The Reinvention of Compositae: Vicki Funk’s Legacy in the Systematics of the Largest Plant Family on Earth
Abstract
Vicki Ann Funk was an American botanist and a leading figure on the taxonomy, systematics, and biogeography of Compositae in the late twentieth and early twenty-first centuries. Of her many contributions to these fields, Vicki’s ardent defense of the value and necessity of strong collections-based research for plant systematics was highly prominent. We will delve into Vicki’s contribution to Compositae systematics and explore how she was often at the forefront of research in the areas of plant phylogenetics, systematics, and collections. She pioneered the use of cladistics in plant systematics in the late 1970s and the early 1980s and helped develop and put in practice novel methods for understanding plant evolutionary relationships. During the next-generation sequencing revolution of the 2010s, she was again at the leading edge developing new tools for studying the world’s largest flowering plant family. Throughout this, she continuously made use of the latest methods and approaches to study evolutionary diversification, biogeography, and classification in Compositae. She spared no effort in supporting small herbaria across the globe, constantly aware (and reminding those around her) of the paramount importance of collections at all levels of plant research. She envisioned producing a global database to track nomenclature in Compositae and saw this massive effort from its origin to its implementation by encouraging contributions from researchers across the globe. In all her work, there was a genuine desire to connect people in pursuit of deeper understanding of the Compositae.
A Botanical Leader and Influencer
Dr. Vicki Ann Funk (fig. 1) received a Bachelor of Science (1969) and Master of Science (1975), working with Dr. Marian Fuller, from Murray State University in Murray, Kentucky. Her master’s thesis was entitled “A Floristic and Geologic Survey of Selected Seeps of Calloway County, Kentucky,” with parts published in the Southern Appalachian Botanical Society journal Castanea and Transactions of the Kentucky Academy of Science (Funk and Fuller 1978, 1980). Vicki completed a PhD in 1980, working with Dr. Tod Stuessy, from Ohio State University, where she studied the systematics of Montanoa Cerv. in Llave and Lex. (Heliantheae, Compositae; Funk 1980). She conducted postdoctoral work at the New York Botanical Garden from 1980 to 1981, working with Dr. Arthur Cronquist. In 1981, Vicki was appointed as a research scientist and curator at the US National Herbarium of the Smithsonian Institution’s National Museum of Natural History, where she studied Compositae and other taxa for almost four decades. Vicki A. Funk conducting fieldwork in Sierra de Famatina, La Rioja province, Argentina. Vicki was fascinated with the phylogenetic position of Famatinanthus, and she made it a priority to study this interesting Compositae in the field.
Vicki’s early influence on plant evolution and taxonomy, combined with her motivation to bring together researchers from different fields, set in motion interactions and collaborations that carried throughout her career. She quickly became a pioneer in her field, leading the charge in the use of modern phylogenetic methods (cladistics) in systematic botany (e.g., Funk and Stuessy 1979). During these early years of cladistics, she was building bridges with leading zoologists, including publishing an edited volume with contributions from both botanists and zoologists. E. O. Wilson highlighted this remarkable collaboration: “The papers collected in this volume are the products of a rather rare event … —a meeting between botanists and zoologists to discuss their common goals and problems” (from the preface to Advances in Cladistics: Proceedings of the First Meeting of the Willi Hennig Society [Funk and Brooks 1981]). Wilson went on to say at the first meeting of the Willi Hennig Society that “it was particularly nice to see so many botanists … indeed the zoologists were almost outnumbered” (Funk and Brooks 1981, p. viii). In the proceedings, scientists published on diverse topics, from plant hybridization to stingrays and their parasites. William Fink (1982, p. 873) wrote in a review of the proceedings, “A noteworthy aspect of this book is the participation of so diverse a group of biologists,” something that would not have happened without Vicki’s leadership.
Further demonstrating Vicki’s ability to push beyond the bounds of contemporary thinking, she developed innovative methods to detect hybridization events in plants in a phylogenetic context, something we still grapple with today despite having whole genomes at our disposal. For example, in a 1985 paper, Vicki described numerous ways to develop hypotheses about hybridizations using phylogenies and detailed, extensive case studies for testing these hypotheses (Funk 1981, 1985). The introduction to her 1985 paper reads as though it could have been written today as Vicki outlined the case for the importance of hybridization and polyploidy to the evolutionary history of plants and the deficiencies of traditional phylogenetic systematics (cladistic methods) for studying the patterns of reticulate evolution (Funk 1985). Recent advances in molecular phylogenomic approaches coupled with novel statistical approaches have uncovered ample evidence for hybridization by systematics and resulted in a renewed interest in hybridization as a driving evolutionary force. Remarkably, the figures drawn by Vicki in her 1980s work are nearly identical to those published in recent papers (e.g., Solís-Lemus et al. 2017).
Vicki’s studies of biogeography were equally innovative. Her volume with Warren Wagner, Hawaiian Biogeography: Evolution on a Hot Spot Archipelago (Wagner and Funk 1995), spurred new studies of island radiations in a biogeographical context and remains essential introductory reading for students of biogeography. For example, Wagner and Funk (1995) defined the famous progression rule, which articulates that clades tend to inhabit older islands first and disperse to younger islands in the order that the islands appear. This book was the result of another great collaboration headed by Vicki in a symposium cosponsored by the American Society of Plant Taxonomists and the Association of Tropical Biology in 1992.
Vicki was responsible for organizing major collaborative projects that generated groundbreaking studies in Compositae and was instrumental in the formation of the International Compositae Alliance (TICA). In 2006, members of TICA met in Barcelona (echoing the Reading and the Kew meetings of 1975 and 1994, respectively), which would eventually result in the publication of the monumental 2009 volume Systematics, Evolution, and Biogeography of Compositae (Funk et al. 2009). At more than 1000 pages, this volume included individual chapters devoted to each tribe recognized at the time, plus palynology, chromosomes, economic uses, and more (fig. 2). The book remains a testament to Vicki’s ability to wrangle a global community of researchers from diverse fields while also serving as a critical resource for anyone studying the family. Cover image of Systematics, Evolution, and Biogeography of Compositae, published in 2009. The massive volume took two years to complete and remains the authority and reference text for this major plant family.
At the height of the next-generation sequencing (NGS) revolution, Vicki again realized the power of new approaches and how they could give insight into the evolution of Compositae. Vicki obtained funding from the Smithsonian to carry out a pilot study testing whether the NGS approach (Hyb-Seq) could resolve evolutionary relationships in the family, with ultimate success (Mandel et al. 2014; Mandel 2021). For several years, Vicki gathered samples, both through collection trips and from her extensive network of collaborators, into the penultimate NGS study of the family: all subfamilies covered, all but two recognized tribes, and over 200 genera (Mandel et al. 2019). This work resulted in new evolutionary relationships and prompted new classifications, for example, increasing the number of subfamilies from 13 to 16, all which were described in a special issue of Taxon (Susanna et al. 2020), again demonstrating Vicki’s powerful ability to bring people together doing leading-edge work.
Vicki’s Impact on Collections-Based Research
Vicki’s great virtues included her ability to identify challenges to be solved, to integrate fields of knowledge, and, above all, to foresee new ones to be developed. Everything that had to do with Compositae was interesting to her, and everything was considered from the perspective of how to take the research to the next level. Enthusiastic by nature, she would stop at nothing to pursue her goals. To every problem Vicki applied her logic of reducing it to the simplest form in such a way that the solution, however hard to implement, was also readily identifiable. There was always a hypothesis to test!
Given this way of approaching things in life, and her training focused on the use of botanical collections, it was inevitable that Vicki placed collections as one of the central elements of her work. She constantly sought ways to promote the use of botanical collections themselves and support their maintenance and development in all the places she visited. Vicki understood the importance and relevance of botanical collections as the ultimate source of information. An excellent legacy in this sense was her compilation of the potential uses of herbarium samples, which details a comprehensive list covering the wide spectrum of human activities that in one way or another benefit from the use of collections (Funk 2004).
Botanical collections are essential to connect any discovery that is made about a plant with the universe of knowledge generated by the scientific community, past and present. Vicki’s passion for collections manifested itself in every place she went; no matter where she was, she always made sure to make at least one collection. However, even more important was her willingness to connect with and support local botanists, especially those who were just starting their careers and chose to study Compositae. These visits to local herbaria resulted in networking and involvement of people in the synantherologist2 community—everybody became connected.
Vicki loved fieldwork and lived by the maxim “My worst day in the field is always better than my best day in the office.” Desert and mountainous areas are where Compositae are most diverse, so it was no surprise that those were also her favorite places for fieldwork. First Mexico, during her PhD years in search of her beloved Montanoa, then South Africa, and eventually the Andes (especially Patagonia) were among her favorite destinations. She was a compulsive collector, always planning the next trip. On field trips, she was methodical, always analyzing the best course to follow to maximize the trip’s benefits, knowing the value of the resources at stake in preparing for even a simple field trip. Her preferred role on a collecting trip was that of the navigator, reviewing maps and attending to the itinerary. Once arrived at the collection site, she would strategically walk around the area, and after locating a species of interest, she sat next to it to study it carefully, taking photos and recording data. According to her own testimony, those moments alone with the plants, in the field, were some of the ones she enjoyed the most. Then at night came the second round of data collection to expand on the information recorded for each specimen, the postprocessing, and the recording of extra data.
Vicki was dedicated to demonstrating the use of herbarium collections and their value as an essential source of information on plant morphology and anatomy. She further emphasized the importance of mining information from herbarium labels in biodiversity studies because those data serve as direct evidence of species distributions through time. For example, data about species distributions allow us to understand the geographic patterns of diversity of the groups, which can inform conservation priorities for rare or threatened taxa. Finally, herbarium specimens properly identified using basic morphology are the bedrock of artificial intelligence and deep learning approaches to species identification.
All this information obtained from herbarium specimens is referenced thanks to the existence of a nomenclatural system that allows the correct communication of the results. Very early in the twenty-first century Vicki had her eye on creating a nomenclatural database for Compositae, which got its start through the participation of multiple stakeholders but notably thanks to Christina Flann and Ilse Breitwieser. This database, originally known as the Global Compositae Checklist, migrated across systems and today resides in the World Register of Marine Species infrastructure under the name of Global Compositae Database (see
The Reinvigoration of a Global Network
The enthusiasm shown by Vicki in supporting and promoting all kinds of interactions among researchers and institutions to advance knowledge of Compositae lives on through her collaborators and trainees. TICA continues to be active (over 400 members), following the legacy of Vicki by promoting monthly meetings, the TICA-Talks, where researchers from all over and at any career stage share the results of their research. One of the positive outcomes of post-COVID life has been the greater virtual connectivity that allows interactions that were previously only relegated to less frequent real-life encounters.
Her legacy also lives on through the continuation of the Compositae Newsletter, which published 50 issues from 1975 until 2012. The newsletter was the only channel dedicated exclusively to publishing works on Compositae and to serving the community of synantherologists. In the last issue, representatives from TICA, led by Vicki, indicated their unanimous desire that a venue to exclusively publish things from Compositae should continue and that it should be in electronic format. The new iteration, Capitulum (fig. 3; see Cover of the first Capitulum issue, published in 2021. Capitulum is the new iteration of Compositae Newsletter, a peer-reviewed publication dedicated exclusively to Compositae research. The journal is managed by the International Compositae Alliance and accepts a variety of article types, from original research to reviews. In the same spirit of its predecessor, Capitulum is distributed freely and can be accessed online at
All these ongoing projects (fig. 4), from (1) resolving phylogeny at deeper levels using genomic methods to understand the main evolutionary events in the family, (2) having a free tool that allows access to the list of species present, (3) strengthening the community of synantherologists through an invigorated TICA, and (4) the resurrection of a publication venue to publish all things Compositae, such as Capitulum, represent clear forms where the legacy and spirit of this incredible scientist that we honor today with this special issue persist in a very tangible way—as Vicki used to say, to infinity and beyond! The four pillars of Vicki’s scientific legacy. From phylogenomics to databasing, from disseminating research advances to uniting a community across the world, the legacy of Vicki lives on across a myriad of projects.
We would first like to thank the International Journal of Plant Sciences, especially Chris Caruso and James Ellis, for giving us the opportunity to publish this set of papers in a special issue honoring our great friend and colleague. We would like to thank all those who participated in this special issue and the symposium at Botany 2022 in Anchorage, Alaska, “A Botanist at the Extreme: Honoring the Great Contributions of Dr. Vicki A. Funk.” Through her spirit, enthusiasm, and passion, Vicki lives on through all of us who knew her and even those who never had the opportunity to meet her.
Notes
2 Synantherologist, one who studies Compositae, from “syn” and “anther,” meaning anthers fused as in the androecium in Compositae.
Literature Cited
Fink WL 1982 Cladistics in action. Science 216:872–873. Funk VA 1980 Systematics of Montanoa Cerv. in Llave and Ilex. Compositae, Heliantheae. PhD diss. Ohio State University, Columbus. ——— 1981 Special concerns in estimating plant phylogenies. Pages 73–86 in VA Funk, DR Brooks, eds. Advances in cladistics: proceedings of the first meeting of the Willi Hennig Society. New York Botanical Garden, Bronx. ——— 1985 Phylogenetic patterns and hybridization. Ann Mo Bot Gard 72:681–715. ——— 2004 100 uses for an herbarium (well at least 72). ASPT Newsl 17:17–19. Funk VA, DR Brooks 1981 Advances in cladistics: proceedings of the first meeting of the Willi Hennig Society. New York Botanical Garden, Bronx. Funk VA, MJ Fuller 1978 A floristic survey of the seeps of Calloway County, Kentucky. Castanea 1978:162–172. ——— 1980 Analysis of the distribution of southeastern taxa in seeps of Calloway County, Kentucky. Trans Ky Acad Sci 43:162–172. Funk VA, TF Stuessy 1979 Cladistics for the practicing plant taxonomist. Syst Bot 3:159–178. Funk VA, A Susanna, TF Stuessy, RJ Bayer 2009 Systematics, evolution, and biogeography of Compositae. IAPT, Vienna. Mandel JR 2021 What’s all the hype about Hyb-Seq? Capitulum 1:44–47. Mandel JR, RB Dikow, VA Funk, RR Masalia, SE Staton, A Kozik, RW Michelmore, LH Rieseberg, JM Burke 2014 A target enrichment method for gathering phylogenetic information from hundreds of loci: an example from the Compositae. Appl Plant Sci 2:300085. Mandel JR, RB Dikow, CM Siniscalchi, R Thapa, LE Watson, VA Funk 2019 A fully resolved backbone phylogeny reveals numerous dispersals and explosive diversifications throughout the history of Asteraceae. Proc Natl Acad Sci USA 116:14083–14088. Solís-Lemus C, P Bastide, C Ané 2017 PhyloNetworks: a package for phylogenetic networks. Mol Biol Evol 34:3292–3298. Susanna A, BG Baldwin, RJ Bayer, JM Bonifacino, N Garcia-Jacas, SC Keeley, JR Mandel, S Ortiz, H Robinson, TF Stuessy 2020 The classification of the Compositae: a tribute to Vicki Ann Funk (1947–2019). Taxon 69:807–814. Wagner WL, VA Funk 1995 Hawaiian biogeography: evolution on a hot spot archipelago. Smithsonian Institution, Washington, DC.