The Torreya (Torreya taxifolia) is Missing its Megafaunal Disperser

The torreya (Torreya taxifolia), also known as the stinking cedar because its crushed needles give off a strong resin odor, is a relic species thought to have been more widespread during warm climatic phases of the Pleistocene.  It likely diverged from an ancestor that was even more widespread during the Miocene when warm moist forests occurred all across North America and Asia.  T. taxifolia  is an extremely rare species confined to just the east side of the Apalachicola and Flint Rivers, while a closely related sister species (T. californica) is native to California where it is found in several disjunct populations. 

Pleistocene Ice Ages fostered the spread of arid grassland environments that were unsuitable for torreyas.  Under these conditions the torreya retreated to moist refugia on steep ravines of the Apalachicola and Flint Rivers. Connie Barlow, author of the below referenced book, thinks the torreya  formerly expanded its range as far north as the southern Appalachians, following the end of Ice Ages.  They are better adapted to live in an Appalachian cove forest rather than the environments surrounding their current range.  She hypothesizes the torreya’s current rarity is the result of its disperser’s extinction.  She suspects the giant tortoises (Hesperotestudo crassicutata and H. incisa) ate the torreya cones and defecated the seeds intact.  As the climate warmed following the end of Ice Ages, the tortoise’s range expanded and torreya trees spread in correspondence with this range expansion.  She believes the tortoises were the torreya’s main disperser. Squirrels can disperse the seeds but they are more likely to eat and destroy them, and other mammals are all potentially more likely to destroy the seeds with their teeth when they consume the cones. Tortoises don’t have teeth.  Furthermore, torreya cones contain turpene which is toxic to mammals but not to reptiles.  Now that tortoises are extinct, the torreya is stuck within a tiny range where it is probably going to succomb to fungal diseases. 

Barlow’s hypothesis will be difficult to support with concrete evidence–plant macrofossil remains from warm climatic phases of the Pleistocene are rare in this region.

Connie Barlow and her husband with a very rare Torreya tree.  She hypothesizes that its rarity today is due to the extinction of its most probable disperser–the giant tortoise.

Torreya taxifolia range map.png

Torreya taxifolia range map.

Torreya trees grow in natural communities the late naturalist, Charles Wharton, referred to as “torreya ravines.”  These are cool moist micro-environments also known as steepheads, and they only occur on the east side of the rivers.  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.

Wharton catalogued Torreya Ravines in his book The Natural Environments of Georgia written in 1978.  A more recent updated version of that book (The Natural Communities of Georgia) written by several authors and published last year does not mention torreya ravines.  I fear this means torreya trees may already be extinct in Georgia.  Wild torreya trees can still be found in Torreya State Park in Florida.

Mature torreya trees grow to 60 feet tall, but today few wild torreyas exceed 6 feet before dying back due to fungal disease.  Torreya trees have been transplanted to the Biltmore Estate in Asheville, North Carolina where they are doing much better than the wild trees.  Torreya trees growing on the Biltmore Estate survived a freeze of -30 F.  This shows they are capable of surviving in more northerly latitudes, and this supports Barlow’s hypothesis.


Barlow, Connie

The Ghosts of Evolution

Basic Books 2000

Wharton, Charles

The Natural Environments of Georgia

Georgia Department of Natural Resources 1978


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