ENTS,
I am wondering about these reiterations. We have talked
about it before,
but everything is not clear in my mind. A reitteration is what
exactly? It
appears to be a secondary trunk formed ntially from an upturned
limb... How
do they form and why? What species of trees are know to have
these
reiterations? I know this white pine, some of the hemlocks in
the Tsuga
Search project, Many of the giant conifers from Bob Van Pelts
book. Are
there examples of reiterations in trees that are not conifers?
Please
enlighten me on the subject.
Ed Frank
[Don Bertolette, March 12, 2009]
Ed-
I suppose it might vary from species to species, region to region,
but in general it has to do with disturbance and a trees response.
The re-iterations you mention are most typical in conifers, as they
typically exhibit 'apical dominance'. Which is to say, the tree
wants to grow into the sunlight. When some manner of disturbance
causes damage to the apical tip, like a significant ice storm, a
wind event that breaks the top out, and/or other such single tree
calamaties.
The tree's response, somewhat dependent on genetic predisposition,
often times is to route the trees energy to the next best branch. I
have seen a number of species do this several times (tree lives long
enough, emerges high enough above the forest canopy, it gets taken
back, sometimes, time after time). I don't think I have a photo
anymore, but there are a number of "candelabra trees" in the
redwoods of northern California that may be the most extreme example
of this, in part due to their longevity (number of opportunities for
coastal exposure to Pacific storms), and in part due to their energy
allocation strategies.
-Don
[Will Blozan, March 12, 2009]
Ed,
Here is an excerpt from the Tsuga Search Project that should
help:
Wood distribution
In total, this project sampled 3,340.1 m (10,955 ft) of eastern
hemlock
trunks (N=53) with a cumulative wood volume of 1,672.3 m3 (59,053
ft3)[1].
The modeled trees did not always conform to a simplistic, single
trunk form
however, and required measuring some challenging structures (See
Reiterations and bifurcations below). In all trees sampled,
"conventional"
main trunk wood composed anywhere from 63.7% to 100% of total tree
volume.
Bifurcations (forks of vertical leaders) accounted for a maximum of
36.3% of
total wood volume, considerably more than a maximum of 10.05% for
reiterations (limbs upturned into secondary trunks). Bifurcations
were much
larger; even though a total of 633.1 m (2,076.6 ft) of reiterations
were
measured, total bifurcation volume (42.9 m3; 1514.2 ft3) was 240%
more with
only two-thirds of the cumulative path length. Path lengths (length
of all
measured wood structures) of individual trees ranged from simply the
length
of the trunk to a maximum of 200.7 m (658.4 ft). See Appendix 4:
Wood
distribution of subject trees for more detail.
Reiterations and bifurcations
Reiterations are replicated trunks formed by the tree to exploit
a light
gap. Structure and growth of reiterations are markedly different
from normal
descendent or ascendant branches. Reiterations are included in trunk
volume
calculations as they are part of permanent, branch bearing
structures of the
tree. Their origin arises from the sprouting of normal branches and
can be
initiated by a light gap from the side or from crown damage above.
The
apically dominant portion of these structures are often much younger
and
model-conforming as in a young tree. The cross-section changes to a
more
oval shape as reaction wood is built up to support the added weight
from
increased growth and sap demands as the reiteration gains vigor. The
buttressing allows a more permanent attachment than a conventional
branch
due to support on the lower portion being less likely to fail from
injury to
the upper surface. This also allows wounds with associated decay to
persist
and form a suitable substrate for vascular epiphytes (Sillett, Van
Pelt
2007)
Studies in redwood forests indicate that structural diversity in
the canopy
is strongly linked with aerial biological diversity. Reiterations
generate
niches not found anywhere else in the forest (Sillett, Van Pelt
2007). Our
observations indicated moss mats, epiphytic plants and lichen
gardens to be
associated with these large structures. Large reiterations and
breaks in the
trunks were hosts to shade intolerant shrubs and wind disseminated
trees
such as Rubus and Betula. Complex, reiterated crowns could be
refugia for
those species lying in wait for a light gap or breach in the
rhododendron.
Of the thirty trees discussed in this project, eleven trees had
large
reiterations. Four of these were tall trees and seven were large
trees (two
trees are replicated). A total of thirty-seven reiterations were
measured,
ten of which grew on the Usis Hemlock, the most on a single tree in
the
project. Total reiterated trunk length measured on the Usis Hemlock
totaled
over 147.8 m (485 ft) for a tree total of 200.7 m (658.5 ft) of
trunk.
Likewise, the Cheoah Hemlock had a total of 185.6 m (608.7 ft) of
measured
trunk length in reiterations and bifurcations. These structures are
likely
to greatly increase the crown area of the tree, and bolster wood
production.
Reiterations on the subject trees (no repeated trees) totaled
14.7 m3 (518.8
ft3) in volume, 527.0 m (1,728.6 ft) in cumulative length, and were
found to
compose anywhere from 0.5% to 10% of total tree volume, averaging
4.2% for
all trees. Volume of individual reiterations reached a maximum of
1.78 m3
(62.9 ft3) and a maximum structure length of 18.96 m (62.2 ft). On
average,
the reiterations were 31% larger and 12% longer on the large subject
trees
than on the tall subjects. Fusions between reiterations were
documented in
two trees, forming a strong interlocked crown with braced
bifurcations that
likely reduce mechanical failure of non-conventional crowns.
"Transplanted"
branches were noted in the Cheoah Hemlock. This occurs when a
grafted branch
severs its connection from its origin. This phenomenon is well
documented in
the coast redwoods of California, and is encountered in dense
hemlock hedges
as well. The Cheoah Hemlock and the Usis Hemlock both exhibited
reiteration
fusions; one of which started at 14.3 m (47 ft) and fused to another
reiteration 9.7 m (31.9 ft) higher in the tree. The Cheoah Hemlock
had a
branch that crossed the inner canopy while grafting to three
separate
structures.
Height of origin was noted for every measured reiteration. The
lowest
occurred at 12.59 m (41.3 ft) and the highest at 38.35 m (125.8 ft)
above
mid-slope. The highest recorded reiteration was over 25 cm (12 in)
in
diameter and was encountered in the Usis Hemlock. This tree also
contained
the highest cumulative and relative volume of reiterations; 4.36 m3
(154.3
ft----------3) and 10.05% respectively. The most massive reiteration
was
found in the Cheoah Hemlock and scaled 1.78 m3 (62.9 ft3). This
individual
structure alone contained a path length of 34.33 m (112.6 ft) of
measured
trunks. A graphing of the origin heights below likely reveals a
random
height of origin as the events initiating the formation of a
reiteration
would likewise be random. Data gathered from a control population of
non-superlative trees are needed for comparison.
Of the thirty subject trees, three had large bifurcations that
warranted
frame mapping along the section(s) of trunk fusion. These massive,
fused
areas were up to 1.6 m (5.25 ft) across. Two other subject trees
were
bifurcated but did not exhibit reiterations. Of the subject trees,
only the
Cheoah Hemlock was both reiterated and bifurcated. This tree,
supporting a
bifurcation more than 83 feet long, was also nearly entirely cloaked
in live
crown from base to top. The longest and largest bifurcation occurred
on the
Yonaguska Hemlock; 28.75 m (94.3 ft) long and 5.37m3 (189.6 ft3)
respectively. See example of a frame-mapped bifurcation fusion
below:
Frame mapping example from the Yonaguska Hemlock bifurcation fusion
All the remaining trees in the project had a single trunk; one of
which was
broken at 31.74 m (104.1 ft). This tree (the Headless Giant) - as
well as
several others - had the vast majority of the crown originating on
reiterations with very few non-reiterated branches. Several
reiterations
reiterated again (second order) with one fourth order reiteration
noted in
the Usis Hemlock. This immense tree had a fusion of two giant
reiterations
that supported a complex limb system supporting multiple reiterated
sprouts
- some originating from the same limb like a row of small trees
extending
upwards of ~12 m (~40 ft) from the trunk origin.
Will F. Blozan
President, Eastern Native Tree Society
President, Appalachian Arborists, Inc.
Continued
at:
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