Susan Lynn Williams: the Life of an Exceptional Scholar, Leader, and Friend (1951–2018)

Susan Lynn Williams (1951–2018) was an exceptional marine ecologist whose research focused broadly on the ecology of benthic nearshore environments dominated by seagrasses, seaweeds, and coral reefs. She took an empirical approach founded in techniques of physiological ecology. Susan was committed to applying her research results to ocean management through outreach to decision-makers and resource managers. Susan’s career included research throughout the USA in tropical, temperate, and polar regions, but she specialized in tropical marine ecology. Susan’s scholarship, leadership, and friendship touched many people, leading to this multi-authored paper. Susan’s scholarship was multi-faceted, and she excelled in scientific discovery, integration of scientific results, application of science for conservation, and teaching, especially as a mentor to undergraduate and graduate students and postdoctoral scholars. Susan served in a variety of leadership positions throughout her career. She embodied all facets of leadership; leading by example, listening to others, committing to the “long haul,” maintaining trust, and creating a platform for all to shine. Susan was an important role model for women in science. Susan was also a loyal friend, maintaining friendships for many decades. Susan loved cooking and entertaining with friends. This paper provides an overview of the accomplishments of Susan in the broad categories of scholarship, leadership, and friendship.


Introduction
Susan William's academic career began at the University of Michigan. She started out in an engineering program, but switched to biology when she discovered her true calling, graduating as a Biology major with a Bachelor of Science degree in 1972. Susan moved to Fairbanks, Alaska, where she conducted her Master's research at the Institute of Marine Science, University of Alaska. Her research was supported by the at the Fairleigh Dickinson's West Indies Laboratory in St. Croix, US Virgin Islands. Susan married Bruce Nyden in 1986 in St. Croix, and they remained married until her death. She joined the research faculty at Friday Harbor Laboratories, University of Washington (1986)(1987)(1988)(1989)(1990), and then moved to San Diego State University where she was based from 1990 to 2000. Susan then took on the Directorship of the Bodega Marine Laboratory, University of California, Davis, and remained an active faculty member there. Susan's research and teaching showed no signs of slowing. True to her calling as an educator, Susan was driving with a car full of live marine organisms from Bodega Bay to deliver a lecture at UC Davis when her life was tragically ended in 2018.
Over the course of her career, Susan received numerous honors and awards including a Fulbright Award . Susan was selected for these awards at a diversity of institutions, not because she sought them, rather because she was an incredibly hard working, caring, and deeply committed scientist who served as an amazing role model to young women scientists.
Susan excelled at all aspects of academia, including obtaining research funding, obtaining science infrastructure funding, presenting at scientific conferences, teaching and mentoring, publishing peer reviewed papers (over 100), reviewing papers (50+ journals), and delivering numerous public talks and media interactions.
Susan conducted a large portion of her research underwater (Fig. 2). She helped pioneer the generation of researchers who made observations and conducted experiments in subtidal marine habitats. Susan was an expert SCUBA diver, and was inducted posthumously into the Women's Divers Hall of Fame. She was an aquanaut four times, living in an underwater habitat. Susan was certified as a SCUBA diver for 45 years, beginning in 1972. Susan continued to conduct underwater research projects, most recently focused on understanding the drivers of success in restoring coral habitats (e.g., Williams et al. 2017), in Indonesia. Susan would dive for pleasure as well as for research. Her former mentor John Ogden wrote the following: "Several years ago, Nancy [Ogden] and I joined her, Bruce [Nyden] and others on a 10day coral reef diving cruise in the Lesser Sunda Islands of Indonesia on a beautiful traditional Bugis phenisi schooner. Susan was energetic and studious, diving four times a day and building a long species list of reef fishes and inverts. But I remember most her joyous smile when she was among the Indonesian crew and villagers that we visited. She found something personal and magical in the people and the culture." Susan loved field work, and was not one to complain about being uncomfortable. One of her former students, Amanda Newsom, told the following story: "Early one dark, stormy morning in (I think) late fall, when the lab was mostly silent and empty still, I was coming in and saw this petite form marching down the hall towards me in a bright yellow rain slicker, boots, and coveralls. She was weighed down with equipment, and she looked up at me as we drew closer. There was an enormous grin on her face and her eyes were sparkling in a way I hadn't seen before. She announced, 'I'm going out ... to do field work!'."

Scholarship
Susan Lynn Williams was a scientist who embodied the ethos of conducting original research and making that research relevant for the betterment of society. In a seminal work,  Boyer (1990) defined scholarship in four categories: discovery, integration, application, and teaching. Susan Williams excelled in all four categories, each described in the following sections.
Discovery Susan actively conducted original research in both the laboratory and the field, and she was instrumental in making unique discoveries. Susan represented the generational shift to conducting much of her field research underwater; her predecessors were largely restricted to the intertidal zone or the laboratory. Susan did conduct intertidal and laboratory research, but she also was an accomplished SCUBA diver, and she spent endless hours underwater in all sorts of conditions. Susan's underwater observations backed up with laboratory studies led her to challenge existing paradigms throughout her career.
Susan used radioisotopes in her Masters' research to measure carbon uptake by seagrasses (Williams and McRoy 1982) and then used stable isotopes to track the uptake of sediment nutrients by coenocytic green algae (Williams 1984;Williams and Fisher 1985). Coenocytic green algae are giant single cells with multiple nuclei that move via cytoplasmic streaming. Susan's PhD research upended the paradigm that algae absorbed water column nutrients exclusively. Early on, Susan recognized the importance of seagrass detritus in marine food webs (e.g., Suchanek et al. 1984) as well as the importance of tropical seagrass as a food source for megaherbivores (e.g., Thayer et al. 1984;Williams 1988a). During and immediately after her dissertation research, Susan studied the deepwater environments off St. Croix (e.g., Williams 1988bWilliams , 1990Williams and Carpenter 1988), which required extensive SCUBA diving combined with experimental manipulations and state-ofthe-art scientific techniques. Susan compared the circadian rhythms of a seagrass and a green alga using in situ light manipulations (Williams and Dennison 1990). Susan began and maintained a long and productive collaboration with Bob Carpenter, and they co-authored papers on coral reef algal turfs over several decades (e.g., Carpenter 1988, 1990;Williams 1993, 2007).
Moving to the west coast, Susan began looking at the genetic diversity and structure of seagrasses (e.g., Williams and Davis 1996;Williams and Orth 1998;Williams 2001). She was one of the first scientists to recognize the emergence of Ruppia in disturbed seagrass communities (e.g., Johnson et al. 2003). But then Susan's research added a new component, studying the effects of introduced species, starting with studies of an Asian mussel in San Diego Bay (Reusch and Williams 1999), followed by studies of the invasion of a marine macroalga, Caulerpa, in southern California (e.g., Williams and Grosholz 2002;Williams and Schroeder 2004). Susan became an invasive species polymath, publishing over twenty papers on aspects of (a) invasive species and native seagrass interactions, (b) post-invasion management, (c) invasion vectors and their management, (d) invasions and climate change, and (e) synthesizing ecological aspects of invasive species (Williams and Smith, 2007).
Integration Susan was a key member of the Seagrass Trajectories Working Group at the National Center for Ecological Analysis and Synthesis (NCEAS). In addition to Susan's important individual contribution, as an advocate of women in science, she also recruited two young female University of California graduate students, Randall Hughes and Suzanne Olyarnik to join the working group. Several seminal papers resulted from this synthesis effort, including a Bioscience paper "A global crisis for seagrass ecosystems" (Orth et al. 2006) and a Proceedings of the National Academy of Sciences paper "Accelerating loss of seagrasses across the globe threatens coastal ecosystems" (Waycott et al. 2009). Each of these papers has received over 2000 citations and represent the most cited seagrass papers in the world. At the special session as a tribute to Susan Williams at the Coastal and Estuarine Research Federation conference in 2019, almost all of Susan's co-authors from this NCEAS project attended, a testament to her lasting impact.
Susan did comprehensive reviews of various topics throughout her career, including the impact of invasive species (Lodge et al. 2006;Williams and Grosholz 2008;Ojaveer et al. 2018), the impact of invasive species on seagrasses (Williams 2007), trophic transfers from seagrasses (Heck Jr et al. 2008), and plastics in seafood (Rochman et al. 2015).
Application US Congressman Jared Huffman submitted a tribute to Susan to the Congressional Record that included the following statements: Dr. Williams' scientific research on coastal ecology and her activism surrounding the expansion of marine sanctuaries have left an indelible impact on the world. Throughout her career, Dr. Williams' research underlined the connection between the health of oceans and the communities surrounding them. Her work on the ecology of nearshore marine ecosystems, seagrass, coral reefs, and invasive species helped illustrate the impact of warming oceans on coastal environments and yielded strategies for mitigating those impacts. Her work was heralded by the international scientific community, and provided the foundation for critical changes to state and federal policies addressing the management of coastal environments. Dr. Williams became a valued advisor for state and federal officials seeking to protect the California coast. Nowhere was this more apparent than through her work in significantly expanding two national marine sanctuaries off the coast of Northern California: the Cordell Bank National Marine Sanctuary, and the Gulf of the Farallones, now known as the Greater Farallones National Marine Sanctuary. Both sanctuary boundary expansions were proposed in legislation in 2005, and were ultimately expanded to more than twice their original size by the Obama administration in 2015. Throughout that ten-year process, Dr. Williams was the driving force that propelled the policy forward. (162 Cong. Rec. E737 2018) Susan made multiple trips to Sacramento, CA, and Washington, D.C., to provide testimony or to brief congressional delegations. Susan worked tirelessly to provide the scientific basis for the expansion of marine sanctuaries and she often made these trips to Washington, D.C., at her own expense while juggling her many teaching, research and administrative duties. Susan also made the case for the inclusion of biodiversity in habitat restoration (Hughes et al. 2017) and she kept working to influence management and policy, even when she sometimes felt as though she was "Shouting into a hurricane".
Susan's latest research interests and science application efforts were in Indonesia, and she was deeply involved in a fruitful collaboration with Indonesian scientists and the Mars Incorporated's coral reef restoration program. This experience in the Coral Triangle, a global biodiversity hot spot, caused Susan to reevaluate her priorities and shift her perspectives (e.g., Williams 2013). Susan made multiple trips to Indonesia, often taking graduate students along with her and conducting outreach (e.g., Sur et al. 2018) in addition to focusing on capacity building and research. Former student Holly Hanson Henderson said the following: "I carry the lessons and approach to science that I learned from Susan with me in my work every day. Susan was unwavering in her belief that science needs to be thoughtful and rigorous, and that it is our obligation as scientists to go out into the world to make a difference." Susan wrote about how species richness accelerates marine ecosystem restoration in the coral triangle (Williams et al. 2017). Susan became the lead scientific advisor to Mars on a large-scale coral restoration project, documented in a paper published posthumously (Williams et al. 2019). She also helped design a second trial, which is still ongoing and is the world's largest controlled coral reef restoration experiment, scheduled to finish in early 2021.

Teaching
Susan's former students were polled and they generated a word cloud to describe her. Words to describe Susan included the following: brilliant, sparkling, incisive, creative, trailblazer, generous, influential, passionate, direct, genuine, diplomatic, frank, tough, human, supportive, impactful, incredible, and honest (Fig. 3).
Bobby Espinoza said this about Susan's mentorship: "All along, Susan supported me, as a person. Most other advisors would have abandoned me and my project soon after it morphed into something so distant from their own research interests. Not Susan. And she has continued to support me throughout my career." Bengt Allen said the following: "As I look back over the past 20 years since I left her lab, I've more or less been successful by continuing to do all of the things she had me doing as a new graduate student." Amanda Newsom said this about Susan's impact: "I don't think I would have pursued or merited the work I do now without that shift Susan cultivated in me towards valuing myself and my truth. As a trailblazer for women in our field, Susan had to learn how to value herself in order to be the empowering presence she became. I think her influence was what finally made me understand my own worth as a professional. She demanded that I develop a thicker skin and a higher opinion of myself, which is a difficult thing to do, but I think she accomplished it with many of her students." Randall Hughes said that the Ralph Waldo Emerson quote fit Susan's mentorship; "Our chief want in life is somebody who will make us do what we can." In that vein, Susan was the advisor for the following graduate students: Lucina Hwang, Seth Yarish, and Eric Telemaque (Stony Brook Univ.); Mary Ruckelshaus (Univ. Washington); Michelle Chow, Tim Lu, Bobby Espinoza, Russ DiFioni, Chris Davis, Donna Ross, Patrick Ewanchuk, Amy Sewell, Alex Cheroske, Bengt Allen, Holly Hanson Henderson, Megan Cooper, and Carolyn Lieberman (San Diego State Univ.); and Laura Rodriguez, Heidi Weiskel, Cascade Sorte, Amanda Newsom, Amber Szoboszlai, Grace Ha, and Katie DuBois (Univ. California, Davis). But this list is far from complete-Susan was often a "drive-by" advisor who provided many more students and postdocs with input and career advice. Susan provided important input to Matthew Bracken, Jeffrey Wright, Christopher Harley, Raquel Muñiz-Salazar, Brendon Larson, Cynthia Hays, Randall Hughes, Jae Pasari, Laura Reynolds, and Brent Hughes (Univ. of California, Davis) and Florence Thomas, Thorsten Reusch, Jorge Terrados, and Terrie Klinger (San Diego State Univ.). Susan also provided K-12, undergraduate, and graduate mentoring to numerous students through programs like the Research Experiences for Undergraduates, National Science Foundation Graduate K-12 program (CAMEOS at University of California, Davis).
Susan taught a wide diversity of courses over her career, including Seagrass Biology and Ecology, Marine Ecology, Biological Invasions, Climate Change in the Nearshore Ocean, Marine Primary Producer Physiological Ecology, Life in the Sea, Principl es of Ecology, Marine Environmental Issues, Phycology, Ecology and the Environment, Evolution and Ecology, Aquatic Ecology, Biogeochemistry of Coral Reefs, Algology, Estuarine Botany, Advanced Phycology, Ecology of Coral Reefs, and Tropical Coastal Marine Ecology. Susan developed numerous syllabi, lectures, and laboratories for individually designed courses. Susan even co-authored a paper that outlines ways for research scientists to more actively participate in K-12 education (e.g., Komoroske et al. 2015).
Susan participated in the Aldo Leopold leadership training program, along with Rosamond Naylor and others. Susan excelled in the five Leopold leadership attributes: 1. Leading by example, demonstrating personal leadership. Susan was usually the hardest working member of any team. Her former student Cascade Sorte said the following: "Have any of you heard the suggestion that to impress your supervisor, you should start work before them and/or stay at work after them? Well, that is literally impossible when you work with Susan. Her work ethic and influence are incredible." Susan never asked others to do what she would not do herself. She conducted field work in all conditions and never shirked the drudgery of clean up or breaking down experiments. Susan lived her life in an exemplary manner, always conscious of those around her who had less privileged lives.
2. Listening to colleagues, students and community members. Susan was a great listener. She walked into a room to say "There you are", rather than "Here I am." She was respectful to others, and was someone who took the time to listen to all the people around her. She had an open door policy and various students, faculty, and staff often took the opportunity to use Susan as a sounding board and a friendly ear.
3. Committing to the "long haul." Susan was a consistent and committed scientist. She spent years and decades on various projects, resulting in some seminal publications that will be difficult to replicate. For example, Susan published a major work in Ecological Monographs that stemmed from hundreds of hours underwater (Williams 1990) that was the culmination of years of effort. Brent Hughes described the persistence that Susan encouraged after multiple unsuccessful attempts to obtain funding for a seagrass restoration project in Elkhorn Slough. The ultimately funded and successful restoration project is now known as "Susan's Garden." 4. Developing and holding trust. Susan was a trusted confidant to many people. She was direct and honest at all times. Her former student Amber Szoboszlai said the following: "Of all the professors I met with from California to Washington, Susan was the only one who was direct and honest about her assessment of my application. Her recommendation illuminated a pathway to grad school for students traditionally seen as deficient in their requirements. Although Susan's honesty was at times hard to receive, her transparency and willingness to 'tell it like it is' were genuine and impactful for many of her students."

5.
Creating a platform for all to shine. Susan was the best audience. Leila Hamdan said, "Susan was always sitting front and center and nodding encouragingly." Former student Holly Hanson Henderson said the following about Susan's cohort of graduate students: "We supported each other. We bought swim team coats to wear after diving and had them embroidered with 'Team Williams'." Susan was painfully aware of the challenges of being a woman marine scientist. Her former student Megan Johnson Cooper said that Susan told her the following: "Being a female scientist is important. We often need to be stronger than the men in the room."

Friendship
Science is often thought about in terms of ideas, data, and hypotheses. But science is performed by humans, and this dimension of science is just as important as the data and theories. When Susan's friends and colleagues learned of her passing, our collective sense of loss was keenly felt. Many of us appreciated Susan and her humor, humility, care, values, and her wonderful smile. Susan's career was largely based at semi-remote field laboratories. One of the things that occur at field laboratories is the establishment of intense personal relationships. People working at field laboratories tend to both work and socialize together. In addition, field trips can be strenuous experiences that test people's fortitude, and this leads to a realization of your abilities and the abilities of your colleagues. Thus, Susan's personal relationships with colleagues were often deeper than typical academic relationships.
Susan accumulated many friends and she maintained some four and five decade friendships. People like Bob and Hedy Carpenter, Bill Dennison, Jud Kenworthy, John and Nancy Ogden, and Fred Short were long-time friends and colleagues. For Susan, the friends that she made along her life journey enriched her life, along with her family, in particular her sister Holly, brother-in-law Art, and niece Kaitlin.
Susan loved to cook and share meals with friends. Her former student Alex Cheroske said this about the impact of her cooking: "I tell people that by the end of my first year of grad school, I had a subscription to the journal Ecology and Bon Appetit magazine. Susan was a mentor both in the field and in the kitchen and I took copious notes while I was observing her work in both realms." Bengt Allen said the following: "Personally, aside from providing continued support and guidance long after I graduated from her lab, the other major impact Susan had on my life was through her cooking. Susan used to have lab parties at her house once a semester or so, where she would cook just amazing food. My then girlfriend (now wife of almost 20 years) and I would marvel at her custom kitchen with 6-burner stove, copper pots, and built-in cookbook library. Her passion for cooking as a vocation outside of work was contagiouswe regularly cook using recipes that Susan provided us for some of our favorite dishes we first tried at her house (e.g., vodka pasta with prosciutto, creamy chicken jalapeno quesadillas, and habanero salsa)." Cascade Sorte wrote the following: "This empowerment and blurring of the lines of work and 'life'two of the messages that I think Susan personifiesare also messages that can help diversify our field, by keeping people like myselfwho are deeply committed to both family and sciencein the field. When, just over one month after starting my graduate work in her lab, I announced to Susan that I was expecting, she could not have responded more positively. Not oncenot one single timehas Susan seemed to question my scientific ambition or ability to juggle my family and work responsibilities."

Conclusion
Susan left a lasting impression on the people she encountered throughout her life, as was evidenced in the outpouring of feelings and thoughts stimulated by her death. She was Roberta Marinelli's "North Star" and Tessa Hill said that Susan was "A Whole Person". Susan's former students and mentees said that Susan taught them to be "Big and Brave" and commented on the sticky note she had in her office that said "You Can Do It". Susan Williams created a legacy of impacts through her scholarship, leadership and friendship.
Mary Oliver (1935-2019) wrote a wonderful poem "Pipefish" which was read by Susan's former mentee A. Randall Hughes at her memorial at U.C. Davis. This poem is a fitting tribute to Susan: In the green and purple weeds called Zostera, loosely swinging in the shallows, I waded, I reached my hands in that most human of gestures-to find, to see, to hold whatever it is that's there-and what came up wasn't much but it glittered and struggled, and it had eyes, and a body like a wand, it had pouting lips. No longer, all of it, than any of my fingers, it wanted away from my strangeness, it wanted to go back into that waving forest so quick and wet. I forget when this happened, how many years ago I opened my hands-like a promise I would keep my whole life, and have-and let it go. I tell you this in case you have yet to wade into the green and purple shallows where the diminutive pipefish wants to go on living. I tell you this against everything you areyour human heart, your hands passing over the world, gathering and closing, so dry and slow.
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