mixed plot is shown in Figure 2. At each of the three sites one replicate plot of each treatment was established (seven treatments times three replicates = 21 plots). All plots were located between 1 and 4 metres from the adjacent forest. At 4 16 and 28 months following planting the roots were sampled and examined for mycorrhizal formation at 4X magnification. Ectomycorrhizal roots were categorized into morphological groups based on distinct macroscopic (Figure 3) and microscopic features. Simpson s diversity index was calculated to describe the ectomycorrhizal community associated with the seedlings at the various treatments. Diversity indices such as Simpson s incorporate the two components of diversity: richness (the number of species in a defined sampling unit) and evenness (the proportional abundance of these species). High evenness is equated with high diversity. FIGURE 2 Mixed plot at Adams Lake one year after planting. Results Ectomycorrhizal Community The mycorrhizal community associated with Douglas-fir was characterized by the presence of few dominant mycorrhizae (Figure 4). The dominant mycorrhizae included Thelephora-like E-strain I and II Cenococcum-like Mycelium radicis atrovirens (MRA) and Rhizopogonlike morphotypes. Twenty-eight months following planting five of the six dominant types of ectomycorrhizae encountered were common to both Douglas-fir and paper birch. Of the dominant types encountered only Rhizopogon was restricted to one host (Douglas-fir). Overall 91% of mycorrhizal paper birch roots were colonized by fungi that were also associated with Douglas-fir. Fifty-six percent of mycorrhizal Douglas-fir roots were formed by fungi that also colonized paper birch. The number of ectomycorrhizal morphotypes associated with paper birch increased over the three years of study. In 1992 nine types were identified whereas 2 types were identified in 1993 and 26 types in 1994 (Figure 5). A similar increase in the number of fungal associates was FIGURE 3 Thelephora-like mycorrhizae on paper birch. 2

3 Malakwa 92 Hidden Lake 92 Adams Lake 92 Malakwa 93 Hidden Lake 93 Adams Lake 93 Malakwa 94 Hidden Lake 94 Adams Lake Percent of root tips identified for the Douglas-fir seedlings. Nine types were encountered in types in 1993 and 32 types in Ectomycorrhizal Diversity Interplanting Douglas-fir with paper birch resulted in an increase in the evenness component of the diversity Thelephora E-strain I E-strain II Cenococcum MRA Rhizopogon FIGURE 4 The proportions of the six dominant mycorrhizal types on roots of planted Douglas-fir sampled in and of ectomycorrhizae associated with Douglas-fir seedlings (Figure 6) but it did not affect the number of mycorrhizal types per seedling. The diversity of ectomycorrhizae associated with birch remained high in both single-species and mixed plots. Planting density did not affect either component of diversity. Interpretation Ectomycorrhizal Diversity Sixteen and 28 months after planting interplanting Douglas-fir with paper birch increased the evenness of ectomycorrhizal types on Douglas-fir compared with planting Douglas-fir in pure stands. This mixture effect resulted from an increase in the abundance of subdominant mycorrhizae such as the Cenococcum-like and MRA morphotypes on Douglas-fir. In contrast Douglas-fir seedlings grown in pure stands were colonized predominantly by one ectomycorrhizal type such as the Thelephora-like or Rhizopogon-like morphotypes. Seedlings associated with a diverse array of ectomycorrhizal fungi are thought to benefit from the range of functions that the different fungi perform. Therefore planting in mixtures can be regarded as contributing both to seedling success and biodiversity. There are a number of factors that may have resulted in the increased evenness of ectomycorrhizal fungi associated with Douglasfir seedlings. First ectomycorrhizal Douglas-fir Paper birch Douglas-fir Number of mycorrhizal morphotypes Paper birch Evenness Evenness FIGURE 5 The number of mycorrhizal types encountered on the roots of Douglas-fir and paper birch 4 (1992) 16 (1993) and 28 (1994) months after planting. FIGURE 6 Evenness per seedling for the mycorrhizal communities of paper birch and Douglas-fir planted in mixed (dark bars) or single-species (stippled bars) plots for 4 (1992) 16 (1993) and 28 (1994) months. Evenness was higher for Douglas-fir when grown in mixture with paper birch than when grown alone in 1993 (P =.7) and 1994 (P =.9). 3

4 fungi vary in their level of host specificity. It is possible that certain fungi that are commonly restricted to one host (e.g. paper birch) become capable of colonizing a second host (e.g. Douglas-fir) when the mycelium is already established with its primary host. An additional or alternative factor contributing to an increase in evenness for Douglasfir may be due to chemical and/or biological changes in the soil caused by paper birch. Paper birch improves the nitrogen and ph status of soils and hosts a unique microflora including numerous species of bacteria. Many such rhizosphere bacteria have recently been identified as mycorrhization helper bacteria which can promote ectomycorrhizal colonization (Garbaye 1994). Some bacteria may also play an important role in the uptake of certain nutrients. For example certain strains of Bacillus sp. associated with tuberculate ectomycorrhizae can fix nitrogen thereby influencing nitrogen dynamics in the soil (Li et al. 1992). Shared Ectomycorrhizal Fungi and Nutrient Transfer The finding that paper birch and Douglas-fir share ectomycorrhizal fungi indicates that they can be linked by a common mycelium and that carbon nutrients or water could transfer between the two species. In a separate field study carried out at Adams Lake Simard et al. (1997) showed that 7% of fixed carbon was transferred between paper birch and Douglas-fir through their mycorrhizal linkages. There was a 6% net carbon gain by Douglas-fir which was increased with shading. It is speculated that interspecific hyphal linkages and nutrient transfer can reduce competition between these two species in the field. Reduced competition may allow the persistence of Douglas-fir in otherwise inhospitable environments thereby increasing plant species diversity. Other studies have shown that biodiversity contributes to productivity and resilience of ecosystems (Tilman et al. 1996). Ecology of Pioneer Fungi Succession occurs in ectomycorrhizal fungal communities much the same as it does in plant communities. Early-stage or pioneer ectomycorrhizal fungi are characterized by rapid dispersal and the ability to colonize from spores or fragments of mycelium. As such these fungi are commonly observed associated with naturally regenerating or planted seedlings in newly disturbed areas. Examples of early-stage fungi include all of the dominant fungi encountered in this study: Thelephora-like Cenococcum-like MRA Rhizopogonlike and E-strain morphotypes. It is interesting to note the presence of Thelephora-like and MRA mycorrhizae (common greenhouse contaminants) associated with planted seedlings. Our study showed that Thelephora and MRA inoculated seedlings after they were outplanted not in the greenhouse suggesting that large effective sources of inoculum for these fungi exist in clearcuts. In contrast to early-stage mycorrhizal fungi late-stage fungi are characterized by their inability to colonize from spores and a requirement for an intact mycelial connection with a live host for colonization. Therefore in the absence of refuge plants the inoculum available to seedlings will be restricted to earlystage fungi capable of colonizing from spores or fragments of mycelia. Examples of late-stage fungi include Lactarius and Russula which occurred infrequently (1 4% of mycorrhizal roots) over the three years of this study. Despite their infrequent occurrence there was apparently a viable source of inoculum for Lactarius and Russula present in the clearcuts. It is possible that other broadleaves present on site such as trembling aspen or black cottonwood were colonized by these fungi and that the extending mycelium provided a viable inoculum source for the planted seedlings. Interestingly under greenhouse conditions these fungi did form ectomycorrhizae with seedlings; this indicates that in some cases colonization can occur from isolated fragments of inoculum. Conclusions and Management Implications 1. This study shows that interplanting Douglas-fir with paper birch will increase below-ground biodiversity by increasing the evenness component of ectomycorrhizal diversity. 2. The observation that five of the six dominant types of ectomycorrhizae were common to both Douglas-fir and paper birch indicates the potential for carbon transfer between the two species via a common mycelium. A shared mycorrhizal mycelium is speculated to reduce competition between the two species encourage co-existence and therefore favour plant species diversity. 3. The retention of broadleaves such as paper birch will help to achieve the biodiversity objectives outlined in the Forest Practices Code by increasing tree species diversity structural diversity and the diversity below-ground. Text by Shannon Hagerman Soil Biology Consultant Bond Road Winfield B.C. V4V 1J5 4

5 Acknowledgements Thanks to several anonymous reviewers who provided technical review of the primary journal article by Jones et al and Kerry McLean who provided technical assistance. Funding for the research was provided by the Hardwood Management Subprogram of the Canada-British Columbia Partnership Agreement on Forest Resource Development ( ). Funding for this extension project was provided by Forest Renewal B.C. Literature Cited Garbaye J Tansley review no. 76. Helper bacteria: a new dimension to the mycorrhizal symbiosis. The New Phytologist 128: Jones M.D. D.M. Durall S.M.K. Harniman D.C. Classen and S.W. Simard Ectomycorrhizal diversity on Betula papyrifera and Pseudotsuga menziesii seedlings grown in the greenhouse or outplanted in single-species and mixed plots in southern British Columbia. Canadian Journal of Forest Research 27: Li C.Y. H.B. Massicotte and L.V.H. Moore Nitrogen-fixing Bacillus sp. associated with Douglas-fir tuberculate ectomycorrhizae. Plant and Soil 14: Morrison D. G.W. Wallis and L.C. Weir Control of Armillaria and Phellinus root diseases: 2- year results from Skimikin stump removal experiment. Canadian Forest Service Pacific Forestry Centre. Inf. Rep. BC-X-32. Simard S.W Douglas-fir in northern ecosystems: interactions with paper birch. In Ecology and management of Douglas-fir at the northern limits of its range. Workshop Proceedings September 3 and October Fort St. James B.C. J.D. Lousier and W. Kessler (editors). University of Northern British Columbia Press Prince George B.C. In press. Simard S.W. D.A. Perry M.D. Jones D.D. Myrold D.M. Durall and R. Molina Net transfer of carbon between tree species with shared ectomycorrhizal fungi. Nature 388: Simard S.W. and A. Vyse Paper birch: weed or crop tree in the Interior Cedar Hemlock forest of south British Columbia. In Interior Cedar Hemlock White Pine Forests: Ecology and Management. Symposium Proceedings March Spokane Wash. D.M. Baumgartner J.E. Lotan and J.R. Tonn (editors). Department of Natural Resource Sciences Washington State University Pullman Wash. Tilman D. D. Wedin and J. Knops Productivity and sustainability influence biodiversity in grassland ecosystems. Nature 379:

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