Bud Lanyon, Fellow and Past President of the American Ornithologists' Union, left us on June 7, 2017, after a long battle with cancer. He was an extraordinary ornithologist, a giant on whose shoulders many of us still stand. A superb field biologist, his legacy includes pioneering studies of the use of vocalizations to diagnose cryptic species of meadowlarks and flycatchers. He authored 102 publications and was awarded the AOU's highest honor, the Brewster Medal, in 1968. Above all, he was a positive and gracious role model who showed us the way through his rigorous solutions to complex problems, his keen intellect, and his unforgettable, dry sense of humor.
Born in Norwalk, Connecticut, on June 10, 1926, Bud grew up in the countryside near Hanover, New Hampshire, home of Dartmouth College. His early memories of the spring return of meadowlark songs presaged his later research on them. In seventh grade, at age 13, Bud met Richard Weaver, a naturalist at Dartmouth. Weaver focused Bud's teenage energy on birds, local ornithology, and the AOU. Bud excelled in high school, both academically and as an athlete. He was president of his class and recipient of Hanover High School's Balfour Medal for the outstanding senior student. Following graduation, Bud enlisted in the navy and served as a radio operator from 1944 to 1946. Undergraduate years at Cornell University (1946–1950) followed. There, Donald Griffin, Arthur Allen, and Paul Kellogg, among others, stimulated Bud's lifelong interest in recording bird songs.
While serving as an interpretive specialist for the National Capital Parks in Washington, D.C., during the summer of 1947, Bud met Vernia Elizabeth (Vickie) Hall. They married on January 29, 1951 (her birthday), in the nation's capital.
Following graduation from Cornell in 1950, Bud launched his research career with award-winning graduate studies of meadowlarks (Sturnella) under John T. Emlen at the University of Wisconsin. Whether Eastern Meadowlarks and Western Meadowlarks were subspecies or sibling species was hotly debated then. They had different primary songs, but geographic variation and possible hybridization defied morphological resolution of the question. Bud's integrated study of meadowlark behavior and ecology in southern Wisconsin was one of the first to test the biological species concept in a zone of contact and possible sympatry. He discovered that the meadowlarks were interspecifically territorial, then a new observation. He documented the absence of hybridization, the role of female choice as a premating isolating mechanism, and the sterility of rare F1 hybrids. In their groundbreaking study of the ontogeny of meadowlark vocalizations, he and Vickie pioneered the techniques for hand rearing songbirds, including the use of a novel dietary formula that is still in use. Bud proved that the different songs of Eastern and Western meadowlarks were learned, but that the chatter calls, which were key behavioral isolating mechanisms, were innate. Bud's research on meadowlarks spanned 30 years (1952–1982), with expansions to meadowlark populations in Central and South America. His unpublished monograph On a Lark or Two details this extraordinary research program.
Following completion of his Ph.D. in 1955, Bud held one-year teaching positions at the University of Arizona (1955–1956) and Miami University of Ohio (1956–1957). Having joined the AOU in 1947 while at Cornell, he now regularly presented his meadowlark research results at AOU meetings. Dean Amadon took notice and offered him a position at the American Museum of Natural History (AMNH) with the goal of expanding the expertise of the scientific staff to include behavior and technology. Bud joined the AMNH staff in 1957 and remained there for the rest of his career. One of his initial roles was as director of the museum's new Kalbfleisch Research Station in Huntington, Long Island. At Kalbfleisch, he mentored a generation of field biologists funded by one of the first National Science Foundation undergraduate research programs. Vickie played the role of “mom” for 10 to 12 undergraduates each summer. Bud also built a rich field research program, which included the transitions of bird communities that accompanied the succession of farmland into woodland. Sixteen years later, their two children having fledged, Bud and Vickie moved to the city. He chaired a vibrant Department of Ornithology at AMNH from 1980 until his retirement in 1988. Among his leadership roles, Bud spearheaded the training of graduate students by AMNH curators affiliated with the City University of New York. Also under his leadership, the Chapman Fellowship Program became a major source of funds for student research and for competitive postdoctoral positions at the museum.
His meadowlark research had demonstrated that Bud was undaunted by the technical challenges of field studies on cryptic bird species from 50 years previous, and following completion of his Ph.D. research he expanded his research from meadowlarks to flycatcher (Myiarchus) species of the Western Hemisphere. He started with the three species that coexist in southern Arizona by describing their vocal repertoires objectively with first-generation sonographs. He and Vickie then confirmed that vocalizations of hand-reared nestlings were innate, and that dawn songs of adult male Myiarchus were highly conserved characters for species diagnosis.
During his Myiarchus studies, Bud also pioneered the use of reciprocal playback experiments for resolving issues of species taxonomy. Using heavy, primitive tape recorders and speakers, he designed protocols that gave birds the choice between the songs of two different species broadcast from speakers at two locations. After the subject made its choice by responding to the playback from one speaker, Bud reversed the presentations to elicit a confirming reorientation. We now take playback of vocalizations for granted in field studies and global birdwatching.
In addition to the all-important vocal characters, Bud integrated analyses of subtle morphological differences in the color of tail feathers and mouth linings. He carefully paired vocal repertoires with voucher specimens to resolve the identities of historical specimens in museum collections. Step by logical step, with explicit objectives, and with support from colleagues in the many countries he visited, Bud expanded his taxonomic resolutions to Myiarchus species in Mexico and Central America, then to those of the Caribbean, then to all remaining members of the genus in South America. Dozens of expeditions and hundreds of playback experiments later, he defined the identities and geographic variants of all 22 species in Myiarchus. This program was a model of applying the scientific method in field research based on objectively testing clear hypotheses one at a time. Mark Robbins speaks for all of us in his admiration of our colleague: “I consider, to this day, his monograph on South American Myiarchus to be the single best piece of field work ever. If one understands the logistics involved in that incomparable work, you come away in awe of Bud's determination.”
The higher classification of tyrant flycatchers framed the final major chapter in Bud's remarkable research program. The resolution of myiarchine flycatchers as a major clade of the tyrant flycatchers required attention to the rest of this vast family, all without the benefit of access to DNA data. So, Bud applied the new doctrines of cladistic analysis to the classification of the flycatchers using characters of the skull and syrinx. Field studies of the enigmatic Flammulated Flycatcher (Deltarhynchus flammulatus) confirmed that it had a myiarchine nest as well as skull morphology. Mourners (Rhytipterna) had myiarchine skull characters, but a different nest. In all, he examined the skulls of 88 of 90 tyrannid genera, syringes of all the genera, and nest behavior of 81 of the genera to build the first phylogenetic classification of these suboscines.
Upon retirement, Bud celebrated his 30-year research commitment to the flycatchers by applying for a vanity license plate: MYRKUS. His request was denied at first because the clerk in charge thought it might be pornographic! But Bud prevailed, as usual.
The AOU was Bud's main professional society, starting with his first visit as a teenager in 1946. He served the society in many roles: Council member, committee chair, vice president, and president (1976–1978). He was a natural leader and diplomat. His infectious humor served him well. No attendee will forget his memorable banquet talk in 1977 (and again in 1983 at the AOU's centennial celebration). Rather than a heavy review of some timely topic, Bud opted for a humor-filled reading of Joel Allen's feisty letters to colleagues in the founding years of the AOU. Proper etiquette and diplomacy required judicious editing of Allen's rich expletives and character assassinations. So Bud substituted a loud mechanical click for each deletion, all eliciting a chorus of laughter from his colleagues. On another occasion (Winnipeg 1975), Bud moderated the paper session in which his son Scott was to present his first AOU paper. Bud deliberately mangled Scott's surname to the delight of all except young Scott, who survived that debut to become president himself (2014–2016), the first father-and-son tandem to lead the AOU.
After retirement, Bud and Vickie moved to the northern Adirondacks of New York, where they, and Scott, built a log cabin by hand on the upper slopes of Little Porter Mountain. From their summer camp, dubbed “Porter House Stake,” Bud quickly became the valued “birdman” of the Keene Valley community. He recruited local citizen scientists to monitor arrival and departure dates of local birds. He also led modern explorations of the poorly known avifauna of Adirondack Park, often by canoe into remote backcountry. Together he and Vickie patiently braved the swarms of black flies to locate the first recorded nest of a Palm Warbler in New York State. With the approach of winter they migrated as “snowbirds” from the Adirondacks to Arizona only to return, like the birds they loved, in the spring. Adventures in their camper trailers mirrored their youthful years in a trailer in Wisconsin. After Vickie died in 2004, Bud settled down in the Victorian, a retirement home on Cape Cod.
Three of Bud's special interests close this tribute to him. First, and not surprisingly given his lifelong interest in new technologies, Apple computers infused Bud's life as soon as they became a household innovation. Bud was a natural computer guru who loved to explore the creative opportunities afforded him by his Macs. He confessed once that he probably would have become a computer programmer rather than an ornithologist if that had been an option. Second, an athlete himself, Bud loved sports. As an avid fan of the Red Sox, Celtics, and Patriots, he followed their wins and losses loyally as a primary pleasure in his final years. Lastly, Bud's privately published Confessions of an Octogenarian, an Autobiography of a Professional Bird-Watcher (2006) details his rich career as husband, father, and ornithologist. Fittingly, Bud dedicated it “in loving memory to Vickie my dear wife, best friend and devoted companion for 53 years.” They were a team, warm and generous to all.
Carrying on Bud's legacy are his son, Scott (vice provost and dean of graduate education and professor, University of Minnesota); his daughter, Cyndy (information systems specialist, Woods Hole Oceanographic Institution); and his four grandchildren, Jeffrey Chandler, Jonathan Chandler, Ashley Lanyon, and Cassandra Lanyon.