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C=IE, St=Warszawa, O=Ditorpolish Bot, Ou=St CONTENTS 1. Introduction ............................................................................................................................................................... 5 1.1. Fungal succession and old-fields ....................................................................................................................... 5 1.2. Scheme of spontaneous secondary succession on abandoned fields near the Białowieża old-growth forest ....................................................................................................................................................... 8 2. Subject and aims of study ....................................................................................................................................... 12 3. Study area ................................................................................................................................................................ 13 4. Material and methods ............................................................................................................................................. 15 5. Results ...................................................................................................................................................................... 25 5.1. Fungi in permanent plots ................................................................................................................................ 25 5.2. Changes in species and sporocarp distribution along successional gradient .............................................. 34 5.2.1. Species richness and diversity ............................................................................................................... 34 5.2.2. Species abundance patterns .................................................................................................................. 39 5.2.3. Spatial frequency patterns .................................................................................................................... 53 5.2.4. Ectomycorrhizal carpophore production ............................................................................................ 53 5.3. Similarity structure of macrofungi .................................................................................................................. 56 5.4. Quantitative changes in ectomycorrhizal roots along successional gradient ............................................. 60 6. Discussion ................................................................................................................................................................ 61 6.1. Macrofungi in the scheme of spontaneous secondary succession on the abandoned fields near Białowieża ................................................................................................................................................ 62 6.2. Patterns of fungal succession in post-agricultural land ................................................................................ 66 7. Summary of results and conclusions ...................................................................................................................... 81 8. References ............................................................................................................................................................... 85 Appendix 1. Soil analysis in the permanent plots ..................................................................................................... 95 Appendix 2. Vegetation description in the permanent plots ................................................................................. 101 Appendix 3. Macrofungi in the plots representing subsequent phases of vegetation succession in the old-fields near the Białowieża old-growth forest .................................................................. 115 Appendix 4. Results of k-mean analysis of fungi .................................................................................................... 141 Appendix 5. List of Basidiomycota in the old-field successional series near the Białowieża old-growth forest (Jelonka reserve and Czechy Orlańskie reserve) with notes on distribution and substrates ...................................................................................................................................... 145 ABStract Izabela Ka ł u c K a . Macrofungi in the secondary succession on the abandoned farmland near the Białowieża old- growth forest. Monogr. Bot., Vol. 99, 155 pp., 2009. the paper deals with the succession of basidiomycete macrofungi on abandoned farmland. The research was carried out in the unique complex of old fields of different age located on the southwest edge of the Białowieża old-growth forest (NE Poland). These lands undergo spontaneous secondary succession leading to formation of a continental fresh pine forest. Regular observations of basidiomata in 16 permanent plots (11 plots of 1000 m2, 5 plots of 400 m2) representing initial, optimal and terminal stage of vegetation development from psammophil- ous grassland to a pine forest with 100-year-old tree stand were conducted for three years. Soil and vegetation analyses were also carried out as well as quantitative examination of ectomycorrhizae in root samples. the series yielded 300 species of macromycetes (including 6 collective taxa) found in permanent plots plus 18 species outside the plots. The species composition, spatial distribution of sporocarps, and quantitative struc- ture of fungal communities changed along the successional gradient. Increasing diversity of fungi resulted from persistence of species typical of open areas and species associated with younger trees, as well as accumulation of species associated with older trees and more developed stands. Some species typical of younger phases were re- placed with species arriving later in the chronosequence. Species usually occurred long before and after attaining maximum abundance and frequency. Species associated with certain periods of vegetation development could be distinguished, as well as species accompanying the phytocoenoses regardless of their developmental stage. Patterns in structural changes of the fungal community, e.g., in number of species, abundance, frequency and production of carpophores, could also be recognised. They were different in ectomycorrhizal and in saprotrophic fungi. The occurrence of the former was influenced mainly by the presence of symbiotic trees and their age, stand structure and soil properties, while occurrence of the latter was affected by microclimatic factors and by diversity of accessible substrates. Changes in fungal communities were consistent with vegetation changes, but were also specific to this group of organisms. Key words: old-field succession, fungal succession, ectomycorrhizal fungi, saprotrophic fungi, sporocarp production, NE Poland 1. INTRODUCTION the influence of fungi on the formation and functioning of ecosystems is more complex and diverse than usually appreciated. Fungi take part in all major ecological processes, either in direct or indirect ways, and the existence of a great variety of other organisms and whole ecosystems depends on their activity. As decomposers, they drive the global carbon cycle by preventing an excessive accumulation of organic matter and its breakdown into the compounds which can be utilized by other organisms. As mycorrhizal partners fungi enhance plants water uptake and mineral nutrition, they supply regulatory substances and protect against pathogens and environmental stress. They interconnect individual plants via common mycorrhizal networks and build extensive sink-source relationships affecting the process of primary production as well as the composition and dynamics of vegeta- tion. Fungal endophytes inhabiting tissues of most plants can considerably influence their condition and alter adaptation capabilities. As lichen compounds, fungi contribute to the colonization of most pioneer habitats on earth and they provide a shelter for the important nitrogen-fixing organisms – cyanobacteria. As parasites they strongly influence popula- tions of their hosts and act as biological control. Also, fungi serve as a habitat or food for different organisms including bacteria, other fungi, nematodes, insects and mammals (Ra y n e R 1992; TR a p p e & Lu o m a 1992; Di g h T o n 2003). According to RAYNE R (1992) In short, fungi demand the attention of anyone eager to understand the complex interactions and feedback governing life processes in the biosphere. 1.1. Fungal succession and old-fields One of the key life processes is succession. It is a process of initiation and development of a community (and whole biocoenosis) within the confines of a potential biochore and relatively uniform environmental conditions (Fa L i ń s K i 1998). Succession of both plants and fungi can be defined as a directional change in the composition, relative abundance and spatial pattern of species comprising communities (FR ANKLAND 1992, 1998). In strict mycological sense it is a sequential occupation of the same site by thalli (normally myc- elia), either of different fungi, or of different associations of fungi (Ra y n e R & To dd 1979). The term fungal community is frequently used in this context meaning assemblage of fungi living together in the same time and space (Be g o n et al. 1996) or occurring within a certain uniform habitat (aR n o ld s 1992a). two different types of fungal succession can be distinguished depending
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