Fungi and Fungus-Like Organisms in a Temperate Deciduous Forest Canopy
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Fungi and Fungus-like Organisms in a Temperate Deciduous Forest Canopy Der Fakult¨at fur¨ Biowissenschaften, Pharmazie und Psychologie der Universit¨at Leipzig eingereichte KUMULATIVE DISSERTATION zur Erlangung des akademischen Grades Doctor rerum naturalium (Dr. rer. nat.) vorgelegt von Diplom Biologe, UNTERSEHER, Martin geboren am 05. Mai 1975 in Malsch, Karlsruhe Preface and acknowledgements This dissertation is of cumulative type and consists of five papers dealing with canopy research at the Leipzig Canopy Crane (LAK) research facility. The papers have been published in or accepted by both national and international journals. Two contributions from newsletters are also included. Four of the five peer-reviewed papers and one short article deal with the investigation of diversity and ecology of wood decay fungi and fungus-like organisms. The remaining two contributions about tree frogs (Hyla arborea) on trees in up to 30 metres in height exemplify the importance of incidental observations during the fieldwork. As crypto- gams are not considered in the articles, their contents will not be discussed further. The papers are listed in consequential order in the second part of the thesis. The overview of this PhD thesis consists of a review of the publications. A compact description of the methods applied in the field of the Leipzig crane site and in the la- boratory is given as well as a spanning discussion of the main results of this work. Studying treetops of the world’s forests and the life inside is a comparatively new area of environmental science. As mycology is still linked poorly with canopy research, a brief summary of past and present scientific activities in forest canopies follows this preface, hoping that it will help mycologists and other groups interested in environmental science getting familiar with foggers, climbing gear, and canopy cranes. To reduce the gap of scientific knowledge concerning fungi in tree crowns, the studies within this PhD thesis have been designed as a starting point for the first long-term investigations to assess the diversity and ecology of fungi and fungal organisms in the canopy of a floodplain forest. Studies about leaf-inhabiting endophytic fungi are current- ly performed at the Leipzig crane site and a molecular characterisation of lignicolous fungi is planned for the future. The aims of the present work were (i) to create a species list of lignicolous fungi, slime molds and myxomycete-like organisms occuring in the canopy, (ii) to analyse beta diversity, and (iii) to critically comment on the influence of ecological factors on the species composition of these organisms in tree crowns. With respect to research about wood decay fungi, all field and laboratory work (e.g. sampling, microscopy, identification of fungi) as well as data interpretation, statistics, and paper writing have been done by myself with initial help from colleagues and my supervisors Prof. Dr. W. Morawetz and Dr. P. Otto. Ophir Tal was entirely responsible for gathering, processing, and interpreting meteo- rological and small-scale climatic data. The study of slime molds and myxomycete-like organisms was done in cooperation with Prof. Dr. M. Schnittler from the Botanical Institute at the Ernst Moritz Arndt University in Greifswald, Germany. I was responsible for the field work and for applying multivariate analyses to the data. Cultivation and identification of the organisms, addi- tional statistical evaluations, and most of the paper writing was conducted in Greifswald. As this work will be published solely under my name, it is my duty and pleasure to mention those persons and institutions who contributed most to the successful comple- tion of my PhD thesis. Wilfried Morawetz, head of the Department of Systematic Botany and Botanical Gar- den, was the person who initiated the LAK Project in 2001 and agreed to launch my- cological studies one year later. From the beginning on I have been fully integrated into his ‘canopy team’. As the Leipzig Canopy Crane is an object of high public interest, he mentioned my work in many interviews, and seeing myself regularly in newspapers and even in national telecasts greatly helped maintaining my motivation when times were hard and difficult. I am grateful to his lasting promotion of this work. Peter Otto also agreed to be a supervisor of my doctoral thesis. He formulated the first proposal for this topic which has been a sound fundament to start working with canopy fungi. Peter J. Horchler, the former coordinator of the LAK Project, played an important role as he introduced me to multivariate statistics and provided many informations, thoughts, and data of other studies within the crane project. Until he left the LAK it was a pleasure to share the workroom with him. Without naming every single colleague and person of the department, I can say that I enjoyed an open-hearted, friendly, and very helpful atmosphere every day during my PhD thesis. Financial support for my studies has been kindly provided through projects UFZ- 16/2000 and UFZ-04/2004 from the UFZ Centre for Environmental Research Leipzig - Halle in the Helmholtz-Association (UFZ) and from the Friends of the Botanical Garden Leipzig. I am deeply indebted to my girlfriend, my longtime companion, and mother of our son Jacob, Carmen Schmitt. With her patience and thoughtfulness for the many nights and leisure-times that I spent for my research, she accounted like no other person for my work. I also thank my parents for bringing me up the way they did and for providing everlasting support. Table of contents Preface and acknowledgements . i A short introduction of canopy research . 1 1 Overview of the PhD thesis 10 2 Articles 24 2.1 Studies of the diversity of lignicolous fungi in the canopy of a floodplain forest in Leipzig, Saxony . 25 2.2 Species richness and substrate specificity of lignicolous fungi in the canopy of a temperate, mixed deciduous forest . 34 2.3 Influence of small-scale conditions on the diversity of wood decay fungi in a temperate, mixed deciduous forest canopy . 51 2.4 Species richness and ecological characterization of myxomycetes and myx- omycete-like organisms in the canopy of a temperate deciduous forest . 66 2.5 Diversity of lignicolous fungi in the canopy of a deciduous forest in Leipzig, Germany . 80 2.6 Hyla, high! . 84 2.7 High above – sitting sites of the tree frog (Hyla arborea L.) in tree crowns of the Leipzig floodplain forest . 87 3 Summary 90 4 Curriculum vitae 96 5 Erkl¨arung (Declaration, in German only) 99 iii A short introduction of canopy research A short introduction of canopy research As there exist a number of comprehensive synopses of canopy research (Lowman & Nadkarni 1995; Stork, Adis & Didham 1997; Linsenmair et al. 2001; Mitchell, Secoy & Jackson 2002; Basset et al. 2003; and Lowman & Rinker 2004), as well as free online material (http://www.stri.org/english/research/facilities/ terrestrial/cranes/canopy crane network.php) the following introduction to ca- nopy science will be comparatively short. Although research in the treetops is of great public interest with impressiv documentaries in television (e.g. http://www.zdf.de/ ZDFde/inhalt/19/0,1872,2150003,00.html or http://www.bbc.co.uk/sn/tvradio/ programmes/horizon/madagascar.shtml) and full of emotional sensations with many books and articles broaching the issue of canopy science in a more popular way (e.g. Mitchell 1989; Lowman 1999; Halle´ 2001; Nadkarni 2004; and http://www. heraldtribune.com/apps/pbcs.dll/article?AID=/20050731/COLUMNIST18/507310 439), I will restrict the following section mostly to scientific facts. It is nearly impossible to define an exact starting point for canopy research because the hidden life in the treetops attracted naturalists such as Alexander von Humboldt for a long time. One of the first successful attempts to explore the rainforest roof was made in 1929 during an Oxford University expedition to Moraballi Creek in British Guiana. With an enormous assortment of military equipment such as rocket-firing ma- chines, line-throwing guns or rope ladders, and the help of many natives, the explorers managed to reach several crowns without felling the trees and to install simple, tempo- rary platforms (Mitchell 1989, pp. 23-25). Apart from this rare example, serious investigations of forest canopies started in the second half of the 20th century not until new methods of reaching the treetops were successfully established. With permanent platforms or towers it was then possible to conduct longer or even permanent studies in tree crowns. The most famous canopy project at that time were certainly the studies on the steel tower in Mpanga Forest in Uganda. Built by the East African Virus Research Institute, it followed the trends of studying mosquitoes and other tropical biting insects since the secret of yellow fever cycle was disclosed in the mid 20th century by Jorge Boshell and others (e.g. Bugher et al. 1944; Galindo & Trapido 1955). The studies of Corbet, Haddow and others (Corbet; Haddow & Corbet; Dirmhirn, all 1961) revolutionized the perception of ecological processes and abiotic parameters high above the forest floor. Almost two decades later canopy scientists used further technologies adapted to bio- logical research in tropical forests and launched the next generation of canopy science. One certainly could cite the initial works of Perry (1978), Erwin & Scott (1980) and Erwin (1982) as milestones in canopy research. For the first time, Donald Perry used climbing gear that was slightly modified from alpine techniques to meet the demands page 1 A short introduction of canopy research of tropical forest canopies. He and his colleagues built cobweb-like nettings high above the ground on which they managed to move inside the canopy up and down, and from treetop to treetop.