주목잎비자나무-Torreya taxifolia Arn.

과명	Taxaceae (주목과)	속명	Torreya (비자나무속)
전체학명	[정명] Torreya taxifolia Arn.	추천명	주목잎비자나무
이명		외국	Cedar Stinking,Stinking cedar

추천명변경: 주목잎비자 -> 주목잎비자나무

Torreya taxifolia, commonly known as Florida torreya or stinking-cedar, but also sometimes as Florida nutmeg or gopher wood, is an endangered subcanopy tree of the yew family, Taxaceae. It is native to only a small glacial refugium in the southeastern United States, at the state border region of northern Florida and southwestern Georgia.

Species discovery

In 1821 colonial control of the Florida Territory shifted from Spain to the United States. Plantation owners and their slaves began to move into the territory, exacerbating conflicts with the native peoples and the existing population of runaway slaves. One such plantation owner was the patriarch of the Croom family, who in 1826 purchased land around the town of Tallahassee. When he died in 1829, his two sons inherited and invested further in the region, buying up or leasing other plantations.[9]

 

Plaque on the tree's discovery by Hardy Bryan Croom (Torreya State Park, Florida)

One of the two sons was Hardy Bryan Croom. Croom had studied law and became a state senator in North Carolina in his early thirties, but he also devoted time to exploring the sciences. Among other scientific interests, he described himself as fond of botany. He assembled a small personal herbarium and authored a monograph on the carnivorous plant genus Sarracenia.

While exploring along the east side of the Apalachicola River, Croom noted that the flora was quite different from elsewhere in the Tallahassee region. Beginning in 1833 he sent dried specimens to herbaria in the north, including that of another glacial relict woody plant found in the same, limited area. That plant is now known as Florida yew (Taxus floridana), and it is also critically endangered. In 1834 Croom initiated correspondence with the botanist John Torrey. Torrey carried forward studies of the species that would eventually carry his name after the tragic death at sea not only of Croom but also of his wife and children. Their shipwreck occurred during a hurricane in 1837, off the coast of North Carolina.

Taxonomy and naming

In 1838 this novel species was described by George Arnott Walker-Arnott from specimens sent to John Torrey and collected in Florida by Hardy Bryan Croom.

Arnott commemorated Torrey in the generic epithet. The etymology of the specific epithet is from Latin taxus, meaning 'yew', and folium, meaning 'leaf': i.e., 'yew-leaved'. Other species of Torreya have longer, less yew-like leaves, but this is not the reason that it was given this name, as the other species were described after this one.

The University of North Carolina Herbarium has a single specimen, originally from the Jesup Herbarium of Dartmouth College, sent by Croom in 1833 from the "Apalache River" (now, Apalachicola River). Curiously, it was first labelled as "Taxus montana Willd.", a South American tree, which was then later changed to Podocarpus taxifolia from southern New Zealand, and finally relabelled as Torreya taxifolia.

Taxonomic and naming changes also occurred when European botanists analyzed herbarium samples. The species was moved to the junior synonym Caryotaxus taxifolia in 1865 by Johann Baptist Henkel and Wilhelm Christian Hochstetter in their monograph on the conifers of the world, Synopsis der Nadelhölzer. In 1873 Karl Heinrich Emil Koch moved the species to Foetataxus taxifolia. In 1891 Edward Lee Greene validated Constantine Samuel Rafinesque-Schmaltz's generic epithet Tumion and erroneously moved this species there as Tumion taxifolium.

 

Plate from Nuttall's The North American sylva, published 1849. It is captioned: "A branch of the male plant, natural size. a. Male amentum. b. Back view of one of the stamens magnified. c. Female ament and ovule, magnified. d. Section of the ripe seed. e. Germinating seed."

In Thomas Nuttall's entry about Torreya taxifolia in his book about American trees, which was published in 1849 although it had been for the most part completed in 1841, he relates that in the correspondence Torrey had sent him, mention had been made of specimens of another species of taxoid tree which had been sent to him by Croom from the same region. To this plant Nuttall "doubtfully attaches the name" Taxus montana, somewhat of a nomen nudum, because Nuttall never actually described the plant besides quoting a summary description from Torrey's letter to him. Nuttall is doubtful about the taxon, because according to him it seems "scarcely distinct" from T. brevifolia of the Pacific Northwest. Following the publication of this work, however, he was attributed as the author of this scientific name. By 1865 this name was misapplied to Torreya taxifolia under the name Torreya montana. Henkel and Hochstetter synonymised this taxon with T. taxifolia in their work mentioned above. According to the Index Kewensis this was in error; the name Taxus montana had actually already been given to a species, now Prumnopitys montana, described (validated, in fact) in 1806 by Carl Ludwig Willdenow from specimens collected by Alexander von Humboldt and Aimé Bonpland on their famous scientific exploration of the Americas, and Nuttall had in fact referred to Willdenow's species. John Nelson, in his more utilitarian as opposed to scientific 1866 horticultural handbook of firs and pines for growing in Britain, introduced the name Foetataxus montana to write about Torreya taxifolia, apparently unaware of the German publication the previous year. In fact, all these sources were wrong, for Nuttall states that he found a newer specimen of Croom's, of the same taxon, in the Herbarium of the Academy of Natural Sciences of Philadelphia labelled as Taxus floridana! Despite that the original synonymy with T. floridana, all these names are still maintained in the synonymy of Torreya taxifolia in some modern databases as of 2020.

Type species of genus Torreya

 

Torreya taxifolia compared with two Asian species and Florida yew

Torreya taxifolia is the type species of the genus Torreya, owing to the timing of its entering herbaria used in western science. The genus has far greater representation in east Asia and also in the mountains of California (Torreya californica) than in its relictual range in Florida.

Family level classification has been controversial, but with genetic analysis it is generally placed in the yew family, Taxaceae. Its closest relative within the yew family is genus Amentotaxus. The genus Cephalotaxus also used to be considered a close relative, but it is now classified within an entirely different family, Cupressaceae.

Vegetative structure among Torreya species is very similar, as seen in the image above. All samples were taken and photographed onsite at Cox Arboretum in Georgia (US). All were young saplings growing in similar light and soil conditions. Florida yew, Taxus floridana, is easily distinguished from Florida torreya by touch: while both genera have pointed leaf tips, the yew tip is soft while torreya is so hardened it easily punctures skin.

Common names

 

Torreya taxifolia new growth 

 

Ripe torreya seeds, October 2013, Mt. Olive, North Carolina

 

One of the original 1939 plantings of Florida Torreya at the Biltmore Estate near Asheville, NC (2004).

The United States government official page listing the endangered status of Torreya taxifolia shows only one common name, Florida torreya. The current (2020) version of the official recovery plan for the species lists three common names, in order: Florida torreya, Florida nutmeg, and stinking cedar. The current (1993) page for this species of the U.S. Department of Agriculture Fire Effects Information System, lists six common names, in this order: Florida torreya, Florida nutmeg, gopherwood, polecat wood, Savin, stinking cedar.

Internationally, the common names listed for Torreya taxifolia are, in order, Florida Nutmeg Tree, Florida Torreya, Gopherwood, and Stinking Cedar. This is in accordance with the latest (2010) update of the species page of The IUCN Red List of Threatened Species. The Integrated Taxonomic Information System lists two common names: Florida nutmeg and Florida torreya.

There is a long history of documented common names that are also paired with descriptions of the species features that may have given rise to such names. One of the first prominent botanists whose documentation of the species includes common names was Thomas Nuttall. Writing in the early 1840s, he proposed the name "yew-leaved torreya." He noted that locally the species was known as "stinking cedar." He ascribed this name to the "strong and peculiar odour" of the timber, especially when it is "bruised or burnt". He also mentioned that the seed, covered in an aril, is approximately the size of a nutmeg.

In 1865 the German botanists Johann Baptist Henkel and Wilhelm Christian Hochstetter noted that the Americans called the tree "stinking cedar" and "wild nutmeg". They explained that the name "nutmeg" is derived from the bone-hard shelled and acorn-sized seeds, which are covered in an aril somewhat similar to that of true nutmeg. They also described that when the leaves are crushed they exude a pungent and disagreeable odour, which is why the local Americans used the name "stinking cedar". They themselves called the plant "Torrey's Nuss-Eibe", which translates into English as "Torrey's nut-yew." According to the British gardening writer John Nelson in 1866, Torreya species in general were known as "stinking cedars" or "stinking nutmegs" by the locals. Recommending the name "mountain yew" for British use, Nelson was unaware that this Florida species grows almost at sea level — and thus nowhere near any mountains Of all species of its genus, Florida torreya is the only one whose historically native habitat is not in mountainous terrain.

 

Portrait of Asa Gray (1810-1888), by George P. A. Healy

In the spring of 1875, Harvard botanist Asa Gray embarked on a trip to the panhandle of Florida, to "make a pious pilgrimage to the secluded native haunts of that rarest of trees, the Torreya taxifolia". The trees observed by Gray during that trip grew up to a meter in circumference and 20 meters tall. Pertaining to its common name, he wrote:

"The people of the district knew it by the name of 'Stinking Cedar' or 'Savine' — the unsavory adjective referring to a peculiar unpleasant smell which the wounded bark exhales. The timber is valued for fence-posts and the like, and is said to be as durable as red cedar. I may add that, in consequence of the stir we made about it, the people are learning to call it Torreya. They are proud of having a tree which, as they have rightly been told, grows nowhere else in the world."

Description

Technical description

Florida torreya will grow as a symmetrical cone-shaped conifer, with leafy branches remaining at ground level, if horticulturally planted and maintained in full sun. LEFT: October 2018 Fred Bess shows Connie Barlow, founder of Torreya Guardians, one of the two male trees he planted in his front yard in Cleveland, Ohio. RIGHT: Close-up of seeds in one of his two female trees.

Torreya taxifolia is an evergreen tree that may reach heights of 18 metres (59 ft) with an 80 centimetres (31 in) diameter trunk, although it typically grew to 9–12 metres (30–39 ft) tall and 30–50 centimetres (12–20 in) in diameter, and most stands today are composed of immature trees of less than 3 metres (9.8 ft) tall. The crown is open and conical in overall shape, with whorled branches. These branches are spreading to slightly drooping. The bark of two-year-old branches are coloured yellowish-green, yellowish-brown or grey. On mature trees the bark is only about 0.5 inches (1.3 cm) thick and is irregularly divided by shallow fissures.

The stiff, needle-like leaves are sharp to the touch and are arranged in two ranks on the branches. Images on this page show their dimensions; sizes vary depending on access to sunlight and the tree's overall health. The leaves are glossy green above and light green below, with a very slightly sunken grayish stripe of stomata on either side of the midrib on the underside, and slightly round in transverse profile on the topside. The leaves have an unpleasant, strongly pungent, resinous odor when crushed.

LEFT: October 2013 the ability of a Florida torreya to bear both male and female cones on the same tree was documented at the home of A.J. Bullard in Mt. Olive, NC. RIGHT: Mature seeds are large and flesh-covered.

Genus Torreya is subdioecious, with separate male and female plants that may however include branches bearing cones of the opposite sex. The male (pollen) cones resemble those of a common yew, but are much larger and have imbricated scales (bracts) at their base. They are 5–7 mm long, grouped in lines along the underside of a shoot. The female cones when young are more angular than the male (as seen in the photo to the right). They are grouped two to five together near the tip of a branchlet. Tiny at first, they mature in about 18 months to a drupe-like structure with the single large nut-like seed (photo at right) surrounded by a fleshy covering called an aril, 2.5–3.5 centimetres (0.98–1.38 in) long including aril, about the size of a nutmeg. The aril is green but becomes streaked with purple as it matures, advancing to orange or fully purple in late fall. Unlike true yews, in which the aril forms a "cup" around the seed, in this plant the aril completely encloses the seed, leaving only a minute perforation at the end. When the aril is removed, the seed bears a striking resemblance to a large acorn.

stems searching for sunlight in different directions.

 

 

 

 

Distribution

Wild population

 

Map showing the counties and locations of populations of Florida torreya along the Apalachicola River. The lone population in Jackson County is marked with an X, as it no longer exists.

Torreya taxifolia is restricted to limestone bluffs and ravines along the east bank of the Apalachicola River in the central part of the northern Florida Panhandle and immediately adjacent southernmost Georgia. There used to be one small colony west of the Apalachicola at Dog Pond in Jackson County,[34][48][49] but it no longer exists.

This endemic tree grows along the Apalachicola River just south of the confluence of the Chattahoochee River and the Flint River, which drain the southern Blue Ridge Mountains, a subrange of the Appalachian Mountains. It occurs only in the Florida counties of Gadsden and Liberty and extends one mile into Decatur County, Georgia. The area in which it naturally occurs is 203 square kilometres (50,000 acres), stretching 35 kilometres (22 mi) along the Apalachicola River.

Alvan Wentworth Chapman, a leading botanist of his time in the southeastern states, wrote in 1885 that "Florida torreya" was found only in "widely separated clumps or groves" and appeared to be "exclusively confined" to the cliffs along the east shore of the river and to "the precipitous sides of the ravines" that dissected those cliffs. "It is never seen in the low ground along the river, nor on the elevated plateau east of it, nor, indeed, on level ground anywhere."

Prehistoric distribution

 

A champion size California torreya in Samuel Taylor State Park, north of San Francisco (2022). For scale, see the person touching the tree.

 

Peak glacial refuges and their famous relict trees, near river-system outlets in eastern North America. Critchfield Spruce went extinct before botanical explorers arrived. Florida torreya is critically endangered. The Franklin Tree (Franklinia alatamaha) went extinct in the wild soon after samples were collected and propagated in Philadelphia, PA.

Torreya taxifolia is a member of an ancient genus at least 160 million years old. Fossil evidence indicates genus Torreya was widely distributed across the Northern Hemisphere beginning in the Jurassic period and as recently as the Pliocene. Like other globally disjunct genera of the Arcto-Tertiary Geoflora, Torreya was forced southward during the cooling of the Plio-Pleistocene epochs. In North America, the genus found refuge as numerous, but small and isolated, populations in California's coastal mountains and the west slope of the Sierras, which still constitute the native range of Torreya californica. Glacial advance was far greater on the eastern side of North America. But there, no mountains offered refuge as far south as the genus apparently needed to retreat. Hence, the only refuge that maintained Torreya through the coldest episodes of the Pleistocene yet also during the warming Holocene were ravines along the east shore of the Apalachicola River of northern Florida. Today, this location entails the entirety of the native range of Torreya taxifolia, thus making it a paleoendemic.

Each of the U.S. Fish and Wildlife Service official documents pertaining to Torreya taxifolia portrays this species as a glacial relict. The 1984 official listing of the species as endangered included this description:

"The Florida torreya and other endemics of the Apalachicola River system have received much attention from scientists and local residents. The relictual nature of this area accounts for the presence of many unique species (James, 1967). During recent glaciations, species migrated southward by way of the Apalachicola River system, which served as a refugium during cooling periods. The Apalachicola River is the only Deep River system that has its headwaters in the southern Appalachian Mountains. With the receding of the glaciers, cool moist conditions persisted on the bluffs and ravines of the Apalachicola River after climatic change rendered the surrounding area much drier and warmer."

The initial recovery plan in 1986 gave more details on the prehistory, stating, "Torreya is a genus of four or five species from Florida and Georgia, California, China, and Japan. The present geographic distribution of the genus is similar to the distributions of several other plant genera. The distributions, together with fossil evidence, suggest that these genera had wide distributions during the Tertiary Period that were subsequently reduced by climatic changes during the Quaternary (James 1961, Delcourt and Delcourt 1975)."

The most recent (2020) recovery plan update states, "Based on fossil records, we can speculate that the geographical range of T. taxifolia included North Carolina and perhaps, it was forced south by glaciers, and when they retreated, it became isolated in small areas of the southeastern United States." This 2020 federal document describes the fossil history in this way: "Fossil records of Torreya are limited to seeds, leaves, and secondary wood of the Upper Cretaceous (Boeshore and Gray 1936, Chaney 1950). The records indicated that the distribution of the genus in past geological times was much wider than the present distribution. A fossil named T. antigua, which has some characteristics in common with T. taxifolia and T. californica, was described from the Mid-Cretaceous of North Carolina and was also collected from the near MacBride's Ford, Georgia (Boeshore and Gray 1936)."

Fossil pollen of genus Torreya and other genera within Taxaceae is generally deemed indistinguishable, one from another, and also from genera within families Taxodiaceae and Cupressaceae. Therefore, it is difficult to support past presence or absence of such genera in geographic locales where macrofossil plant material is rare or absent, even if substantial pollen (as in Quaternary bogs) is available. Nonetheless, because the Torreya genus entails sister species of a strongly disjunct distribution pattern of geographic ranges throughout the Northern Hemisphere, with macrofossil evidence at northerly latitudes during warm episodes of the Tertiary Period, it is clear that the eastern North American species of genus Torreya occupied more northerly habitats during many millions of years of history prior to the Quaternary glaciation.

American beech, Fagus grandifolia, is in its southernmost range in Torreya State Park. In this December 2014 photo in the park, the papery brown leaves of a beech still cling to its branches.

The relictual character of the historically native range of this conifer species has been recognized for more than a century. The 1884 Annual Report of the Commissioner of Agriculture (USA) characterizes Torreya taxifolia (for which "Stinking Yew" and "Savin" are listed as common names): "No doubt the Torreya is a relic of a past epoch, when it may have had a wide range at the time when the elephant and mastodon were denizens of this country."

Botanist Henry C. Cowles visited the area in 1904 and his observations were quoted at length by two colleagues, John M. Coulter and W.J.G. Land in a 1905 paper in Botanical Gazette. Cowles observed that torreya in the wild, "were associated with a remarkable and somewhat extensive group of northern mesophytic plants, and the conclusion is irresistible that Torreya is a northern plant of the most pronounced mesophytic tendencies, and to be associated with such forms as the beech-maple-hemlock forms of our northern woods, our most mesophytic type of association."

Torreya taxifolia - Wikipedia

https://youtu.be/4ui-cqSfY3o