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Florida Nutmeg Tree

Torreya taxifolia Arn.

Comments

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Torreya taxifolia is a rare endemic mainly along the Apalachicola River.

Populations of Torreya taxifolia were thriving until the 1950s, but since then they have been decimated by fungal disease (R. L. Godfrey and H. Kurz 1962). Only nonreproductive stump sprouts remain in the wild. The Florida torreya was listed as federally endangered in 1984 under the U.S. Endangered Species Act, and efforts are underway to reestablish this once thriving species in its native habitat (L. R. McMahan 1989).

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Flora of North America Vol. 2 in eFloras.org, Missouri Botanical Garden. Accessed Nov 12, 2008.
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Description

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Trees to 13(--18) m; trunk to 8 dm diam.; crown rather open-conical. Branches spreading to slightly drooping; 2-year-old branches yellowish green, yellowish brown, or gray. Leaves 1.5--3.8 cm, abaxial side with 2 scarcely impressed, grayish bands of stomates, rounded on adaxial side, emitting fetid odor when crushed. Pollen cones pale yellow. Seed (including aril) 2.5--3.5 cm; aril glaucous, dark green, streaked with purple.
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Flora of North America Vol. 2 in eFloras.org, Missouri Botanical Garden. Accessed Nov 12, 2008.
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Distribution

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Fla., Ga.
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Flora of North America Vol. 2 in eFloras.org, Missouri Botanical Garden. Accessed Nov 12, 2008.
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Habitat

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River bluffs, slopes, and moist ravines; of conservation concern; 15--30m.
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Flora of North America Vol. 2 in eFloras.org, Missouri Botanical Garden. Accessed Nov 12, 2008.
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Synonym

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Tumion taxifolium (Arnott) Greene
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Common Names

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More info for the terms: natural, tree

Florida nutmeg
gopherwood
Florida torreya
polecat wood
Savin
stinking cedar


TAXONOMY:
The scientific name of Florida nutmeg is Torreya taxifolia Arn.
It is a member of the yew family (Taxaceae) [11,17]. There are no
recognized subspecies, varieties, or forms [2].



LIFE FORM:
Tree

FEDERAL LEGAL STATUS:
Endangered [17]

OTHER STATUS:
Florida nutmeg is state-listed as threatened [24].






DISTRIBUTION AND OCCURRENCE
SPECIES: Torreya taxifolia
GENERAL DISTRIBUTION:
Florida nutmeg is endemic to three counties in northern Florida
(Liberty, Gadsden, and Jackson) and extends 1 mile into Decatur County,
Georgia [2,11]. The natural range of this species extends along the
limestone bluffs on the eastern bank of the Apalachicola River and its
tributaries for a 40-mile (64-km) stretch [14]. There is a small colony
of 60 trees approximately 6 miles west of the river at a site known as
Dog Pond in Jackson County [2,11]. Florida nutmeg is not an abundant
species, and local occurrence is widely scattered along the Apalachicola
River [9,11]. There is a small, introduced population of trees located
in Asheville, North Carolina, on the Biltmore Estate [14].
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bibliographic citation
Esser, Lora L. 1993. Torreya taxifolia. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/

Conservation Status

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Florida nutmeg is state-listed as threatened [24].
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Esser, Lora L. 1993. Torreya taxifolia. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/

Distribution

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More info for the term: natural

Florida nutmeg is endemic to three counties in northern Florida
(Liberty, Gadsden, and Jackson) and extends 1 mile into Decatur County,
Georgia [2,11]. The natural range of this species extends along the
limestone bluffs on the eastern bank of the Apalachicola River and its
tributaries for a 40-mile (64-km) stretch [14]. There is a small colony
of 60 trees approximately 6 miles west of the river at a site known as
Dog Pond in Jackson County [2,11]. Florida nutmeg is not an abundant
species, and local occurrence is widely scattered along the Apalachicola
River [9,11]. There is a small, introduced population of trees located
in Asheville, North Carolina, on the Biltmore Estate [14].



Distribution of Florida nutmeg. Map courtesy of USDA, NRCS. 2018. The PLANTS Database.
National Plant Data Team, Greensboro, NC [2018, June 21] [16].

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Esser, Lora L. 1993. Torreya taxifolia. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/

Fire Management Considerations

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More info for the terms: fire suppression, forest, natural

Schwartz, a biologist with the Florida Nature Conservancy, suggested
that in the past, smoke may have acted as a natural fungicide,
suppressing the fungi now infecting Forida torreya. Ground fires
resulting from lightning strikes were a constant feature of the region's
longleaf pine forests until recently. Smoke drifting from these upland
fires settled in the ravines where Florida nutmeg grew. This may have
kept the fungal spore load low. After fire suppression, the spore load
may have reached a critical mass, resulting in the present outbreak
[21]. In August and October of 1987, 2,670 acres (1,080 ha) of a
longleaf pine-slash pine (P. elliottii) forest were burned. Two of the
eleven fungal pathogens identified in stricken Florida nutmeg were
suppressed by smoke [13]. The Tall Timbers Research Station in
Tallahassee, Florida, is currently researching the effects of smoke on
the fungi that infect Florida nutmeg. The research is as yet
unpublished [23].
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bibliographic citation
Esser, Lora L. 1993. Torreya taxifolia. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/

Key Plant Community Associations

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Florida nutmeg is associated with oak-tupelo-cypress
(Quercus-Nyssa-Cupressus) and oak-pine (Quercus-Pinus) forests on the
eastern bank of the Apalachicola River [14]. The longleaf
pine/wiregrass (P. palustris/Aristida stricta) sandhill community is
upslope from these forests [1,21].
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bibliographic citation
Esser, Lora L. 1993. Torreya taxifolia. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/

Life Form

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More info for the term: tree

Tree
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Esser, Lora L. 1993. Torreya taxifolia. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/

Management considerations

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More info for the terms: natural, tree

Florida nutmeg is almost extinct in its natural range [9]. In 1988 the
Center for Plant Conservation stated that Florida nutmeg faces such a
serious and immediate threat of extinction that it will be gone in 10
years unless concerted conservation steps are taken [4]. An intricate
array of circumstances threatens Florida nutmeg. The population is
reduced because of habitat destruction by inundation and logging and
fungal pathogens that kill young trees before they reach sexual maturity
[4,21].

Disease: Florida nutmeg populations are drastically reduced by stem
and needle blights [2]. The fungi responsible for these blights have
been identified as members of the genera Physalospora and Macrophoma.
As many as 11 species of fungi attack Florida nutmeg [9,13]. How the
infection begins is unknown. It may begin with fungi attacking the tree
while the fungi are in their sexual reproductive cycle [14].

Sudden exposure to full sunlight following logging of other tree species
may stress Florida nutmeg, leading to susceptibility to fungal invasion
[12].

Fungicide: Infected Florida nutmeg trees treated with the commercial
fungicide Maneb recovered markedly and produced new growth with little
or no infection [12,14].

Pests: Feral pigs uproot and destroy Florida nutmeg seedlings [20].
Deer preferrentially select Florida nutmeg saplings as antler rubbing
posts, and sometimes kill saplings while rubbing their antlers [21].
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bibliographic citation
Esser, Lora L. 1993. Torreya taxifolia. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/

Phenology

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More info on this topic.

More info for the term: aril

Reproductive structures emerge in March and April [11]. Seeds ripen
from August to October and are released from September to November
[11,14]. Midsummer aril ripening has been reported for Florida nutmeg,
but is not typical [11]. Needles persist for 3 to 4 years [20].
license
cc-publicdomain
bibliographic citation
Esser, Lora L. 1993. Torreya taxifolia. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/

Post-fire Regeneration

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More info for the terms: geophyte, root sucker

Tree with adventitious-bud root crown/soboliferous species root sucker
Geophyte, growing points deep in soil
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bibliographic citation
Esser, Lora L. 1993. Torreya taxifolia. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/

Taxonomy

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The scientific name of Florida nutmeg is Torreya taxifolia Arn.
It is a member of the yew family (Taxaceae) [11,17]. There are no
recognized subspecies, varieties, or forms [2].
license
cc-publicdomain
bibliographic citation
Esser, Lora L. 1993. Torreya taxifolia. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/

U.S. Federal Legal Status

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Endangered [17]
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bibliographic citation
Esser, Lora L. 1993. Torreya taxifolia. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/

Associated Forest Cover

provided by Silvics of North America
Florida torreya is not included among the forest cover types established by the Society of American Foresters but is commonly known to be among the oak-gum-cypress or oak-pine types. In 1919, it made up about 4 percent of the forest along the Apalachicola River. The most commonly associated species are beech (Fagus grandifolia), yellow-poplar (Liriodendron tulipifera), American holly (Ilex opaca), Florida maple (Acer barbatum), loblolly pine (Pinus taeda), spruce pine (P. glabra), white oak (Quercus alba), eastern hophornbeam (Ostrya virginiana), and sweetgum (Liquidambar styraciflua). Shrubs and lianas associated with Florida torreya are poison-ivy (Toxicodendron radicans), greenbriar (Smilax spp.), crossvine (Bignonia capreolata), yaupon (Ilex vomitoria), Florida yew (Taxus floridana), blackberry and dewberry (Rubus spp.). Forbs, grasses, and sedges include sedges (Carex spp.), panic grass (Panicum spp.), partridgeberry (Mitchella repens), little sweet Betsy (Trillium cuneatum), giant cane (Arundinaria gigantea), and American climbing fern (Lygodium palmatum) (4,5,6).

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Climate

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The climate of the region in the Florida Panhandle where Torreya taxifolia grows is warm and humid, with a wet summer and dry fall and winter. Average rainfall is 810 to 860 mm (32 to 34 in) during the growing season from April 1 to September 30, while the average yearly rainfall totals 1420 mm (56 in). The growing season averages 270 days. Killing frosts usually occur between November 25 and February 28. The average January temperature is 12° C (54° F), while the average July temperature is 27° C (81° F). Occasional cold waves in the winter bring a minimum temperature of -9° to -7° C (15° to 20° F). A low of -19° C (-2° F) has been recorded in Tallahassee, 64 km (40 mi) east of the Apalachicola River (10).

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Damaging Agents

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Godfrey and Kurz examined populations of Torreya taxifolia in 1962 and observed that many trees were infected by fungi that cause a stem and needle blight. They reported that many areas where the species previously thrived contained only a few skeletal trunks, some with abortive sprouts at their bases. The fungi responsible for the blight appear to be Physalospora spp. and Macrophoma spp., but investigators have not determined precisely how the fungal agents act to cause the blight. There is speculation that a sexual stage of the causal agent may be necessary to establish the infection and that the condition or age of the tree may be the important factor. Researchers found that the commercial fungicide Maneb at a concentration of 671 g per 378 liters (1.5 lb per 100 gal) applied at weekly intervals resulted in good control over 9 weeks of treatment, and that treated trees recovered markedly and produced new growth with little or no infection (1,5).

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Flowering and Fruiting

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Florida torreya is dioecious. Female flowers are produced in March and April and the ovule develops in a sessile, arillate structure. At the end of the second season, the fertilized ovule forms a single, nearly globose gray-blue fruit 2.5 to 4.1 cm (1.0 to 1.6 in) long, 1.9 to 3.6 cm (0.75 to 1.4 in) wide, which matures as early as August or as late as early November. Staminate cones are also initiated in March and April. These are small, globular-ovate, and bear four pollen sacs on each scale. Torreya taxifolia first produces male and female cones at age 20 (2,8,9).

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Genetics

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No population differences have been observed in this species. No natural hybrids occur because this species is separated from its nearest North American relative, T. californica, by more than 2090 km (1,300 mi).

If Florida torreya is to be preserved, it will be necessary to isolate and propagate blight-resistant trees. Such genetic material may be propagated from Torreya taxifolia cuttings because they root readily. Once the seedlings are well established, they may be outplanted in suitable habitats along the Apalachicola River.

A number of diseased Florida torreya up to 9 m (30 ft) in height are growing in the Maclay State Gardens, FL. There are 14 disease-free specimens on the Biltmore Estate, Asheville, NC, that are more than 40 years old and up to 12 m (40 ft) tall but it is unlikely that they are blight resistant. Seeds and cuttings from the Biltmore Estate should be used to perpetuate disease-free trees inside and outside the species' natural range. All of the species at the Maclay Gardens as well as any accessible trees in their natural habitat should be treated for fungal infection if Torreya taxifolia is not to become extinct.

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Growth and Yield

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The bark on mature trees is about 1.3 cm (0.5 in) thick and irregularly divided by shallow fissures. The dark-brown outer bark often is tinged with orange while the inner bark is yellow. The twigs are green and stiff. The leaves are green, lustrous, stiff, and pungent. The common name, stinking-cedar, is derived from the disagreeable odor given off when any part of the tree is bruised.

Florida torreya is a small tree with whorled branches, reaching 12 m (40 ft) in height and 30 to 50 cm (12 to 20 in) in d.b.h. Its habitat on steep bluffs and its small stature and low population have made its exploitation impractical, and the species has never been commercially important (3,8).

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Reaction to Competition

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Florida torreya appears to grow better in full sunlight at the Maclay State Gardens than in the dense shade of its natural habitat. It may most accurately be classed as tolerant of shade in its native habitat. No information on competition is available, however.

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Rooting Habit

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Florida torreya seedlings have a well-branched taproot. A 5-cm (2-in) seedling produces a taproot 5 cm (2 in) long. No information is available on root growth and the development of mature trees.

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Seed Production and Dissemination

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Little is known about germination of Torreya taxifolia seeds; they may germinate without stratification. According to R. Bowden at the Maclay State Gardens in Tallahassee, FL, some seeds germinated when placed in rich, damp topsoil. At Maclay, Bowden is currently investigating the physiological requirements for Torreya taxifolia germination. He has obtained 80 percent germination of 35 seeds by placing them in wet sphagnum moss (2). Germination is hypogeal. Torreya taxifolia requires an after-ripening period before germination as does its closest American relative, T. californica (8,9).

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Seedling Development

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Little is known about seedling development because few seedlings have been produced in the wild since the species was infected by a blight in the late 1950's. Perhaps mycorrhizae are beneficial to seedling establishment and growth. Seedlings in their natural habitat have developed in the deep shade of hardwoods and pines.

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Soils and Topography

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Florida torreya is restricted to steep, deeply shaded limestone slopes and wooded ravines. Soils in these areas most likely fall within the orders Alfisols and Mollisols.

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Special Uses

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Florida torreya has finely grained wood that is light, hard, strong, and durable. Its specific gravity is 0.5145. Because of its durability, it was formerly used for fence posts. Florida torreya have been cut for Christmas trees, but in 1980 and 1981 there were only a few Florida torreya tall enough to be used for this purpose (7). Observation indicates that animals frequently eat torreya seeds (8,9).

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Vegetative Reproduction

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Florida torreya can perpetuate itself vegetatively by producing sprouts at the base of the parent tree, although, in almost every instance, only one sprout survives after several years. Probably every existing Florida torreya in its present native habitat is a product of vegetative reproduction.

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Distribution

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Florida torreya grows naturally in three counties in Florida, Gadsden, Liberty, and Jackson. It is also found in southern Decatur County, GA, just north of Chattahoochee, FL. The natural range of this species extends along the limestone bluffs for a 64-km (40-mi) stretch on the eastern bank of the Apalachicola River and its tributaries from Chattahoochee south to Torreya State Park in northern Liberty County, FL. One population exists approximately 11 km (7 mi) west of the Apalachicola River in the vicinity of Ocheessee Pond in Jackson County, FL.


- The native range of Florida torreya.

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Brief Summary

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Taxaceae -- Yew family

Richard Stalter

Florida torreya (Torreya taxifolia) is an endangered species. This small rare tree is nearly extinct in the wild, threatened by a fungal disease of the stem. Known locally as stinking-cedar because of the pungent odor given off when the leaves are crushed, it was first discovered in 1833 by H. B. Croom near the Aspalaga Crossing on the Apalachicola River. Its rarity limited its use except locally for fenceposts and Christmas trees. The largest living specimen is in North Carolina and measures about 89 cm (35 in) in d.b.h., 14 m (45 ft) in height, with a crown spread of 12 m (40 ft).

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Physical Description

provided by USDA PLANTS text
Tree, Evergreen, Dioecious, Habit erect, Trees without or rarely having knees, Tree with bark rough or scaly, Young shoots 3-dimensional, Buds not resinous, Leaves needle-like, Leaves alternate, Needle-like leaf margins entire (use magnification), Leaf apex acute, Leaves < 5 cm long, Leaves < 10 cm long, Leaves not blue-green, Leaves white-striped, Scale leaves without raised glands, Needle-like leaves flat, Needle-like leaves not twisted, Needle-like leaf habit erect, Needle-like leaves per fascicle mostly 1, Twigs glabrous, Twigs not viscid, Twigs without peg-like projections or large fascicles after needles fall, Seeds within berry-like cone, Seeds within scales, Aril dark green, Berry-like cones reddish, Bracts of seed cone included, Seeds tan, Seeds brown, Seeds wingless.
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Torreya taxifolia

provided by wikipedia EN

Torreya taxifolia, commonly known as the Florida nutmeg,[1][7][8] Florida torreya,[1][6][7][9][10][11][12][13][14][15] gopher wood[1][6][7][9][10][12][14][15] or stinking-cedar[6][7][9][10][11][12][13][14][15] is an endangered tree of the yew family, Taxaceae,[7][11] found in the Southeastern United States, at the state border region of northern Florida and southwestern Georgia.[1]

T. taxifolia became one of the first federally listed endangered plant species in the United States in 1984;[9][16] the IUCN has listed the species as critically endangered since 1998.[1][17] In 2010 98% of the mature trees of the species were believed to be have been destroyed due to a poorly understood fungal blight as well as inundation due to dams and destruction by deer using trees as antler rubbing posts.[1][7]

Taxonomy

In 1821 the United States had managed to conquer the Florida Territory from Spain after years of surreptitiously trying to usurp Spanish rule, and begun the process of ethnically cleansing the land of its previous inhabitants. Plantation owners and their stocks of Negro slaves had begun to move south to colonise and exploit the new land. One among them was the patriarch of the wealthy Croom family, who in 1826 purchased land around the town of Tallahassee. When he died in 1829, his two sons inherited his holdings and resolved to invest further in the region, buying up or leasing numerous plantations and eventually becoming the largest landowners in the area. They had all of their slaves shipped from their North Carolina plantations to labour at clearing the land, cultivating maize and cotton, and building giant mansions for their slave-masters. One of the two sons was Hardy Bryan Croom, who besides having studied law, being state senator in North Carolina since his early thirties, and his duties running the family and managing his properties, devoted some time to exploring the sciences.[18][19][20][21] Among other scientific interests,[20] he describes himself as fond of botany, had assembled a small personal herbarium[22] and posthumously authored a monograph on the carnivorous plant genus Sarracenia.[20]

East across the Apalache River from the first estate of his brother Bryan Croom in Gadsden, who had acquired it in 1826 and coined it 'Rocky Comfort',[20][21] (another source maintains Hardy Bryan Croom leased the property in 1832), he noted that the flora was quite unique compared to elsewhere in the Tallahassee region,[21] and sent specimens to the herbaria in the north from 1833,[18][23] and corresponded with the botanist John Torrey about the plants here from 1834 onwards.[22] Among the exsiccata he sent north in 1833 were samples of a type of yew,[18] which in his 1834 first letter of reply to an inquiring Torrey he describes as having "little doubt" was the Taxus baccata, the common yew of Europe. His description of the red berries appears to confuse this tree with another rare, local and hitherto undescribed species, T. floridana.[22]

Torrey realised that Croom was quite mistaken, and that the specimens represented a species new to science.[23] Croom and his entire family drowned in a shipwreck off the coast of North Carolina in 1837.[18][19][20][21] The novel species was eventually described by George Arnott Walker Arnott in April 1838 from specimens sent to Torrey and collected in Florida by Croom.[23][24]

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.[3] In 1873 Karl Heinrich Emil Koch moved the species to Foetataxus taxifolia.[25] In 1891 Edward Lee Greene validated Constantine Samuel Rafinesque-Schmaltz's generic epithet Tumion and erroneously moved this species there as Tumion taxifolium.[26][27]

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.[23] Following the publication of this work, however, he was attributed as the author of this scientific name.[3][4] 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.[3] 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.[4] 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.[5][28] 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![23] 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.[2]

The University of North Carolina Herbarium has a single specimen, originally from the Jesup Herbarium of Dartmouth College, sent hither by Croom from the "Apalache River" in 1833. 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.[18]

Higher classification

It is the type species of the genus Torreya. The species, and the genus Torreya in general, has also been placed in the botanical family Cephalotaxaceae.[29]

Etymology

Arnott commemorated Torrey in the generic epithet.[13][23][15] The etymology of the specific epithet is from Latin taxus, meaning 'yew', and folium, meaning 'leaf': i.e., 'yew-leaved'.[12][30] 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.[12]

Common names

It is likely most often known locally by the common name stinking-cedar.[6][7][9][10][11][12][13][14][15] Another local name used is gopher wood.[1][6][7][9][10][12][14][15] Nuttall, writing in the early 1840s, coins the name "yew-leaved torreya" for it, but describes that in the land where it grows in the 1830s, it was known as "stinking cedar", which he ascribes to the "strong and peculiar odour" of the timber, especially when it is "bruised or burnt". He also mentions the seed, covered in the aril, are approximately the size of a nutmeg.[23] In 1865 the German botanists Henkel and Hochstetter note that the Americans called the tree "stinking cedar" and "wild nutmeg". They mention 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 describe 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 call the plant "Torrey's Nuss-Eibe", which translates as "Torrey's nut-yew" in English.[3] According to the gardening writer Nelson in 1866, Torreya species in general were known as "stinking cedars" or "stinking nutmegs" by the locals, though he himself recommended the name "strong-odoured yew" as preferred for British use. This specific species was called "stinking cedar" by the Americans according to him, although he recommended the name "mountain yew" for Britain -this is a calque of the (incorrect) Latin name he was using, he was apparently unaware of the fact that this species grows almost at sea level and nowhere near any mountains![5] In fact, of all species of Torreya, it is only the species which is never found in mountainous areas.

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Seed cone

Although the vernacular name Florida torreya was formally recommended by most conservation works and internet databases,[6][7][9][10][11][12][13][14][15] this was changed to Florida nutmeg in the late 2010s.[7][8] Other names for this tree are savin,[7][11][15] polecat wood,[7][15] yew-leaved torreya,[15][23] foetid yew,[15] stinking cedar,[1][31] mountain yew (Britain)[5] or stinking yew (applied to the Californian species Torreya californica).[5]

Description

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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."[23]

Torreya taxifolia is an evergreen tree that may reach heights of 18 metres (59 ft) with an 80 centimetres (31 in) diameter trunk,[9] although it typically grew to 9–12 metres (30–39 ft) tall and 30–50 centimetres (12–20 in) in diameter,[7] and most stands today are composed of immature trees of less than 3 metres (9.8 ft) tall.[17] The crown is open and conical in overall shape,[9] with whorled branches.[10] These branches are spreading to slightly drooping. The bark of two-year old branches are coloured yellowish-green, yellowish-brown or grey.[9] The young branchlets divide into threes.[23]

The stiff, needle-like leaves are sharp to the touch,[10] 1.5–3.8 centimetres (0.59–1.50 in) long[9] and 3 mm broad. These are arranged in two ranks on the branches[13] and are coloured glossy green above and light green below,[10] with a very slightly sunken grayish stripe of stomata on either side of the midrib on the underside,[9][10] and slightly round in transverse profile on the topside.[9] The leaves have an unpleasant, strongly pungent, resinous odor when crushed.[9][10]

It is dioecious, with separate male and female plants.[1] The male (pollen) cones resemble those of a common yew, but are much larger and have imbricated scales (bracts) at their base.[23] They are 5–7 mm long, grouped in lines along the underside of a shoot. The female (seed) cones are single or grouped two to five together on a short stem; minute at first, they mature in about 18 months to a drupe-like structure with the single large nut-like seed surrounded by a fleshy covering called an aril, 2.5–3.5 centimetres (0.98–1.38 in) long including aril,[9] about the size of a nutmeg.[23] The aril is glaucous and coloured dark green and streaked with purple at full maturity in the fall.[9] 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. The aril is fleshy in consistency, like a fruit.[23] When the aril is removed, the seed bears a striking resemblance to a large acorn.[3][23]

Distribution

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Foilage

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,[1] near the town of Chattahoochee, Florida;[15] there is also a small colony west of the Apalachicola at Dog Pond in Jackson County.[7][32] It grows along the Apalachicola River just south of the confluence of the Chattahoochee River and the Flint River, which reach northwards through Columbus and Atlanta to drain the southern Blue Ridge Mountains, a subrange of the Appalachian Mountains.[23] It only occurs in the Florida counties of Gadsden, Jackson and Liberty, and extends one mile into Decatur County, Georgia[7] (another source states the northernmost wild individual currently growing in Georgia is within 200 metres (660 ft) of the Florida state line). The outlying population west of the Apalachicola, in Jackson County, has always been small: In 1938 H. Kurz reported it to be restricted to one stand with a few trees,[33] in 1974 this was reported as consisting of 60 trees,[7] and in 1989 Mark William Schwartz and R. Nicholson reported it to consist of five trees.[33]

The area in which it naturally occurs is 203 square kilometres (50,000 acres), stretching 35 kilometres (22 mi) along the Apalachicola River.[33]

There is a small introduced population on the Biltmore Estate in Asheville, North Carolina, where it was planted as an ornamental plant.[7]

Prehistoric distribution

A number of sources state that this species has been recorded from fossils which suggest that at one time it grew over much of the eastern United States and that it has existed for 165 million years;[15][31] this is incorrect.[34] Only two fossil examples are known of Torreya in eastern North America. The first was a species with dense, spirally arranged leaves described as Tunion carolinianum by Edward W. Berry in 1908 from the Mid-Cretaceous of North Carolina,[35] the second is only known from a single piece of fossilised wood from the Upper Cretaceous, also from North Carolina, which has been described as Torreya antiqua.[36]

Ecology

It grows at altitudes of 15–30 metres (49–98 ft), mostly on wooded ravines, bluffs and steep,[1][9][11] north-facing slopes.[1] These ravines have nearly permanent seeps. It also occurs in the bottoms of ravines and adjacent floodplains.[11] It grows in the shade under the canopy of larger trees.[1][7][11] The soils of this region are calcareous,[11][23] moist, dark-coloured, sandy loams.[11]

It occurs in two types of habitats restricted to the river drainage, an oak-tupelo-cypress forest or an oak-pine forest.[7] These forests are mostly deciduous, but evergreen hardwoods and conifers are also common.[11] R. M. Harper, who travelled throughout northern Florida by horse and train to document the compositions of various forests, stated in 1914 that the most frequent species in the Apalachicola ravines were Magnolia grandiflora (9.5%), spruce pine (Pinus glabra, 5.6%, American beech (Fagus grandifolia, 4.1%) and the understory Torreya taxifolia (4.0%) and Ilex opaca (3.5%).[33] According to the IUCN the large trees in this habitat are Fagus grandifolia, tulip tree (Liriodendron tulipifera), Acer barbatum, sweetgum (Liquidambar styraciflua), white oak (Quercus alba), and occasionally loblolly pine (Pinus taeda) and Pinus glabra.[1] Magnolias also grow in these woods.[11] Often these woods are hung with vines such as Smilax species and crossvine (Bignonia capreolata).[1] Another rare conifer, the Florida yew (Taxus floridana), occasionally grows with Torreya taxifolia.[1][37] A rare salamander found in these ravines is Desmognathus apalachicolae. The ravines boast the southernmost parts of the ranges of a number of more northern species such as the plants Hydrangea quercifolia, Epigaea repens and Kalmia latifolia and the copperhead snake Agkistrodon contortrix.[37] Upland around these woodlands are flatwoods, which are a longleaf pine/wiregrass (Pinus palustris/Aristida stricta) sandhill plant community.[7][37] It is uncommon in its native habitats, with individuals spaced far apart.[7]

Various animals eat the seeds.[7] Rodents favour the seeds.[11] Squirrels apparently remove the seeds, which they eat, from the arils, which they do not eat, and often store them in caches for the winter, where they may sprout into new trees.[38]

In a 2001 newsletter, the writer Connie Barlow suggested that T. taxifolia may be an evolutionary anachronism similar to the Osage orange (Maclura pomifera) and Kentucky coffeetree (Gymnocladus dioicus), which are thought to have been dispersed by a now-extinct animal. She made this suggestion based on the fact that this species is rare, that it grew well in the cooler temperatures of North Carolina, and her belief that the species was only found on the east bank of the Apalachicola and was unable to disperse to the west bank with the animals she thought might be the main dispersal agents today, squirrels. She theorised than during the last interglacial (the geologically short periods between ice ages) and the proceeding ones, the species grew further north. According to her theories, because there might have been an unknown and now extinct animal in the past which was better suited to eat and defecate the seeds, or otherwise move the seeds, the species was now "stuck" behind the Apalachicola which served as a refuge habitat for a relict population during the long periods of glacial conditions which characterised the Pleistocene. She further supposed that because the arils contain terpenes which are usually toxic for mammals, and the seed's shell is so thin that mammal molars would crush the seeds, that the purported extinct animal might not be a mammal and instead suggested some unknown species of large tortoise as the extinct ecological partner.[38]

Uses

It is has been far too rare to harvest commercially since the 1950s, but it has a beautiful, yellow-coloured, close-grained wood. The timber is lightweight, hard, strong and highly durable.[7] In the 19th century, the tree was harvested for wood that was used as fence posts,[7][11][12] shingles,[11][12] cabinets,[7] Christmas trees,[1][11][7] firewood,[11][12] and as a fuel for riverboats on the Apalachicola River.[1] Fences made of this wood in the 1910s were still good in the 1970s.[7] In the 1830s it was locally abundant enough for the trees to be harvested to be sawn into planks, which were much used in the construction of the village of Aspalaga Landing. In this era it was also recommended as making excellent posts for fencing, not being liable to attack by insects.[23]

When the trunks are damaged, the trees yield a small quantity of pasty, viscous, blood-red turpentine, which can be dissolved in alcohol, but has a very powerful and unpleasant odour.[23]

Cultivation

The tree is well-represented in cultivation, and is widely planted outside of its native range as an ornamental, and is found in private gardens, arboreta and botanical gardens.[11][12]

It was first imported to grow in Europe in 1840. Large trees were to be found in Germany by the 1860s.[3] It is tolerably hardy, but grows very slowly in Britain, and as such was only recommended for collectors.[5]

It has occasionally been planted as a landscape tree around Tallahassee, and one such specimen in Lee, Florida has achieved 30 feet (9.1 m).[11] Some large specimens are grown elsewhere in botanical gardens. The champion tree of the species is in a private garden in Norlina, North Carolina, having a height of 45 feet (14 m), a trunk diameter of 88 centimetres (35 in) dbh and a canopy width of 40 feet (12 m) in 1996.[11][12] It has been said to endure winter temperatures of −31 °C (−24 °F) in North Carolina without much problem.[12][38]

Propagation

Cuttings of Torreya shoot-tips often show fungal contamination. Experimental studies with weekly sprayed applications of a combinations of systemic fungicides, such as thiophanate-methyl combined with zinc and Maneb or thiabendazole, were able to eliminate the fungi from stock plants after four weeks, although these preparations were not US government approved as of 1987.[15]

Tissue culture methods of propagation were being investigated as of 1987.[15]

Conservation

The first person to notice that the trees were dying was the forester L. T. Nieland in 1938, although he never wrote a formal paper about it. In the same year Kurz made a detailed study of the ecology of species and mentioned that there was no danger to the species vanishing from its habitat if lumbering would cease.[13] In 1954 Kurz and R. K. Godfrey surveyed the population and noticed no symptoms of decline. However, in 1962 Godfrey and Kurz, having surveyed the species again, reported that the natural stands of this species, which was historically never widely distributed, were dying off and extinction seemed assured, if not already accomplished.[13][39] By 1962, only non-reproductive sprouts regrowing from the top-killed stumps remained in the wild,[12][15][39] a situation which has persisted throughout the 20th century.[9][11] They attributed this to deforestation due to the lumber trade.[13][39] However, samples of the afflicted trees were sent to the University of Florida, where Erdman West suggested the decline appeared due to a fungal blight of some kind. In a 1967 article S. A. Alfieri Jr. et al. of the Florida Department of Agriculture presented their research of the phenomenon. The disease presents itself as a small, yellowish spot on the needles (leaves) -one or more per leaf, which soon spreads throughout the leaf until it dies, turns brown and falls off. Subsequently the disease spreads into the twig, and all of the needles fall off, leaving only a tuft of new growth at the tip. The disease spreads in stages, first affecting portions, later the entire tree. Severely infected trees eventually loose most of their needles and twigs. Fungal fruiting structures grow on wholly necrotic tissue on the underside of the leaves, twigs and buds. In some cases, the stems of affected or dead trees also showed fungal canker. Young seedlings are also affected, but less severely than older trees. Trees growing in the shade are more affected than those in full sunlight.[13]

In 1985 the phytopathologist Nabih Elias El-Gholl, also of the Florida Department of Agriculture, was able to prove that there were at least two pathogens involved; the leaf-spot disease presented as light greyish-green spots, which later became tan-coloured, and were up to 8.4mm long and as wide as the needle, with brown, irregularly-shaped, 2.4mm long × 2mm wide, necrotic centres and were able to infect the plants in the absence of wounds within three days of inoculation. Initially only a few spots, maximally four, developed per plant. After some two weeks the infected needles died and fell off -by this time the undersides had developed spore structures,[40] which were primarily found along the two grey bands of stomata, but may also occur on the twigs.[15]

The canker disease presents as elongated fusiform swellings on the stems which eventually may break open. They are often found at the base of the trees.[15]

In 1993 Schwartz, who has spent much of his career researching the conservation of this species, and Sharon M. Hermann, reported on the progression of the disease. They monitored a census population of some 100 trees for four years, in this time 10% died, mostly the smaller individuals. Most plants consisted of multi-stemmed, regenerating stumps; the mean length of the longest stems were less than one meter. Less than half of the plants showed any growth in length during these four years, and 32% lost their primary stem, which meant the mean size of individuals within the population was decreasing. Death was not strongly related to specific location or if the plants showed disease symptoms, although stem mortality was higher with trees showing a higher severity of foliar pathogens.[32] In 2000 Schwartz et al. reported that despite Godfrey and Kurz's 1962 prediction, a small population size, continued population losses, and no known seed production for decades, extinction of T. taxifolia had not yet occurred, and although the population would likely steadily decline, it was unlikely to go extinct within the next 50 years.[33]

Status

This was one of the first federally listed endangered plant species in the United States in 1984 (Federal Register 1/23/84).[9][15][16] It is one of only two species of conifer protected under the Endangered Species Act.[12] It was listed as "threatened" within the state of Florida by 1987 (Florida Statutes Section 581.185),[15] again in 1994 by the Florida Game and Fresh Water Fish Commission,[7] and in 1998 the status in Florida was changed to "endangered".[8][10] In 2002 it was added to the list of "endangered" plants of Georgia.[8][11] It has been a "critically endangered" species on the IUCN Red List since 1998, in 2000 this was said to be due to estimated 98% decline in mature individuals since the early 1950s.[1][17]

Population

In 2000 Schwartz and colleagues calculated that before the start of the decline, the original population of T. taxifolia may have been 300,000 between 650,000 plants, with 357,500 individual trees being the preferred estimate, based on a historic abundance of 14.2% of dominant ravine trees in 1914, or 30 trees/ha, and a historic distribution of some 203 square kilometres (50,000 acres).[33] Only 27 trees were counted in Georgia in 1981.[17] In 1993 Schwartz estimated that there were only 1500 trees in the wild.[11][32] Based on the extrapolation of a survey of five stands of the 2000 survey, attenuated with assumptions that these stands were the most dense, Schwartz et al. calculated the population in its native habitat to be between 500 and 4,000 individuals, with none capable of reproduction. The most likely population was calculated to be 1,361 individuals, not including the population west of the Apalachicola. Using their estimate of the pre-decline population, they thus calculated a minimum 98.5% decline in population over the last 50 years.[33] The IUCN estimated that the population is 999 mature individuals in its native habitat (although, confusingly, it also states the population to consist of between 500 and 600 trees elsewhere in the assessment), of which less than ten were known to produce male or female cones.[1] It stated that the population continued to decrease,[1] although the decline may be reversible should the causes of infection be better understood.[1][33] IUCN population viability analyses indicate that extinction within its native range is inevitable.[1]

The IUCN stated the total extent of occurrence is estimated to be about 200 square kilometres (49,000 acres),[1] this is the same as the historical distribution. Schwartz states that these has been no logging in its habitat since the 1950s and that this species has experienced no habitat loss: the vast majority of the original ravine forest habitat remains.[33]

Threats

The most significant current threat to the species is the continued reproductive failure associated with fungal pathogens.[1][14] Individuals do not reach reproductive size before being top-killed.[1] Almost no plants are able to reach a reproductive size, and where seeds do form, these are soft, crumbly and not viable.[7] Up to ten species of fungi have been found growing in infected Torreya taxifolia.[13][14][15][41]

Alfieri et al. were able to isolate five species of fungi in 1967 from the stems and leaves, but failed to isolate any of the fungi from the cankered stems on the media they were using, potato dextrose agar. They reported that the disease appeared to implicate species tentatively identified as Physalospora and Macrophoma, but also isolated the fungi Rhizoctonia solani, Sclerotium rolfsii and a Sphaeropsis species from the infected tissue.[13] They attempted to reproduce the infections by spraying the isolates, as well as pulverised stem and leaf litter, on seedlings grown in greenhouses from unaffected ornamental trees in local parks, but their experiments failed to induce the disease in their seedlings.[13] Nonetheless reports from 1975 and 1988 continued to identify the responsible fungi as these previous two species.[7] In 1985 El-Gholl was finally able to prove Koch's postulates with Gibberella baccata (as Fusarium lateritium, an anamorph isolate) as the agent responsible for the leaf spot pathogen, but he was unable to replicate the disease causing the stem cankers.[14][40][42] In 1987 Alfieri et al. reported on experiments with this fungus, as well as another two more species isolated from the infected plants, Xylocoremium flabelliforme (the anamorph state of Xylaria cubensis) and the Macrophoma which they had re-identified as a Phyllosticta species, the anamorph form of Guignardia. They were able to reproduce El-Gholl's results, but again failed to identify the canker producing agent, the other two isolates proved not to cause any disease.[14][15] In a 1996 article Lee et al. mentioned that the endophyte they were studying, Pestalotiopsis microspora might be the cause of the decline of the species, because a strain they isolated produced a hitherto unknown cytotoxin they named "torreyanic acid", a dimeric quinone.[43] P. microspora is a usually commensal fungus commonly found within the tissues of many plant species, and is only rarely a pathogen -in these cases it is an opportunistic pathogen.[14] However, recent research has identified a previously unknown species of Fusarium may be the cause.[1]

By 1987 the canker disease had spread to plants cultivated outside the native range such as in the University of Florida campus in Gainesville and in the Alfred B. Maclay Gardens State Park in Tallahassee.[15]

According to one older theory logging of shade trees may stress individuals of this species, which does not like being suddenly exposed to full sunlight, conducing infection.[7][13][14]

Another threat to the population is destruction by deer.[1] Deer preferentially select young trees of this species to rub their new antlers on, sometimes killing them. Feral pigs may also uproot and destroy seedlings.[7] Another possible cause of the historical decline may be changes in the environment due to fire suppression, and changes in water tables linked to the construction of dams.[1] Specifically, many trees were killed when the land was flooded[7] in the construction of the Jim Woodruff Dam impounding Lake Seminole.[31]

According to one writer, the population of the species may have been impacted by postglacial global warming, as it may possibly be better adapted to the cooler climate found in Florida during the last ice age. It may not have been able to move north due to poor dispersal abilities. This is based on the fact that the introduced population on the Biltmore estate in North Carolina appears to be doing fine, and is even reproducing naturally -squirrels plant the seeds in the flower beds and these germinate.[38]

Actions

Fungicide treatment with Maneb has been shown to be remarkably effective, with plants showing renewed growth afterwards with little to no fungal infection. It has been applied since 1967,[7][13] but as of 1987 it is not specifically labelled for use in Torreya.[15]

A theory perhaps first put forward in 1990 by Greg Seamon, land manager at the Apalachicola Bluffs and Ravines Preserve, was that Torreya taxifolia may be somehow suffering from the suppression of fire in the upland longleaf pine-wiregrass sandhill ecosystems. However, after regular fires were again instituted in the 1990s in the preserve, this appeared not to have an effect, with the species experiencing an increased three percent annual mortality at the turn of the century.[37]

Two years after declaring the species an endangered species, in 1986, the U.S. Fish and Wildlife Service published a recovery plan. The chief tactic was to produce a genetically diverse collection of trees for reproduction and reintroduction into the wild. Other priorities were to protect the habitat of remaining populations, and to study the disease and methods of propagation.[11]

Plants in general attract very little funding compared to vertebrates, this tree is no different. The US federal government spent only $1,500 on conservation of the species in 1993, placing it 884th out of 926 endangered species. Schwartz et al. stated that recovery is possible, but this will require a greater level of commitment and a dedicated increase in funding.[33]

Small experiments by Schwartz and Hermann in the 1990s found that planting cuttings in the native habitat is possible. Schwartz et al. recommended population augmentation by planting volunteers isolated by more than 100m from extant stands to reduce chance of contamination.[33]

The tree is well-represented in cultivation, and is widely planted for conservation purposes outside of its native range.[11][12] Plants outside of its historic native distribution are undamaged by fungal disease and produce viable seed.[11][12][33] In Tallahassee for example, not far from the native range, a number of large trees had been grown for almost a century in the Alfred B. Maclay Gardens without problems and had been successfully propagated here since the 1960s,[13] although the trees in this park became infected by the canker disease by 1987.[15] It is found in arboreta, botanical gardens and elsewhere throughout the world.[11][12] In the Netherlands, for example, it is grown in the Arboretum Trompenburg, the Arboretum Oudenbosch, the Hortus Haren, the Hortus Botanicus Amsterdam[44] and the Pinetum Blijdenstein.[45]

In the United States the Center for Plant Conservation maintains the species in its National Collection of Endangered Plants; the Arnold Arboretum of Harvard University is the primary custodian of this species.[11] The Atlanta Botanical Garden and the State Botanical Garden of Georgia are actively propagating the plant for conservation purposes. These are members of the Georgia Plant Conservation Alliance, which is protecting the species by growing seedlings and cuttings, and planting these both in situ and ex situ.[11][31][46] Plants are also grown at the United States Botanic Garden.[31]

In her intriguing but somewhat unsubstantiated 2001 newsletter article Barlow synthesized her suppositions to suggest that the only way to conserve the species was for humans to engage in assisted migration, transplanting across great distances. She noted that, in essence, humans are now the gardeners artificially preserving most isolated islands of "wilderness" amongst a sea of human development, if we like it or not.[38] In this she was not unique, in 1994 Foote and Jones had already written that the survival of this species was likely dependent upon cultivation in gardens.[11] The idea is that T. taxifolia for some reason was unable to migrate north from its "ice age pocket refuge" in northern Florida.[47] This inspired the formation of a group, Torreya Guardians, which are attempting to rewild Torreya taxifolia trees in cooler mountain areas in the southern Appalachians in an attempt to aid the species.[48] Some conservationists consider T. taxifolia as the lead candidate for assisted migration.[47] This project has proved contentious.[49]

Protected areas

The natural populations are largely protected within the Torreya State Park[12][13] and at the Nature Conservancy's Apalachicola Bluffs and Ravines Preserve (first parcels purchased in 1982 to protect the ravines, before the species was officially listed as endangered).[37]

References and external links

  1. ^ a b c d e f g h i j k l m n o p q r s t u v w x y z aa ab ac ad Spector, T.; Determann, R. & Gardner, M. (10 August 2010). "Torreya taxifolia". IUCN Red List of Threatened Species. IUCN. 2011: e.T30968A9585489. doi:10.2305/IUCN.UK.2011-2.RLTS.T30968A9585489.en. Retrieved August 15, 2014. Listed as Critically Endangered (CR A2ace v3.1)|date= / |doi= mismatch
  2. ^ a b "Torreya taxifolia Arn". Plants of the World Online. Board of Trustees of the Royal Botanic Gardens, Kew. 2020. Retrieved 16 March 2020.
  3. ^ a b c d e f g Henkel, Johann Baptist; Hochstetter, Wilhelm Christian (1865). Synopsis der nadelhölzer, deren charakteristischen merkmale nebst andeutungen über ihre cultur und ausdauer in Deutschlands klima (in German). Stuttgart: Verlag der J. G. Cottaschen Buchhandlung. p. 367, 368. doi:10.5962/bhl.title.15349.
  4. ^ a b c "Taxus montana". International Plant Names Index. Royal Botanic Gardens, Kew. 2020. Retrieved 16 March 2020.
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  6. ^ a b c d e f g Wunderlin, Richard P.; Hansen, Bruce F.; Franck, Alan R.; Essig, F. B. (17 March 2020). "Torreya taxifolia - Species Page". ISB Atlas of Florida Plants. Institute for Systematic Botany, University of South Florida, Tampa. Retrieved 18 March 2020.
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Torreya taxifolia: Brief Summary

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Torreya taxifolia, commonly known as the Florida nutmeg, Florida torreya, gopher wood or stinking-cedar is an endangered tree of the yew family, Taxaceae, found in the Southeastern United States, at the state border region of northern Florida and southwestern Georgia.

T. taxifolia became one of the first federally listed endangered plant species in the United States in 1984; the IUCN has listed the species as critically endangered since 1998. In 2010 98% of the mature trees of the species were believed to be have been destroyed due to a poorly understood fungal blight as well as inundation due to dams and destruction by deer using trees as antler rubbing posts.

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