Grace 'Betty' Lotowycz, shown in her WWII Women Airforce Service...

Grace 'Betty' Lotowycz, shown in her WWII Women Airforce Service Pilot uniform, died April 8, 2016, at age 99. She was among 1,074 WASP pilots, the first women to fly for the U.S. military. Credit: Lotowycz Family

Most people knew Grace “Betty” Lotowycz as a woman who would leap at the sight of a few-flowered nutsedge, or gush over the unicorn-shaped inflorescence of wild chamaelirium luteum.

But few knew that the co-founder of the Long Island Botanical Society and longtime curator at the Planting Fields Arboretum had once been among a band of pioneering women who flew into aviation lore during World War II, and who were mostly erased from history just as quickly.

Lotowycz, 99, who lived in Mill Neck until 2003, died April 8 at a retirement home in Louisville, Colorado.

But not before living a series of adventures that helped redefine femininity in America.

Born in Manhattan, she took up rock climbing and mountaineering in the 1930s, while attending upstate Vassar College.

She graduated with a degree in botany in 1938, in an era in which less than 4 percent of American women were college-educated, and went climbing in the Swiss Alps with the newly formed Experiment in International Living, an organization seeking to advance peace through student-exchange programs.

Upon her return, she began to focus on her botanical career, working at the Brooklyn Botanic Garden and as an editorial assistant at Life Magazine.

But when the War Department, later renamed the Department of Defense, began hiring licensed women pilots during World War II to free up men for combat, Lotowycz found an aviation program in Pennsylvania and learned to fly.

She was accepted into the Women Airforce Service Pilots program, formed in 1943 to provide female pilots for noncombat military flight duties in the United States. The 1,074 WASP pilots, the first women ever to fly for the U.S. military, served as test pilots, towed target dummies and ferried planes among military bases.

The program was both exhilarating and dangerous — 38 WASP pilots died in service — and drew women who would not be dissuaded by society’s male-oriented strictures. Fellow WASP pilot Carla Horowitz, 93, of Manhattan said one of her fellow pilots hid BBs in her clothing to make a weight requirement.

“Were we daring? Yes,” said Horowitz, who trained with Lotowycz. “We had all had a little flying experience, and we loved it.”

But almost as soon as the program began, it was over. Hounded by the skepticism of male military leaders, the WASP program was disbanded by the end of 1944, less than two years after it had begun. In a further indignity, records of the WASP program were sealed for more than 30 years, obscuring the women’s contributions from history.

The records were finally opened in 1977, when a Pentagon announcement that it would begin training women military pilots for the first time in history drew outraged protest.

President Jimmy Carter signed legislation that year granting former WASPs veteran status — although with limited benefits — despite opposition from the American Legion and other veterans groups. In 2009, President Barack Obama awarded the WASP pilots the Congressional Gold Medal at a White House ceremony, saying, “The Women Airforce Service Pilots courageously answered their country’s call in a time of need.”

But family members said Lotowycz, who attended the ceremony in a wheelchair, did not dwell on her military service, or history’s 33-year snub.

“My mother moved on from that and didn’t live in the past,” said her daughter, Helen Rising of Brightwaters.

Lotowycz met Wladimir “Bill” Lotowycz, himself a World War II sub-hunter pilot, while the two worked for Pan American World Airways’ airfreight division. They were married in 1946. When he was offered a transfer to Damascus, Syria, she packed up their two children and moved there, until civil unrest forced them to go home.

They eventually settled in Mill Neck. In 1962, she began working at Planting Fields, where she established a herbarium of preserved plants that currently numbers about 10,000 specimens.

At the age of 88, she co-authored a book, ‘‘Illustrated Field Guide to Shrubs and Woody Vines of Long Island.’’ Co-author Barbara Conolly described Lotowycz as a botanist who often vocalized her enthusiasm upon finds.

In addition to Rising, Lotowycz is survived by daughters, Sophia Stoller of Boulder, Colorado, Mary Ball of upstate Locke, and Ann Lotowycz of Mill Neck.

Her ashes will be interred at a private ceremony in June at St. John’s of Lattingtown, in Locust Valley.

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