About Torreya taxifolia

Genus Torreya is a primitive member of the yew family (Taxaceae). Six species of this genus are known worldwide: 3 in China, one in Japan and Korea, one in California, and one in the Florida panhandle. The Florida species is by far the most imperiled and is the subject of our concern.

"Comparisons of rbcl chloroplast DNA sequences involving T. californica, T. grandis, T. jackii, T. nucifera, and T. taxifolia indicated that Florida torreya is very distinct from other species, and is most closely related to T. californica and T. grandis (Price 1999). In addition, the DNA sequences suggested that the closest generic relative is the Asian Amentotaxus." — "Torreya taxifolia 5-year review (by US Fish & Wildlife Service)

LEFT: A fleshy sarcotesta surrounds the single large seed of T. tax
RIGHT: Connie Barlow with STATE CHAMPION Torreya californica near Santa Cruz CA, 2005.

Genus Torreya is a mountain species in all locations except Florida. That Florida torreya exists as a remnant population in a well-known "peak glacial refuge" is an indicator that its pre-glacial native range was the Appalachian Mountains (at least the southern portions).

... The Japanese Walnut is very like the American Butternut, while, rather curiously, the Japanese Thuja [red cedar] and the two Chamaecyparis, the Piceas [spruce] and Abies [fir], resemble species of Pacific North America, a region whose flora has little affinity with that of eastern Asia. Torreya is common to the two regions; in America it is one of the most local of all our trees, while in Japan it is abundant in the mountainous regions of the central and southern parts of the empire. — "Notes on the Forest Flora of Japan", by C.S. Sargent, 2000, Arnoldia.
Access fact sheets by American Conifer Society on each species:

     T. nucifera     T. grandis     T. fargesii     T. jackii     T. californica     T. taxifolia

The above fact sheets contain the above maps of where the native ranges of each species would transcribe onto the USDA plant zones (which are determined only by "annual extreme minimum temperature" — and thus have nothing to do with high-temperature limits, rainfall limits, or effects of competitive exclusion in natural settings). Notice how far north T. nucifera would reside; in fact, that species has long been planted in botanical gardens of northern states. And notice where T. taxifolia would be found.

Note: The T. nucifera page above characterizes this species as having an ability that Torreya Guardians documented with our T. taxifolia: "It is a subdioecious plant, with individual trees producing either mostly male or mostly female cones, but usually with at least some cones of the other sex present." A webpage presents the uses and food-processing of Torreya nucifera in Japan.

The Missouri Botanical Garden stewards Torreya taxifolia. Its species-specific Plant Finder page reads, "Native to USDA Zone 8, but probably winter hardy to Zone 5.... Some mature trees which have been planted outside the native range of this tree have grown well. Trees within the native range are under attack from a fungal blight (perhaps a species of Fusarium) which threatens to drive this tree to extinction."

Florida Torreya is a subcanopy species. Excerpt from the 1986 "Florida Torreya Recovery Plan", U.S. Fish & Wildlife Service:

"... The Florida torreya is an understory tree of mature beech-magnolia-pine forests (hammocks) (Harper 1914). The canopy trees are mostly deciduous, but evergreen hardwoods [Southern Magnolia] and conifers are also fairly common. These areas have diffuse sunlight in summer, and a relatively open canopy in winter (Kurz 1938b, Brock 1983).... Other species of Torreya appear to have similar habitat requirements (Burke 1975). Torreya californica is found 'on moist, shaded slopes and along water courses' (Abrams 1940) [see also California Torreya webpage.] Torreya nucifera is an understory element of beech forests in Japan (Ishizuka 1974). Torreya grandis occurs in mixed forests of southeastern China (Lee 1973). Fossils of the genus occur in assemblages of other species indicative of mesophytic forests (Knowlton 1919, Leopold and Macginitie 1972, Raven and Axelrod 1978)..."
     "... Growth following germination is slow. Eight to 12-year-old torreya trees are generally 6-8 feet tall. They become sexually mature when 10 feet or taller (Bowden 1981). Under optimal conditions, growth continues after maturation, attaining heights of 60 feet (Reinsmith 1934). The largest existing tree is one that was moved to Norlina, North Carolina in 1840. It is 45 feet tall with a basal diameter of 34 inches (Turnage 1983)." [Note: It was growing in full sunlight and later declined, being delisted as "national champion" of its species in 2016.]

Torreya taxifolia (often referred to as T. tax or Florida torreya) is an evergreen conifer tree historically found only along a 65 kilometer stretch of the Apalachicola River of northern Florida and the adjacent sliver of southern Georgia. It favors the cool and shady ravines, known as "steepheads," that dissect the high bluffs of the river's east shore. Despite its current extreme endemism, the species was once a prominent mid- and under-story member of its forest community, which includes an odd mix of northern and southern species: towering beech and hickory next to tall evergreen magnolia, and surrounded by stubby needle palm. Note: Click to learn more about the the unique steephead habitats, including the other cool-adapted plants that are now restricted to those locations, some of which are threatened or endangered.

LEFT: "Historically native" range of Torreya taxifolia (along Apalachicola River), marked by orange.

RIGHT: Healthy growth on a young T. taxifolia, following "assisted migration" of this endangered conifer to the mountains of North Carolina.

• Below is how the 1986 recovery plan for Torreya taxifolia (its first) speaks of Florida Torreya's distribution. This plan is online and, in some ways, contains more background information than does the current recovery plan.

   VIDEO: Site Visits to Florida's Endangered Torreya and Yew Trees

Connie Barlow presents 15 years of baseline photos and videos she recorded of Torreya taxifolia and Taxus floridana in their historically native range in Torreya State Park in northern Florida. Photos of spectacular California Torreya trees, recorded by Barlow in 2005, show the potential for Florida Torreya recovery efforts to strive for. Fred Bess shows (in 2014 video) 2 Asian conifers (Cephalotaxus and Cunninghamia) used in landscaping that are Torreya look-alikes. Paleoecological evidence that Florida's Torreya was "left behind" in its peak glacial refuge supports "assisted migration" actions.

63 minutes - assembled & published, January 2016

TORREYA SEEDS CANNOT BE STORED. Torreya's seed is recalcitrant and cannot be stored except via cryo-preservation following laboratory manipulation of tissue culture via "somatic embryogenesis". The thousands of seeds currently being produced ex situ must therefore be used for plantings or will be lost. They cannot be inexpensively stored. In 2018, a paper published in Nature Plants confirmed that a large proportion of plants (especially endangered plants and notably trees) have recalcitrant seeds that cannot be stored: "Seed banking not an option for many threatened plants".

"A Pious Pilgrimage" by Botanist Asa Gray in 1875

      In the spring of 1875, distinguished Harvard botanist Asa Gray embarked upon 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 grew up to a meter in circumference and were as much as 20 meters tall.

"... One young tree, brought or sent by Mr. Croom himself, has been kept alive at New York [Central Park] — showing its aptitude for a colder climate than that of which it is a native — and has been more or less multiplied by cuttings..." article in PDF

EXCERPTS FROM "A PIOUS PILGRIMAGE": "... 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...
    "The largest [Torreya] tree I saw grew near the bottom of a deep ravine; its trunk just above the base measured almost four feet in circumference, and was proportionally tall. But it was dominated by the noblest Magnolia grandiflora I ever set eyes on, with trunk seven and a half feet in girth....Seedlings and young trees are not uncommon, and some old stumps were sprouting from the base, in the manner of the California Redwood.

A.W. Chapman's Torreya Experience, 1885


LEFT: Map published in 1885 of Torreya's native range, in report by A.W. Chapman

RIGHT: Map of 20th century range of Torreya taxifolia, from "Defining Indigenous Species: An Introduction" by Mark W. Schwartz, chapter in Assessment and Management of Plant Invasions, 1997.


Online access to Chapman's report in the April 1885 Botanical Gazette

EXCERPT FROM CHAPMAN: ... Torreya taxifolia "occupies a narrow strip of land extending along the east bank of the Apalachicola River from Chattahooche on the north to Alum Bluff on the south, a distance of about twenty miles, and forming a continuous forest, but in detached and often widely separated clumps or groves, generally mingled with, or overshadowed by, magnolia, oaks, and other native trees. . . It is a wild, hilly region, abounding in rocky cliffs and deep sandy ravines ("spring-heads") and unlike in scenery and vegetation any other part of the low country known to me. To these cliffs, and to the precipitous sides of the ravines, the tree appears to be exclusively confined; for 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. Hence, although the suggestion may appear a startling one, were the trees of the whole region growing side by side in one body, I estimate that an area of a few hundred acres would suffice to contain all of them.
     ... But its chief value is due to the remarkable durability of its wood when exposed to the vicissitudes of climate; for it is credibly reported that some fences constructed of it sixty years ago still remain in sound condition. In consequence of this peculiarity it is now extensively employed by the inhabitants of the surrounding country for posts, shingles, and other exposed constructions. In view of these facts, the future of our Torreya is a matter calculated to excite very grave apprehensions. A tree possessed of such valuable qualities, occupying an area so limited in extent, and in the midst of a population where the old rule of "Let him take who has the power, And let him keep who can" has unlimited sway, is destined, it is to be feared, to ultimate extinction.
     Let us indulge the hope that the interest which is beginning to be manifested in regard to the preservation of our forests generally, may result in measures statutory or otherwise for its preservation.

The Strange Mix of Vegetation Along the Apalachicola River


LEFT: hickory trunk in front of evergreen magnolia canopy, with trunk of American Beech at center.

RIGHT: Several shrubby palm species beneath a large beech in Florida's Torreya State Park, along the Apalachicola River. The botanical mix in the "steephead ravines" in this park is a treasure because the same site served, just 18,000 years ago, as a crucial "pocket reserve" in which the botanical richness of today's southern and central Appalachians took refuge at the peak of the last glacial advance.

   VIDEO: "Glacial Stragglers": Here the Apalachicola is presented as a peak glacial refuge.

FLORIDA ANISE is featured as a species that didn't get very far north post-glacial (timecode 12:48).

FLORIDA YEW (13:50) and FLORIDA TORREYA (14:25) are featured as species that were entirely left behind. (Scientifically, "glacial relicts".)

Quote by Alex Eilts: "Ironically, they are now trapped in the same place that sheltered them in the past. But now as the the climate warms, their refuge — turned prison — may lead to their extinction."

Note: Mark Gelbart hypothesizes that Mastodons, rather than tortoises (or squirrels) were the dispersers of Torreya taxifolia during the Pleistocene. See also his blog contending that Torreya is missing its megafaunal disperser. In the latter blogpost, Gelbart draws from a 1978 book by Charles Wharton, The Natural Environments of Georgia, in which the Torreya's very limited habitat in Florida is described as "torreya ravines." Associated plants are listed: "The dominant trees in a torreya ravine are red maple, southern sugar maple, beech, magnolia, basswood, elm, torreya, and sabal palm. Most of these species have northern affinities and are more commonly found in Appalachian cove forests. Other plants found in torreya ravines also represent species of northern affinities such as strawberry bush, hydrangea, and redbud. Wharton found torreya growing with beech, sourwood, and plum in the Faceville Ravine on the Flint River."

ASSOCIATED SPECIES: "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)." — Silvics Manual, Volume 1: Conifers (Click on Florida Torreya).

Learn why T. tax is at the brink of EXTINCTION

EDITOR'S NOTE: This 1905 publication contains the first suggestion that Torreya's preferred habitat lies northward of its endemic Florida range.  Access online the entire report.


Note that in this 1905 report, the author posits that

"It is 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."

PHOTO ABOVE: In 2013 AJ Bullard demonstrated on his Torreya taxifolia tree in Mt. Olive North Carolina that this species actually will produce both female cones (top branchlet) and male cones (middle branchlet) on a single individual. Lower left branchlet shows vegetative buds.

In an email to Lee Barnes on 9/29/16 Frank Callahan wrote of his mature Florida Torreya trees in Medford OR: "Both of these trees exhibit male and female 'flowers', which is unusual for this taxon." A 1904 issue of The New Phytologist provided strong evidence of male and female reproductive specimens on the same individual of Torreya californica:

   ... The tree of Torreya califoniica at Orton Longueville bears both staminate cones and ovules. Mr. Harding has kindly given me the following account of their distribution. " The ovules and staminate flowers are not confined to one side of the tree, but for some years I have noticed that the ovules are more abundant on the side facing the north, also a sprinkling of male that side as well; but the male is certainly more abundant (four times as many) on the south side. There are less (about half as many) ovules on the south side."

The above paper is very detailed re reproductive structures and timing of development. Here are a few extracts of note:

The genus Torreya now consists of four species of restricted distribution, inhabiting respectively Japan, China, Florida, and California. In Cretaceous times it was much more widely spread, being also recorded from Greenland, France, Bohemia, and other districts. Such a history suggests that the genus, which has been comparatively little studied, is an old one, and might be expected to shew primitive characters.

... The peculiar ruminated endosperm of the seed in its second year of development has been described by Professor F . W . Oliver in this journal and he has drawn attention to its similarity to that of certain paleozoic forms. The vascular anatomy of the seeds, which is unique and isolated among recent plants, and also presents analogies with fossil types, has been discussed by the same author in a paper published in the Annals of Botany in 1903' and based on a lecture delivered before Section K of the British Association at the Belfast Meeting in 1902. These topics will be dealt with in greater detail in a future memoir. A further contribution to our knowledge of the plant was made by Mrs. A. G. Tansley (Miss Edith Chick), who published an account of the structure of the seedlings in the NEW PHVTOLOGIST for May 1902 (Vol. 11. p. 83). The young plants shewed some strikingly primitive characters; centripetal wood was found in the cotyledons, which were lobed and adhered together like those of Ginkgo and the Cycads...

...The male cones appear as minute buds in the axils of the leaves on the part of the shoot belonging to the current year, but they may remain dormant for a long time. For instance on one branch gathered this summer no cones had been developed on the parts of the axis corresponding to the years 1904, 1903 and 1902, while a considerable crop of cones occured on the 1901 wood. On another branch ripe cones were found on the 1903 and 1902 wood. At the base of the cone there are a variable number of pairs of decussating bracts. These get more scarious and filmy as we pass up the axis, and one or more of the uppermost pairs have fimbriated margins. During the winter which precedes their ripening season the young cones are completely ensheathed in their bracts, and it is not till the following spring that the bracts separate at the tip and disclose the sporophylls (Figs. 1, 2, 3). ... In dividing into two cells whilst still enclosed in the pollen-sac the pollen-grains of Torreya agree with those of Cephalotaxus and differ from those of Taxus...

The ovules of Torreya californica occur on the shoots of the current year, especially near the base.... [after the winter] Before the end of April the integument has over-topped the nucellus and the arillus has begun to appear... At the time of pollination, three or four weeks later, a drop of liquid exudes from the micropyle, and in this the pollen grains are caught....

"A Remarkable Colony of Northern Plants Along the Apalachicola River, Florida, and Its Significance"

by H. C. Cowles, 1905

Report of the Eighth International Geographic Congress
Held in the United States


In this association one finds two of our most notable endemic plants — Torreya and Croomia. It seems likely, then that we should regard Torreya taxifolia as a northern mesophytic left stranded to-day only in Florida.

It presumably is one of the plants that failed to follow up the last retreat of the Pleistocene ice, and is preserved here perhaps because of exceptionally favorable topographic conditions.

Another archival piece: Frank Thone contributed a short section on "The Apalachicola River endemics" in the 1926 edition of Naturalist's Guide to the Ameericas:

"There is a colony of Torrey taxifolia, together with a number of other endemic and disjunct species along the bluffs of the Apalachicola River between Chattahoochee and Bristol. Are well drained, but at the same time thoroughly moist, because of the abundant seepage water and the low evaporation rates that prevail. Magnolia occurs among the trees, and scrub palmettos in the undergrowth are plentiful. Torreya grows all over the slopes of the ravines, from stream-bank to rim, wherever there is any shade. It is mostly an undergrowth tree, seldom reaching a height of more than 20 or 25 ft. or a trunk diameter of more than 6 in."


   ABOVE LEFT: vegetative buds.

ABOVE RIGHT: male pollen-producing buds.
(photos by Fred Bess, spring 2016)

LEFT: The male structures release pollen sequentially over an extended period — which could be a crucial adaptation for this species that generally has only male or female structures per each tree. (photo by Clint Bancroft)

      FAR LEFT: Deer tick offers scale for the size of Torreya leaves, which show off their dangerously sharp points (27 May 2016).

LEFT: New-growth leaves are vulnerable until their tips harden. This specimen suffered total herbivory/dieback, but is putting forth a doublet growth from near its base (1 October 2016). The upper will swing into a vertical position, becoming the new main stem, and its twin will become the first lateral branch.

Photos by Daein Ballard, New Hampshire

    Lee Barnes, one of the founding Torreya Guardians (and a native North Carolinian), measures the growth of "Thoreau" T. taxifolia tree, four years after it was planted at Corneille Bryan Native Plant Garden, in Junaluska NC. (photo by Connie Barlow, May 2012). In spring of 2014, Lee rediscovered in his archives a paper he wrote for a graduate course in horticultural science in 1983. You can access a pdf of that paper: "Morphology of Torreya taxifolia".

In 2013, Torreya Guardians began trying to learn how best to encourage symbiotic fungi (mycorrhizae) to grow amidst Torreya's roots in our plantings, so Lee's 1983 observation is prescient: "Seedlings produce extensive root systems before much top growth. Also, numerous individuals have noted that Torreya spp. grow naturally slow. It has also been observed that many species with thick roots grow slowly unless inoculated with mycorrhizae. Currently, investigations are underway to determine if mycorrhizal inoculation will increase the growth rate of rooted cuttings and micropropagated plants." (p. 12)

Note: Four unpublished papers by Lee Barnes and his doctoral dissertation are in the "Literature cited" list of the 1986 "Florida Torreya Recovery Plan.

    Genus Torreya in online Encyclopedia of Life

The Encyclopedia of Life online has a lot of photos of genus Torreya — especially the one Californian and several Asian native species. For example, the photo left of ripe seeds of California Torreya confirms that this sister species has the same seed shape and color as Torreya Guardians have documented of North Carolina plantings of the Florida Torreya. (If you click on the "original" link associated with each photo, you will sometimes find not only the original photo but detailed information on date and place.)

ABOVE: Click on images for links to these Torreya taxifolia archive pages of Hooker's Icones Plantarum, 1840.

PHOTO LEFT: A big Chinese torreya tree is seen at Huaqiao Village of Banqiao Township of Hangzhou City, east China's Zhejiang Province, Nov. 3, 2017. There are more than 300,000 torreya trees in Banqiao Township and planting of the nutritious torreya has increased local growers' income. (Xinhua/Xu Yu)

PHOTO RIGHT: Farmers hold hands around a thousand-year old Chinese torreya tree in Zhaojia Town of Zhuji City, east of China's Zhejiang Province, Nov. 7, 2017. Zhuji City is a major producer of Chinese torreya with the total output reaching over 2,500 tons in 2016.

  "Environmental Status of the Stinking Cedar, Torreya taxifolia", by Richard Stalter and Steve Dial, 1984, Bartonia 50: 40-42.


Annual Report of the Commissioner of Agriculture 1884, Report of the Botanist, Geo. Vasey, pp. 126-27.
Included within this section is a report by A. H. Curtis of field trip findings.

    Apalachicola River Endemics

Section of Naturalist's Guide to the Americas, 1926, Ecological Society of America.

Excerpt from

50th Anniversary celebration of career of John Torrey

The American Naturalist

p. 45


Click above for a 2-minute video of Torreya State Park in n. Florida.

WWW www.TorreyaGuardians.org

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Could Torreya Take the Place of Eastern Hemlock?

Annotated List of Papers/Reports Online re Assisted Migration