Cryptogams as indicator organisms in ecology and conservation biology DISSERTATION ZUR ERLANGUNG DES DOKTORGRADES DER NATURWISSENSCHAFTEN (DR. RER. NAT.) DER FAKULTÄT FÜR BIOLOGIE UND VORKLINISCHE MEDIZIN DER UNIVERSITÄT REGENSBURG vorgelegt von Josef Simmel aus Mitterfels im Jahr 2016 Das Promotionsgesuch wurde eingereicht am: 28.07.2016 Die Arbeit wurde angeleitet von: Prof. Dr. Peter Poschlod Declaration of manuscripts The below-mentioned chapter of the present PhD thesis is largely identical with a manuscript already submitted for publication. All other chapters either have not yet been submitted or are not intended for separate publication. *Chapter 2 was published in FUNGAL ECOLOGY with the thesis’ author being the main author: Simmel, J. , Bässler, C., Poschlod, P.: Ellenberg indicator values for macromycetes – a methodological approach and first applications. Fungal Ecology 26 (2017). DOI: 10.1016/j.funeco.2016.09.004 Table of contents CONTENTS PAGE Summary ……………………………………………………………….. ii Acknowledgements ……………………………………………………………….. … v Motto ……………………………………………………………….. vi Chapter 1 – General introduction ………………………………………………… 1 Chapter 2 – Ellenberg indicator values for macromycetes – a methodological approach and first applications ...……………………………………... 6 Chapter 3 – Lifestyle and threat of macromycetes – are there any functional traits 25 correlated with? ..…....………………………………………………… Chapter 4 – Does the cryptogam vegetation of calcareous grasslands reflect land- use history? ….………………………………………………………... 37 Chapt er 5 – The cryptogam vegetation of grasslands after 37 years of management and succession – bryophytes, lichens, and macromycetes of the “grassland management project” of Baden-Wuerttemberg …………... 50 Chapter 6 – Shifts in species composition of macromycete assemblages in the nature reserve “Sippenauer Moor” – an assessment using species identities, Ellenberg indicator values, and functional traits ….…...…... 68 Chapter 7 – Conclusions and perspectives …….……………………….……...…. 79 References ………………………………………………………....…….. 83 Table Appendix ……………………………………………………………….. 103 i Summary Summary The present thesis was designed to compare the influence of land-use effects on three groups of cryptogam species, namely the bryophytes, lichens, and macromycetes. Further topics were to newly develop an indicator value system for macromycetes and to compile a broad overview of functional traits applicable to macromycetes. These research topics are intended to (further) stimulate ecological research using macromycetes. In Chapter 2, I present a methodology for Ellenberg indicator values (EIVs) for macromycetes, including a new indicator value scale, i.e., the substrate openness O. Furthermore, the hemeroby concept is incorporated for the first time in the system of indicator values. Based on the newly developed methodology, I compiled EIVs for 10 parameter scales for a set of nearly 640 macromycete species. To test the applicability of EIVs for this species group, I analysed the data set by dividing the species by Red List classes or by lifestyle groups. The EIVs light intensity, substrate nutrient availability, substrate openness, and hemeroby related to these two classifications significantly differed. Critically endangered species on average have distinctly higher demands regarding light and substrate openness than not or less strongly threatened ones, which in turn are more tolerant to human impact and have higher demands in nutrient availability. Mycorrhizal species on average have higher demands on substrate openness and are less tolerant to high nutrient levels than saprobiotic or parasitic species. This pattern clearly highlights the points of threat for many macromycete species. Chapter 3 persued a similar approach like Chapter 2. Using the same macromycete species set, I compiled data for 31 functional traits that cover a broad range of features of, e.g., fruit body morphology, hymenial structure, spore morphology, and propagule dispersal. In a comparative way, Red List classification and lifestyle groups were used to analyse the species set and to get an insight which traits may be connected with the ecology or endangerment of species. Red List classification accounted for significant differences in three traits, and the lifestyle types in 28 traits. I describe the differentiations and discuss them against the background of ecological and morphological research. Chapter 4 was designed to compare the influence of land-use history on vascular plants and cryptogams. I compared the results of previous studies on the vascular plant cover of ancient and recent sites of dry calcareous grasslands to the cryptogam vegetation of these sites. I also ii Summary applied Ellenberg indicator values and the indicator species concept. Species numbers and Ellenberg indicator values were quite similar in ancient and recent grasslands. Nevertheless, we could identify indicator species for both grassland types, with Cladonia furcata ssp. subrangiformis and Hygrocybe persistens var. persistens as strongest indicators of ancient grasslands, and Rhytidiadelphus squarrosus as strongest indicator of recent grasslands. The vascular plant vegetation of the recent grasslands also included arable weeds and crop species, being residuals of the former land-use type. This pattern is very useful in distinguishing recent from ancient sites. However, we found no counterpart for the cryptogam vegetation. Thus, land-use history seems to have less influence on the composition of the cryptogam vegetation in grasslands. In Chapter 5, I present the first survey of the cryptogam vegetation of the grassland management project of Baden-Wuerttemberg. Using eight study sites the cryptogam vegetation found in the management and successional plots is compared. After a project term of 37 years, I assessed the effects of different mulching, mowing, and grazing methods as well as of undisturbed succession on the cryptogam vegetation, and give recommendations for the management and establishment of species-rich sites. The most species-rich bryophyte vegetation was found for the management types mulching every third year, mowing twice per year, and, for small acrocarpous species only, controlled burning. Macromycete species richness was highest in successional plots. These, despite the comparably short time for development, also yielded a surprisingly rich epiphytic vegetation. To enhance species diversity it is recommended to leave old trees and to newly create situations of high structural diversity by connecting wooded stands with grassland sites differing in the intensity of maintenance. Chapter 6 is a holistic approach applying the three concepts of species identity, Ellenberg indicator values, and functional traits to analyse the influence of management and past draining on the macromycete funga of a calcareous fen, the Sippenauer Moor. I assessed changes by comparing the present mapping data with data from a study carried out in 1998 and 1999. While species numbers were similar for the Sphagnum patches, we found a considerable loss of species and of red listed species for the remaining fen area, and only very few species were found in both studies. Ellenberg indicator values and functional traits did not yield significant differences. However, we found a considerable increase in the number of ubiquitous species. These changes in species composition most probably are caused by an insufficiant management, tree encroachment, and after-effects of the past draining. We iii Summary recommend to apply a more adapted management to prevent further species losses and to maintain the high quality of the fen. iv Acknowledgements Acknowledgements During my PhD time, many kind people helped me in many ways – by sharing their knowledge, their experience, their technical know-how, or ‘simply’ their time with me. I’d like to say “Thank you very much!” to all of you! First of all, I want to thank my supervisor Prof. Dr. Peter Poschlod, for guiding me on a safe path through my PhD studies, for letting me work on ‘my’ fungi, and for maintaining such a friendly and homely atmosphere – both intra- and extra-professionally. It was a great time, and I’m especially thankful for your understanding during my times of Lyme borreliosis. I’m also very grateful for the sustained support and scientific input I received from my mentors, Prof. Dr. Christoph Reisch and Dr. Claus Bässler, and I thank you very much for all kinds of help. Several people guided me on the way to further expand my knowledge of the species and ecology of fungi and other cryptogams, and therefore I especially want to thank Dr. Helmut Besl (†), Prof. em. Dr. Andreas Bresinsky, Max Kronfeldner (†), Hans Halbwachs, Wolfgang von Brackel, and Helmut Zitzmann. I always could rely on the help of Günter Kolb and Sabine Fischer, and I’m very much obliged for this. Furthermore, I’d like to thank the past and present Viktor-von-Scheffel-members (Verena Busch, Annika Sezi, and Sara Saeedi), my past and present office colleagues (Christina Meindl, Juliane Drobnik, Daniela Listl, Franziska Kaulfuß, and Judith Lang), and my past and present colleagues, especially Maria Hanauer and Philipp Glaab. Last but not least, I want to thank my family and friends, who always were there for me when I needed someone. v Motto “Fungi are the grand recyclers of the planet and the vanguard species in habitat restoration .” Paul Stamets “Kryptogamen, meine Herren, meine Damen , das sind Pflanzen ohne Samen ; Doch die samenlosen Dinger wickeln Weise um den