About Torreya taxifolia

Narrative Summary of Torreya's History and Growth Characteristics in PDF
Created by Connie Barlow in November 2020 for new torreya seed planters
The most up-to-date (while short, 4 pages) summary presentation on this website

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."

   PHOTO LEFT: Harvesting the seeds of Torreya grandis in China, full article here.

• 3-minute Chinese VIDEO on the processing and harvesting of Torreya grandis seeds.

Editor's note: Because the seeds are harvested green and when oblong, it is possible that harvesting pre-maturation ensures that the shell beneath the fleshy sarcotesta remains tender enough to chew. The shell hardens in fully mature seeds, whose fleshy sarcotesta turns a purple and orange color.

COMPARISON of live Torreya species branchlets.


Above photos by Connie Barlow:
LEFT: Cox Arboretum, Canton GA, elevation 1200 feet, NW Georgia, March 2019
RIGHT: Yosemite National Park, California, May 2005 (full report)

CAUTIONARY NOTE: Barlow's experience onsight with live specimens (horticulture and wild in USA) suggests to her that the branchlet forms vary more widely within a Torreya species than between them. Even the same genotype will express differently, depending on habitat conditions (especially thrival and access to sunlight).

Genus Torreya was part of the Arcto-Tertiary Geoflora (circumpolar) prior to the Quaternary cooling.

Notice Torreya in the following two lists, which appeared in the 1988 book by Jonathan D. Sauer, Plant Migrations: The Dynamics of Geographic Patterning in Seed Plant Species. Europe and California lost the most tree species of the warmer geoflora during the Plio-Pleistone. East Asia retained the most, with eastern North America coming in second place. Genus Torreya survived in 3 of the 4 geographic regions (missing in Europe now).

SOURCE: Appendix to "Hemisphere-scale differences in conifer evolutionary dynamics", by Andrew B. Leslie et al., 2010, Proceedings of the National Academy of Sciences.

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.]


PHOTOS: Left was the Norlina NC Tree in 1975; growth form in full sun. Right is the largest of the 3 original plantings at Caroline Dormon Nature Preserve in central Louisiana; growth form beneath a deciduous canopy.

CALIFORNIA TORREYA INFORMATION helpful for understanding Florida Torreya: USFS "Fire Effects" species page on Torreya californica contains useful information.

EXTRACT: Male California nutmeg bear their microsporophylls within strobili. In contrast, the ovules of female trees are not contained within strobili but are solitary [16]. Male strobili begin growth the year prior to flowering, while females trees develop ovules in one growing season [21]. Torreyas are wind pollinated [16]. Male trees must normally be within 75 to 90 feet (23-27 m) of female trees in order to effect pollination [24]. Seed production is erratic. Good seed crops may be followed by crop failure the following year [10]. Seeds mature in 2 years [19]. Being heavy, seeds usually fall near the parent plant; wind dissemination is rare [17]. Seed predation by Steller's and scrub jay is high [10]. Seeds require a 9- to 12-month stratification period before germination [21]. In one study, seeds stratified for 3 months before planting took an additional 9 months to germinate under greenhouse conditions. Ninety-two percent of seedlings germinated at that time. [15]. Temperature regimes during the stratification period were not noted. Seeds sometimes germinate without stratification but do so slowly [21].

Growth of trees in the understory is slow [10]. Sudworth [24] reported trees from 4 to 8 inches (10-20 cm) in diameter were 60 to 110 years of age, while those from 12 to 18 inches (30-46 cm) in diameter were 170 to 265 years old.. The growth rate needs further study, however, as rates of over 1 foot (30 cm) per year have been reported in cultivars [3]. Preliminary data obtained from tree-ring counts of saplings on the El Dorado National Forest shows some trees attained heights of 4.8 feet (1.5 m) in 28 years [10].

California nutmeg sprouts from the roots, root crown, and bole following damage to aboveground portions of the tree [3,10,19]. Some nutmegs reproduce by layering [21], but the layering capacity of California nutmeg is unknown.

SUCCESSIONAL STATUS: California nutmeg is very shade tolerant [9] and is found in late seral and climax communities [3]. Following disturbance such as fire or logging, sprouts growing from surviving perennating buds appear in initial communities [10].

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.


2015 EXPERIMENT LAUNCHED by Jason Richardson, PhD Student, Department of Integrative Biology, University of South Florida.
Feb 3, 2015, Jason Richardson wrote to Lee Barnes: "... I would like a pack of seeds. I assume Connie or Jack had me added to this list. If you are unfamiliar, I am a PhD student at the University of South Florida. For my dissertation, I am examining the effects gopher tortoises have on plant community composition, and also how they affect germination of seeds in their diet. I am particularly interested in how they may enhance germination of rare and endangered plant seeds, and if they may be used in conservation efforts for such plants, and thus I emailed the Torreya Guardians about obtaining seeds." February 9: "I will be quantitatively testing germination rate and percentage as well as qualitatively examining the seed coat structure via electron microscopy. These two methods will be done both for ingested and non ingested seeds. I had not considered the stratification effects, could I get a pack of 20 of both stratified and non? More is always better for statistical analyses, but I think this will be a good start to see where it leads."
     Note by Connie: Lee sent Jason seeds fr the 2014 harvest, but they were already cleaned of the fleshy covering that would be what tortoises are interested in. JACK SENT 2015 SEEDS with green flesh.
2020 REPORT OF RESULTS by Jason Richardson via email to Connie Barlow:
July 21, 2020, Jason Richardson wrote to Connie Barlow: "Seeds were unable to be consumed by gopher tortoises (too large). Any possible chelonian seed disperser is probably extinct. Seeds were fed to a sulcata tortoise at Zoo Tampa. Some were crushed but we did manage to collect a good number of them passed seemingly intact. I unfortunately had no germination success for these seeds, nor for the unpassed seeds. I have since graduated and am no longer working in science. If you are interested I can give you some other researchers who may be interested in trying to replicate or further this work."
     Supplemental information by Connie Barlow: (1) Because Torreya seeds usually require 2 winters of cold stratification before germinating, it is possible that less time and/or insufficient cold were available for study prior to the need to publish results. (2) Richardson's PhD thesis can be found in full
here; torreya seed experiments are not mentioned in it, however, "Seeds from the two fleshy-fruited species, O. humifosa and P. angustifolia, germinated in significantly greater proportions and faster after gut passage than seeds that did not pass through the gut." (3) Wikipedia entry on Sulcata Tortoise: "The African spurred tortoise (Centrochelys sulcata), also called the sulcata tortoise, is a species of tortoise, which inhabits the southern edge of the Sahara desert, in Africa. It is the third-largest species of tortoise in the world, the largest species of mainland tortoise, and the only extant species in the genus Centrochelys."



Crucial aspects for understanding the natural history of Florida Torreya is to bear in mind the GEOGRAPHY of the southeastern USA (specifically, direction and distance of major river flows) and FLOATING CAPACITY OF THE LARGE SEED. These topics are covered in more detail on our webpage that focuses on the PALEOECOLOGY OF FLORIDA TORREYA.

EXCERPTS: ... 7A. Perhaps T. taxifolia did successfully migrate back and forth between the Apalachicola and the Appalachians throughout previous climate cycles in the Pleistocene — but the arrival of humans in the Holocene interglacial period became an obstacle. Paul S. Martin in 2004 posited that fire-setting by humans could have disrupted forest continuity along the river corridor. Connie Barlow in 2004 suggested that over-hunting of squirrels (the current dispersal agent) by first peoples settled along the Apalachicola River could have contributed, or perhaps over-hunting of a large tortoise species that might also have distributed seeds. The wikipedia entry on Torreya taxifolia cites Connie Barlow's "evolutionary anachronisms" paper as positing an extinct tortoise or other reptilian disperser. Note: Mark Gelbart offers sound arguments for mastodons to have been the primary seed disperser of Torreya, in a 2014 blogpost: "Mastodons, not Giant Tortoises, were the Probable Dispersal Agent of Torreya Seeds".

7B. An alternative hypothesis is that the large-seed of Torreya (which is sometimes capable of floating for several days) easily caught a fast and obstacle-free river ride southward from the Appalachian Mountains by way of the Chattahoochee River at the onset of cooling during the Pliocene or Pleistocene. But there was no river-flow way to return north during any of the interglacial warmings. In the first half of the 20th century, the add-on of human-caused warming finally surmounted the physiological threshold of genus Torreya, and it was no longer able to ward off more than a half-dozen native diseases. The fact that the Florida Yew, Taxus floridana is also a subcanopy rare endemic limited to the same relictual range as Florida Torreya suggests that river-flow assistance may truly be limiting. After all, the seed of Florida Yew is bird-dispersed; dispersal by birds offers faster and greater long-distance distribution than can squirrels or tortoises.

The Florida Torreya and the Atlanta Botanical Garden, by David Ruland, in Conifer Quarterly, pp. 10-14 (2007). Note: As of 2019, David Ruland is Greenhouse Manager for Atlanta Botanical GArden.

EXCERPTS: ... According to fossil records, the Florida torreya is estimated to be over 165 million years old. Like many other conifers with such an impressive age, it was once scattered throughout the northern hemisphere. Scientists theorize the species was driven south by glaciers that once covered the northern latitudes. When the glaciers retreated, the Florida torreya was left isolated in small microhabitats of the southeastern United States, where it thrived for thousands of years.

... Many of the original Arnold Arboretum cuttings have matured into cone and seed producing trees [at Atlanta Botanical Garden] that, in total, form over 500 viable seeds per year on average. These plants are grown in the ABG "seed orchard" and propagules produced from these seeds have been used to facilitate the next phase in the recovery of this species.

... In 2002 ABG initiated a collaborative project with Florida State Park Service that involved reintroduction of seedlings into ravines at Torreya State Park (TSP), where Torreya taxifolia has been extirpated.... Great efforts are made to ensure that introduced plants are not planted in ravines where existing plants occur. The plants are bare-rooted prior to placing in the native soil. Four treatments are used on the outplanting: fungicide, fertilizer only, fertilizer and lime, and control. These experimental transplants will help determine the optimal treatment, if any, that is needed for future success reintroducing this species. A total of 200 seedlings have been outplanted in TSP and the survival rates so far are encouraging.

... Despite the successes of the conservation program, the Florida torreya faces a long road ahead to recovery. Even if wild populations were capable of producing viable seed, the Florida torreya would seem incapable of expanding its limited range due to a lack of a natural dispersal agent. The aforementioned squirrel has proven to be capable of seed dispersal but almost certainly is not the original prime disperser. It was most likely a large extinct animal, although speculation on such matters in the plant world is endless.

... The concern over the Florida torreya's inability to reclaim its former habitats has given rise to a movement among conservationists called "assisted migration." The basic idea is to see populations of T. taxifolia moved further north into more hospitable climates. It would be encouraged to integrate naturally, thereby securing the tree in the wild again. Torreya taxifolia does thrive in areas such as Asheville, North Carolina and even much further north. Indeed, as is the case with many types of plants, the cool night / warm day temperature differential would seem to be conducive to a healthier tree. The Atlanta Botanical Garden is not a proponent of such measures. It is prudent to establish safe-guarded populations in cooler climates within the confines of cultivated or human disturbed areas, not in pristine natural habitats. Therefore these plants can be further evaluated in a botanical garden setting and seed development encouraged without creating further ecological disturbance.

The Florida torreya is a glacial relic, seemingly stranded in an increasingly hostile niche without any natural means of escape or survival. This tree would certainly be doomed without the intercession of concerned individuals and institutions....

• October 2019 / Connie Barlow / NEW VIDEO by Tallahassee Public TV station on Hurricane Michael Damage

Six-minute video titled Torreya State Park After Hurricane Michael: One Year Later was produced by WFSU, the public TV station affiliated with Florida State University in Tallahassee. The video begins with a look at the two unlikely survivors of the hurricane where the entrance road ends in a parking lot. Both Gregory House and a planted little grove of torreya trees at the lawn edge survived, the tall trees fallen all around them.

   For viewers and readers familiar with the paleoecological foundation undergirding the drive for "assisted migration" poleward of the glacial relict Torreya tree, the video offers a few hints of the steephead ravine ecosystem similarities in the park to habitats now found in the southern Appalachians. The actions of Torreya Guardians are of course not mentioned. But the accompanying essay does say this:
"In the 1950s, a fungal blight wiped out a population of about 600,000 Torreya taxifolia in the region. The Florida Park Service, Nature Conservancy, and the Atlanta Botanical Garden have been working to revive the Florida torreya, a species whose future may lie in its likely ancestral home of North Carolina, where planted trees have thrived disease free."

• August 2019 / Connie Barlow / 2018 paper ranks genus Torreya as both very rare and ancient

   Kevin M. Potter, Dept. Forestry and Environmental Resources of North Carolina State University, published a paper in the May 2018 online issue of the journal Biological Conservation:

"Do United States protected areas effectively conserve forest tree rarity and evolutionary distinctiveness?"

Connie Barlow added red type and arrows to the original figure, left.

Notice that the sister species in Florida and California of genus Torreya are among the rarest of 352 tree species native to North America.

These two Torreya species also are among the most ancient tree lineages.

Together, rarity and age of origin call out for the highest levels of conservation attention.

• May 2019 / Connie Barlow / New paper useful for understanding male/female flexibility in Torreya

During our site visit to the mature Florida Torreya trees in Louisiana, our guides recounted their experience with the largest specimen beginning as male and then starting to produce some female buds on various branches — culminating in seeds that fell and germinated beneath the parent tree. A 2019 paper on Striped Maple of eastern North America (a subcanopy species, just as is Torreya) can help us understand how to observe and possibly predict an individual's ability and propensity to begin producing seeds. Access full text.

   EXCERPTS: ... Male-dominated sex ratios occurred consistently across study sites and the 4 years that sex expression was monitored. Approximately one-third of trees [studied as single branches cut and grown in a lab conditions] changed during any 2-year period. The five most common transitions were, in descending order of frequency: from non-reproductive to male, male to full or partial female flowering, female to dead, and from partial to full female flowering....

... We have shown that in the sexually plastic tree Acer pensylvanicum a variety of factors influence expressed sex. Chief among them are previous sex and the health of an individual. Although the general theory regarding ESD in dioecious plants has indicated that females are often found in relatively better condition and at larger sizes, we find the opposite pattern in this species.... We show that mortality is disproportionately high in females....

   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.

1890 Editorial Reports Common Name Switch to Torrey-Tree

MAY 7, 1890 issue of Garden and Forest, page 222 excerpt:

"...Trees, which are usually of importance to man or sufficiently conspicuous to attract his attention, obtain naturally local vernacular names before science imposes others on them, and the common names once engrafted on a language, almost always hold their own among the people of the country where the trees are found. It was a matter, therefore, of much interest and some surprise to hear recently, in western Florida, the Torreya taxifolia, one of the rarest of all our trees, spoken of generally as the 'Torrey-tree'; and to find that Stinking Cedar, the unattractive name by which this tree was first known to the inhabitants of western Florida, was gradually being replaced by that of on of the Nestors of American botany.
     The reason for this change is found perhaps in the fact that this tree, from its rarity, the interest attached to the geographical distribution of the small genus to which it belongs, and the reverence which his successors have always felt for the name of John Torrey, has several times been visited in its remote and isolated stations on the banks of the Appalachicola by men of science from distant parts of the country. When the people of the region, therefore, found that men of mature years and apparently in the enjoyment of all their faculties had journeyed thousands of miles merely to look at a tree which they had always considered as valuable only because it furnished indestructible material for fence posts, their own interest and curiosity became aroused; and therefore hearing these eccentric strangers talking always about Torrey and Torreya, the name has gradually become fixed, and now 'Torrey-tree' may often be heard in at least two or three counties of west Florida."

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







"John Torrey: A Biographical Notice by Asa Gray

American Journal of Science and Arts

EXCERPT ON DISCOVERIES OF 4 TORREYA SPECIES: "... Almost in his youth a genus was dedicated to him by his correspondent, Sprengel: this proved to be a Clerodendron, misunderstood. A second, proposed by Rafinesque, was founded on an artificial dismemberment of Cyperus. The ground was clear, therefore, when, thirty or forty years ago, a new and remarkable evergreen tree was discovered in our own Southern States, which it was at once determined should bear Dr. Torrey's name. More recently a congener was found in the noble forests of California. Another species had already been recognized in Japan, and lately a fourth in the mountains of Northern China. All four of them have been introduced and are greatly prized as ornamental trees in Europe. So that, all round the world, Torreya taxifolia, Torreya California, Torreya nucifera, and Torreya grandis — as well as his own important contributions to botany, of which they are a memorial — should keep our associate's memory as green as their own perpetual verdure."



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

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Could Torreya Take the Place of Eastern Hemlock?

Annotated List of Papers/Reports Online re Assisted Migration