John J. Skvarla (1935–2014)

John J. Skvarla (*30 March 1935, †2 March 2010) can be numbered among the great actuo-palynologists of the twentieth century. His work is a lasting contribution to our knowledge of the ultrastructure of recent pollen of many families of flowering plants. His findings have had a considerable influence on plant systematists in evaluating classification and relationships in the many groups that he studied. His early work in the late 1960s at the University of Texas at Austin with Don Larson was pioneering in the application of transmission electron microscopy (TEM) to the ultrastructure of pollen and its development. While at Austin, John established a close relationship with Billy Turner, an expert on the Compositae. This developed into John’s extensive investigation of the pollen morphology of that family, of which he became the world authority, working with many collaborators in recent years, notably with Vicki Funk, Alexandra Wortley and Stephen Blackmore. His research developed and expanded greatly when he moved to the University of Oklahoma at Norman, where he spent the majority of his career. The laboratory, first equipped with a fine Philips 200 TEM was a pride and joy to John, who almost solely operated it for some 20 years. This facility began the foundation of a palynological laboratory that was pre-eminent in the world for nearly half a century.

John was a willing collaborator, keen to work with and to help anyone, who shared an interest in palynology. He developed extensive international networks, spending a sabbatical in 1971–1972 at the Gunnar Erdtman Palynological Laboratory in Stockholm, Sweden, working with John Rowley, the then Director. In Stockholm, he collaborated with a number of other people, notably Joseph Praglowski, with John Pettitt from the British Museum (Natural History) and Orville Dahl of the University of Pennsylvania as visiting colleagues during his stay. On his return journey home, he spent some weeks at Kew collecting pollen samples and discussing with Charles Jeffrey, the Kew authority on Asteraceae. Here, he met Keith Ferguson and they became friends, which later developed into an invitation for Keith to spend some time in Norman working with John. In 1977, Kew granted Keith six months leave after much assistance from John who influenced Pat Brenan, then Keeper of the Herbarium and later Director, Jack Heslop-Harrison, who was Director, and most importantly, Peter Raven. Peter Raven (Missouri Botanical Garden) was joint organiser of the first International Legume Conference held at Kew in 1978 and supported the visit, both through his influence and financial help; this visit began a major joint investigation of the pollen morphology of Fabaceae subfamily Papilonoideae.

John visited London and Kew again in 1974 to participate in the Linnean Society symposium, ‘The Evolutionary Significance of the Exine,’ where he delivered a paper co-authored with Joseph Praglowski and Peter Raven on the pollen morphology and its significance in the systematics of Onagraceae. The quality of the electron micrographs and especially the TEM micrographs of the exine stratification, detail of the joining of the tetrads and the remarkable thin sections of the viscin threads remain superlative. Further papers on Onagraceae, in collaboration with Peter Raven, followed. A collaboration with Joan Nowicke of the Smithsonian Institution’s National Museum of Natural History began work on ‘Centrospermae’ pollen and a series of large papers followed of outstanding quality, both in illustration and interpretation.

In 1975, John returned to the UK to take part in the International Compositae Conference, which was convened to bring together new lines of evidence from phytochemistry and micromorphology. John presented a synthesis of his work with Billy Turner and Don Larson on the family, describing three fundamental patterns of exine stratification in the family. However, at that time, the palynological evidence seemed to be at odds with all of the widely accepted classifications of the family, leading to the conclusion, now known to be mistaken that exine ultrastructure was simply too variable to be an important character in the systematics of the family.

Meanwhile, the laboratory at Norman grew. A small scanning electron microscope was added to the facilities, funded by one of the half dozen or more large successful NSF (National Science Foundation) grant proposals that John wrote. His students included Alice Wittenbach, Varsha Patel, Ed Vezey and Mike Farabee. There were many visitors. John Rowley was perhaps the most regular visitor, and numerous joint publications resulted. A number of visitors came for training, including Peggy Bolick, then from Austin, Stefanie Ickert-Bond studying at Tempe when she first visited, and Audrey Walker from Amherst. Keith Ferguson’s stay resulted in a collaboration continuing for some 15 years, and he became a regular visitor to Norman. Masamichi Takahashi spent a year in the Laboratory in 1987–1988 and a number of important joint papers on pollen grain development resulted. John also collaborated with his Oklahoma University colleague Lois Pfiester and co-authored a number of papers that established sexuality in the dinoflagellates, the last phylum for which sexuality was to be described.

Skvarla’s work was consistently of the highest standard. He carried out almost all of his own TEM examinations and he spent hours in the darkroom preparing the images to illustrate his publications and lectures. He was once overheard saying to a collaborator, ‘don’t worry about being nervous about delivering your paper, the pictures will speak for themselves’. When preparing a joint paper for publication, he liked to sit down with the collaborator and read the manuscript aloud weighing each sentence to be sure that the meaning was clear and it carried the information he wished to impart.

The University of Oklahoma, with a $500 000 donation from the Noble Foundation in 1978, built a dedicated Electron Microscopy Unit adjacent to Skvarla’s laboratory and he became the founding Director of what would become the university’s core imaging facility. Upon its completion, he stepped aside to make way for Charles Daghlian in the hopes that Daghlian would become tenured and remain in Norman. This move is indicative of his generosity. Shortly afterwards, Daghlian was enticed by other opportunities, however, and Scott Russell was appointed Director, and the two laboratories collaborated with joint facilities. John was honoured with a George Lynn Cross Research Professorship in 1980, which continued to provide financial support for John’s active research until his death.

Throughout, a mainstay of the Skvarla Laboratory and later of the University Electron Microscopy Unit, was Bill Chissoe. Bill was a Lew Wentz undergraduate student in Skvarla’s laboratory after military service in the US Navy in the late 1960s. He became a master of ultramictrotomy and embedded and thin sectioned specimens for John for over 40 years, continuing to work in the Electron Microscopy Unit whenever John needed samples prepared, even after retirement. Additionally, Bill’s training in electronics in the forces proved invaluable. As well as collaborator and technician, he was a good friend and colleague of John’s over many years. After Bill’s retirement and in response to the advent of digital imaging, Greg Strout became a trusted helper, whose perfectionistic approach to excellent images matched John’s passion and attention to detail.

In 2005, when Vicki Funk asked Stephen Blackmore to prepare an overview of Compositae pollen for The International Compositae Alliance (TICA), he replied that this would be impossible without involving John. The final team also included Harold Robinson and Alexandra Wortley, and the major conclusion of the study was that the major types of exine stratification, described by Skvarla and Larson in the 1970s, were, in fact, highly congruent with the phylogeny of the family. Thus, the full importance of John’s pioneering work only came to be known after 40 years, when the classification of Compositae based on the Supertree compiled by Vicki Funk and her collaborators was available. John took this on board in a very modest way, suggesting that he had always thought the data must mean something.

His enormous contribution to palynology was recognised in 2008 when Vicki Funk arranged an international symposium in his honour during the Botanical Society of America conference in Vancouver, Canada.

John was a kind and generous man giving his time and facilities freely to students and colleagues alike. He was always self-effacing and reluctant to seek the limelight – a family man keenly following his children’s progress both at school and at sports and proud of his wife Donna’s senor librarian position and her national importance in quilt work. He was also a keen fisherman and enjoyed the summer home that he and Donna built in Colorado, where he could fish at will. A devoted Catholic, he and Donna were pillars of the Catholic Church in Norman.

John Skvarla was born 30 March 1935, in Passaic, New Jersey, to Joseph E. Skvarla and Emma Wolsko Skvarla. He earned his bachelor’s degree in geology from Upsala College in East Orange, New Jersey, a master’s degree from Miami University in Oxford, Ohio, and a PhD in Botany from the University of Texas at Austin. He is survived by his wife Donna whom he met in Austin and two daughters and two grandchildren. Two children predeceased him.