Historic Groves of Torreya Trees

Long-term Experiments in Assisted Migration

Evidence of Northward (1) Thrival and (2) Non-Invasiveness

Report by Connie Barlow

August 2018 (last update March 2021)

   SUMMARY: Based on observations and documentation by Torreya Guardians of "historic groves" (where one or more trees were planted north of Florida before the 1984 designation of Florida Torreya as an endangered species), it is reasonable to conclude that Torreya taxifolia is non-invasive and can thrive in locations substantially north of its peak glacial refuge.

• BACKGROUND: The Torreya Guardians in 2004 were just three: Connie Barlow, Paul S. Martin, and Lee Barnes. We were the original guardians in spirit, but our own planting of seeds and seedlings would begin in earnest four years later, when Jack Johnston joined the team.

A side activity to our own plantings and seed distribution is visiting and documenting historic groves of Torreya trees planted in the first half of the 20th century — and well north of the constricted native range in the Florida panhandle.

   Only recently have we come to recognize how important it is to document these historic groves. Such groves inadvertently now serve as long-term experiments for determining in northward states:
1. whether Florida Torreya can survive (even thrive / naturalize) when assisted in migrating northward.

2. whether Florida Torreya is so well adapted to cooler climates that it morphs from "critically endangered" to "invasive."

PHOTO: Jack Johnston at the century-old Harbison Grove, Highlands NC, 2015.

• Click for ANNOTATED LIST OF ACADEMIC PAPERS that use horticultural plantings to identify species that are already severely lagging in Holocene range shifts, including failure to disperse beyond peak Pleistocene relict ranges.

* * * * *

PURPOSE: This page was created August 2018 to highlight how field examinations of historic groves of Florida Torreya (planted north of its peak glacial range in Florida) confirm that
(1) Florida Torreya can thrive and naturalize in northward states.
(2) It will not become invasive if assisted to migrate northward.
     Lack of documentation establishing the combination of thrival and non-invasiveness were the key obstacles to adding an assisted migration action to the management plan for Florida Torreya during the plan update in 2010. The official advisor who objected to such management action is listed as Ms. Tova Spector (Florida Park Service). Her objection is quoted on p. 25 of the 2010 Recovery Plan Update. She stated:
"The reason for not moving Torreya taxifolia outside of its range was addressed by Schwartz (2005). Moving Torreya outside of its range would alter the natural community where it is introduced. In addition the species may be susceptible to decline from factors in the introduced location. Instead trees should be safeguarded in botanical collections until the causal agent(s) for its decline can be mitigated in its historical range."
Because the recovery plan will be updated in 2019, Connie Barlow stepped forward to aggregate the combination of thrival + noninvasiveness evidence established in Torreya Guardians' documentation of historic groves. As well, because Ms. Spector based her 2010 recommendation on a paper by Mark W. Schwartz, 2005, Barlow suggests that the 2019 plan advisors be aware that the Schwartz paper was the second half of a 2-part "Forum" publication, in the Winter 2005 issue of Wild Earth magazine. The first half, not cited by Ms. Spector, argued the pro-assisted-migration viewpoint for Torreya taxifolia. It was written by Barlow and Paul S. Martin, then emeritus professor at the University of Arizona (Pleistocene ecology). Both papers are listed below, along with a 31-coauthor guideline document for "managed relocation" led by Mark W. Schwartz after 3 years of study by a coalition of scientists recruited during a 2008 meeting of the Ecological Society of America. These 3 documents, as well as direct communication with Mark W. Schwartz as to whether managed relocation of T. taxifolia might now be established as safe as well as necessary would be crucial grounding for the 2019 recovery plan advisors.
• Connie Barlow and Paul S. Martin, "Bring Torreya taxifolia North Now", Wild Earth, Winter 2005.

• Mark W. Schwartz, "Conservationists Should Not Move Torreya taxifolia", Wild Earth, Winter 2005.

• Mark W. Schwartz et al, "Managed Relocation: Integrating the Scientific, Regulatory, and Ethical Challenges", BioScience, August 2012.

For an annotated list of papers and news reports on the assisted migration / managed relocation debate, visit this Assisted Migration Scholarly Links webpage.

The table extract below from the USF&WS official Recovery Plan Ad Hoc Report Results documents that an "Inventory of Plantings at Botanical Gardens" is listed as a potential action but that such an inventory has not yet been initiated by institutions associated with the official recovery plan. (Hence, the importance of Torreya Guardians undertaking this effort voluntarily.)


Horticultural Plantings
of Florida Torreya

Sites documented north of Florida
by Torreya Guardians

   A. Naturalized Groves (offspring onsite)

A1. Harbison House, Highlands NC

A2. Biltmore Gardens, Asheville NC

A3. Norlina Tree, Norlina NC

A4. Kennedy Home, Clinton NC

A5. Bullard Home, Mount Olive NC

A6. Caroline Dormon Preserve, Saline LA

B. Mature Trees Producing Seeds

B1. Bess Home, Cleveland OH

B2. Callahan Home, Medford OR

B3. Cheekwood Estate, Nashville TN

C. Mature Trees Not Producing Seeds

C1. Henry Foundation, Gladwyne PA

C2. Spring Grove Cemetery, Cincinnati OH

C3. Chattahoochee River, Columbus GA

C4. Yinger Tree, York PA

C5. Kalmia Gardens, Hartsville SC

D. International Plantings Producing Seeds

List of all EX SITU PLANTINGS (since 1850), 24 pages in PDF, by Paul Camire, 2018.

   Types of plantings/sites documented, 25 pages in PDF:

  • Historic Nurseries that sold Torreya taxifolia: 11
  • Current Nurseries with occasional inventories for intrastate sale: 9
  • U.S. government collections: 3
  • State government / Public Grounds: 6
  • Private homes / gardens: 23
  • International Collections: 14
  • Torreya Guardians seed production (private lands): 2

    Note: There are likely many more mature Torreya taxifolia in existence. Over the years there have been many promoters of the species. Burl Turnage, A.J. Bullard, Bill Alexander, Lee Barnes, Jack Johnston and Frank Callahan have distributed seeds/ seedlings widely in order to save the Florida Torreya. After Croom's discovery of the species in 1833, it became a desired plant at several Victorian-era estates, in the north, from at least 1840. There are bound to be more discoveries of plants in the future, and any additions should be made to this list. Contact the author or Torreya Guardians.

  • PURPOSE OF THE EX SITU LIST: Paul Camire writes, "This list is meant to bring awareness that T. taxifolia has been in cultivation a long time. There is also a possibility that specimens from these nurseries, or descended from these trees, could appear near any of these locations."

    Northern locales of historic nurseries include: Yorkville NY, Rochester NY, Flushing NY, Elizabethtown NJ, Philadelphia PA, Cincinnati OH, Govanstown MD.

    A. Naturalized Groves (offspring onsite)

    A1. Harbison House, Highlands NC  (planted ca. 1920; 6 original trees and >10 seedlings/saplings)

    Jack Johnston measures girth, 2015.

       Webpage - Site Visits and History

    Because this grove and surrounding regrowth forest was left undisturbed for most or all of the grove's century of growth, it is the best site for studying how this tree grows and reproduces in a wild forest. The overtopping deciduous canopy creates deep summertime shade. Absence of other conifers offers full sun in the leafless months.

    Because the trees were planted less than 10 feet from one another, two growth characteristics assist the trees in maximizing photosynthesis: (1) prolific basal sprouts where the trunks face outward from the grove: (2) Lowest branches stretch outward along the ground as far as 20 feet. This suggests the grove may have had full sun originally, but then was overtopped by native Buckeye and Tuliptree, which encouraged the torreyas to reorient growth horizontally.


    1. NON-INVASIVE. This century-old grove melded into the forest, with approx. 10 volunteer seedlings and saplings nearby.

    2. NEEDS HELP MIGRATING WITH CLIMATE CHANGE. Photo left shows a large sapling 40 yards distant from the parent grove. It is the farthest outlier. Such confirms that this large-seeded tree has very low capacity to migrate with climate change. This may be why the four sister species of genus Torreya all live in mountain habitats — where trees can retreat upslope, around to a north-facing aspect, or into a deep shady canyon when climate warms.

    3. NATURALIZES EASILY (at 3,300 foot elevation in western North Carolina).

    4. NO EVIDENCE OF DISEASE. Site visits by Torreya Guardians in 2006, 2015, 2017, and 2021 revealed no discernible evidence of diseased branches on the originals, nor of struggling saplings or seedlings.

    A2. Biltmore Gardens, Asheville NC   (planted 1929; 9 of 12 originals survive and 9 volunteer saplings)

    Michael Dowd measures girth, 2015.

       Webpage - Site Visits and History

    In 1939 Chauncey Beadle supplied the Biltmore Estate with a dozen Torreya taxifolia seeds or specimens collected in Florida — prior to any understanding of climate change and endangered species. Now this 90-year-old grove and its offspring are precious for securing the wellbeing of the species and for demonstrating that (with little human help) North Carolina is an ideal habitat for escaping the lethality of the diseases of a now too-warm Florida.

    Bill Alexander (retired Biltmore Landscape Historian) was the first to advocate (in the 1990s) that Florida Torreya could best be served by moving the species north. He also was the first to donate seeds to Torreya Guardians (via Lee Barnes).

    Note: During the 2004 storms, half of the originals were top-damaged by falling White Pine canopy trees; three were removed. Sun-scalding was relieved only when new, UV-resistant leaves grew. Throughout the grove of originals, herbicides were used to remove poison ivy; a thick layer of long-leaf pine needles followed as ground cover.


    1. BILTMORE TREES SHOW NO SIGN OF INVASIVENESS. The 90-year old grove demonstrates the ability of squirrels to distribute seeds some distance away from the grove (including across the paved road), but numbers and range are far from qualifying as "invasive."

    2. TRUNK GIRTH GROWTH MEASUREMENTS: Despite substantial top-damage from two hurricanes September 2004, there is measurable trunk growth between 2006 and 2015 (see the trunk girth chart).

    3. BILTMORE TREES WITHSTAND FUSARIUM INFECTIONS. The original (1986) recovery plan for Florida Torreya documents that the canker-forming pathogen has, in fact, been producing non-lethal cankers in the Biltmore Grove since (at least) 1986. (p.3):

    The Biltmore trees (32 years later) are still withstanding the Fusarium, as confirmed in a 20 February 2018 email to Connie Barlow from Jason Andrew Smith (University of Florida forest pathologist):
    "There is no doubt in my mind that the primary driver in the mortality of the trees [in Florida] is the pathogen. It is reasonable to assume that it is easily moved around.... We did confirm that the trees at Biltmore Estate in Asheville already are infected, for example ..."
    Therefore, 32 years of documentation of pathogen non-lethal presence is very strong evidence that the Fusarium blight (whether native or exotic in origin) becomes lethal only when climate deterioration severely stresses the Torreya species and/or strengthens the winter survival or summer pathogenicity of the disease agent."
    1. TEST FOR FUSARIUM AT ALL MATURE GROVES. Forest pathology researchers at University of Florida could instruct Torreya Guardians (or site-specific local institutions) on how to (a) detect Fusarium cankers and (b) collect and send specimens to the UF pathology lab. If the Fusarium is present in other groves of mature trees, but not trending toward lethal consequences, then there is no need at this time to genetically engineer Fusarium resistence into the Torreya taxifolia genome. Genetic engineering (using CRISPR technology) was reported as a recommended next step in March 2018 (following a "Torreya Symposium") here and here. Instead ...

    2. ADD ASSISTED MIGRATION TO THE RECOVERY PLAN. There is ample evidence that "assisted migration" into the Appalachian Mountains and other northward regions is the best approach for rapidly moving the species toward "recovery" (hence, de-listing). Historic groves (as reported here) have satisfied "precautionary principle" concerns in that

    (a) the species is now proven to thrive (and naturalize) in northward states and
    (b) the species is non-invasive.

    A3. Norlina Tree, Norlina NC   (planted ca. 1860; many seedling offspring collected and distributed during the 20th century, including for generating new seedlings sold by Woodlanders in Aitken, SC)

    Webpage of the NORLINA TREE

    ABOVE: Photo by Lee Barnes, 1984


    ABOVE: Photo by Paul Camire, 2021

    LEARNINGS: The Norlina Tree is well over 150 years old. Because it has produced seeds (and volunteer seedlings in the surrounds) that have given rise to many of the younger horticultural plantings in North Carolina and northward, it is of central importance. Indeed, it had been listed as the National Champion Torreya taxifolia until its recent stark decline. Many regional experts had assumed it was dead. But Paul Camire (Michigan Torreya Guardian) visited this tree in December 2021. Via text and photos, Paul reveals not only that this tree is still alive but also (a) that spilt herbicide containers stored in its shade likely accounted for its stark decline, and (b) the vegetative details of its gradual return to the world of the living.

    A4. Kennedy Home, Clinton NC   (planted 1850s; 1 of 2 originals remain; seedlings are prolific but mowed or weeded out regularly)

    Michael Dowd uses stick to knock seeds from sunny
    side of mature Torreya tree (31 October 2013).

       Webpage - Site Visits and History

    A. J. Bullard (of Mt. Olive, NC) documented this tree and its larger associate in 1995. As many as 5,000 seeds were collected by Bullard from the pair of trees in a single year. In 1998 a storm blew down a long-leaf pine which crushed the larger, sunnier associate.

    In 2013 (October 31), Connie Barlow collected fallen seeds and dug up six seedlings (delivered to Jeff Morris). In 2014 Jeff Morris observed more seedlings in distant flower beds. He also noticed that a 6-foot-tall basal sprout had grown from the stump of the old blow-down. Whether or not that basal is a continuing pollen source is unknown; nonetheless, the remaining original tree still produces viable seeds.

    The remaining tree is overtopped on the southward side by a deciduous Tilia and a Swamp Chestnut Oak. During the winter a same-size Southern Magnolia (evergreen) blocks much of the southward sunlight.

    JEFF MORRIS reported in 2014: " ... There are two 4-ft-tall volunteers that the late Mr. Kennedy moved from under the now-gone tree, and planted in the backyard some years back. It is possible that they provide pollen to the 'fruiting' Torreya. There is at least one additional volunteer about 3-ft-tall along the driveway, bordering a neighbor's property. Finally, we learned that there used to be two additional mature Torreya taxifolia trees, larger than either of the Kennedy's pair, about a block away. They produced fruit for a number of years until the property last changed hands (within the past six years or so). The new owner didn't know what they were, and had them cut down before even occupying the residence. But they likely produced volunteers, even within only a hundred or so feet, in shaded areas. On our next visit, we need to look carefully at prospects in the neighborhood."

    PHOTOS BELOW: (1) Seeds ripen to orange and purple, (2) squirrels remove the fleshy husks, then bury seeds, (3) volunteer seedling amidst leaves of the neighboring Chestnut Oak and Southern Magnolia.


    1. SQUIRRELS CANNOT HARVEST HANGING SEEDS. Early on, Barlow visited Atlanta Botanical Garden and saw the pile of wire cages used to prevent squirrels from harvesting seeds. All the ABG "trees" at the time were rooted branchlets — so very short, thick, and shrubby. In contrast, trees grown from seed have droopy long branches — with exceedingly sharp needles — and the seeds are borne only at the tips (one growing season back). It was evident at the Clinton tree that squirrels had to wait for the seeds to fall. A.J. Bullard reported that having a Swamp Chestnut Oak nearby helped: squirrels preferred the acorns to the torreya seeds.

    2. DISCOVERY OF MYCORRHIZAE. Connie delivered seeds and seedlings from her 2013 trip to Jeff Morris. Later, Jeff reported (17 November 2013):

    "When I was transplanting the six seedlings that Connie gave me on November 3rd, I made an observation that I had not paid attention to before: mycorrhizal root nodules, similar to those I have seen on Cephalotaxus and Podocarpus seedlings in the past."
    3. SUBDIOECIOUS*. This tree (as well as the two saplings that Bullard grew on his own property that are offspring of the pair of Clinton trees) provided the first opportunity to document that a single tree can produce both male and female cones. Notably, female cones were visible only on branches of the least shady side of the tree (in this case, the northward side, away from the southward neighboring trees).

    * Torreya nucifera has previously been documented as "subdioecious." A 2008 report provides background on the subdioecious adaptation in plants. Since it occurs mostly by way of the male gender producing female reproductive organs, Barlow wonders if it is a way for the male to (a) test for the existence of nearby male rivals, and (b) to ensure reproductive success in the happenstance that its own wind-carried pollen is ineffective at reaching one or more females.

    A5. Bullard Home, Mount Olive NC  (grown fr seed ca. 1990; 2 specimens; one or both monoecious)

    UPDATE 2021: Joe Facendola in his November 2021 seed-collecting trip to this site (1,480 seeds collected) was authorized by Mrs. Bullard to dig up and collect 3 seedlings that squirrels had apparently planted in unmowed areas. This discovery qualifies the Bullard site as ramping up to FULL NATURALIZATION status.

    Webpage - Site Visit and History

    A.J. Bullard with ripe hanging seeds (31 October 2013).

       A.J. Bullard (introduced above as early documenter of the Clinton NC trees) planted seeds from the Clinton grove at his home in nearby Mt. Olive. The bushiness of the small-stature trees is notable (both have single trunks, so are indeed grown from seed).

    In a 6 December 2013 phone conversation with A.J. Bullard, Connie Barlow learned the history of the two specimens. In approx. 1995 A.J. and his cousin dug up (with the permission of the owner) 75 seedlings that were growing in the unmowed areas around the Clinton NC grove. The seedlings were primarily found along the hedgerows and in the flower garden. He gave away (or traded for other plants to conserve on his property) 73 of the 75 seedlings. He planted just two on his own land, each then about 3 feet tall. He estimates that they probably sprouted from a seed crop around 1988-90.

    It is impossible to know whether these two specimens contain significant genes from the tallest (since, blown-down) Torreya on the Clinton NC property, but because A.J. estimates from his 1995 visit that the tallest tree produced about 2/3 of the entire seed crop (of perhaps more than 5,000 seeds from the two trees combined), it is likely that these two specimens contain genes no longer found in the seedlings and seeds that I (Connie Barlow) collected beneath the one remaining (smaller) Clinton Torreya in October 2013.


    A.J. Bullard with his two dense, squat Torreya trees. Darlington (Laurel) Oak is visible overhead.

      1. SUN MAXIMIZES SEED-PRODUCTION BUT SQUAT GROWTH FORM. Connie recalls that these Torreyas had a tall planted oak for a neighbor southward on the lawn, which was the only other tree. Although A.J. said that fox squirrels occupy the tree, I saw none, and there were a lot of very ripe unharvested seeds beneath the torreyas. Thus, from the standpoint of maximizing seed production, the two Mt. Olive trees indicate torreyas can withstand lots of sun, if introduced to sun very early so that a squat, brushy growth form results, rather than a tall growth form. A.J. reported that he watered these two only immediately after planting; no watering in later years — which may be an excellent practice to urge this tap-rooted tree to grow its roots deep rather than outward.

    2. TIME FOR MATURATION. Seed-grown, with no over-topping canopy neighbors (other than one south-shading tree) documents that it takes 18-20 years for a tree in central (non-mountainous) North Carolina to produce its first seeds.

    A6. Caroline Dormon Nature Preserve, Saline LA   (planted circa 1950; 3 originals and seven seedlings)

    November 2018 Torreya Guardians Clint Bancroft and Connie Barlow received a guided tour of the Torreya trees by the preserve stewards, Rick Johnson and his adult son David. The two largest of the same-age trees (planted circa 1950) looked lush and healthy. The smallest one had died suddenly in 2017, following a severe drought. The precise cause(s) of death have not been discerned, but no evidence of water-mold cankers were seen on the branches.

    Webpage - Site Visit and History


    1. LONG GROUND-RUNNING BRANCH ADAPTATION. Photo left shows a low branch curving down and running along the ground about 30 feet, with multiple diverging subbranches creating a vast sheet of photosynthetic capacity. Slow-growing Torreya is easily over-topped by neighboring fast-growing pines and hardwoods, so this branch is stretching toward the sunny, maintained wide path. Torreya Guardians have documented this unusual adaptation also at the Biltmore Grove and Harbison House.

    2. SUBDIOECIOUS - MALE TREE PRODUCES SOME SEEDS. This century one of the three trees (long established as male) began producing several to 15 seeds, but not every year. All four seeds that ripened in 2018 were donated to Torreya Guardians.

    3. LACK OF SEED DISPERSAL. Fewer than a dozen seedlings have been documented to date, and all established directly beneath the canopy, indicating probable absence of squirrel involvement. The stewards have transplanted such seedlings mostly within a nearby grove of young Bigleaf Magnolia. One that will be transplanted in a future year is marked by a pink flag in photo left. It established exactly at an outer edge of the long, ground-running branch. (The seedling's leaves are a lighter shade of green than the older leaves of the ground-running branch.)

    4. NATURALIZED AND NON-INVASIVE. Documentation of seedlings, coupled with the absence of dispersal, fulfills two crucial criteria for safety of ex situ plantings: (a) The species can thrive and reproduce outside of Florida, yet (b) Torreya is incapable of becoming a noxious weed.

    B. Mature Trees Producing Seeds

    B1. Bess Home, Cleveland OH  - 4 potted specimens purchased (onsite in South Carolina) from Woodlanders Nursery in 2009 and 2010 (3 of which were seed-grown and the fourth a rooted branchlet).

    Webpage - Site Visits and History

    ABOVE: November 2017 photos - The first seeds were produced from potted seedlings planted 9 and 10 years earlier. Five seeds were produced on the right-most tree among the three seen here. In 2018, that same tree produced 19 seeds — and the shrubby rooted branchlet specimen nearby produced its first 4 seeds. The remaining two trees produce pollen each year. (The trees are given no artificial wind protection in winter nor water during summer droughts.)


    Photo above: Fred Bess shows visitor (Connie Barlow)
    his Torreya trees, 2 October 2018.

       1. COLD HARDINESS. The original potted seedlings that had grown outdoors in South Carolina struggled in the early years with polar winds as cold as -17F (see the site's webpage). But it seems that leaves produced in recent years are now fully adapted. This is the farthest north known production of seeds.

    2. SEED PRODUCTION REQUIRES FULL SUN. As with California Torreya, seeds on Florida Torreya are produced only on the full-sun side of the tree.

    3. RAPID INCREASE IN SEEDS. The female tree produced 4 seeds in 2017, then 19 seeds in 2018, and 168 by 2021.

    Note: Because the plantings are surrounded by mowed lawn (crucial for maintaining sunlight on this slow-growing tree), and because Fred Bess is determined to harvest the ripe seeds before the squirrels do, unaided "naturalization" into the surrounds is unlikely.

    VIDEO: "Seeds of Florida Torreya Produced in Ohio"

    Connie Barlow filmed this 2 October 2018 site visit. The VIDEO features important findings, including:

       1. These trees have put forth leaves well acclimated to severe cold spells in Ohio. On the windward side of the tree, branch tips are occasionally killed, but a ring of new growth results and the tree becomes plusher and thus even more wind-proof.

    2. Seeds are produced only on the branches that receive nearly full sun. (Connie notes from her 2005 site visit to wild California Torreya habitat that this seems to be a standard of the genus.)

    B2. Callahan Home, Medford OR  (2 specimens planted from seeds in 1995); no naturalization is possible, owing to pavement and mowed lawn.

    Photo above: Frank Callahan, holding a seed, next to his pair of Florida Torreyas.

       Webpage - Site Visit and History

    Torreya Guardians learned of Frank Callahan's pair of Florida Torreya trees in Medford, Oregon, when he contacted us to help him find good homes for a bumper crop of seeds autumn 2016.

    In an email to Lee Barnes on 9/29/16 Frank wrote: "We are in the Sunset Garden Book as Zone 7, the trees have survived -6 degrees F. in Medford w/o damage. Both of these trees exhibit male and female 'flowers', which is unusual for this taxon."

    Note: This above seed-producing pair is periodically watered and no naturalization occurs because the lawn is mowed up to the trees (with pavement on the other side). The summer dry season and intense sunlight and heat of inland Oregon would likely kill unwatered Torreya trees. However, along the northern coast of California, Torreya californica is found subcanopy — usually beneath a year-round evergreen canopy of Redwoods and Douglas-firs.


    1. DISCERNING WHEN SUN IS TOO EXTREME. The lower left corner of the photo above shows the most luxurious, biggest-leaved branch of all: It extends under the patio roof (photographer Barlow is standing under that roof), so it never gets any direct sunlight. Both seed-bearing specimens were planted on the north-facing side of the house. The furthest tree from the patio is experiencing sun scald at its top (the summers have gotten super hot and dry in Oregon). Both are also getting scalded where the open west-facing white pavement reflects late-afternoon sun and heat directly onto the trees. Again, this location in full westerly summer afternoon sun has brought forth a very dense, brushy growth form.
       PHOTO LEFT: Sun-scalded leaves of rooted branchlet shrubby torreyas that Frank donated to a city park. Sun-scalding happened because two diseased pine trees (shading south and west) were cut down the previous year.

    2. CAUTION ABOUT PROLIFIC SEED PRODUCTION. It is well known that tree species under great stress will sometimes produce a bumper crop of seeds (in a final attempt to reproduce vigorously if it senses possible demise.) We'll need to track these trees in future years to learn of their health.

    3. CONFIRMATION THAT ROOTED BRANCHLETS WILL NOT YIELD SINGLE-STEM TREES. Frank also planted rooted branchlets in a nearby park. The multi-stem shrubby growth form there confirms that Florida Torreya will not adopt a tree form when derived from a rooted branchlet — although some maintain that multi-year vertical staking will eventually yield a single leader.

    B3. Cheekwood Estate / Howe Garden, Nashville TN  - This site has not yet been visited and photographed by Torreya Guardians, but our volunteer compiler (Paul Camire) of a 27-page report on "Florida Torreya Ex-situ Specimens posts this information on page 8:

    Cheekwood received three, 2-year-old Torreyas from the U.S. National Arboretum in 1985. Two of the three trees are still thriving, one male and one female, with first seed production occurring 24 years after planting. The largest specimen measured at 18" circumference, 35' tall and a 20' spread in 2012. Both specimens are now measuring in at 40'+ tall. The two largest trees can be located in the Howe Garden. Seeds/cuttings were collected in 2009 and propagated with assistance from Dr. Croom. Cheekwood also received 6 seedlings from Atlanta Botanical Garden in 2000, located by the Development House, that are also thriving and producing seed. Many thanks to Cheekwood Plant Collections Manager, Shanna Jones, for taking the time to give a full history on the Cheekwood Torreyas. 1200 Forrest Park Dr, Nashville, TN 37205

    C. Mature Trees Not Producing Seeds

    C1. Henry Foundation, Gladwyne PA  (planted ca. 1940-50s); two mature male specimens; no seeds or seedlings

    Photo by Paul Camire.

       Webpage - Site Visits and History

    The two torreyas on the grounds have been surviving in Pennsylvania since Mary Henry planted them in the 1940s to 1950s. One is believed to have been grown from seeds and the other was brought in as a seedling.

    August 2018 Torreya Guardian Paul Camire visited the site. Prior to his visit he did historical research on the site and contacted the site's director (a grand-daughter of the botanist who planted the trees, Mary Gibson Henry). While onsite he also reviewed archived documents.

    Both specimens have the growth form of seed-grown main stems, with early basal growth that also went skyward.

    Because the trees have taken the tall growth form, only careful scrutiny (with binoculars) would reveal whether any of the full-sun branches are capable of producing female cones. Nonetheless, no volunteer seedlings have yet been discovered in the surrounds (which includes re-growth wild forest).

    KEY LEARNINGS: The site is never watered and the specimens are in the company of over-topping deciduous trees (see photo aove). Thus it is in an ideal location for initiating careful observations of how the trees contend with weather conditions in this semi-wild, farthest-north site of fully mature specimens. It would be useful if every autumn, the torreya pair could be scrutinized in quest of a few possible seed-bearing branches.

    C2. Spring Grove Cemetery, Cincinnati OH  (planted 1962); single shrubby specimen grown from rooted branchlet; male; no offspring.

    Torreya Guardian Fred Bess visited tree January 2017.

       Webpage - Site Visits and History

    In 2016 the American Forests Champion Tree Register made this Spring Grove Cemetery Florida Torreya specimen the new national champion of the species, following the confirmed death or demise of the previous champion, the Norlina NC tree.

    Our Torreya Guardian from Cleveland, Fred Bess, visited the tree January 2016 and concluded:

    "It seems obvious that it is cutting-grown [not from seed], as it is an oversized bush."

    UPDATE 2021: 2021 UPDATE: FLORIDA TORREYA (Torreya taxifolia) is no longer listed as a species on the national champion site. Perhaps it was removed because, as a cutting-grown plant, this Ohio specimen lacked the single stem of a seed-grown Torreya — and single stem is part of the core definition of what distinguishes a tree from a shrub. However, this shrubby cemetery specimen is still listed on State of Ohio Champion Trees website, but of course listed within the set of "Non-Native Big Tree Champions." There it records the measurements, plus "Nominator: Brian Heinz and Spring Grove Cemetery."


    1. COLD HARDINESS. This relatively unprotected tree appears to have survived well some gruesome winters. It is therefore important evidence of cold-hardiness. That the "national champion" Florida Torreya is now found in Ohio (previously, the champion tree was in Norlina NC) is yet another indication that assisted migration would serve this species very well.

    2. This specimen is registered as male, so if no female cones are ever seen on it, this may suggest the following: Perhaps a tree grown from a rooted branchlet cut from a male branch is incapable of producing any female branches. In contrast, we have good evidence that seed-grown trees are so capable.

    RECOMMENDATION. This tree deserves a female companion! Because gender cannot be determined in seeds or seedlings, 4 or 5 of seeds/seedlings will need to be planted nearby in order to pair up with the genetics of this tree. From an esthetic perspective, however, planting one or two rooted branchlets clipped from female branches might be ideal. As well, rooted female branchlets will produce female cones much faster than a seedling will.

    C3. Chattahoochee River, Columbus GA  (1 male remaining of 3 originals in neighborhood, planted ca. 1890s).

    Webpage - Site Visits and History

    Columbus, GA is on the east bank of a free-flowing section of the Chattahoochee River. The Chattahoochee River would have been the key conduit for southward displacement of this large-seeded species from the s. Appalachians during peak glacial times of the Pleistocene.

    Until the first decade of the 21st century, three old torreya trees were still found on residential properties in a historically old neighborhood that bordered the river. As of 2015, only one of the three trees still stands. Its top is dead and a huge section of lower bark has been stripped away (lightning?) on the street-side of the tree. Clint Bancroft and Jack Johnston measured the tree in 2016: circumference = 80 inches.


    Clint reports: "The owner was aware of his tree's fame and also told me that one of the three original Columbus Torreyas was located in the yard to the right of his house. Although healthy, it was cut down a few years ago by the person who bought the house.

    PHOTO left top: In 2015, Connie Barlow and Michael Dowd visited the tree in February 2015. Michael is admiring the old Southern Magnolia in the front yard of this historic house (1892). The Torreya (a male) is visible on the other side of the yard. Note: Torreya Guardians have taken cuttings from apical tips of basals to ensure (a) genetic safeguarding and (b) that clonal offspring will take the tree form.

    PHOTO left bottom: Dr. Kim Coder (Warnell School of Forestry, University of Georgia) published a richly-illustrated pdf of Torreya taxifolia in Columbus, GA. The photo left is captioned, "Male and female trees side-by-side in historic residential neighborhood."


    1. TORREYA CAN GROW LONG-TERM IN FULL SUN. (But we lack of history as to whether these front-yard trees were artificially watered during droughts.) The key is to never swiftly alter the lighting regime, as long-lived evergreen leaves take years to adjust via new growth.


       Standing on the porch (photo left), one sees the lowest branches of the Torreya on the right; Southern Magnolia branches to the left. Across the road is a city park that parallels a free-flowing section of the Chattahoochee River (visible left of the gazebo). Because some ripe Torreya seeds are "floaters" (and remain so for several days in water), the species could have launched seeds into s. Appalachian creeks as the climate cooled, and in only perhaps ten generations established riverside populations all the way to the Apalachicola River in Florida.

    As the glaciers retreated, however, only squirrels (and perhaps now-extinct giant tortoises) could have helped Torreya move north. Too slow, too far, to return this species to its mountain habitat.

    C4. Yinger Home, York PA  (one specimen planted in late 1970s from rooted branchlet)

    Note: Paul Camire tracked down this information in 2018 and contacted Barry Yinger.

       EMAIL RESPONSE FROM BARRY YINGER: "Yes, I planted a Florida Torreya at my farm in York County, PA in the late 1970s. It was given to me by the late nurseryman Tom Dodd Jr. of Semmes AL, who propagated it as a cutting from a plant collected in Florida. Apparently it was a cutting from a lateral branch because it never developed a leader as you see with seed-grown plants.

    "My farm was in the cold part of Zone 6 for many years and is now mid-Zone 6 to 6b. However, in January 1994 we had the lowest recorded temperatures ever. It was about -23 F the first night and -12 the next. The high during the day in between was -4 F. I lost a number of trees and shrubs, but the Torreya was not damaged beyond a little discoloration of some of the leaves. I never measured the tree, but after nearly 40 years it was about 20 feet tall and nearly as wide. If it had been rooted from a terminal cutting it would have been a lot taller. As it was, it grew as a big blob. I will try to find a photo for you. I sold my farm in 2016."


    C5. Kalmia Gardens of Coker College, Hartsville SC  (one specimen planted in late 1970s from rooted branchlet)

    Note: Thanks to Dan Hill (Assistant Director of Kalmia Gardens) and Evan D. Cook (previously at Moore Farms Botanical Garden in Lake City SC, which has 4 young T. taxifolia specimens) for providing information. Kalmia Gardens website.

    There are two tall trees at this location. However, the main stem of one died recently, but it has a small basal sprout.

         The remaining tree is pictured in the two photos left. The inner photo shows the dark evergreen foliage up to its top at a distance. The left-most photo shows a mature main stem with several basals.

    This tree is unique as it is the first Torreya taxifolia to be documented with a burl. The burl has multiple shoots of terminal growth extending from it.

    No information is readily available on year of planting nor acquisition source. A site visit report would be very helpful as to presence or absence of reproductive structures and whether any seed-born offspring are detected in the unmowed surrounds.

    D. International Plantings Producing Seeds

    • BELGIUM: Kalmthout Arboretum  (planted 1910); a single specimen; 300+ seeds in 2011

    Article: "Torreya taxifolia produces seeds in Kalmthout Arboretum", International Dendrology Society, 2012

    Torreya Guardians generally doesn't try to find and document seed production of horticultural plantings on other continents. But the article above offers some important observations.

    BACKGROUND: "... It was brought here by the curator of the collections Antoine Kort around 1910 from a nursery close to the Italian Lago Maggiore. The plant grows in south-facing ground at the edge of the estate, close to the historic Vangeertenhof. A good ten years ago, the surrounding plants were removed so that this very special tree could get a full quota of light and air. A low wall protects the base of the trunk from bright sunlight and strong winds. Since then, the circumference of the tree has almost doubled. Another superb example of this cultivar grows in the Allard Arboretum in Angers (France)."

    The giant Torreya taxifolia at
    Allard Arboretum, Angers France.
       (1) CAN POLLEN FROM OTHER TORREYA SPECIES FERTILIZE FLORIDA TORREYA? Seed production here "is quite exceptional when you think that this species is actually dioecious and that there is only one plant in Kalmthout... Elsewhere in the garden is an old example of Torreya nucifera and a few young examples of Torreya californica. Perhaps the fruits are the result of cross-fertilisation. But then again, perhaps not. In his manual of cultivated conifers, Gerd Krussmann announced that Torreya can also be monoecious on rare occasions."

    (2) NATURALIZATION. "The volunteers of our seed team collected a total of around 250 seeds; the others were left on the tree and who knows, perhaps we will see a spontaneous burst of seedlings one of these days. The seed coats (sarcotesta) of the seeds that had been collected were immediately removed and the seeds were put into damp turf for safekeeping. We sowed a small percentage of them ourselves; the rest were distributed to other botanical gardens and arboretums. In the rest year after sowing, a single seed had germinated. Then almost all the other seeds germinated in the second year — a hopeful and promising sign."

    NOTE: View the lengthy List of All Ex Situ Plantings that Torreya Guardian Paul Camire compiled. Near the end of that pdf you will find the list of 14 international plantings that he was able to discover online.


    Garden Plants Get a Head Start on Climate Change, by Sebastiann Van der Veken et al., May 2008, in Frontiers in Ecology and the Environment.

    EXCERPTS: Conservation biologists are concerned that climate change will cause widespread extinctions because limited capacity for migration could compromise species' ability to adjust to geographic shifts in habitat condition. However, commercial plant nurseries may provide a head start for northward range shifts among some plant species. To investigate this possibility, we compared the natural ranges of 357 native European plant species with their commercial ranges, based on 246 plant nurseries throughout Europe. In 73% of native species, commercial northern range limits exceeded natural northern range limits, with a mean difference of approx. 1,000 km. With migration rates of 0.1 to 5 km per year required for geographic ranges to track climate change over the next century, we expect nurseries and gardens to provide a substantial head start on such migration for many native plants. While conservation biologists actively debate whether we should intentionally provide "assisted migration", it is clear that we have already done so for a large number of species. ... ... Of the 534 ornamental species sold in Britain during the 19th century and examined by Dehnen-Schmutz et al. (2007), 27% were subsequently found growing outside of cultivation, and 30% of these had established populations, clearly demonstrating the potential for horticultural plants to spread into non-cultivated habitats (see Sullivan et al.2005).
    The role of botanical gardens in climate change research, by Richard B. Primack and Abraham J. Miller-Rushing, 2009, Tansley Review in New Phytologist.
    EXCERPTS: Botanical gardens have a unique set of resources that allows them to host important climate change research projects not easily undertaken elsewhere. These resources include controlled growing conditions, living collections with broad taxonomic representation, meticulous record-keeping, networks spanning wide geographic areas, and knowledgeable staff. Indeed, botanical gardens have already contributed significantly to our understanding of biological responses to climate change, particularly the effects of temperature on the timing of flowering and leaf-out.... The evidence reviewed here shows that botanical gardens can serve as particularly valuable sources of long-term data to describe how plants are responding to climate change.... Of special concern, we suggest that species be monitored to determine whether previously hardy species are no longer able to grow at particular botanical gardens. Additionally, researchers should note species that were previously difficult to grow at a site that are able to thrive in the changing climatic conditions.
    "Plants' native distributions do not reflect climatic tolerance", by Tierney Bocsi et al., 2016, Diversity and Distributions
    EXCERPTS: In an era of rapid climate change, conservation biogeography often focuses in particular on risks to rare native species. Our analyses suggest that species native ranges often underestimate their climatic tolerance.... Most species have not been broadly introduced outside of their native ranges. As a result, our understanding of climatic tolerance limits for the bulk of species can only be inferred from occurrences within their native ranges. Accordingly, ecological forecasting studies assessing extinction risk and supporting conservation planning are typically based on native occurrences.

      Our results show that native range data strongly underestimate the climatic conditions where many species could occur. The vast majority (86%) of species had adventive occurrences in climate conditions that were not encompassed by native range distribution data. Range models expanded substantially when adventive occurrences were included, and particularly when those adventive occurrences had novel climate conditions.

    LEFT: Fraser Fir, how adventive occurence expands ecological niche. (All maps in Supplement)

    Thus, the broader range models are likely due to expansion of the realized niche rather than a model response to the addition of more data points. The corresponding expansion of niche models by 35% geographically suggests that an assumption of climatic equilibrium in the native range may be optimistic for many native species.... This study supports the findings of Early & Sax (2014) that niche shifts are similarly prevalent amongst non-invasive species. This accumulation of evidence strongly suggests that the native distribution of a species is a poor proxy for climatic tolerance. Moreover, our results suggest a pattern of greater niche expansion amongst plant species with smaller native range sizes.... Many species would already be able to successfully establish outside of their native ranges, even before climate changes. The latter finding could support conservation strategies like managed relocation, particularly for slow-growing, long-lived species like trees that might require planting well in advance of shifting climate conditions. Given that many plant species appear limited by dispersal ability, it is unlikely that they will be able to keep up with climate change unaided.

    "Limited Range-Filling Among Endemic Forest Herbs of Eastern North America and Its Implications for Conservation With Climate Change" - by Stephanie K. Erlandson, Jesse Bellemare, and David A. Moeller, 8 December 2021, Frontiers in Ecology and Evolution.
    EXCERPTS: Species with low dispersal and poor range-filling capacity may be especially threatened by modern climate change because they are unlikely to migrate quickly enough to keep pace with changing climate (Thomas, 2011). In some cases, their realized and potential distributions might shift apart entirely, exposing them to high risk of extinction (Sax et al., 2013). In this context, many endemics might be particularly susceptible, as their distributions often reflect past marginalization by climate change (e.g., Pleistocene glaciations), and their current distributions are often restricted to unusual habitats (high elevations, north slopes) in southern areas that are surrounded by warmer, unsuitable habitat (Ohlemüller et al., 2008). Conservation of such endemics may require assisted migration (McLachlan et al., 2007). However, in the case of endemic forest herbs of the Southern Appalachian Mountains, it has been unclear whether suitable habitat already exists outside their native ranges. Our models based on current climates predicted suitable habitat north of the range margins of three endemics, suggesting that assisted migration could be successful currently and that single, larger-scale translocations might be feasible, rather than “many small steps” moving incrementally northward over decades... Notably, Diphylleia cymosa and Shortia galacifolia are known to survive, reproduce, and recruit new adults in parts of New England where they have been transplanted decades ago, making these patterns plausible (J. Bellemare, pers. obs.).
         ... In the case of the endemics investigated here, our Last Glacial Maxium projections suggest that they could have persisted near their present-day distributions in the Southern Appalachian Mountains (SAM) even during the colder climatic conditions of the Pleistocene. This pattern is consistent with horticultural observations that narrow endemics of the SAM often survive and reproduce in gardens far to the north. However, this Pleistocene context might not suggest a similar likelihood for pre-existing tolerance to the significantly warmer conditions projected for the future given that they would be unprecedented in the recent geological past. In the absence of rapid evolution or preexisting tolerances, poleward migration might be necessary for the long-term survival of the endemic species investigated here. However, natural dispersal to northern areas for endemic species may be particularly unlikely, given their apparent lack of substantial northward migration during the Holocene.... Given the severity of reductions in suitable habitat within the range and the disjunct nature of suitable habitat outside the range, assisted migration may be an important strategy for long-term conservation (Hoegh-Guldberg et al., 2008)... Our results suggest a history of dispersal limitation following the last glacial maximum along with an environmental barrier to northward migration. Conservation of endemic species would likely require intervention and assisted migration to suitable habitat in northern New England and Canada.
    "Climatic criteria for successful introduction of Quercus species identified by use of Arboretum data", by Corrie Lynn Madsen et al., March 2021, in Forestry.
    EXCERPTS: Climate change is projected to have a major influence on forest tree populations and composition. Translocation of species outside their historic range has been suggested to maintain healthy forests and tree species. The introduction of exotic species into botanical gardens and arboretums worldwide demonstrates the ability of many trees to grow outside their natural habitat and may play an important part in avoiding climate driven extinction if grown in a matching climate. However, it remains to be determined which climatic factors are the most important predictors of climatic match. In this study we use information from the arboretum in Horsholm, Denmark, to analyse differences in performance of translocated Oak (Quercus) and show how data from tree collections can be used to predict success of assisted migration.
         ... We find that arboretum data contain biogeographic information that may help interpret factors involved in climatic adaptation, and assist in selecting suitable source areas for assisted migration. The full potentials can only be realized, however, if data from several arboreta are combined. Here we determined criteria for successful migration under current conditions, but what will they be under future conditions? Establishing a network of collections ('metacollections' as coined by Dosmann 2006) across Europe or the world would make it possible to expand the scope of analyses considerably by verifying whether the same parameters are decisive for success under different conditions, and by allowing more detailed studies of individual species or genera. This could potentially lead to a new understanding of assisted migration under climate change.
    "Reconsideration of the native range of the Chinese Swamp Cypress (Glyptostrobus pensilis) based on new insights from historic, remnant and planted populations", by Jinlog Zhang and Gunter A Fischer, 2021, Global Ecology and Conservation.
    EXCERPTS: ... Tree populations in human-modifed landscapes are usually considered less important compared to "natural" or "wild" populations and consequently are rarely collected comprehensively during scientific plant surveys. Indeed, no matter whether the species were planted to enhance natural populations, used for ex situ conservation or as roadside trees in cities, their potential importance to conservation is underestimated. In fact, if trees are planted in habitats with comparatively low anthropogenic disturbance such as nature reserves or in greening projects in rural areas, such as roadsides, highway slopes, edges of agricultural fields or gallery forests along streams and rivers, the species may start to reproduce and spread by themselves. Nonetheless, such planted tree stands, even when they are located within the species historic and/or present native range, are usually considered cultivated and not 'natural' or 'wild'. If such stands are able to reproduce and grow without human assistance over a longer period of time, then they could be considered as 'rewilded' under the concept of Carver et al. (2021).... For a rare and undercollected species, which has experienced a significant decline in range size and habitat degradation, taking historical records such as buried ancient trees and "planted" trees would supplement the museum collections and hence improve species distribution modelling. Based on the output of species distribution modelling, we argue that the native range of the Chinese Swamp Cypress should be much broader. Consequently, naturalized or rewilded populations within the modelled range should be considered native and receive adequate protection.

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