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''Shortia galacifolia'', the Oconee bells or acony bell, is a rare North American plant in the family
Diapensiaceae Diapensiaceae is a small family of flowering plants, which includes 15 species in 6 genera. The genera include '' Berneuxia'' Decne. (1 species), '' Diapensia'' L. (5 species), '' Galax'' Sims (1 species), '' Pyxidanthera'' Michx. (2 species), '' ...
found in the southern
Appalachian Mountains The Appalachian Mountains, often called the Appalachians, are a mountain range in eastern to northeastern North America. The term "Appalachian" refers to several different regions associated with the mountain range, and its surrounding terrain ...
, concentrated in the tri-state border region of
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,
North Carolina North Carolina ( ) is a U.S. state, state in the Southeastern United States, Southeastern region of the United States. It is bordered by Virginia to the north, the Atlantic Ocean to the east, South Carolina to the south, Georgia (U.S. stat ...
, and
South Carolina South Carolina ( ) is a U.S. state, state in the Southeastern United States, Southeastern region of the United States. It borders North Carolina to the north and northeast, the Atlantic Ocean to the southeast, and Georgia (U.S. state), Georg ...
. Additional populations (some of them naturalized) have been found in
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Tennessee Tennessee (, ), officially the State of Tennessee, is a landlocked U.S. state, state in the Southeastern United States, Southeastern region of the United States. It borders Kentucky to the north, Virginia to the northeast, North Carolina t ...
,
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, and
Massachusetts Massachusetts ( ; ), officially the Commonwealth of Massachusetts, is a U.S. state, state in the New England region of the Northeastern United States. It borders the Atlantic Ocean and the Gulf of Maine to its east, Connecticut and Rhode ...
. ''Shortia galacifolia'' has also been reported in Japan, as have three other species of ''Shortia''.


Description

Creamy-white flowers appear from late March to May. Seedlings appear in late July and early August. This is about 6 weeks after the maturation of the capsule.
Stolon In biology, a stolon ( from Latin ''wikt:stolo, stolō'', genitive ''stolōnis'' – "branch"), also known as a runner, is a horizontal connection between parts of an organism. It may be part of the organism, or of its skeleton. Typically, animal ...
s have shallow roots. It prefers loamy soil with some sand and/or clay. It prefers areas where the annual rainfall is . Plants can tolerate bright light for only 2–3 hours per day at midday. The leaves have an orbicular shape with wavy edges plus a heart-shaped base. Seeds are slender and oval, with a light to medium brown color. Young plants are usually within of mature plants, but have been found as far away as . The stalks grow high and the plants prefer shade and soils with a high
humus In classical soil science, humus is the dark organic matter in soil that is formed by the decomposition of plant and animal matter. It is a kind of soil organic matter. It is rich in nutrients and retains moisture in the soil. Humus is the Lati ...
content.


Habitat

Within its small range, ''S. galacifolia'' is invariably found along rivers and in gorges where the land is sloping and shows evidence of natural or man-made disturbance: mud slides, erosion, trees knocked down by wind, logging, etc. ''Shortia galacifolia'' often forms a dense mat that may prevent seeds of other species from embedding in the soil and germinating. Its decayed vegetative matter may also have a toxic effect on other species. Consequently, it is often found as the only or one of few species of ground cover in a given area. It is found at elevations from . The pattern of elevation distribution varies widely from one watershed to another. Tree species and genera commonly associated with ''S. galacifolia'' are: ''
Tsuga canadensis ''Tsuga canadensis'', also known as eastern hemlock, eastern hemlock-spruce, or Canadian hemlock, and in the French-speaking regions of Canada as ''pruche du Canada'', is a Pinophyta, coniferous tree native plant, native to eastern North America. ...
'' (eastern hemlock), ''
Betula lenta ''Betula lenta'' (sweet birch, also known as black birch, cherry birch, mahogany birch, or spice birch) is a species of birch native to eastern North America. Description ''Betula lenta'' is a medium-sized deciduous tree reaching tall, except ...
'' (sweet birch), ''
Acer rubrum ''Acer rubrum'', the red maple, also known as swamp maple, water maple, or soft maple, is one of the most common and widespread deciduous trees of eastern and central North America. The U.S. Forest Service recognizes it as the most abundant nati ...
'' (red maple), ''
Liriodendron tulipifera ''Liriodendron tulipifera''—known as the tulip tree, American tulip tree, tulipwood, tuliptree, tulip poplar, whitewood, fiddletree, lynn-tree, hickory-poplar, and yellow-poplar—is the North American representative of the two-species genus ...
'' (yellow poplar), ''
Liquidambar styraciflua ''Liquidambar styraciflua'', commonly known as the American sweetgum among other names, is a deciduous tree in the genus ''Liquidambar'' native to warm temperate areas of eastern North America and tropical montane regions of Mexico and Central A ...
'' (sweetgum), and ''
Fagus Beech (genus ''Fagus'') is a genus of deciduous trees in the family Fagaceae, native to subtropical (accessory forest element) and temperate (as dominant element of Mesophyte, mesophytic forests) Eurasia and North America. There are 14 accepted ...
'' (beech genus). It is less commonly seen with: '' Quercus prinus'' (chestnut oak), ''
Quercus alba ''Quercus alba'', the white oak, is one of the preeminent hardwoods of eastern and central North America. It is a long-lived oak, native to eastern and central North America and found from Minnesota, Ontario, Quebec, and southern Maine south as ...
'' (white oak), ''
Pinus strobus ''Pinus strobus'', commonly called the eastern white pine, northern white pine, white pine, Weymouth pine (British), and soft pine is a large pine native to eastern North America. It occurs from Newfoundland (island), Newfoundland, Canada, west ...
'' (eastern white pine), '' Oxydendrum'' (sourwood), and ''
Robinia ''Robinia'' is a genus of flowering plants in the family Fabaceae, tribe Robinieae, native to North America. Commonly known as locusts, they are deciduous trees and shrubs growing tall. The leaves are pinnate with 7–21 oval leaflets. The flowe ...
'' (locust tree). Common understory species include '' Rhododendron maximum'' (greater rhododendron) in moister conditions and ''
Kalmia latifolia ''Kalmia latifolia'', the mountain laurel, calico-bush, or spoonwood, is a flowering plant and one of the 10 species in the genus of Kalmia belonging to the heath(er) family Ericaceae. It is native to the eastern United States. Its range stretch ...
'' (mountain laurel) in drier conditions. Several factors have made ''S. galacifolia'' an
endemic Endemism is the state of a species being found only in a single defined geographic location, such as an island, state, nation, country or other defined zone; organisms that are indigenous to a place are not endemic to it if they are also foun ...
relict A relict is a surviving remnant of a natural phenomenon. Biology A relict (or relic) is an organism that at an earlier time was abundant in a large area but now occurs at only one or a few small areas. Geology and geomorphology In geology, a r ...
species. It reproduces successfully only in disturbed areas. Light and soil conditions that are beneficial to ''S. galacifolia'' change to its detriment as forest canopy and understory get re-established. ''Shortia galacifolia'' has been successfully cultivated as far north as Grand-Métis, Quebec.


Gray's search for a specimen

''Shortia galacifolia'' is a relict evergreen
herb Herbs are a widely distributed and widespread group of plants, excluding vegetables, with savory or aromatic properties that are used for flavoring and garnishing food, for medicinal purposes, or for fragrances. Culinary use typically distingu ...
which long bewitched
Asa Gray Asa Gray (November 18, 1810 – January 30, 1888) is considered the most important American botany, botanist of the 19th century. His ''Darwiniana'' (1876) was considered an important explanation of how religion and science were not necessaril ...
, the eminent American
botanist Botany, also called plant science, is the branch of natural science and biology studying plants, especially Plant anatomy, their anatomy, Plant taxonomy, taxonomy, and Plant ecology, ecology. A botanist or plant scientist is a scientist who s ...
, a saga detailed in the paper "Asa Gray and his Quest for ''Shortia galacifolia"''. During a month in Paris from mid-March to mid-April 1839, Gray had seen a fragment of the plant in the
Paris Paris () is the Capital city, capital and List of communes in France with over 20,000 inhabitants, largest city of France. With an estimated population of 2,048,472 residents in January 2025 in an area of more than , Paris is the List of ci ...
herbarium,
Jardin des Plantes The Jardin des Plantes (, ), also known as the Jardin des Plantes de Paris () when distinguished from other ''jardins des plantes'' in other cities, is the main botanical garden in France. Jardin des Plantes is the official name in the present da ...
. He had long sought it in the wild in the mountains of
North Carolina North Carolina ( ) is a U.S. state, state in the Southeastern United States, Southeastern region of the United States. It is bordered by Virginia to the north, the Atlantic Ocean to the east, South Carolina to the south, Georgia (U.S. stat ...
. Gray's diary entry for April 8, 1839 records him seeing the specimen, which he believed was a new genus. That specimen had been discovered by
André Michaux André Michaux (' → ahn- mee-; sometimes Anglicisation, anglicised as Andrew Michaud; 8 March 174611 October 1802) was a French botanist and explorer. He is most noted for his study of North American flora. In addition Michaux collected specime ...
, who identified the place where he found the specimen as "High Mountains of Carolina." There has been disagreement as to whether Michaux's original collection site was in
Transylvania County, North Carolina Transylvania County is a County (United States), county in the U.S. state of North Carolina. As of the 2020 United States census, 2020 census the population is 32,986. Its county seat is Brevard, North Carolina, Brevard. Transylvania County co ...
at the confluence of the Horsepasture and Toxaway rivers, or in
Oconee County, South Carolina Oconee County is the westernmost County (United States), county in the U.S. state of South Carolina. As of the 2020 United States census, 2020 census, the population was 78,607. Its county seat is Walhalla, South Carolina, Walhalla and its lar ...
along the
Keowee River The Keowee River is created by the confluence of the Toxaway River and the Whitewater River in northern Oconee County, South Carolina. The confluence is today submerged beneath the waters of Lake Jocassee, a reservoir created by Lake Jocasse ...
at Jocassee. At the time it was thought to be one of the last living specimens of the plant, with fruits but no flowers. Much of the area around Jocassee and the Keowee River was covered by up to of water after the Jocassee Dam was completed in 1973. In Michaux's journal for December 8–11, 1788, he says he found the specimen near the headwaters of the Keowee, near where two rivers join together. Prior to the plant's rediscovery, Gray made several unsuccessful trips to this region, the last one in 1876. A specimen of ''S. galacifolia'' was not rediscovered until May 1877 on the banks of the Catawba River in McDowell County, North Carolina by George McQueen Hyams (1861–1932). His father, M. E. Hyams (1819–1891), collected medicinal herbs and sold them to a
Baltimore, Maryland Baltimore is the List of municipalities in Maryland, most populous city in the U.S. state of Maryland. With a population of 585,708 at the 2020 United States census, 2020 census and estimated at 568,271 in 2024, it is the List of United States ...
drug firm. The Hyams did not know what they had found. Eighteen months later they sent the specimen to lawyer and botanist
Joseph Whipple Congdon Joseph Whipple Congdon (April 13, 1834 – April 5, 1910) was a lawyer by trade who contributed significantly to early botanical exploration in California, particularly in the Yosemite region, where he resided in Mariposa from 1882 until 1905. C ...
, who contacted Gray, telling Gray that he felt he had found ''Shortia''. For the preceding 39 years, Gray had been enraptured by his search for a living example in the field. When a specimen was finally placed in Gray's hand as proof, he exclaimed: "Now let me sing my ''
nunc dimittis The Nunc dimittis (), also known as the Song of Simeon or the Canticle of Simeon, is a canticle taken from the second chapter of the Gospel of Luke, verses 29 to 32. Its Latin name comes from its incipit, the opening words, of the Vulgate transl ...
''." Gray wrote about the rediscovery to his colleague William M. Canby on October 21, 1878. Gray wrote to the elder Hyams, on October 27, 1878, telling him the great news. He noted that because the Hyamses had waited eighteen months before contacting Congdon, they had missed the chance to have their discovery included in a new botanical book published in the meantime. In spring 1879 Gray led an expedition, in which the Hyamses assisted, to the spot where ''S. galacifolia'' had been found. Gray's final trip to this region was in 1884. In Gray's diary entry for April 8, 1839, he named the genus after Charles Wilkins Short, because the plant was native to America in a region close to where Short lived in Kentucky. Short and Gray never met, but they corresponded with one another frequently. Short never saw a live nor dried specimen of his namesake genus. The ''galacifolia'' part of the name means "galax-like leaves" because its evergreen leaves are shaped like leaves in the genus '' Galax.'' Gray made his first field trip to find a wild specimen from late June to late July 1841. Gray never saw ''Shortia'' in its native habitat while it was in bloom. He also never visited the area of the Keowee River. The 2011 U.S. postage stamp honoring Gray also depicts this species.


References


Further reading

*
Gray, Asa. 1878. ''Shortia galacifolia'' rediscovered. Botanical Gazette 4(1):106-109.
{{Taxonbar, from=Q867748 Diapensiaceae Endemic flora of the United States Flora of the Appalachian Mountains Flora of the Southeastern United States Oconee County, South Carolina Endangered flora of the United States Plants described in 1842 Taxa named by John Torrey Taxa named by Asa Gray