Robin S. Waples—Recipient of the 2018 Molecular Ecology Prize

Most of us can probably point to one or two individuals who have had a crucial influence on our career development and chosen field of study. For me, one of those instrumental individuals is Robin Waples. I met Robin in 1991 when I was a graduate student at the University of Oregon. Two of my UO mentors, Michael Lynch and Russell Lande, had encouraged me to contact him as I began seeking employment in quantitative conservation biology. Robin was then a geneticist at the NOAA National Marine Fisheries Service's Northwest Fisheries Science Center (NWFSC) in Seattle, where he still works as a Senior Scientist. He had arrived at the Center in 1986 to start a National Research Council postdoctoral fellowship with Fred Utter, a pioneer in fish genetics. Robin collaborated with Fred on population structure and gene conservation in Pacific salmon, launching a storied career as an NWFSC researcher working at the interface of science and natural resource policy. Robin generously created a position for me to focus on genetics in fish, and apart from a few diversions along the way, I have been working with him on this and related topics ever since. Robert Snowden Waples, Jr., was born 18 January 1947 in Berkeley, California. He attended Palo Alto High School, where he excelled in swimming; he went on to Yale to major in American Studies and to swim, competing with the likes of future Olympians Don Schollander, John Nelson, and Mark Spitz. A few little known facts: a 3-year varsity all-American in freestyle sprints at Yale, Robin made the US swimming squad for the Pan American Games in Winnipeg, Manitoba, in 1967. In 1968, he swam the 400-yard freestyle relay as Team Captain, helping Yale to finish second in the NCAA Championships that year. With characteristic self-effacing humour, when I asked him about his best event, he replied “the 75-yard butterfly.” After graduating in 1969 with a B.A. in American Studies, Robin took several years to travel and explore various interests, spending much of his time on or under the water in the tropics. He taught junior high school English in Hawaii, coached water polo and worked as a lifeguard, a wine steward and at a lumber mill in Cairns, Australia. After a decade and some crash coursework in biology, he entered graduate school at the Scripps Institution of Oceanography and did a Ph.D. in Marine Biology on the population genetics of marine fishes with the ichthyologist Richard Rosenblatt. His dissertation involved applying horizontal starch gel electrophoresis and histochemical staining for 26 enzyme systems to analyse gene flow in 10 species of marine shore fishes collected from four areas in southern California and Baja California, Mexico (Waples, 1987). His subsequent move to Seattle to study with Fred Utter was an evolution of this work involving some adaptive radiation on his part. Although Robin's early research involved taxonomy and population genetics of subtropical marine fishes, since coming to the Center much of his work has concentrated on salmon genetics and conservation. Before assuming the role of an NWFSC Senior Scientist in 2000, he led its Conservation Biology Division, during which time he was at the forefront of constructing a framework for identifying conservation units using genetic, life history and ecological data (Waples, 1991a, 1995, 2006), determining the conservation status of these units for Pacific salmon and steelhead, and assessing overall risk to units that include many populations (Allendorf & Waples, 1996; Waples, Adams, Bohnsack, & Taylor, 2007; Waples, McClure, Wainwright, McElhany, & Lawson, 2010; Waples, Winans, Utter, & Mahnken, 1990). His work required integrating complex biology with often arcane legal and policy considerations, and it led to a federal policy to define species of Pacific salmon under the US Endangered Species Act (ESA). Under his leadership, Division scientists conducted a series of comprehensive reviews of the coastwide status of Pacific salmonids under the ESA.11 https://www.westcoast.fisheries.noaa.gov/publications/status_reviews/salmon_steelhead/salmon_steelhead_esa_status_reviews.html From 1999 to 2003, he was the NWFSC's scientific lead for salmon recovery planning. Robin's research output is prodigious: He has over 220 publications since 1986 and his work has been cited over 23,550 times (15 of his papers have been cited 500 times or more). More impressive than these numbers, however, are the breadth, depth and impact of his research on applied conservation. His investigations are remarkably diverse considering that he is regarded by many as a fish geneticist, an impression that belies the fact that a major theme of his research is applying evolutionary and ecological principles to real-world problems in conservation and management. This diversity is evident in both topic and taxon. Broadly speaking, much of his work has involved adapting population genetics theory to applications involving species in the wild, particularly those of conservation concern. An early but sustained interest of his is in characterizing population structure in multiple species experiencing high gene flow (Waples, 1987, 1998; Waples & Gaggiotti, 2006). One of his better known contributions to population genetics has been the development of methods to estimate effective population size from genetic data that account for specific life history features, such as complex age structure with semelparity and overlapping generations (Waples, 1989, 2002; Waples, Antao, & Luikart, 2014; Waples, Grewe, Bravington, Hillary, & Feutry, 2018; Waples, Larson, & Waples, 2016; Waples, Luikart, Faulkner, & Tallmon, 2013; Waples & Yokota, 2007). He has advanced innovative approaches to the analysis of genetic mixture and admixture in mixed-stock fisheries (Anderson, Waples, & Kalinowski, 2008; Smouse, Waples, & Tworek, 1990; Waples & Smouse, 1990) which have been widely implemented in the management of Pacific salmon harvest. Robin has also published extensively on evaluating genetic risks and benefits of fish propagation (e.g., salmon hatcheries and marine fish aquaculture; Waples, 1991b, 1999; Waples & Drake, 2004; Waples, Hindar, Karlsson, & Hard, 2016), on integrating genetic and demographic factors into analysis of age structure and spatial structure (Waples, Scribner, et al., 2018) and on monitoring genetic and demographic population structure in managed species (Milligan et al., 2018; Schwartz, Luikart, & Waples, 2007; Tallmon et al., 2010). Recently, he has begun to explore the evolutionary responses of natural populations to human-altered environments (Waples et al., 2017) and the potential of genomics to assist conservation efforts (Hendricks et al., 2018; Waples & Lindley, 2018). Although much of his work with NOAA has focused on genetics and conservation of salmon and other migratory fishes, Robin has also conducted related studies on cetaceans (Waples, 2011) and on terrestrial mammals (Waples, Scribner, et al., 2018), and he has even investigated inbreeding and its effects on performance in thoroughbred racehorses. In addition to the Molecular Ecology Prize, Robin has received several other prestigious awards. The federal government recognized his innovative scientific leadership in advancing the implementation of the ESA by honouring him with a NOAA Administrator's Award in 1992, and bestowed upon him and his team a series of U.S. Department of Commerce Group Medals (a Silver in 1999, a Bronze in 1996 and two Bronzes in 2003), in recognition of their work on conservation of Pacific salmon and endangered southern resident killer whales. Robin earned a NOAA Distinguished Career Award in 2008 for his groundbreaking research in conservation genetics that advanced protection of genetic diversity in marine organisms. The same year, he received the William E. Ricker Fishery Conservation Award from the American Fisheries Society. In 2013, Robin was given the Edward T. LaRoe III Memorial Award from the Society for Conservation Biology. He was elected President of the American Genetic Association and elected to the Washington State Academy of Sciences in 2014, and inducted into the American Fisheries Society Genetics Section's Hall of Excellence in 2015. He has twice given the Jack W. Jones Memorial Lecture to the Fisheries Society of the British Isles (in 2001 and 2016), and during 2003–2004, he was a visiting scientist at university laboratories in France and the United States. He serves as an editor for several scientific journals and on numerous national and international scientific panels and steering committees dedicated to conserving marine organisms—from fish to reptiles to whales—around the world. Throughout his career, Robin has been highly supportive of students and young scientists. As well as participating on numerous graduate student committees domestically, he has served as an Opponent for Doctoral Dissertations at universities in Denmark, Ireland, Norway and Sweden. For over a decade, he served as Director of the NWFSC's Internal Grants Program, which has provided more than $2.4 million in grants for innovative research projects by Center staff, especially junior scientists. …it is important to make every effort to understand the ecology and life history of the target species. In the absence of such information, not only will a researcher be unable to make quantitative adjustments for possible sources of bias, she will not even be in a position to know whether bias has occurred, nor will she be able to develop effective strategies to minimize sampling error. This is the kind of wise counsel that has stayed with me as I have tried to reach conclusions from my own studies. Surely it is sound advice for any biologist drawing inferences about evolution or ecology in nature. But his contributions are not limited to the literature: One of Robin's most appreciated qualities is his eagerness to collaborate and to help one think a problem through (even if we ultimately have to consult Bill Hill, Russ Lande or my dog-eared copy of Wright's “Evolution and the Genetics of Populations”). The Molecular Ecology Prize is a fitting tribute to Robin and his body of work, which continues to have such an impact on a generation of geneticists and leaves a remarkably rich legacy for conservation biologists. I am grateful to Loren Rieseberg and Karen Chambers for the opportunity to write this brief sketch of Robin. Mike Ford, Krista Nichols and Linda Park have my thanks for their reviews of the manuscript. I am indebted to Robin for his steady encouragement and friendship over the years.

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