Josef Hafellner – a Life Amongst Lichens and Their Parasites

Herzogia 29 (2) Teil 1, 2016: 213 –234 213 Josef Hafellner – a life amongst lichens and their parasites Martin Grube, Walter Obermayer*, Helmut Mayrhofer & Toby Spribille Abstract: Grube, M., Obermayer, W., Mayrhofer, H. & Spribille, T. 2016. Josef Hafellner – a life amongst lichens and their parasites. – Herzogia 29: 213 –234. The impact of Josef Hafellners scientific work on the systematics of lichenized ascomycetes and on the knowledge on lichenicolous fungi is discussed and the value of his extensive collections (c. 100,000 lichens including c. 8,000 lichenicolous fungi) is highlighted. A bibliography (1975 –2016) is presented. Zusammenfassung: Grube, M., Obermayer, W., Mayrhofer, H. & Spribille, T. 2016. Josef Hafellner – ein Leben inmitten von Flechten und ihren Parasiten. – Herzogia 29: 213 –234. Der Einfluss von Josef Hafellners wissenschaftlichen Arbeiten auf die Systematik von lichenisierten Ascomyceten und auf die Kenntnis von flechtenbewohnenden Pilzen wird diskutiert und auf den Wert seiner umfangreichen Aufsammlungen (ca. 100.000 Belege, davon ca. 8.000 Belege lichenicole Pilze) hingewiesen. Eine Publikationsliste (von 1975 bis 2016) wird vorgelegt. Key words: Bibliography, eponyms, lichenicolous fungi, European Alps. Introduction Few scientists make such a mark on their science that they can be said to change their field, and fewer still are involved in setting up the framework for an entire branch of natural history. One such person is Josef Hafellner, to whom we dedicate the present contribution. Conscious that he is – notwithstanding his recent retirement – still very much fully active as a researcher, we wish here to highlight his impact on our science to date. Josef Hafellner has dedicated his scientific career to the classification and taxonomy of fungi, in particular lichen-forming ascomycetes and the fungi inhabiting lichen symbioses (known as lichenicolous fungi). Consistent with this central theme, he has also contributed a large body of work to lichen floristics, mainly within the eastern parts of the European Alps. Here we provide a short overview of Hafellner’s contribution to the systematics and taxonomy of lichens, of his scientific output resulting from numerous field trips, and of the results coming from one of his favorite hobby horses, the study of lichenicolous fungi. Studies on systematics and taxonomy of lichens Josef Hafellner began studying under Josef Poelt soon after the latter was appointed chair of the department of Systematic Botany in Graz in 1972. Under the influence of Poelt, Josef * Corresponding author 214 Herzogia 29 (2) Teil 1, 2016 Hafellner soon became enthusiastic for mycology, despite his early interests in plant systematics. With his first publication, about the ascus structures in the genus Thelocarpon (Poelt & Hafellner 1975), he already visited two of the topics that would later become mainstays of his research: the ascus structural characters as central to ascomycete classification, and the classification of lichenicolous fungi. He continued this research in his doctoral thesis about the genus Karschia, which at the time was circumscribed as a heterogenous assemblage of fungi with different life styles, including both lichenized and lichenicolous species (Hafellner 1979). He went on to demonstrate his broad competence of lichen systematics with a mono- graph of the primarily tropical lichen genus Letrouitia (Hafellner 1981), which doubled as his “habilitation thesis” and gave him the possibility to pursue a scientific career in the German-speaking academic culture of those times. Perhaps the central work of Josef Hafellner’s scientific career and a milestone in lichenology was his contribution to the Festschrift for Josef Poelt (Hafellner 1984b). It represented a comprehensive taxonomic study of the families Lecanoraceae and Lecideaceae, which at the time were interpreted as two large and heterogenous families of lichenized ascomycetes, in part still echoing the antiquated concept of Zahlbruckner. Although their phenotypic hetero- geneity was widely recognized by contemporaries, the two families remained taxonomically untouched. As Josef Hafellner was convinced that the characters of the reproductive structures were the most significant for classification, he undertook to provide a new classification of phenotypic groups that reflected natural relationships. His new concept recognized no less than 48 families instead of the former two and led to a radically new classification that was based on fertile structures, particularly based on fine differences in ascus structures that are stainable with Lugol’s solution. When Hafellner established this taxonomic framework the knowledge about phenotypic diversity of lichens was far less developed than it is today, and the new concept therefore provided a solid basis for classification of lichens. At that time, however, his concept was both revolutionary and controversial. Two of the authors (MG and HM) remember the intense discussions about the many new or reinstated families at the International Botanical Congress in Berlin 1987. However, as time passed, an increasing number of the families proposed by Josef Hafellner were accepted by peer lichen systematists. When PCR and DNA sequencing was introduced to phylogenetic analysis of lichens only a few years later, this also served as a starting point for testing these taxonomic ideas and the sig- nificance of ascus characters. Today, this paper counts among the most cited papers in lichenological systematics, with over 320 citations as of summer 2016, according to Google Scholar. During career as a taxonomist, Josef Hafellner introduced more than six hundred scientific names, and his scientific output comprises 289 publications (see publication list at the end of this account). His contributions to lichenology have meanwhile been honored by using his name directly, or as an anagram, in naming of four genera and seventeen species, as listed below: a) Seventeen names of species in honour of Josef Hafellner (bold = described in the present Festschrift): Arthonia pepei Etayo & Pérez-Ortega; Biatora hafellneri Rodr.Flakus & Printzen; Biatoropsis hafellneri Millanes, Diederich, M.Westb. & Wedin; Caloplaca hafellneri S.Y. Kondr. & Kärnefelt; Capronia hafellneri Nograsek; Capronia josefhafell- neri Zhurb. & Etayo; Dactylospora hafellneriana Sérus.; Hypotrachyna hafellneri Elix, T.H.Nash & Sipman; Letrouitia hafellneri S.Y.Kondr. & Elix; Megalaria hafellneriana Kantvilas; Opegrapha hafellneri E.Zimm., Etayo & F.Berger; Schismatomma hafellneri Egea & Torrente; Stigmidium hafellneri Zhurb.; Triblidium hafellneri Magnes; Trichoconis Grube, Obermayer, Mayrhofer & Spribille: Josef Hafellner – a life amongst lichens and their parasites 215 hafellneri U.Braun, Khodos., Darmostuk & Diederich; Xanthoparmelia hafellneri Elix; Xanthoria hafellneri S.Y.Kondr. & Kärnefelt. b) Four names of genera in honour of Josef Hafellner (bold = described in the present Festschrift): Fellhanera Vězda (106 taxa); Hafellia Kalb, H.Mayrhofer & Scheid. (32 taxa; later synonymized with Buellia); Hafellnera Houmeau & Cl.Roux (one taxon; later synonymized with Schaereria); Henfellra Halıcı, D.Hawksw., Z.Kocak. & M.Kocak. (one taxon). In addition, Hafellner’s surname frequently crops up as a secondary derivation from exist- ing scientific names (Lichenostigma fellhanerae, Malcolmiella fellhaneroides, Malmidea fell haneroides, Phaeosporobolus fellhanerae and seven taxa of Fellhaneropsis). Even a chemical compound (hafellic acid) is derived from ‘Hafellner’ due to its occurence in the genus Hafellia. As the 289 publications imply, Hafellner has been a prolific author, and while many of these papers were built on his own effort alone, not a few were also carried out in the context of his many collaborations. Hafellner was never especially fond of attending conferences, but he spent long hours interacting with the many scientific visitors to the Institute of Plant Sciences in Graz and has worked together in the field with many of the leading figures in 20th and 21st century lichenology. The many authors that contributed to the present Festschrift are a testa- ment to years of friendships and/or scientific joint endeavors with Josef Hafellner. Materials for studies on lichen biodiversity Studies of lichen biodiversity, also documented by his extensive collections, are a further mainstay of Josef Hafellner’s lichenological activity. It is almost impossible to list all of the expeditions, which resulted in many lichen floristic accounts and have kept the present lichen curator in Graz (WO) quite busy. To date, Josef Hafellner has collected and determined more than 100,000 specimens from all over the world, most of which are housed in the herbarium GZU. Hafellner’s first specimen, dated 29 June 1974, labeled as number ‘1' (Fig. 4), was determined by him as Acarospora chlorophana [for more information on ‘specimen number one’ see caption in Fig. 4]. Later he accepted this species in Pleopsidium, a genus which he re-established in recognition of the differences in chemistry and ascus type from Acarospora (Hafellner 1993a). Specimen number one, as it turns out, was to be the beginning of a long and storied collecting career spanning five continents. a) Excursions to extra-European countries and areas outside the European Alps Josef Hafellner conducted numerous lichenological forays in the Macaronesian islands (for published results see Hafellner 1995b, 1999a, 2002d, 2005a, 2008b), but other important lichenological

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