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2014 In Memoriam: Robert Lloyd Rausch—A Life in Nature and Field Biology, 1921–2012 Eric P. Hoberg Department of Agriculture, Agricultural Research Service, [email protected]

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J. Parasitol., 100(4), 2014, pp. 547–552 Ó American Society of Parasitologists 2014

IN MEMORIAM

Robert Lloyd Rausch—A Life in Nature and Field Biology 1921–2012

‘‘For myself, I express gratitude for the opportunity to investigate some given ecosystem of which he is part must be a fundamental attribute of zoonotic diseases on the arctic coast. . . I received as well the kind man’’ (Rausch, 1985). hospitality and assistance of the Inupiat,˜ in Barrow and Wainwright and He embraced this philosophy and traveled the globe, where his science other villages on the coast and in Anaktuvuk Pass in the Brooks Range. touched the lives of many, contributing directly to our understanding of They shared with me their knowledge of the fauna and flora of the Arctic, pathogens and diseases of wildlife and people and ultimately defining the transmitted among themselves for generations. They were and are my interfaces for humans and the biosphere. Robert Rausch was forever mentors.’’ curious about our world and always accessible to relate a recent insight or Robert L. Rausch (2001) a unique story, revealing the intensity and passion of his science and depth of his humor. He was recognized in many ways—as a preeminent Robert Lloyd Rausch was born in Marion, Ohio on 20 July 1921 and parasitologist and mammalogist, as an extraordinary and fearless field passed away at his home on Bainbridge Island, Washington on 6 October biologist, as a generous mentor, as a colleague, and as a dear friend for 2012. In his 91 years and over more than 6 decades of a renowned career, those fortunate to have known him well. Robert L. Rausch was a student in the broadest sense of field biology and When we think about Robert Rausch, it is Alaska that comes to the a naturalist in the expansive sense of another century always seeking to forefront, but there were years of training that prepared him for the understand the intricate and often hidden connections for diversity and pathways that led to far corners of the north and beyond. Robert Rausch history in our complex world. Highlighting these points in his Presidential was educated at the Ohio State University, receiving a B.A. in zoology and entomology in 1942 and a D.V.M. in 1945. Later, an M.Sc. followed at the address to the American Society of Parasitologists in 1984 he reflected, Michigan State University in 1946, and a Ph.D. at the University of ‘‘. . .The ability to distinguish, name, and categorize the components of a Wisconsin in 1949, both focusing on parasitology and wildlife manage- ment. His arrival in Alaska in late 1948 followed the establishment of the DOI: 10.1645/14-561.1 Arctic Health Research Center (AHRC) of the U.S. Public Health Service

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FIGURE 1. Robert Rausch in the Brooks Range, Alaska ca. 1949 with George llano (right) during survey and inventory for vertebrates and parasites (photo: Archives, University of Alaska Museum). and an adventurous journey up the recently cleared Alcan Highway. He of science (Rausch, 1951). As he observed in his understated manner, was in many ways on terra incognita, and had a mandate to investigate ‘‘...we had the benefit of the indispensable help of the Inupiat,˜ who shared zoonotic diseases and related health problems of the indigenous peoples of their unique knowledge, and whose guidance in the field prevented ill- Alaska at a time of considerable cultural transformation across northern advised actions on the part of persons unfamiliar with the arctic communities. There was limited knowledge and understanding of what we environment.’’ After arriving in Alaska in 1948, Robert Rausch was now call pathogen biodiversity, and he was immediately immersed in field learning quickly, and spent much of the ensuing 3 years in the field, studies that took him to Barrow and St. Lawrence Island and into the traveling with the remaining nomadic bands, gaining the respect of his heart of the Brooks Range (Rausch, 2001), a path of biological mentors and acquiring the special skills and tools for survival in the exploration across starkly beautiful and deceptively complex arctic extremes of the northlands (Rausch, 2001). In April 1949 he traveled with ecosystems. In parasitology, his name is intimately linked to the far companions across the North Slope and Brooks Range by dogsled, and northern environments of Alaska and eastern Siberia (Fig. 1). provided the first documentation of the nomadic lifestyle of the caribou- Robert Rausch had a special place in his heart for the Inupiat˜ of the dependent Inupiat˜ of the inland region, whose descendants are now settled Arctic Coastal Plain and Anaktuvuk Pass in the Brooks Range of Alaska, at Anaktuvuk Pass (Rausch, 1951). He worked in a time and place that establishing enduring friendships through living on the land in the pursuit was unique, and in a region of the world already undergoing substantial IN MEMORIAM 549 cultural and environmental transition. Years later, Robert Rausch was the were central in defining the ecological context for transmission of first recipient of the Arctic Science Prize (1984) established by the North multilocularis and distribution of alveolar hydatid disease Slope Borough to honor those who have made critical contributions to in northern communities. Prior to the definitive studies by Robert Rausch northern biology and the well-being of people in those sensitive and Everett Schiller in 1951, there was no consensus about the environments on which increasingly fragile cultures remain dependent. independence of E. multilocularis and (Rausch Robert Rausch began his northern biomedical work as Senior Assistant and Schiller, 1951, 1954; Tappe et al., 2010). These studies established the Scientist in the Public Health Service, Anchorage, in the Territory of modern foundations for exploring diversity within this important group of Alaska, and later became Chief of the Infectious Disease Section at the zoonotic taeniids. Further, effective measures for control emerged from AHRC in Fairbanks (1967–1974). Along the way and later were academic investigations, initiated in early 1949. A basic realization was the focal posts at the University of Alaska (1967–1975), the Department of nature of zoonotic E. multilocularis that he and his research team at the Veterinary Microbiology in the Western College of Veterinary Medicine, AHRC had identified on St. Lawrence Island and adjacent areas of the University of Saskatchewan (1975–1978), and in the Departments of north slope of Arctic Alaska. Dr. Rausch and colleagues were Pathobiology and Comparative Medicine, University of Washington instrumental in revealing the ecological context for transmission and (1978–2012), where he most recently held an appointment as Professor perpetuation of alveolar and cystic hydatid disease in northern commu- Emeritus. He served in an advisory capacity to numerous national and nities and polycystic hydatid disease in South America (Rausch and Fay, international committees for the World Health Organization, National 2002; D’Alessandro and Rausch, 2008). Science Foundation, and the National Academy of Sciences, addressing Further in the public health arena, Robert Rausch led investigations issues related to public health, zoonotic diseases, and polar biology. Dr. that defined the diversity of Diphyllobothrium tapeworms and their Rausch was the President for the American Society of Parasitologists circulation among people in western Alaska (Rausch, 1954; Rausch et al., (ASP) (1984) and participated on the Council for the Wildlife Disease 1967; Rausch and Hilliard, 1970; Rausch et al., 2010). These studies Association (WDA) (1976–1978). Among his numerous and notable resolved the interactions and relationships of strictly marine and honors were the Henry Baldwin Ward Medal (1961) and Eminent freshwater cycles for species of Diphyllobothrium and the ecological Parasitologist for the ASP (2011), the Meritorious Service Medal mechanisms involved for the transfer and exchange of parasites through bestowed by the U.S. Public Health Service (1965), the K. F. Meyer fishes and estuarine systems which influence the distribution of human Award from the American Veterinary Epidemiology Association (1979), infection (Rausch and Adams, 2000). Concurrently, concerns over the the Distinguished Service Awards from the ASP (2001) and the WDA occurrence and transmission of Trichinella nematodes occupied a theme in (1983), and the Public Service Award (2009) from the American projects linked to public health (Rausch et al., 1956; Rausch, 1970, 1972). Veterinary Medical Association. Honorary degrees were bestowed by The latter culminated in a 2008 paper defining the ecological conditions of the University of Saskatchewan (1985), University of Alaska (1987) and transmission for Trichinella among marine and the role of the Universitaet Zeurich (1992), and he was recognized as the accelerated climate change as a determinant of changing patterns of Distinguished Alumnus for the Ohio State University (1994). Addition- infection and potential for human exposure for this zoonotic helminth ally, the Robert and Virginia Rausch Visiting Professorship was (Rausch et al., 2008). Among many examples across terrestrial and aquatic established in 1978 in the Western College of Veterinary Medicine to environments, that paper exemplifies the considerable scope and depth of promote a discussion and education in parasitology and related concepts ecological knowledge that Robert Rausch brought to all facets of his for veterinary and human medicine. research. The importance of contributions by Robert Rausch addressing Over a career spanning 60 years, Robert Rausch was at the forefront of zoonotic helminths in aquatic and terrestrial environments especially research exploring systematics, , biogeography, epidemiology, across the northern latitudes cannot be overestimated (e.g., Jenkins et al., and pathology of helminth parasites among vertebrate hosts. Dr. Rausch 2013). always emphasized what we now regard as an interdisciplinary approach, A considerable legacy of field collections in parasitology and incorporating often disparate observations about place, hosts, and mammalogy lives on through the archives and specimens of the Museum parasites into an expansive view of the biosphere. The result has been of Southwestern Biology, University of New Mexico in Albuquerque, and an extraordinarily detailed and rich picture of biodiversity and the myriad the recently established Division of Parasitology. The Robert and Virginia factors involved as determinants of biotic structure. Parasitology as an Rausch Helminthological Collections (RHC) consist of an estimated integrative discipline is the core message and lesson which continues to 60,000 lots of specimens (wet and permanently mounted) including 40– emerge from diverse studies and insights by Robert Rausch, who was 1 of 50,000 whole mounts and additional histological materials on slides the most perceptive observers of natural history to have graced our representing 60 yr of active research. Curation and conservation of this science. As his students, we were constantly challenged to go beyond vital resource of specimens, information, and literature is underway superficiality, to ask large fundamental questions, to access and use a deep (National Science Foundation [NSF], Division of Biological Infrastruc- global literature, and to chart the directions necessary to provide deeper ture 1057383, ‘‘Integration and Curation of the Robert and Virginia evolutionary, historical, biogeographical and ecological perspectives for Rausch Helminthological Collection: A Resource for Science and Society our observations. in the MSB Division of Parasitology’’). The RHC is considered to be 1 of In papers numbering over 300, Robert Rausch developed important the 3 largest parasite collections in the Western Hemisphere, and is the insights about the systematics, taxonomy, and diagnostics of cyclophyl- first to have databases integrated in the fully interactive Arctos platform lidean tapeworms (e.g., , Echinococcus, and Paranoplocephala), the (see Cook et al., 2005) (http://arctos.database.museum/SpecimenSearch. diphyllobothriids and such nematodes as Trichinella. In the spirit of his cfm). In parallel to the parasite collections there are 4,000 host voucher integrative field biology, there were equally significant taxonomic, specimens in the Division of Mammalogy at the MSB from the RL and biogeographic, and ecological studies of mammals, particularly sorico- VR Rausch Collections. Overall the RHC is a critical resource of national morphs, rodents, and carnivores, all representing critical components of interest in areas addressing integrated studies of emerging infectious broader host–parasite assemblages. His work on mammalian groups, from diseases, zoonotic parasites and pathogens, accelerating climate change, the largest of the brown bears to the most miniscule of shrews, including and general issues of biodiversity assessment, and further serves as an recognition of several previously unknown species (e.g., Rausch, 1963; historical record and baseline for sensitive environments in dynamic Rausch et al., 2007), parallels those in parasitology. Classic and seminal change (e.g., Hoberg et al., 2013). These specimens and records are further observations are frequent among the 50 contributions designated under linked to those of the U.S. National Parasite Collection (USNPC), which the series Studies on the Helminth Fauna of Alaska, which resulted from holds 133 type series and 639 lots of parasites collected by Robert and extensive faunal surveys designed to document diversity. Characterized by Virginia Rausch. meticulous methods and by great innovation and considerable depth, The legacy of the RHC and a lifetime of research by Robert Rausch are these studies articulated and explored important hypotheses designed to evident and exemplified by the expansion of recent studies exploring the reveal a complex web of emphasizing the interface for people, diversity and history of high altitude faunas (e.g., Hoberg et al., 2012). On other mammalian hosts, and our environment. a regional basis, parasites of rodents and carnivores in Alaska, and more Considering taeniids, his studies had a substantial impact on our broadly in the Arctic were a primary focus of parasitological survey and understanding among species of Echinococcus and hydatid disease research by Robert Rausch (e.g., Rausch, 1952, 1957, 1976). Studies on throughout the world (e.g., Rausch, 1967, 2003; Rausch and Bernstein, lagomorphs and the Schizorchis tapeworms (Rausch and Ohbayashi, 1974; 1972; D’Alessandro and Rausch, 2008). Robert Rausch and colleagues Rausch and Smirnova, 1984) set the stage for recent detailed phylogeo- 550 THE JOURNAL OF PARASITOLOGY, VOL. 100, NO. 4, AUGUST 2014

FIGURE 2. Robert and Virginia Rausch and American colleague Kenneth A. Nieland (seated) during field work in far eastern Siberia, on the Taigonos Peninsula, Magadanskaya Oblast, Russia in 1977 (photo: V. L. Kontrimavichus). graphic investigations of parasite faunas of pikas (Galbreath, 2009). Robert L. Rausch was a pioneer in every sense and has become an Further, ongoing studies of tapeworms in arvicoline and other rodents, integral part of the history and landscape of Alaska and the north. Lasting especially among species of Paranoplocephala and Anoplocephaloides, have friendships were established with the villagers at Anaktuvuk Pass in the relied heavily on specimens held in the RHC and USNPC that were central Brooks Range and on the remote shores of St. Lawrence Island in collected in Siberia, Alaska and the continental United States (e.g., the northern Bering Sea. He was often accompanied on these journeys by Haukisalmi et al., 2001, 2002; Wickstrom¨ et al. 2003; Haukisalmi et al., Virginia Rausch, his wife and partner in science through 61 yr. 2006; Haukisalmi and Rausch, 2007; Haukisalmi et al., 2008, 2009; Explorations took Robert and Virginia across many varied corners of Haukisalmi, 2009). A proliferation of new genera and species has emerged, the planet including Siberia, Mongolia, China, Japan, and South America, which is now establishing an expansive view of diversity among arvicoline where research and teaching were the mechanisms for important cultural parasite faunas across the Holarctic (Haukisalmi, 2009). The RHC has and intellectual exchange. They were the first Americans and westerners in also figured very prominently in revealing diversity and cryptic complexes the 1970s invited to travel across remote and formerly closed areas of of species for the hymenolepidid genus Arostrilepis distributed among northeastern Siberia (Fig. 2). Working with Academician Vytus Kon- trimavichus, Dr. Svetlana Bondarenko, Dr. Gennady Atrashkevich, Dr. Holarctic arvicolines. Where once a single widespread and morphologi- Arsenii Krechmar, Mrs. Galena Krechmar, Dr. Daniil Berman, and cally variable species was recognized, integrated studies combining colleagues of the Institute of Biological Problems of the North in morphology and molecular methods now reveal a minimum of 13 species Magadan, Russian inventories of mammalian parasites were conducted (e.g., Makarikov et al., 2013); materials collected by Robert and Virginia around the Chaunsk Gulf, Taigonos Peninsula, and Aborigen Field R. Rausch in Chukotka and Alaska were instrumental in these studies of Station on the Kolyma River. These journeys were instrumental in diversity. In this regard, the decade-long Beringian Coevolution Project opening access to the Russian Far East for many investigators from North (BCP, funded through NSF grants DEB-Biotic Survey and Inventories America and were significant for driving studies of diversity of the 0196095, 0415668 and DEB-Biodiversity Discovery and Analysis- Holarctic fauna (e.g., Rausch, 1994). The Russian gateway, however, was 1258010-1256832-1256493) emerged directly from conceptual foundations initially opened by Robert Rausch in his participation during the depths of and hypotheses established by Robert Rausch about diversity and the the Cold War in the meetings of the All-Union Conference of processes of faunal assembly linking Eurasia and North America during Helminthology hosted by the Russian Academy of Sciences in Moscow the late Pliocene and Pleistocene (Rausch, 1994). In the true Rausch during 1963 and 1968 when he was able to meet the iconic Academician K. tradition, we continue to explore and reveal the tapestry of biogeography I. Skrjabin and his students, who have had such pervasive impact on and evolution that has shaped the structure of complex assemblages of helminthology throughout the world. Coming full circle, Robert and mammalian hosts and their parasites established at the Beringian nexus Virginia Rausch traveled to Moscow in 2004 for an international across the northern roof of the world (e.g., Hoberg et al., 2012). symposium in parasitology honoring the 125-yr Anniversary of K.I. IN MEMORIAM 551

FIGURE 3. Robert and Virginia Rausch and friends at the Laboratory of Helminthology (GELAN) in Moscow, 2004. Robert and Virginia were participating in the International Symposium ‘‘Main Achievements and Perspectives of Parasitology Development,’’ dedicated to the 125-year anniversary of K. I. Skrjabin and the 60-yr anniversary recognizing the foundation of the Helminthological Laboratory of USSR AS, Institute of Parasitology RAS (14–16 April 2004, Moscow). The meeting was arranged by the Russian Academy of Sciences (RAS), Scientific Council on Parasitology RAS, Parasitological Society, All-Russian Society of Helminthologists and Institute of Parasitology RAS., 2004, at GELAN, Moscow. From left to right: Kirill Galaktionov (Zoological Institute, St. Petersburg and Former President of the Parasitological Society), Robert L. Rausch, Virginia R. Rausch, and Gennady Atrashkevich (Institute for Biological Problems of the North, Magadan), under an historical portrait of K. I. Skrjabin (photo: K. Galaktionov).

Skrjabin and the 60-yr Anniversary recognizing the foundation of the Academy of Sciences, St. Petersburg for providing images of Robert and Helminthological Laboratory and a long tradition of field and laboratory Virginia Rausch during field work and meetings in Chukotka and parasitology in Russia (Fig. 3). Moscow. I thank the University Alaska Archives for allowing access to During his distinguished career, Robert Rausch, a man of great images from early field studies during 1949 in the Brooks Range, Alaska. I integrity and principles, served in his lifetime as a singular ambassador for also thank Dr. Joseph Cook, Director of the Museum of Southwestern parasitology in a broad international arena. Robert was a friend to many, Biology, University of New Mexico, for confirming current activities for was honored to be considered an equal among the nomadic bands of databasing and curation of the Rausch Helminthological Collections. Nunamiut traversing the Brooks Range 60 years ago, and will be long remembered for the strength of his science and his perceptive views of the natural world. LITERATURE CITED

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