Suzanne DeChillo/The New York Times Disease-tolerant trees were traced to this ancient elm in Princeton, N.J. ARTICLE TOOLS Suzanne DeChillo/The New York Times Roger W. Holloway, a wholesale nursery supplier, has used Princeton elm cuttings as the genetic stock for his trees. Suzanne DeChillo/The New York Times Elm trees line Washington Road into Princeton, N.J. The trees, planted in the 1930's, have not succumbed to Dutch elm disease. Suzanne DeChillo/The New York Times Above, a leaf of one of the Princeton elms.

From Sturdy Old Survivor, a Hardier Elm Grows

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"Princeton is an excellent American elm," said Keith S. Warren, director of product development at J. Frank Schmidt & Son, in Boring, Ore., one of the largest commercial nurseries in the country. He said that while some people still resist planting elms for fear of disease, overall acceptance is strong and growing.

"Elms create their own atmosphere," Mr. Warren said. "They really give a different feel to a street."

Elms are a vision of steadfastness, a dream of cathedral arches over city streets, a memory of cool shade on a blistering summer afternoon. For much of American history they had a devoted following whose many members planted the sturdy giants in front yards and along local streets from the Eastern Seaboard to the western prairies.

Then came the disease, imported in a shipment of European logs, and the infatuation turned deadly. Because they were so well loved, and had been planted so bountifully, elm trees turned out to be quite vulnerable. The fungus raced from tree to tree through intertwined roots or on the back of the peripatetic elm bark beetle.

Mr. Holloway remembers the elms from his boyhood and thought they were gone forever until he happened to see one while flipping through a catalog nine years ago.

The tree came from Princeton Nurseries. It used to be located in Princeton, but as land became more valuable, the business was moved to Allentown, N.J. Princeton University now owns most of the land that belonged to the nursery, including the part that includes the stunning row of elms that the nursery planted along Washington Road in the early 1930's.

A few years ago, after Mr. Holloway started experimenting with growing cuttings of the Princeton elm he got through the catalog, he visited Princeton and saw the Washington Road trees himself. "I'd never seen anything like that," he said. "It was just awesome."

Convinced that he had found elms that could resist disease, Mr. Holloway tried to grow them from cuttings; doing so was not easy, but it ensured that the disease tolerance of the parent would be passed on. Failures meant the death of thousands of trees, but he eventually perfected a method, which he declined to reveal.

Along the way, he heard stories about the origins of the Princeton elm, and investigated its background. Several people told him, with great certainty, that the man who headed Princeton Nurseries at the time, William Flemer Jr., used seedlings from the cemetery tree to develop the Princeton elm.

"That's the mother lode," Mr. Holloway said when he revisited the tree in April. Besides its great height and girth, the tree has a brownish-cinnamon tone common to Princeton elms.

"This tree has been here for 300 years while elm trees all around it were dropping like flies," he said. So he checked in with Claude G. Sutphen, 72, the third generation of his family to run Princeton Cemetery. Mr. Sutphen confirmed that yes, he had heard the story linking the big elm and Princeton Nurseries and did not doubt it for a second.

"This was always a big strong tree," Mr. Sutphen said. To prove just how long, he rummaged around his office for a few minutes and came back with an 1854 photograph of Aaron Burr's grave.

The elm is already massive.

The final clue came from the genetic decoding. To make a case for saving the trees during a road-widening project that was eventually scuttled, a Princeton resident sent Mr. Kamalay, the molecular biologist, cuttings from a dozen trees, including the Washington Road elms, the cemetery giant, and other elms, but kept their identities hidden.

Mr. Kamalay prepared DNA profiles of each, then compared them. The findings were incontrovertible. The DNA of the cemetery tree was identical to that of the elms along Washington Road. And their DNA matched that of the elms sold by Princeton Nurseries, which provided the genetic stock for Mr. Holloway.

There is just one problem. The son of the man who planted the elms more than 70 years ago said he was not sure the original came from the cemetery tree.

"Could be," said William Flemer III, now 82, "but if so, my father never mentioned it."

Mr. Flemer, who is retired but keeps his hand in the running of Princeton Nurseries, recalled that his father grew a batch of elm seedlings, and weeded out those with imperfections until just one was left, and that became the Princeton elm.

Mr. Flemer seemed to accept the findings about the cemetery tree, but not to embrace them. Nor did he endorse Mr. Holloway's quest for the mother of all Princeton elms. "If he wants to say the seed came from there, let him," Mr. Flemer said. "There's no one who's going to naysay it."