Thomas Norwood Taylor (1938–2016)

Recently, paleobotany lost one of its prominent practitioners. Thomas N. Taylor, Distinguished Professor at the University of Kansas, died on 28 April 2016 at home at the age of 78. He advanced our knowledge of the plants of the tropical wetlands of the Carboniferous Period (ca. 310 million years ago [Mya]), developed our understanding of the paleobiology of Antarctica during the Permian and Triassic Periods (ca. 290–200 Mya) and led research on the fossil record of fungi. Although fossil fungi were known since the earliest days of paleobotany, they were largely understudied, mainly because of the difficulty of recognizing and interpreting them. Taylor began to work on paleomycology in the 1980s with his PhD student Cynthia A. Wagner and later with his postdoctoral associate Sarah Stubblefield. These interests were further developed during the tenure of an Alexander von Humboldt Senior Research Award, which enabled him to visit and work with Winfried Remy at the University of Münster, Germany in 1994 and again in 1996. The close collaborations that he established with colleagues based in Germany blossomed into a highly productive research program on the fossil record of fungi.

Figure 1. Thomas N. Taylor, 2009 (from Taylor and Krings 2010; photo used with permission from the Society for Sedimentary Geology).

Figure 2. Thomas N. Taylor collecting permineralized Permian chert from the Skaar Ridge locality, Beardmore Glacier area, central Transantarctic Mountains, Antarctica, 2003.

Although fossil fungi have been known since the earliest days of palaeobotany, their abundance and quality have been unjustly minimized. Taylor will be remembered as the author of two of the most important textbooks in the field: three editions of Paleobotany: The Biology and Evolution of Fossil Plants (Taylor et al. 2009) and more recently, Fossil Fungi (Taylor et al. 2015). In addition, he was an author of more than 400 refereed journal articles and book chapters, four edited books, and six symposium volumes. His friend and colleague Gar Rothwell (personal communication 2016) remarked, “His career is a fantastic example of research productivity, research breadth, research innovation and rigor.” His other colleagues and students will remember him as an enthusiastic paleobotanist and a dedicated teacher and mentor. He loved field work and collected fossil plants on six continents. Today, these collections form the core of the University of Kansas paleobotany collections.

Thomas Norwood Taylor was born 14 June 1938 in Lakewood, Ohio. He earned his bachelor’s degree in botany and geology in 1960 at Miami University in Oxford, Ohio, and his PhD in 1964 in paleobotany at the University of Illinois, Urbana-Champaign. After graduating, he was a National Science Foundation postdoctoral fellow at the Yale University from 1964 to 1965. In 1965, he was appointed assistant professor at the University of Illinois at Chicago Circle and was promoted to full professor before his departure in 1972. After a short tenure from 1972 to 1974 at the Ohio University, Taylor moved to the Ohio State University where he served as senior research scientist at the Byrd Polar Research Center and chair of the Department of Botany for 21 years. In 1995, he joined the Department of Ecology and Evolutionary Biology at the University of Kansas as Roy A. Roberts Distinguished Professor. He was also a curator of paleobotany at the University’s Biodiversity Institute and Natural History Museum (FIG. 1).

Taylor had 14 PhD and 11 MS students, many of whom, along with his 27 postdoctoral associates, focused on the flora of the coal-forming swamps of the Carboniferous Period. Under the guidance of Wilson N. Stewart at the University of Illinois and later Theodore Delevoryas of Yale University, he studied fossil plants in coal balls. These large nut-shaped cobbles, formed from carbonates, frequently are found associated with coals of this age preserving the tissues of fossil plants in exquisite detail. One outcome of this early research was a superb monograph on fossil seeds of pteridosperms, which are extinct relatives of gymnosperms (Taylor 1965). Motivated by the wealth of valuable information preserved within coal balls, Taylor went on to establish a highly productive research program at the University of Illinois at Chicago Circle where he accepted his first graduate students in 1967 and his first PhD student in 1975. One aim of this work was to develop a greater knowledge of the reproductive structures of key groups of plants, including the extinct relatives of the horsetails, lycopods, and gymnosperms, that were such important elements of the coal-swamp floras. Taylor also pioneered the use of scanning and transmission electron microscopy in the study of fossil spores and pollen (Taylor 1973). The works ensuing from the coal balls and the electron microscopy represent an enormous contribution to our knowledge of the biology of Carboniferous Period plants (e.g., Millay and Taylor 1979; Taylor 1981, 1982).

Taylor’s interest in fossil fungi developed during the 1980s. Fungi are also a common element of coal balls, but they had been mostly overlooked by paleobontanists who were more interested in the plants. Together with his PhD student Cynthia A. Wagner, Taylor published a paper in Science describing spores similar to those produced by the modern endomycorrhizal fungus Glomus (Wagner and Taylor 1981). This was very early fossil evidence for fungi in the Glomales. “This was the fungus that really got him looking for more projects in mycology,” said Michael A. Millay (personal communication 2017). A productive collaboration with his postdoc Sara Stubblefield resulted in 11 papers published between 1983 and 1988, which together illustrated the diversity of fungal interactions and their importance in Devonian and Carboniferous Period ecosystems. These papers presented evidence for various mutualisms, parasitism, and fungal decay of woods. In a much needed review (Stubblefield and Taylor 1988), they summarized the state of the art, making the case for the existence of a rich fossil fungus record, the surface of which had barely been scratched.

Impressed by the abundance of fungal fossils in permineralized deposits, Taylor’s interests were drawn to ever more ancient ecosystems. In 1992, he began a collaboration on the Rhynie chert with Winfried Remy and German colleagues from the University of Münster, which led to many high profile publications. The Rhynie chert is a 407-My-old hot-spring system, which preserves the remains of an early terrestrial ecosystem fossilized more or less in situ (Edwards et al. 2017). Remy had been working on the Rhynie chert for many years, notably discovering the earliest evidence for the gametophyte generation of land plants (Remy et al. 1993). Taylor, Remy, and their collaborators, especially Hagen Hass, who prepared the materials, published four papers together on zoosporic fungi, focusing in particular on parasitism in a green alga. Endomycorrhizal associations had long been suspected with some of the Rhynie chert plants (Boullard and Lemoigne 1971); the first convincing evidence of a symbiotic association was demonstrated in the plant Aglaophyton majus by the documentation of arbuscules (Remy et al. 1994b; Taylor et al. 1995b). The team also described the oldest recognizable member of the Blastocladiomycota evidenced by sexual reproductive structures (Remy et al. 1994a; Taylor et al. 1994). Following Remy’s death in 1995, Taylor continued to work with colleagues in Münster, who now included Hans Kerp. Notable discoveries of this period are one of the oldest fossil lichens (Taylor et al. 1995a) and the oldest fossil Ascomycota (Taylor et al. 1999).

In the mid-1980s, Taylor’s interests turned to the Antarctic. Earlier, James Morton Schopf had found a Permian peat deposit in the central Transantarctic Mountains during the first season of field work (1969–1970) of the US Antarctic Program (Schopf 1970). Schopf also first reported the occurrence of a Triassic peat within the Fremouw Formation in Antarctica (Schopf 1978). These discoveries were significant because they represented the first examples of permineralized peats from Gondwana, and only the second known examples in rocks of these ages worldwide (Taylor et al. 1989). Taylor followed these discoveries up on an expedition in 1985–1986. These new finds promised to yield insights into the nature of polar forests, which are unique ecosystems without modern analogues. An important outcome was an edited book that drew together knowledge from throughout the fossil record on the paleobiology and environments of Antarctica and their relationships to other elements of Gondwana (Taylor and Taylor 1990). Further expeditions were undertaken from the University of Kansas and together with Edith L. Taylor and numerous graduate students, they greatly expanded our knowledge of the fossil plants and the paleoenvironments from this southernmost continent (FIG. 2). Fungi were an important but not an exclusive feature of Taylor’s work in Antarctica, and again permineralized soils featured prominently. Together with PhD student Carlie Phipps, Taylor documented the evidence of arbuscular mycorrhizae in the roots of the Triassic cycad Antarcticycas (Phipps and Taylor 1996). More recently, with his postdoc Andrew B. Schwendemann and other collaborators (Schwendemann et al. 2011), he described mycorrhizae associated with the root nodules of a conifer from the same site. These fossils are direct evidence for fungal associations in the polar forests of the Early Mesozoic.

In 1999, Taylor began a collaboration with Michael Krings of the University of Munich and Bavarian State Collections of Palaeontology and Geology. Together with PhD students, postdocs, and other colleagues, this work resulted in numerous publications on the fungi of Paleozoic and Mesozoic Era ecosystems. Important discoveries included arbuscular mycorrhizal fungi associated with the roots of tree clubmosses from a Carboniferous coal swamp (Krings et al. 2011b) and the oldest fossil basidiomycete clamp connections (Krings et al. 2011a). Many of their publications, which also included Hans Kerp, documented the roles and the diversity of fungi and fungi-like organisms in the Rhynie chert, providing information about their importance in early terrestrial ecosystems (e.g., Taylor et al. 2005; Krings et al. 2007, 2014, 2016). In 2015, the book Fossil Fungi was published by Taylor in collaboration with Michael Krings and Edith L. Taylor (https://www.elsevier.com/books/fossil-fungi/taylor/978-0-12-387731-4). This work is structured around the major taxonomic groups of fungi, placing the fossils in a systematic framework. It also includes chapters on spores, lichens, and interactions with other organisms. Each chapter introduces fossils with an easy-to-understand presentation of the main characters that define extant members. This is a landmark book, the first encyclopedic work devoted exclusively to fossil fungi.

During his lifetime, Taylor received numerous professional awards for his research, leadership, and teaching. He served on local, state, national, and international committees, volunteered for numerous administrative tasks throughout his career, and specialized in helping young faculty obtain funding for their research. In 1984–1985, he was the conference program chair of the Botanical Society of America. In 1984, he went to the University of Alberta, Edmonton, as a visiting professor for 6 months, and with Ruth Stockey, his former student, organized the Second International Paleobotanical Congress. He was president of the International Organisation of Palaeobotany between 1994 and 1997. Taylor was elected to membership of the National Academy of Sciences in 1994. He said of his election announced April 26 in Washington (DC), “I am humbled by it. You don’t do science for public accolade; you do it because you’re in love with it. But this is one of those nice things that happens to you once in a career” [sic]. Notably, he also served as a member of the National Science Board (NSB) from 2006 to 2012, advising both President George W. Bush and Congress on science policy. At the International Paleobotany Conference in Tokyo, Japan, in 2012, a special symposium was dedicated to him; the presentations were subsequently published in a special edition of the International Journal of Plant Sciences (Krings et al. 2013). The species name of a new fungus was proposed in his honor for his preeminent contribution to the understanding of the biodiversity of fossil fungi and the biological roles of fungi in paleoecosystems. In the year before his death, Taylor was chosen as the Karling Lecturer of the Mycological Society of America, a lecture that, unfortunately, he was unable to deliver.

Taylor had a very wide circle of colleagues and international collaborators. He hosted many scientists in his laboratory. During the early 1980s, he began to work with Sergio Archangelsy and his students in Argentina. The French connection was with Jean Galtier in Montpellier, with whom he co-authored several papers (Krings et al. 2011a, 2011b). Brigitte Meyer-Berthaud and Anne Laure Decombeix (AMAP, Montpellier) were both postdocs in Taylor’s laboratory. One of us (C.S.D.) met Tom Taylor for the first time in 2007 at a paleobotanical meeting in Paris. She was with her husband Désiré-Georges Strullu, who was a plant physiologist and an expert on extant mycorrhizae. They had a pleasant conversation about fungi, and Taylor remarked that he owned a copy of her husband’s book on fungi that he kept on a shelf in his office. This book (Strullu 1985) was the modern reference work for describing the oldest arbuscular mycorrhizae (Remy et al. 1994b). To the mycologist, Taylor will be remembered for his massive contribution to our understanding of fossil fungi and especially for his beautifully written and illustrated book.

Additional information and photographs of Taylor and his collaborators can be found in two touching tributes: http://bhort.bh.cornell.edu/paleo/tom-tailor-tribute2016.pdf and https://www.youtube.com/watch?v=wFOrQZmrjvA

ACKNOWLEDGMENTS

We thank Wilson A. Taylor (University of Wisconsin, Eau Claire), Mike M. Millay (Ohio University), and Gar Rothwell (Oregon State University) for their helpful comments and Rudolph Serbet (Biodiversity Institute and Natural History Museum, University of Kansas) for providing the photographs.