James William Kimbrough, 1934–2017

James William Kimbrough (FIG. 1) was born November 7, 1934, in Eupora, Webster County, Mississippi. He was the eldest son of Felix and Ethel Kimbrough. James, known as Buddy to his family in Mississippi and Jim to his friends in Florida, grew up on a farm and was the eldest of 10 children. He did not enjoy the hard physical labor of farm work and was determined to find a path away from agricultural work. In his later years, he would say that his dislike of farm work was part of why he excelled in academics. Jim graduated from Sunflower Junior College and then went on to receive a bachelor’s degree in science education in 1957 and a master’s degree in botany in 1960 from Mississippi State University in Starkville. Jim began studying fungi as part of his master’s degree, and he published his first research paper on the developmental biology of the powdery mildew Pleochaeta polychaeta (Kimbrough 1963). After completing his studies at Mississippi State University, he moved to Cornell University in 1960 where he began his dissertation on the biology and taxonomy of discomycetes with Richard Korf. He was part of a long line of systematists who received training at Cornell under Korf, and he perpetuated Korf’s love of the discomycetes in his own career. He received his PhD in mycology and plant pathology in 1964.

Figure 1. Dr. James William Kimbrough working at the microscope in his laboratory, ca. 2001.

Later that same year, Jim joined the faculty of the University of Florida (UF) where he was the mycologist for the next 46 years. He was first appointed in the Department of Plant Pathology in 1964, transferred to the Department of Botany in 1966, and then returned to the Department of Plant Pathology from 1989 until his retirement in 2010. Jim was awarded tenure in 1970 and emeritus status in 2010. During his time at UF, Jim had an appointment that included extension, teaching, and research. His extension appointment consisted primarily of identifying fungi for Florida stakeholders, consulting on fungal plant diseases, identifying and consulting on issues related to molds and mildews, identifying mushrooms for the Florida Poison Control, and acting as the director of the mycological herbarium of the Florida Museum of Natural History (FLAS).

Jim was passionately involved in teaching and mentoring at UF. He taught a series of fungal biology classes, including an advanced laboratory mycology course for graduate students and upper division biology students, as well as a popular undergraduate mycology course that was geared toward non-biology majors—“Molds, Mildews, Mushrooms and Man.” His courses were very popular. “Molds, Mildews, Mushrooms and Man” started with seven students in 1994 and grew to an enrollment of approximately 280 students by 2001 (Anonymous 2001). Jim’s teaching accomplishments were recognized by UF in 2001 when he received the Undergraduate Teacher of the Year Award. He was also the thesis advisor for many graduate students, including 17 MSc students (from 1965 to 2000) and 18 PhD students (from 1973 to 2006). During his career at UF, Jim also supervised five postdoctoral researchers, including Meredith Blackwell (emeritus professor at Louisiana State University) and Richard Baird (professor at Mississippi State University). Gerald Benny began as a postdoctoral fellow with Henry Aldrich and stayed at the University of Florida where he worked in the Kimbrough laboratory until Jim’s retirement in 2009.

Jim’s research focused primarily on the systematics and taxonomy of ascomycetes, particularly the taxonomy of Pezizales and Coryneliales but also other groups. Jim had wide interests in fungal biology and published on a range of topics that included termite-associated fungi, Florida plant pathogens, the systematics of Rhizoctonia, and the impact of molds on air quality. By the time he retired, Jim was author on more than 150 peer-reviewed publications. His research group employed the cutting-edge technology of the time—electron microscopy—in a quest to discover phylogenetically informative structures in several fungal groups and explore details of morphology and development. Research in Jim’s laboratory explored the ultrastructure, ontogeny, and development of diverse ascomycetes (e.g., Benny et al. 1978; Kimbrough and Benny 1978; Gibson et al. 1986b) as well as basidiomycetes (e.g., Khan and Kimbrough 1979, 1980a, 1980b) and zygomycetes (e.g., Maia and Kimbrough 1993, 1994).

Over the years, Jim was the author or coauthor of numerous taxonomic novelties, including at least 10 genera (Caccobius, Cleistoiodophanus, Coprobolus, Fitzpatrickella, Hormiscioideus, Lasiothelebolus, Myrmecomyces, Ochotrichobolus, Termitariopsis, Trichobolus) and more than 60 species across several taxonomic groups. Many of these taxa are coprophilous Pezizales. Jim maintained an interest in these coprophilous discomycetes from the time he worked on them during his dissertation until the very end of his career. These taxa were the subjects of many of his structural and developmental studies and often combined culture work with electron microscopy. His research on termite-associated fungi with students and postdoctoral researchers led to the rediscovery of minute fungi that had only been previously observed and described by Roland Thaxter (Kimbrough and Gouger 1968; Blackwell and Kimbrough 1978). This work also led to additional publications on external fungal parasites of termites, including Termitariales (Khan and Kimbrough 1974a, 1974b; Kimbrough and Thorne 1982; Lenz and Kimbrough 1982) and Laboulbeniomycetes (Kimbrough et al. 1972b; Blackwell and Kimbrough 1976a, 1976b).

Jim contributed several important book chapters that reviewed mostly premolecular progress and challenges in ascomycete systematics, including a 1970 classification of the discomycetes (Kimbrough 1970b), a 1982 chapter on “The Discomycetes” for Synopsis and Classification of Living Organisms (Kimbrough 1982), and a chapter on the use of septal ultrastructure in ascomycete systematics as part of the First International Workshop on Ascomycete Systematics (Kimbrough 1994). He also coauthored an overview chapter on discomycetes for The Mycota VII (Pfister and Kimbrough 2001) and a monograph on Octospora (Wang and Kimbrough 1992).

As mentioned previously, electron microscopy was an important tool in the Kimbrough laboratory and was used to answer questions about ascomatal ontogeny, ascus apex structure, ascospore ontogeny, ascospore wall development, and septal pore ultrastructure. Studies in the Kimbrough laboratory on septal pore ultrastructure were possibly the most groundbreaking among their many contributions to fungal biology. Jim and his collaborators documented and interpreted septal pore ultrastructure as phylogenetically informative characters in a range of different fungi, but with particular emphasis on the Pezizales. This work spanned more than 30 years and included many hours of meticulous sectioning of resin-embedded blocks, hours peering through the oculars of a transmission electron microscope, and hours developing film in a darkroom. During that time, Jim and his collaborators explored and documented septal pores from representatives of nine families of the Pezizales. Arguably the most important discoveries from this work were the description of a unique lamellate structure associated with the hyphal septal pores in Pezizales, “the Peziza septal type” (as compared with the “Neurospora-type” and “Sclerotinia-type”), and the description and interpretation of septal pore ultrastructure at the base of young asci as informative at family or higher levels of Pezizales (Kimbrough 1994). He dubbed these septal types as “aleuroid,” “ascoboloid,” “anthracoboid,” “Ascodesmis-type” (first described by Carroll 1967), “mycolachnoid,” “otideoid,” pezizoid,” “pulvinuloid,” and “scutellinioid,” along with interpretations of ascus septal pores found in Hellvellaceae, Morchellaceae, and Sarcoscyphineae. This was critical information in an order of fungi that included the majority of ascomycete truffle species whose phylogenetic affiliations were unknown at the time. With this knowledge, Kimbrough and his students were able to correctly link the truffles Barssia to the Helvellaceae (Kimbrough et al. 1996), Genea to Pyronemataceae (Otideaceae in their circumscription) (Li and Kimbrough 1994), and Hydnobolites to the Pezizaceae (Kimbrough et al. 1991). Not all of Jim’s interpretations turned out to be corroborated by molecular analyses. For example, molecular evidence does not support a close relationship between Coprotus (Leotiales) and Ascodesmis (Pyronemataceae), although their septal pores were interpreted as similar. Along the same lines, Jim hypothesized that the Rhizinaceae should be considered to be Helvellaceae because both have elongate Woronin bodies, but this is not the case (Kimbrough 1994; Hansen et al. 2013). These incongruities may be a matter of interpretation of the transmission electron microscopy (TEM) data and the conservation of septal pore characters at higher taxonomic levels in some taxa. Nevertheless, molecular phylogenetic analyses have generally borne out the phylogenetic utility of septal pore ultrastructure in the Pezizales (Hansen and Pfister 1996; Hansen et al. 2013).

Another notable contribution to ascomycete systematics came from ultrastructural work that graduate student Jack Gibson performed under Jim’s direction. Jack reexamined the hyphae and “chlamydospores” of Glaziella aurantiaca. At that time, Glaziella aurantiaca was considered a monotypic member of Endogonales despite the fact that it was known to have regular septation. Along with confirming regular septation, Gibson documented Woronin bodies and ascomycete-like septa in Glaziella. He showed that the supposed “chlamydospores” were actually unispored asci. Gibson et al. (1986b) hypothesized that Glaziella did not belong in the Endogonales but was instead closer to the Pezizales, although they placed it within a monotypic order. Their hypothesis about its close relationship to Pezizales was later validated when molecular phylogenetics placed Glaziella within the Pezizales (Landvik and Ericksson 1994; Hansen and Pfister 1996; Hansen et al. 2013).

One more illustration of how Jim’s ultrastructural work revealed surprising fungal relationships that have since been corroborated by DNA evidence is from his presidential address to the Mycological Society of America (MSA) in 1981 where he presented his work on Thelebolus (Kimbrough 1981). Thelebolus had previously been placed in the Pezizales (Kimbrough and Korf 1967; Eckblad 1968), but Jim showed that Thelebolus species had a cleistothecial development that was unlike any other group within Pezizales. He also demonstrated that the ascus structure was composed of thick, multilayered walls that were more similar to loculoascomycetes than to discomycetes in Pezizales. Comparisons with other genera showed that the ascus structure of Thelebolus was most similar to Ryparobius and some Ascozonus species. Molecular work later confirmed that Thelebolus, along with Ryparobius and Ascozonus, are placed outside of the Pezizales. Ironically, the exact phylogenetic affiliation of the Thelebolaceae within the Leotiomycetes is yet to be fully resolved (Landvik et al. 1998; de Hoog et al. 2005; Wang et al. 2006). A summary of the information found over so many years of meticulous work was distilled as drawings to characterize the different septal pore types and presented at the First International Workshop on Ascomycete Systematics (Hawksworth 1994). Images of the ultrastructural findings by Jim and his collaborators, along with others who generated similar data from other fungi, have now been compiled in a phylogenetic framework and are available through the Assembling the Fungal Tree of Life (AFTOL) Structural and Biochemical Database (Celio et al. 2006; https://aftol.umn.edu/).

In the later years of his career, Jim worked on the fleshy fungi of Florida. During this time, he coauthored the description of Macrocybe titans, the largest known agaric from North America (described as Tricholoma titans by Bigelow and Kimbrough 1980). His regional field guide, “Common Florida Mushrooms,” is the only major mushroom book to cover the state of Florida (Kimbrough 2000). This book remains an important guide for Florida fungi and is used widely by Florida extension agents, citizen scientists, and graduate students. This work is based partly on the works of William Alphonso Murrill, an important American mycologist and longtime resident of Gainesville (Kimbrough 1973, 2003). Jim was fascinated with the post-New York Botanical Garden life of Murrill. He researched Murrill‘s activities during the Great Depression and cataloged his return to mycology with the help of Florida plant pathologist George Weber. Jim was known to give Gainesville tours, including stops at Murrill’s grave at the Evergreen Cemetery as well as the two houses Murrill built in town.

A practical example of Jim’s work with fleshy fungi occurred during the 1990s and 2000s when Jim and his research group explored new areas of research and extension, including the study and promotion of shiitake mushroom cultivation in Florida (Webb et al. 1988). Branching out further, he also consulted and studied indoor molds and their health effects (Huang and Kimbrough 1995, 1997).

Jim’s research also led to significant international travel and collaboration. Many of the graduate students in the Kimbrough laboratory were from outside the United States, including students and later collaborators from Brazil (Lenore Maia, Universidade Federal de Pernambuco, and Jose Luis Bezerra, Universidade Federal do Recôncavo da Bahia) and Taiwan (C. C. Tu, Taiwan Agricultural Research Institute). Jim also traveled to present his research and visit collaborators across the globe, including trips to Brazil, Canada, England, France, Germany, Taiwan, and Turkey.

Jim earned the respect of his colleagues for his research and was known for his affability and helpfulness. During his long mycological career, Jim was active in the Mycological Society of America and a member from 1963 until the time of his retirement. He served as a counselor (1971–1973), secretary-treasurer (1974–1977), a member of both the Finance Committee (1974–1980) and Travel Awards Committee (1992–2004), vice president (1977–1978), president elect (1978–1979), and president (1979–1980). In recognition of his scholarship, service, and active career, he received several prestigious awards from the MSA, including the Weston Award for Excellence in Teaching in 1996 and the Distinguished Mycologist Award in 2001.

In addition to his exciting academic life, Jim fostered a rich family life. His wife and children accompanied him on numerous cross-country fungus-collecting expeditions. Such trips are evidenced by multiple fungal collections by Jim’s wife, Jane, and all three of their children, Dawn, Jeff, and Jay. These collections from far flung locations, such as the Huron Mountain Club in the upper peninsula of Michigan, were deposited in the FLAS herbarium for future use. Jim was also a deeply devoted Christian who spent significant time and effort in service to his church and faith. He was an active member of the Northwest Baptist Church in Gainesville, where he served as a deacon. He routinely attended church services and was a regular participant in church activities with his family. Jim passed away January 21, 2017, after a struggle of several years with Parkinson’s disease. He is survived by Jane, his wife of 55 years, three children, seven grandchildren, and one great-grandchild.

ACKNOWLEDGMENTS

The comments and documents shared by Dr. Gerald Benny, the Department of Plant Pathology at the University of Florida, and the Kimbrough family were helpful for creating this document and are greatly appreciated. We also thank Donald Pfister, Hal Burdsall, Brandon Matheny, and Meredith Blackwell for their constructive comments and editorial contributions to this memorial.