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arnoldia

Volume 58 Number 1 1998

Arnoldia (ISSN 004-2633; USPS 866-100) is
published quarterly by the Arnold Arboretum of
Harvard University. Second-class postage paid at
Boston, Massachusetts.

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Editorial Committee
Phyllis Andersen
Robert E. Cook
Peter Del Tredici
Gary Roller
Stephen A. Spongberg
Kim E. Tripp

Copyright © 1998. The President and Fellows of
Harvard College

•\iUN 15 iggg

Page

2 An Evolutionary Perspective on Strengths,
Fallacies, and Confusions in the Concept
of Native Plants
Stephen Jay Gould

11 E. D. Merrill, From Maine to Manila
Ida Hay

20 Light in a Bottle: Plant-Collecting in the
Philippines
Rob Nicholson

27 The Ecology and Economics of Elm
Replacement in Harvard Yard
Peter Del Tredici

Cover: For much of this century, a house and an
American elm comprised the basic domestic land-
scape of eastern North America. Photograph by
Peter Del Tredici.

Inside front cover: Livistonia saribus, the Livistone
palm, named by E. D. Merrill, bears unusual bright-
blue fruits. Photograph from the Archives of the
Fairchild Tropical Garden.

Inside back cover: An American elm of classic, vase-
shaped form photographed in the Montreal Botanic
Garden by Peter Del Tredici.

Back cover: Elmer Drew Merrill, later Director of the
Arnold Arboretum, in a detail of a 1902 photograph
seen on page 12.

Erratum: In our last issue. Winter 1997-1998,
volume 57, number 4, two numbers were inverted in
the Arnold Arboretum Weather 5tation Data — 1997.
Last spring's last frost was a low of 29° on April 16.

PETER ir:L TRILDICI

An Evolutionary Perspective on Strengths, Fallacies,
and Confusions in the Concept of Native Plants

Stephen Jay Gould

An important, but widely unappreciated, concept in evolutionary biology draws a clear and
careful distinction between the historical origin and current utility of organic features.
Feathers, for example, could not have originated for flight because five percent of a wing in
the early intermediary stages between small running dinosaurs and birds could not have
served any aerodynamic function (though feathers, derived from reptilian scales, provide
important thermodynamic benefits right away). But feathers were later co-opted to keep
birds aloft in a most exemplary fashion. In like manner, our large brains could not have
evolved in order to permit modern descendants to read and write, though these much later
functions now define an important part of modern utility.

Similarly, the later use of an argument, often in a context foreign or even opposite to the
intent of originators, must he separated from the validity and purposes of initial formula-
tions. Thus, for example, Darwin's theory of natural selection is not diminished because
later racists and warmongers perverted the concept of a "struggle for existence" into a ration-
ale for genocide. However, we must admit a crucial difference between the two cases:
the origin and later use of a biological feature, and the origin and later use of an idea. The
first case involves no conscious intent and cannot be submitted to any moral judgment.
But ideas are developed by human beings for overt purposes, and we have some ethical
responsibility for the consequences of our actions. An inventor may be fully exonerated
for true perversions of his intent (Hitler's use of Darwin), but unfair extensions consistent
with the logic of original purposes do entail some moral demerit (most academic racists of
the nineteenth century did not envision or intend the Holocaust, but some of their ideas did
fuel the "final solution").

I want to examine the concept of "native plants" within this framework, for this notion
encompasses a remarkable mixture of sound biology, invalid ideas, false extensions, ethical
implications, and political usages both intended and unanticipated. Clearly, Nazi ideologues
provided the most chilling uses.^ In advocating native plants along the Reichsautobahnen,
Nazi architects of the Reich's motor highways explicitly compared their proposed restric-
tion to Aryan purification of the people. By this procedure. Reinhold Tiixen hoped "to
cleanse the German landscape of unharmonious foreign substance."^ In 1942 a team of
German botanists made the analogy explicit in calling for the extirpation of Impatiens

Grapevines (Vitus sp.) in northeastern Connecticut. This native is a commonplace of second-growth forest
where its weight causes serious damage to its host trees.

4 Arnoldia 1998 Spring

parviflora, a supposed interloper; "As with the
fight against Bolshevism, our entire Oecidental
eulture is at stake, so with the fight against this
Mongolian invader, an essential element of this
culture, namely, the beauty of our home forest,
is at stake.

At the other extreme of kindly romanticism,
gentle arguments for native plants have stressed
their natural "rightness" in maximally harmo-
nious integration of organism and environment,
a modern invocation of the old doctrine of
genius loci. Consider a few examples from our
generation;

Man makes mistakes; nature doesn't. Plants
growing in their natural habitat look fit and
therefore beautiful. In any undeveloped area you
can find a miraculously appropriate assortment
of plants, each one contributing to the overall
appearance of a unified natural landscape. The
balance is preserved by the ecological conditions
of the place, and the introduction of an alien
plant could destroy this balance.'*

Evolution has produced a harmony that con-
trived gardens defy.®

Or this from President Clinton himself
(though I doubt that he wrote the text person-
ally), in a 1994 memorandum on "environmen-
tally and economically beneficial practices on
federal landscaped grounds"; "The use of native
plants not only protects our natural heritage and
provides wildlife habitat, but also can reduce
fertilizer, pesticide, and irrigation demands and
their associated costs because native plants are
suited to the local environment and climate."*
This general argument, of course, has a long
pedigree, as well illustrated in Jens Jensen's
remark in Our Native Landscape, published in
his 1939 Siftings: "It is often remarked, 'native
plants are coarse.' How humiliating to hear an
American speak so of plants with which the
Great Master has decorated his land! To me no
plant is more refined than that which belongs.
There is no comparison between native plants
and those imported from foreign shores which
are, and shall always remain so, novelties."^

Yet the ease of transition between this
benevolent version and dangerous Volkist
nationalism may be discerned, and quite dra-
matically, in another statement from the same

Jens Jensen, but this time published in a Ger-
man magazine in 1937;

The gardens that I created myself shall ... be in
harmony with their landscape environment and
the racial characteristics of its inhabitants. They
shall express the spirit of America and therefore
shall be free of foreign character as far as pos-
sible. The Latin and the Oriental crept and
creeps more and more over our land, coming
from the South, which is settled by Latin people,
and also from other centers of mixed masses of
immigrants. The Germanic character of our cit-
ies and settlements was overgrown. . . . Latin
spirit has spoiled a lot and still spoils things
every day.*

How slippery the slope between genius loci
(and respect for all the other spirits in their
proper places as well) and "my locus is best,
while others must be uprooted, either as threats
or as unredeemable inferiors." How easy the fal-
lacious transition between a biological argu-
ment and a political campaign.

When biologically based claims have such
a range of political usages (however dubious,
and however unfairly drawn some may be), it
becomes particularly incumbent upon us to
examine the scientific validity of the underlying
arguments, if only to acquire weapons to guard
against usages that properly inspire our ethical
opposition (for if the biological bases are wrong,
then we hold a direct weapon; and if they are
right, then at least we understand the argument
properly, and can accurately drive the wedge
that always separates factual claims from ethi-
cal beliefs).

Any argument for preferring native plants
must rest upon some construction of evolution-
ary theory — a difficult proposition (as we shall
see) because evolution is so widely miscon-
strued and, when properly understood, so diffi-
cult to utilize for the defense of intrinsic native
superiority. This difficulty did not exist in pre-
Darwinian creationist biology, because the old
paradigm of "natural theology" held that God
displays both his existence and his attributes of
benevolence and omniscience in the optimal
design of organic form and the maximal har-
mony of local ecosystems (see William Paley for
the classic statement in one of the most influen-
tial hooks ever written).’ Native must therefore

The Concept of Native Plants 5

Cortaderia jubata (sawgrass), weedy South American cousin of the garden-variety pampas grass, has invaded
the hills of north-coastal California.

be right and best because God made each crea-
ture for its proper place.

But evolutionary theory fractured this equa-
tion of existence with optimality by introducing
the revolutionary idea that all anatomies and
interactions arise as transient products of com-
plex history, not as created optimalities. Evolu-
tionary defenses of native plants rest upon two
quite distinct aspects of the revolutionary para-
digm that Darwin introduced. (I shall argue that
j neither provides an unambiguous rationale, and

[ that many defenders of native plants have

mixed up these two distinct arguments, there-
fore rendering their defense incoherent.)

The Functional Argument Based on
Adaptation

Popular impression regards Darwin's principle
j of natural selection as an optimizing force, lead-

ing to the same end of local perfection that God
^ had supplied directly in older views of natural

I theology. If natural selection works for the best

1 forms and most balanced interactions that could

i

possibly exist in any one spot, then native must
be best for native has been honed to optimality
in the refiner's fire of Darwinian competition.
(In critiquing horticulturists for this misuse of
natural selection, I am not singling out any
group for an unusual or particularly naive mis-
interpretation. This misreading of natural selec-
tion is pervasive in our culture, and also records
a primary fallacy of much professional thinking
as well.'°)

In Siftings, Jens Jensen expressed this com-
mon viewpoint with particular force:

There are trees that belong to low grounds and
those that have adapted themselves to high-
lands. They always thrive best amid the condi-
tions they have chosen for themselves through
many years of selection and elimination. They
tell us that they love to grow here, and only here
will they speak in their fullest measure."

I have often marvelled at the friendliness of
certain plants for each other, which, through
thousands of years of selection and elimination,
have lived in harmonious relation."

6 Arnoldia 1998 Spring

The incoherencies of this superficially attrac-
tive notion may he noted in the forthcoming
admission, in a work of our own generation,
that natural does not always mean lovely. Natu-
ral selection does not preferentially lead to
plants that humans happen to regard as attrac-
tive. Nor do natural systems always yield
rich associations of numerous, well-halanced
species. Plants that we label "weeds" will domi-
nate in many circumstances, however tran-
siently (where "transient" can mean more than
a human lifetime on the natural time scales of
botanical succession). Such weeds are often no
less "native" — in the sense of evolving indig-
enously— than plants of much more restricted
habitat and geography. Moreover, weeds often
form virtual monocultures, choking out more
diverse assemblages than human intervention
could maintain. C. A. Smyser et al. admit all
this, but do not seem to grasp the logical threat
thus entailed against an equation of "natural"
with "right" or "preferable": "You may have
heard of homeowners who simply stopped
mowing or weeding and now call their land-
scapes "natural." The truth is that these so-
called no-work, natural gardens will be long
dominated by exotic weed species, most of
which are pests and look downright ugly.
Eventually, in 50 to 100 years, native plants
will establish themselves and begin to create
an attractive environment."'^ But not all
"weed" species can be called "exotic" in the
sense of being artificially imported from other
geographic areas. Weeds can be indigenous
too, though their geographic ranges tend to
be large, and their means of natural transport
well developed.

The evolutionary fallacy in equating native
with best adapted may be simply stated by
specifying the essence of natural selection as a
causal principle. As Darwin recognized so
clearly, natural selection produces adaptation to
changing local environments — and that is all.
The Darwinian mechanism includes no concept
of general progress or universal betterment.
The "struggle for existence" can only yield local
appropriateness. Moreover, and even more
important for debates about superiority of
native plants, natural selection is only a "better
than" principle, not an optimizing device. That

is, natural selection can only transcend the local
standard and cannot operate toward universal
"improvement" — for once a species prevails
over others at a location, no pressure of natural
selection need arise to promote further adapta-
tion. (Competition within species will continue
to eliminate truly defective individuals and may
promote some refinement by selection of fortu-
itous variants with still more advantageous
traits, but the great majority of successful spe-
cies are highly stable in form and behavior over
long periods of geological time — not because
they are optimal, but because they are locally
prevalent.)

For this reason, many native plants, evolved
by natural selection as adaptive to their regions,
fare poorly against introduced species that never
experienced the local habitat. If natural selec-
tion produced optimality, this most common
situation could never arise, for native forms
would be "best" and would prevail in any com-
petition against intruders. But most Australian
marsupials succumb to placentals imported
from other continents, despite tens of millions
of years of isolation, during which the Austra-
lian natives should have attained irreplaceable
incumbency, if natural selection worked for
optimality rather than merely getting by. And
Homo sapiens, after arising in Africa, seems
able to prevail in any exotic bit of real estate,
almost anywhere in the world!

Thus the first-order rationale for preferring
native plants— that, as locally evolved, they are
best adapted — cannot be sustained. I strongly
suspect that a large majority of well-adapted
natives could be supplanted by some exotic
form that has never experienced the immediate
habitat. In Darwinian terms, this exotic would
be better adapted than the native — though we
may well, on defensible aesthetic or even ethi-
cal grounds, prefer the natives (for nature's fac-
tuality can never enjoin our moral decisions).

We may, I think, grant only one limited point
from evolutionary biology on the subject of
adaptation in native plants. At least we do know
that well-established natives are adequately
adapted, and we can observe their empirical bal-
ances with other local species. We cannot know
what an exotic species will do — and many, and
tragic, are the stories of exotics imported for a

The Concept of Native Plants 1

restricted and benevolent reason that then grew
like kudzu to everyone's disgust and detriment.
We also know that natives grow appropriately —
though not necessarily optimally — in their
environment, while exotics may not fit without
massive human "reconstruction" of habitat, an
intervention that many ecologically minded
people deplore. I confess that nothing strikes
me as so vulgar or inappropriate as a bright
green lawn in front of a mansion in the Arizona
desert, sucking up precious water that already
must be imported from elsewhere. A preference
for natives does foster humility and does coun-
teract human arrogance (always a good thing to
do) — for such preference does provide the only
sure protection against our profound ignorance
of consequences when we import exotics. But
the standard argument — that natives should he
preferred as best adapted — is simply false within
Darwinian theory.

The Geographic Argument Based on
Appropriate Place

This argument is harder to formulate, and less
clearly linked to a Darwinian postulate, but
somehow seems even more deeply embedded
(as a fallacy) into the conventional argument
for preferring native plants. This argument
holds that plants occupy their natural geo-
graphic ranges for reasons of maximal appropri-
ateness. Why, after all, would a plant live only
in this-or-that region of 500 square kilometers
unless this domain acted as its "natural"
home — the place where it, uniquely, and no
other species, fits best. Smyser et ah, for
example, write: "In any area there is always
a type of vegetation that would exist without
being planted or protected. This native vegeta-
tion consists of specific groups of plants that
adapted to specific environmental condi-
tions."^'* But the deepest principle of evolution-
ary biology — the construction of all current
biological phenomena as outcomes of contin-
gent history, rather than optimally manufac-
tured situations — exposes this belief as nonsense.

Organisms do not necessarily, or even gener-
ally, inhabit the geographic area best suited to
their attributes. Since organisms (and their areas
of habitation) are products of a history laced
with chaos, contingency, and genuine random-

ness, current patterns (although workable, or
they would not exist) will rarely express any-
thing close to an optimum, or even a "best pos-
sible on this earth now" — whereas the earlier
notion of natural theology, with direct creation
of best solutions, and no appreciable history
thereafter (or ever), could have validated an idea
of native as best. Consequently, although native
plants must be adequate for their environments,
evolutionary theory grants us no license for
viewing them as the best-adapted inhabitants
conceivable, or even as the best available among
all species on the planet.

An enormous literature in evolutionary biol-
ogy documents the various, and often peculiar,
mechanisms whereby organisms achieve fortu-
itous transport as species spread to regions
beyond their initial point of origin. Darwin him-
self took particular interest in this subject.
During the 1850s, in the years just before publi-
cation of the Origin of Species in 1859, Darwin
wrote several papers on the survival of seeds in
salt water (how long would they float without
sinking? would they still germinate after such a
long bath?). He determined that many seeds
could survive long enough to reach distant con-
tinents by floating across oceans — and that pat-
terns of colonization therefore reflect historical
accidents of available pathways, and not a set of
optimal environments.

Darwin then studied a large range of "rarely
efficient" means of transport beyond simple
floating on the waves: for example, natural rafts
of intertwined logs (often found floating in the
ocean hundreds of miles from river mouths),
mud caked on birds' feet, residence in the gut of
birds with later passage in feces (Darwin and
others studied, and often affirmed, the power of
seeds to germinate after passage through an
intestinal tract). In his usually thorough and
obsessive way, Darwin assiduously collected
information and found more than enough
means of fortuitous transport. He wrote to a
sailor who had been shipwrecked on Kerguelen
Island to find out if he remembered any seeds or
plants growing from driftwood on the beach. He
asked an inhabitant of Hudson Bay if seeds
might he carried on ice floes. He studied the
contents of ducks' stomachs. He was delighted
to receive in the mail a pair of partridges' feet

PETER I>EL TREDICI

8 Arnoldici 1998 Spring

Eucalyptus globulus is an important source of fuel and building material in the altiplano of South America,
where in some cases it is the sole tree. This native of Tasmania and Victoria selfsows and has naturalized
throughout the area.

caked with mud; he rooted through bird drop-
pings. He even followed a suggestion of his
eight-year-old son that they float a dead and
well-fed bird. Darwin wrote in a letter that "a
pigeon has floated for 30 days in salt water with
seeds in crop and they have grown splendidly."
In the end, Darwin found more than enough
mechanisms to move his viable seeds.

"Natives," in short, are the species that hap-
pened to find their way (or evolve in situ], not
the best conceivable for a spot. As with the first
argument about adaptation, the proof that cur-
rent incumbency as "native" does not imply
superiority against potential competitors exists
in abundance among hundreds of imported
interlopers that have displaced natives through-
out the world: eucalyptus in California, kudzu
in the American southeast, rahhits and other
placental mammals in Australia, and humans
just about everywhere.

"Natives" are only those organisms that first
happened to gain and keep a footing. We rightly
decry the elitist and parochial claims of Ameri-

can northeast WASPs to the title of native, but
(however "politically incorrect" the point), the
fashionable status of "Indians" (so-called by
Columbus' error) as "Native Americans" makes
just as little sense in biological terms. "Native
Americans" arrived in a geological yesterday,
some 20,000 years ago (perhaps a bit earlier), on
the geographic fortuity of a pathway across the
Bering Strait. They were no more intrinsically
suited to New World real estate than any other
people. They just happened to arrive first.

In this context, the only conceivable rationale
for the moral or practical superiority of
"natives" (read first-comers) must lie in a
romanticized notion that old inhabitants learn
to live in ecological harmony with surround-
ings, while later interlopers tend to he exploit-
ers. But this notion, however popular among
"new agers," must be dismissed as romantic
drivel. People are people, whatever their techno-
logical status; some learn to live harmoniously
for their own good, and others do not to their
own detriment or destruction. Preindustrial

The Concept of Native Plants 9

people have been iust as rapacious (though not
so quickly perhaps, for lack of tools) as the worst
modern clear-cutters. The Maori people of New
Zealand wiped out a rich fauna of some twenty
moa species within a few hundred years. The
"native" Polynesians of Easter Island wiped out
everything edible or usable (and, in the end, had
no logs to build boats or to raise their famous
statues), and finally turned to self-destruction.

In summary of my entire argument from evo-
lutionary theory, "native" plants cannot be
deemed biologically best in any justifiable way
(note that I am emphatically not speaking about
ethical or aesthetic preference, for science can-
not adjudicate these considerations). "Natives"
are only the plants that happened to arrive first
and be able to flourish (the evolutionary argu-
ment based on geography and history), while
their capacity for flourishing only indicates a
status as "better than" others available, not as
optimal or globally "best suited" (the evolution-
ary argument based on adaptation and natural
selection).

Speaking biologically, the only general
defense that I can concoct for natives — and I
regard this argument as no mean thing — lies in
protection thus afforded against our overween-
ing arrogance. At least we know what natives
will do in an unchanged habitat, for they have
generally been present for a long time and have
therefore stabilized and adapted. We never know
for sure what an imported interloper will
do, and our consciously planted exotics have
"escaped" to disastrous spread and extirpation
of natives (the kudzu model) as often as they
have supplied the intended horticultural or
agricultural benefits.

As a final ethical point (and I raise this issue
as a concerned human being, not as a scientist,
for my profession can offer no direct moral
insight), I do understand the appeal of the ethi-
cal argument that we should leave nature alone
and preserve as much as we can of what existed
and developed before our very recent geological
appearance. Like all evolutionary biologists, I
treasure nature's bounteous diversity of species
(the thought of half a million described species
of beetles — and many more yet undescribed —
fills me with an awe that can only be called rev-
erent). And I do understand that much of this

variety lies in geographic diversity (different
organisms evolved in similar habitats in many
places on our planet, as a result of limits and
accidents of access). I would certainly be horri-
fied to watch the botanical equivalent of
McDonalds' uniform architecture and cuisine
wiping out every local diner in America. Cher-
ishing native plants does allow us to defend and
preserve a maximal amount of local variety.

But we must also acknowledge that strict
"nativism" has an ethical downside inherent in
the notion that "natural" must be right and
best, for such an attitude easily slides to the
Philistinism of denying any role to human intel-
ligence and good taste, thence to the foolish
romanticism of viewing all that humans might
accomplish in nature as "bad" (and how then
must we judge Frederick Law Olmsted's Central
Park), and even (in an ugly perversion) — but
realized in our time by Nazi invocation of nativ-
ist doctrine — to the claim that my "native" is
best and yours only fit for extirpation.

The defense against all these misuses, from
mild to virulent, lies in a profoundly humanis-
tic notion as old as Plato, one that we often
advance in sheepish apology but should rather
honor and cherish: the idea that "art" must be
defined as the caring, tasteful, and intelligent
modification of nature for respectful human
utility. If we can practice this art m partnership
with nature, rather than by exploitation (and if
we also set aside large areas for rigidly minimal
disturbance, so that we never forget, and may
continue to enjoy, what nature accomplished
during nearly all of her history without us), then
we may achieve optimal balance.

People of goodwill may differ on the best
botanical way to capture the "spirit of demo-
cracy"— from one end of maximal "respect" for
nature by using only her unadorned and locally
indigenous ("native") products, to the other of
maximal use of human intelligence and aes-
thetic feeling in sensitive and "respectful" mix-
ing of natives and exotics, just as our human
populations have so benefited from imported
diversity. Jens Jensen extolled the first view:
"When we are willing to give each plant a
chance fully to develop its beauty, so as to give
us all it possesses without any interference,
then, and only then, shall we enjoy ideal land-

10 Arnoldia 1998 Spring

scapes made by man. Is not this the true spirit
of democracy? Can a democrat cripple and mis-
use a plant for the sake of show and pretense?"’^
But is all cultivation — hedgerows? topiary? —
crippling and misuse? The loaded nature of
ethical language lies exposed herein. Let us
consider, in closing, another and opposite defi-
nition of democracy that certainly has the sanc-
tion of ancient usage. J. Wolschke-Bulmahn and
G. Groning cite a stirring and poignant argu-
ment made by Rudolf Borchardt, a lew who later
died trying to escape the Nazis, against the
nativist doctrine as perverted hy Nazi horticul-
turists: "If this kind of garden-owning barbarian
became the rule, then neither a gillyflower nor
a rosemary, neither a peach-tree nor a myrtle
sapling nor a tea-rose would ever have crossed
the Alps. Gardens connect people, times and
latitudes. If these barbarians ruled, the great his-
toric process of acclimatization would never
have begun and today we would horticulturally
still subsist on acorns. . . . The garden of human-
ity is a huge democracy."'®

I cannot state a preference in this wide sweep
of opinions, from pure hands-off romanticism to
thorough overmanagement (though I trust that
most of us would condemn both extremes).
Absolute answers to such ethical and aesthetic
questions do not exist in any case. But we will
not achieve clarity on this issue if we advocate
a knee-jerk equation of "native" with morally
best, and fail to recognize the ethical power of a
contrary view, supporting a sensitive cultiva-
tion of all plants, whatever their geographic ori-
gin, that can enhance nature and bring both
delight and utility to humans. Is it more "demo-
cratic" only to respect organisms in their natu-
ral places (how, then, could any non-African
human respect himself), or shall we persevere in
the great experiment of harmonious and mutu-
ally reinforcing geographic proximity — as the
prophet Isaiah sought in his wondrous vision of
a place where the wolf might dwell with the
lamb and such non-natives as the calf and the
lion might feed together — where "they shall not
hurt nor destroy in all my holy mountain."

Endnotes

* I. Wolschke-Bulmahn and G. Groning, "The Ideology
of the Nature Garden; Nationalistic Trends in Garden
Design in Germany During the Early Twentieth

Century," Journal of Garden History (1992) 12(1):
73-80; G. Groning and I. Wolschke-Bulmahn, "Some
Notes on the Mania for Native Plants in Germany,"
Landscape Journal [1991] 1 1(2): 116-126; ]. Wolschke-
Bulmahn, "Political Landscapes and Technology:
Nazi Germany and the Landscape Design of the
Reichsautobahnen (Reich Motor Highways),"
Selected CELA Annual Conference Papers: Nature
and Technology, Iowa State University, 9-12
September 1995, vol. 7.

^ Quoted in Wolschke-Bulmahn, "Political Land-
scapes," from a 1939 article.

^ Quoted in Groning and Wolschke-Bulmahn, "Native
Plants."

* C. A. Smyser et al., Nature’s Design: A Practical
Guide to Natural Landscaping. Emmaus, PA.,
1982, xi.

® K. Druse and M. Roach, The Natural Habitat
Garden. New York: 1994, viii.

® President William J. Clinton, Memorandum for the
Heads of Executive Departments and Agencies,
Office of the Press Secretary, 26 April 1994.

'' J. lensen. Siftings: The Major Portion of "The
Clearing," and Collected Writings, Chicago, 1956,
45.

® Quoted in Wolschke-Bulmahn, "Political Land-
scapes," 13.

® W. Paley, Natural Theology. London, 1802.

See S. ). Gould and R. C. Lewontin, "The Spandrels of
San Marco and the Panglossian Paradigm: A Critique
of the Adaptationist Programme," Proceedings of the
Royal Society of London B (1979) 205: 581-198; see
S. J. Gould, "Exaptation: A Crucial Tool for an
Evolutionary Psychology," Journal of Social Issues
(1991)47(3); 43-65.

" lensen. Siftings, 47.

Ibid., 59.

Smyser et al.. Nature’s Design, vii.

Ibid., xi.

Jensen, Siftings. 46.

'® Wolschke-Bulmahn and Groning, "The Ideology of
the Nature Garden," 80.

Stephen 1. Gould is professor of geology at Harvard
University, curator of invertebrate paleontology' at the
Museum of Comparative Zoology, and Alexander
Agassiz Professor of Zoology. Among his books and
articles on the history of evolution and related topics are
Ever Since Darwin (1977), The Panda’s Thumb (1980),
The Elamingo’s Smile (1985), Wonderful Life (1989), and
Eight Little Piggies (1993). This article evolved from a
paper presented at the 1994 Studies in Landscape
Architecture symposium at Dumbarton Oaks — "Nature
and Ideology: Natural Garden Design in the Twentieth
Century" — and published in 1997 under the same title,
edited by Joachim Wolschke-Bulmahn.

E. D. Merrill, From Maine to Manila

Ida Hay

Twenty-two years of adventure in Southeast Asia preceded E. D. Merrill's
career as director of several important botanical institutions, among them the
Arnold Arboretum. His knowledge of the flora of Asia and the South Pacific
was encyclopedic, and it was said he could name more species at sight than
any other American taxonomist.

when twenty-six-year-old Elmer Drew Merrill
left New York harbor for Manila on February 22,
1902, he had no idea that he would remain in
the Philippines for the next twenty-two years,
laying the foundation for a botanical inventory
of the archipelago. After accepting a job offer as
botanist with the Insular Bureau of Agriculture,
he had had less than forty-eight hours to arrange
his affairs, pack, and get to the boat. This rough-
and-ready approach, spawned of a rigorous
childhood in rural Maine, was to characterize
Merrill's remarkable life: this would not be the
last time he made a major career change at the
drop of a hat.

From 1935 to 1946, Merrill was director of
the Arnold Arboretum and Administrator of
Harvard University's Botanical Collections,
which included the Botanic Garden, the Gray
Herbarium, the Bussey Institution, the Botani-
cal Museum, the Harvard Forest, the Atkins
Institution, and the Farlow Reference Fibrary
and Herbarium. When he arrived at Harvard, he
had already had sixteen years' experience man-
aging organizations with diverse functions, in
addition to an extraordinary record of scholar-
ship and publication in taxonomic botany.

Merrill was born in 1876 in East Auburn,
Maine, a village of farmers and shoe factory
workers, one of twins. He described his progeni-
tors as simple, hardworking folk who, neverthe-
less, possessed the "pioneer spirit." His
maternal grandfather was a forty-niner who
journeyed to California by way of Panama,
returning to his wife and children in Maine

without having found any gold. Merrill's father
had run away to sea at age fourteen and worked
as a common sailor until he married; he contin-
ued to sign on for extended fishing trips to the
Grand Banks during E. D.'s youth. It was the
work and the pleasures of rural life that shaped
Merrill's character, as he recalled years later:

Swimming, boating, fishing, hunting, tramping
in the woods — many things were more appealing
to us than work, hut when there was work to be
done it always came first."'

Yet even at an early age he often found time to
collect natural history specimens and to press
plants.

Unlike their three older siblings, Elmer and
his twin, Dana, continued their education
beyond the elementary grades, attending high
school in Auburn, three miles distant from their
home. In one of his more telling comments on
his background, Merrill wrote:

Many times in winter we walked the entire dis-
tance to the city in a howling hlizzard only to
find "no sessions" because of the inclement
weather. We came to have a rather scornful opin-
ion of city people, not blaming the children, but
rather the authorities. At times we made the
trip on snowshoes. . . . This school experience
doubtless had its effect in establishing one
quality — that of persistence, a quality to which I
believe I owe most of the success as I attained
in after life.^

After graduating, both young men entered the
Maine State College at Orono, which became
the University of Maine in 1898, the year they

LIBRARY OF THE ARNOLD ARBORETUM

12 Arnoldio 1998 Spring

Merrill, right, and E. B. Copeland, left, with Joseph French and, standing, Henry Osgood, in the bachelor’s mess
in Manila, ca. 1 905. From the time he arrived in the Philippines until he received an appointment as Associate
Professor of Botany in the University of the Philippines in 1912, Merrill spent at least half his time working in
the field. E. B. Copeland, who joined the botanical staff of the Bureau of Science in 1903, was one of Merrill’s
traveling companions. In 1909, accompanied by a group of American schoolteachers, the two climbed to the
summit of Mount Pulog in northern Fuzon, the third known ascent of the mountain by Westerners.

received their degrees. Although they enrolled
as engineering students, they both transferred
to the general science course after a surfeit of
math classes during their first year. During his
remaining undergraduate years, Elmer took as
many biology courses as he could and studied
the classification of flowering plants on his
own since no formal training was offered. Like
most New England botanists of his day, he
tramped and botanized on New Hampshire's
Mount Washington and likewise explored
Mount Katahdin in northern Maine. He later
gave his 2,000-specimen herbarium to the
New England Botanical Club. He also traded a
collection of his pressed plants dating from this
period to Nathaniel Lord Britton for a copy of
Britton and Brown's Illustrated Flora of the
Northern United States. Though neither of

them could have foreseen it, Merrill would one
day succeed Britton as director of the New York
Botanical Garden.

The outbreak of the Spanish-American War
determined Dana Merrill's career choice. He
enlisted in the Maine Volunteer Infantry,
received his diploma in absentia that spring,
and soon headed out to fight in the Philippines.
He remained in the Army after the war and
advanced through the ranks to brigadier general
in 1935.

Elmer remained at Orono for a year after
graduation. While he worked as an assistant in
the Department of Natural Science, he took
additional courses and continued to study sys-
tematic botany on his own. (In 1904, the Uni-
versity of Maine awarded him a master's degree
for this work.) In 1899 he went to work in Wash-

E. D. Merrill 13

ington at the U.S. Department of Agriculture
(USDA) as an assistant agrostologist (a special-
ist in grasses, a family Merrill termed "particu-
larly difficult"). He found the joh rewarding
and appreciated the opportunity to become
more familiar with the literature of plant tax-
onomy, but he was still undecided about a
career. With time on his hands evenings, he
completed a year and a half of medical school.
Then the offer of employment in the Philippines
turned him permanently in the direction of
plant science.

Among the many programs the U.S. govern-
ment started in the Philippines after taking it
over from the Spanish was an Insular Bureau of
Agriculture, opened in 1901, the year before
Merrill was persuaded by his boss at the USDA
to accept the post of botanist there. He had
expected to see his brother Dana when he
arrived in Manila after the two-month voyage,
but in the first of many ironies that would punc-
tuate his life, he found that
his twin had sailed for San
Francisco two weeks earlier.

It would be thirteen years
before the two met again.

Merrill quickly applied his
energies to the challenges of
his new assignment. The
previous two and a half years
of work on the taxonomy of
grasses had expanded his
botanical purview from New
England to Wyoming, Idaho,
and Montana. Compared to
that of the Western grass-
lands and Maine, however,
the flora that he now con-
fronted was exuberant and
vastly complex. Undaunted,
he immediately envisioned a
complete survey of the Phil-
ippine archipelago, 7,000
tropieal islands with exten-
sive, mountainous, old-
growth forest ringed by
lowlands that had been cul-
tivated for centuries.

Resources for studying
this fascinating flora were

almost nonexistent in Manila,- any botanical
specimens and literature that had been
assembled during the long years of Spanish rule
had either burned in the 1898 war or disap-
peared during the disruptive period of American
takeover. Never one to hesitate, Merrill imme-
diately started collecting weeds behind the
vacant house that served as headquarters for
the Bureau of Agriculture. And within a month
he had left on his first collecting expedition, a
six-week trek through the mountains of Luzon
to Aparri on the north coast. For the next eleven
years he would spend nearly half his time in
the field.

Government officials in Manila quickly rec-
ognized Merrill's abilities and gave him an addi-
tional appointment to the Bureau of Forestry,
thereby consolidating botanical research. In
1903 all botanical work was transferred to the
Bureau of Government Laboratories, which in
1906 became the Bureau of Science.

Bureau of Science buildings in Manila, 1916. Although Merrill’s work in the
Philippines commenced in a vacant dwelling rented as headquarters for the
Bureau of Agriculture in 1902, within three years a new facility was constructed
to house the Bureau of Science. Merrill was director of the Bureau from 1919 to
1923. The destruction of these buildings, along with most of their contents
including the herbarium and botany library, during World War 11 was a tragic
episode in Merrill’s career, even though he was director of the Arnold Arboretum
by that time.

14 Arnoldia 1998 Spring

From the outset Merrill spent much time and
energy building the reference library that was
needed to identify the rich flora he found during
his explorations. In 1902 he made a visit to the
85-year-old botanical garden in Buitenzorg, Java
(Bogor). He found the library and herbarium
there very helpful; in addition to identifying the
Philippine plants he had brought along, he was
able to familiarize himself with the botanical
literature of the Malay Archipelago, the great
chain of islands stretching from southern Asia
to northern Australia. Undoubtedly this visit
inspired his efforts to amass similar resources in
Manila: by the time he left the Philippines in
1923, the herbarium had grown from almost
nothing to over 250,000 specimens, comple-
mented by a library he characterized as one of
the most complete in all of Asia.

Adventures in the Field

Merrill's travels in search of plants took him
the length and breadth of the archipelago and
included remote areas where few, if any Filipi-
nos, let alone Westerners, had set foot. One of
these was the summit of Mount Halcon, which
he and a party of forestry and military personnel
reached in November 1906 after twenty days of
arduous, wet climbing. There existed no report
of Westerners having previously attained the
summit of this mountain; and apparently local
Mangyan tribespeople had never ascended
either, for no signs of trails were seen any-
where near the peak and Merrill was sure that
no human could get there without cutting a
trail, so dense was the mossy forest and so steep
the terrain.

Halcon, at 8,500 feet the third highest moun-
tain in the Philippines, is located on northern
Mindoro, one of the most humid areas in the
entire country. Halcon and its subsidiary ranges
capture an enormous amount of precipitation
nearly year-round, and the mountain is continu-
ally shrouded in fog and clouds. During the
ascent Merrill encountered the entire gamut of
rainforest vegetation that he later came to know
so well. Starting from Calpan to the north, the
party soon left behind the coastal lowland with
its mangrove swamps, cultivated crops, and
abundant tropical weeds. They followed river
courses and occasional Mangyan trails through

dense vegetation dominated by huge trees with
canopies so high and thick that only twilight
reached the forest floor. For the most part
this was primary forest with a several-storied,
species-rich mix of trees that included many
Dipterocarpaceae. They also encountered many
areas of secondary forest, the abandoned clear-
ings of the Mangyan people who regularly
cleared a few acres of the old-growth forest,
burned it over, then planted upland rice, corn,
and other crops for a year or two before moving
on to a new area. Once cultivation stopped,
these clearings were rapidly re-vegetated by a
mix of indigenous and introduced plants quite
different from those of the original rainforest.

Travel was extremely difficult. The rivers the
party followed often led them into steep-sided
ravines, forcing them to ford the swift water fre-
quently. Then, after finally reaching the ridges
at the top of the canyon walls, they had to hack
their way slowly through more forest using
bolos, the Filipino equivalent of machetes.
Sometimes the only way to proceed was to chop
their way up 80-degree slopes.

Once they attained 4,000 feet, the vegetation
began to change markedly to that known as
the mossy forest — a diverse mix of smaller trees
including oak, maple, and several Malaysian
genera with many-branched, scraggly habits, as
well as Rhododendron, Vaccinium, Rubus, and
other shrubby genera found in more temperate
regions. Moisture-loving ferns, mosses, and epi-
phytes grew even more profusely here at higher
elevations than they had in the lower forests;
Epiphytic ferns and orchids . . . become more
plentiful and there is a greater diversity in spe-
cies,- mosses are much thicker and more luxu-
riant, enwrapping even the branches and
branchlets of trees and forming a deep, soft, soil
cover, frequently a foot in thickness.^

The going was not easier in the mossy forest,
even though the woody vegetation became more
and more stunted the higher they climbed.
Thickets of gnarled trees and branching shrubs,
covered with epiphytes and intertwined with
vines, allowed no forward progress without first
clearing a trail step by step. The temperature
had dropped considerably, averaging 60 degrees
Fahrenheit in the daytime, and a rainy period
that lasted thirteen days set in.

E. D. Merrill 15

I

Veitchia merrillii (formerly Adonidia merrilliij. The Christmas palm or Manila palm is admired for
its pendulous clusters of crimson fruit, which contrast attractively with its whitish fruit stalks and
sheaths. It was known only from cultivation in the vicinity of Manila when named for Merrill by
Italian palm specialist Odoardo Beccari (1843-1920). Later the Manila palm’s native habitat was
determined to be restricted to Palawan and the Calamianes Islands on the basis of specimens
collected by Merrill and A. D. E. Elmer, one of his colleagues at the Philippine Bureau of Science.

16 Arnoldia 1998 Spring

Surprisingly, when they reached 7,800 feet,
the montane brush gave way to vegetation
Merrill described as open heath, a collection of
tufted grasses broken only occasionally by
stunted trees and shrubs. They quickly tra-
versed this area only to find that the final 500
feet of elevation was covered with thickets
more dense than any they had previously
encountered:

At times as we came to the crest line, the cold
wind would add to our discomfort. . . . Pitcher
plants (Nepenthes) became very abundant, clam-
bering everywhere in the thickets, so that in cut-
ting our way through the underbrush, at frequent
intervals our bolo slashes would upset the equi-
librium of from one to a half a dozen pitchers,
each holding one-half quart or more of water,
which would be precipitated upon us. These irreg-
ular douches were far more disagreeable than the
constant shower bath from the falling rain.'*

In storms worse than ever, Merrill and
another scientist reached the summit, where
clouds obscured the view. They quickly took
barometric readings and left a record of their
visit sealed in a bottle tied to a tree, since there
were no boulders to use for a cairn.

The return trip to the coast took nearly as
long as the ascent. They were delayed by more
storms, and two members of the party became
lost for a while. When the porters were sent
back to retrieve supplies left at lower elevations
and got cut off by rain-swollen rivers, the party
had to forage in the rainforest for a Thanksgiv-
ing "dinner" of broiled wood rats and boiled fern
tips. Merrill commented that "a man can come
nearer to starving to death in a primary tropical
forest than in almost any other part of the
world," since there is little game, and edible
fruit is either too high in the canopy or too
widely spaced for efficient harvesting. It was
some consolation to Merrill that a new species
was later described from the rat skins and skulls
left over from the holiday dinner.

Although this was probably the most strenu-
ous of his field trips, Merrill accepted many
more challenges in his search for the Philippine
flora. On some occasions he walked 36 miles in
a single day. There were precarious landings in
the surf on remote coasts, and the unnerving
experience of collecting plants among the hast-

ily made graves of tribesmen who had resisted
American troops. And at times he risked his
life by staying overnight in remote villages of
the Mountain Province, where headhunters
were reputed to live.

In order to determine the relationships
between the flora of the Philippines and those of
surrounding areas, as well as for help in identi-
fying certain species, Merrill and his associates
at the Bureau of Science also made collecting
trips to Guam, Borneo, Amboina, Indochina,
and China. He acquired additional specimens for
the herbarium collection in Manila by exchang-
ing material from the Philippines for Indo-
Malaysian, Australian, and Polynesian plants.

Publications

Of course, the fieldwork was only the beginning
for Merrill. His observations in the field
and subsequent scrutiny of pressed specimens,
along with intense study of botanical literature.

Elmer Drew Merrill photographed in Manila, 1914.

Janet Steams

Tne Arnold ArLoretum

NEWS

Katherine H. Putnam Research Fellowships Endowed

George Putnam, chairman, Putnam Funds and the Putnam Investment
Company (second from left), and his wife Nancy Putnam, display a photo
taken in turn-of-the-century China by E. H. Wilson; it was presented to
them by the Arboretum at a reception held in their honor. Standing with
the Putnams are James McCarthy (left), director, MCZ, and Robert E.
Cook, director. Arboretum.

Fifty friends of George and Nancy
Putnam gathered in Cambridge
on Friday, May 15, to show their
appreciation for two generous
endowments established by the
Putnams. Two of Harv'ard’s oldest
biological institutions, the Arnold
Arboretum and the Museum of
Comparative Zoology, will benefit
from endowments of $1 million
each to support research and
scholarship.

The endowment at the Arnold
Arboretum will support the
Katharine H. Putnam Research
Fellowships, established in
memory of Mr. Putnam's mother,
an accomplished horticulturist
and long-time supporter of the

Arboretum. The funds will pro-
vide fellowship stipends and
related research and project
expenses for work in horticulture
and botany using the Arboretum's
living collections of trees and
shrubs.

Putnam Fellowships will fund
graduate students, postgraduate
scholars, and mid-career profes-
sionals who wish to experience the
richness of the Arboretum's
resources and engage in research
work that generates new knowl-
edge and practical applications for
horticulture, landscape architec-
ture, and plant conservation.
Fellowship awards will be particu-
larly appropriate for young

research scientists contemplating a
career in public horticulture and
education.

"George Putnam has been a
friend of the Atboretum for many
years," says director Bob Cook.
“With the establishment of the
Putnam Fellowship endowment,
we can continue to offer the most
promising scientists an opportu-
nity to work with our collections
and to gain the kind of practical
experience that is essential fot
leadership nationally. We deeply
appreciate the commitment of
George and Nancy Putnam to
this critical research and the edu-
cational mission of the Arnold
Arboretum.”

What a Difference a
Year Makes

Peter Del T redid. Director of
Li ring Collections

On Monday, March 31, of last
year, I was sitting in Patrick
Willoughby's office in the base-
ment of the Hunnewell building
as he told me that he had decided
to take the job as head of grounds
maintenance at Wellesley College,
and that his last day as Superin-
tendent would be May 1. As I lis-
tened to Patrick talk about his
future, I was looking out the win-
dow at the occasional snowflakes
that had just started to fall. “It
won't stick,” I said, 'none of it has
this winter." Within 24 hours 1

was earing tliose words. Two feet
of sticky wet snow had been
dumped on the city of boston,
stopping traffic, downing power
lines, and smasliing trees. As most
people reading this will remem-
ber, the Arnold Arboretum was
particularly hard hit: over 1 ,8{)()
trees were damaged, of which 200
have been removed.

but now it is one year later and
1 am pleased to report that a new
superintendent, Julie Coop, is well
established in Patrick’s old office,
and most of the storm damage has
been cleaned up. Of particular

interest is the fact that the high
temperature on the day of the
storm was 42 degrees while on
the same date one year later, the
high was 92! What a difference a
year makes.

Spring planting, which was
virtually nonexistent last year, has
gone very smoothly this year.

Over 150 new trees were set out
during April alone. Unlike last
year, this spring there was no
snow to speak of and the weather
was moist and cool. This not
only allowed the grounds crew
to start digging early, but also

to dig and plant throughout the
entire month.

On behalf of all of us at the
Arboretum, 1 want to take advan-
tage of this dubious anniversary to
thank all of our loyal friends and
supporters who generously donated
labor and money to our storm
damage cleanup effort. The dona-
tions not only helped with the
clean-up, but they also lifted the
spirits of the entire Living Collec-
tions staff It's great to know that
people care deeply about the
future of the Arnold Arboretum.
Thank you very much.

CHALLENGE GIFT FOR CHILDREN'S
SCIENCE EDUCATION

The Arboretum has received a challenge gift of S 100, ()()()
from an anonymous donor. The gift has been directed to
Children’s Science Education and is intended to encour-
age others to help endow the Arboretum’s Field Studies
Experiences (FSE) program.

The FSE program brings children directly from Bos-
ton-area classrooms to the Arboretum’s landscape. Stu-
dents work in small groups with aguide while exploring
and discussing specific science cjuestions related to one
of four different themes: Flowers Change; Plants in
Autumn; Native Trees, Native Peoples; and Around the
World in Trees. Each year the program serves 3,000
schoolchildren in grades three through five.

The Arboretum’s goal in the Harvard University
Campaign is to establish an endowment of S2, 250, 000
that will secure funding for children’s science educa-
tion. Of the total goal, 5750,000 will create an endow-
ment for the FSE program. To date, 5439,000 has been
raised, representing 599f of the goal. “The creation of an
endowment for children’s science education is critical
to our education mission,” says director Hob Cook. “It
addresses a pressing need for excellence in science
education and demonstrates our commitment to chil-
dren, our most important resource for the future.”

If you are interested in making a gift to help the
Arboretum qualify for this challenge gift, or would like
a copy of our publication “The Arnold Arboretum — An
Outdoor Classroom,” please contact Lisa M. Hastings,
Director of Development, at 617/524-1718 x 145.

Harvard Announces
Women's Matching Fund

At a recent forum on women and philan-
thropy, Harvard announced an initiative
aimed at encouraging more women to
participate in philanthropy. The Women's
Matching Fund will match any gift
between 525,000 and 5250,000 made by
women to any part of the University.

The Women’s Matching Fund was cre-
ated by National Campaign Chair Rita E.
Hauser as a way for women to maximize
the impact of their gifts to Harvard. Ms.
Hauser established the fund with her own
gift of 55 million and encouraged other
women to bring the fund balance to 515
million. Gifts made by women will be
matched on a dollar-for-dollar basis until
the fund is depleted.

This new fund offers women who are
considering a campaign gift to the Arbo-
retum a unique opportunity to double
their gifts. Gifts qualifying for the match
can be directed to any of the Arboretum’s
campaign priorities: Living Collections
(including landscape maintenance
projects). Children’s Science Education,
International Biodiversity Conservation,
and the new Shrub and Vine Garden.

For more information about the
Women’s Matching Fund, contact Lisa
M. Hastings at 617/5 14-1 ~ 18 x 145, or
Peg Hedstrom at x 113.

2

SPRING 1998

Community Science Connection Goes National

Candace Jnlyan, Director of Education

In 1995 the Arnold Arboretum received a four-
year grant from the National Science Foundation to
develop a program to strengthen elementary science
teaching and learning and to illustrate ways that sci-
ence institutions can work throughout the year with
local classrooms. The result ol this effort is the Com-
munity Science Connection (CSC), a seasonal study
of trees that spans the entire school year.

Participating teachers begin their study of trees
in the spring and summer, learning how' to look
closely for patterns and changes and to make sense of
what they see. In the fall, students begin their work
by identifying individual trees and recording the
dates when color change begins and ends and when
the leaves drop. In the winter, students learn to
“read” the information found on a twig and to use
that knowledge to determine the best growing
year for their schoolyard trees. In the spring, the
study question turns to “What comes out of a bud?”
While the main focus is on working directly with
schoolyard trees, technology provides a connection
among the participating classrooms and the
Arboretum staff A project web page enables partici-
pants to share data and ideas and provides virtual
activities that encourage and support the outdoor
investigations.

The goals of CSC are twofold: to work directly
with local teachers on meaningful ways to study sci-
ence through an investigation of trees and to develop
a model that can be replicated by other institutions
to further their work with local teachers. The first
year of the project focused on the first goal. To date
we have worked with over fifty teachers in the Bos-
ton, Newton, and Brookline schools. In the final year
of the project we will continue to work with local
teachers and begin work with other science institu-
tions. Descanso Gardens, in collaboration with the
science coordinator of the Los Angeles public
schools, will replicate our tree investigations with
California students. (The director of Descanso, Rich-
ard Schulhof, worked on the CSC project during its
first year in his former role as the Arboretum’s direc-
tor of public programs.) Los Angeles students and
their teachers will be corresponding with Massachu-
setts colleagues through the website, comparing
findings from coast to coast. This work will give us
an opportunity to determine the viability of our
model for other institutions.

Another interesting development in this project
emerged during the past year. The Massachusetts
Audubon Society (MAS) was intrigued by our
seasonal investigations and used it as a model to
create a year-long study of vernal pools. During the
1998—1999 school year, groups of teachers from
eastern, central, and western Massachusetts will
begin a coordinated study of vernal pools that will
continue through fall, winter, and spring based on a
curriculum ourline developed by MAS. These inves-
tigations will be supported by the same technology
component as the tree studies, with a data exchange,
opportunities for conversations, and virtual activities
that go along with the actual investigations. All of
the technology portion of the MAS project will
originate from the Arboretum and the virtual activi-
ties will be a joint development venture between
staff from MAS and the Arboretum.

While the conversations of the participating
classes will remain private, all of the activities for
these investigations will be available to interested
individuals through the Arboretum’s web page
(www.arboretum.harvard.edu). We welcome your
comments.

ARNOLD ARBORETUM NEWS

Karen Madsen

On the Grounds

Tom Akin has joined the Arboretum staff as Assistant
Superintendent of Grounds. A canditlate for the master's
degree in |ilant and soil sciences at the Imiversity of
Massachusetts, Amherst, he has worked with the
L’niversity’s Extension Service both as a research
assistant and extension educator. He comes to us from
W'eston Nurseries where he was IFM (Integrated Pest
Management) Coordinator. Tom’s resume also includes
work with the Peace Corps in the Central Africa
Republic, first as an English teacher, then as keeper
of African killer bees. No doubt he will bring all these
experiences to bear at the Arboretum, where his
duties will include the coordination of the summer
horticultural intern program and its associated

training program.

So Long, Fare Well

Jim Ciorman, Arboretum tour leader and
volunteer coordinator nonpareil, is leaving us
for southeastern Pennsylvania and the
Longwood Gardens Ciraduate Program in
public horticultural administration. An
official staff member at the Arboretum since
1992 and an unofficial staff member even
longer, Jim Ciorman and the Arnold
Arboretum have been synonymous for many
in Boston. Wherever his future takes him, we
know he will continue to be the best emissary
we could hope for. Needless to say, we will
miss him, and we wish him all the best.

Annual Fall Plant Sale

Sunday, September 20, 1998
Case Estates, Weston
9:00 am to 1 :00 pm

The date has been set for this year’s Annual Fall Plant Sale,
and we encourage members of the Ftiends of the Arnold
Arboretum and the general public alike to mark their
calendars. As usual, the event will feature a wide array of
unusual trees, shrubs, herbaceous perennials, and more.

Member benefits at the Plant Sale include “members only”
hours from 9:00 to 10:00 am, a free plant of your choice,
and a 10% discount on all plant purchases in the Barn.
Members at the Sustaining Level (SI 00) and above gain
entrance to the Plant Sale Preview at 8:30 and receive
additional free plants. Call the Membership Office at
617/524-1"' 18 X 165 to join or to upgrade your member-
ship and increase your Fall Plant Sale benefits!

Live and Silent Auctions
Straight Sales
Society Row

Education Sessions in the Teaching Garden
Re/reshments

Catalogs will be mailed to members in August, and your
free plant vouchers will arrive in early September.

To volunteer to help out at the Plant Sale, call Kara
Stepanian at 6l7/524-ni8 x 129.

SPRING 1998

Karpn Madspn

E. D. Menill 1 7

A circle of distinguished friends photographed in the 1940s. Seated from left. Merrill, plant explorer and
collector David Fairchild, naturalist and herpetologist Thomas Barbour, and standing, citrus hybridizer
Walter T. Swingle and paleontologist Theodore White, in Barbour’s Florida garden.

became the material for a prodigious output of
publications. He worked assiduously not only
on problems of identification and classification
but on nomenclature and bibliography as well.
In the course of this work, for example, he pub-
lished several papers updating Manuel Blanco's
1837 Flora de Filipinas. His long-term goal was
to produce a complete descriptive flora for the
Philippines, but first many new species had to
be described and published, and their relation-
ships with other plants explained.

"New or Noteworthy Philippine Plants," a
series of some seventeen papers, was published
intermittently from 1904 to 1922. Merrill also
published about twenty revisions of genera or
families as they occur in the Philippines. Alto-
gether, between 1904 and 1929, he authored one
hundred strictly taxonomic papers on the Phil-
ippine flora. Most were published in the Botani-
cal Section of the Philippine Journal of Science,
which Merrill edited from 1907 to 1918.

The publication in 1912 of tbe 500-page A
Flora of Manila was a major step toward his

longer-term goal. Since the 1,007 species it cov-
ered— a small percentage of the total known for
the entire country — were those that inhabited
low altitudes and could be found in most towns,
this work provided a useful guide for the Philip-
pine people.

But the tasks Merrill assigned himself were
not limited to the Philippine flora. In the course
of studying the origins of Philippine plants and
their relationships to the vegetation of neigh-
boring regions, he wrote exhaustive commen-
taries on the work of earlier botanists, including
the pre-Linnean work of Rumphius on the flora
of Amboina in the Moluccas, and the Flora
Cochinchinensis (1790) of Portuguese mission-
ary Juan Louriero; assembled a great deal of
information on the literature of Malaysian
botany, and became an expert on the local
names for plants of Southeast Asia as well as the
biogeography of the region. He also published
papers on the plants of Borneo, Guam, Sumatra,
Hainan, and Papua, often based on tbe many
specimens that he received from those areas.

ARCHIVES OF THE FAIRCHILD TROPICAL C.ARDEN

18 Arnoldia 1998 Spring

Having begun his botanical career at the
USDA and gone to the Far East under the aus-
pices of the Department's divisions of agricul-
ture and forestry, Merrill was ever aware of the
practical aspects of plant science and of the
human influence on the flora. His observations
on introduced weeds, cultivated plants, and
local plant names initiated a lifelong interest in
the origins of agriculture and the migration of
plants in pre-Columbian times. "The American
Element in the Philippine Flora" (1904), "Medi-
cal Survey of the Town of Taytay: The Principal
Foods Utilized by the Natives" (1909), and
"Notes on the Flora of Manila with Special Ref-
erence to the Introduced Element" (1912) are
some of his earliest papers in economic botany.

In all his many publications on the flora,
Merrill rarely failed to comment on the destruc-
tion of forests and other changes in ecosystems
caused by human activities;

The practical extermination of the original veg-
etation of those regions best adapted to agricul-
tural pursuits is a subject that deserves more
consideration than it has received. Unquestion-
ably, many species of plants have been extermi-
nated in various parts of the Malayan region
within the past century as the population has
increased. The areas being devoted to agriculture
are being rapidly enlarged . . . and the consequent
destruction of primeval forests over large areas is
a strong argument in favor of vigorous and inten-
sive botanical exploration of Malaya.®

The enormous trees and shade plants charac-
teristic of the primary forest cannot persist
under the conditions demanded by modem agri-
culture, and they cannot exist in second growth
forest, grasslands, and bamboo thickets that
rapidly encroach on cleared areas that are aban-
doned. . . . We are witnessing in our own genera-
tion the rapid extermination of some of the
noblest types of tropical vegetation . . . ®

When Merrill wrote these words, the popula-
tion of the Philippine Islands was less than that
of greater London; today the population is ten
million greater than that of all the British Isles.

Becoming an Administrator

Merrill would have loved to spend all his time
working in systematic botany, but in 1912 a
series of additional appointments began to
claim much of it. In that year he was appointed

Associate Professor of Botany at the University
of the Philippines; subsequently, his teaching
duties would occupy from 18 to 36 hours per
week. Then, in 1919, he was appointed director
of the Bureau of Science after a six-month stint
as acting director. In this capacity his responsi-
bilities included medicine, public health, chem-
istry, weights and measures, materials testing,
geology, mining, fisheries, zoology, and anthro-
pology, in addition to botany. Although he
accepted the position "with diffidence and
reluctance," he found in himself a talent for
handling problems in fields widely divergent
from his own, and his executive ability quickly
won him respect. It is perhaps not surprising
that the botanist whose identical twin became
a brigadier general turned out to have a knack
for administration.

But his new role cut even more severely into
the time available for preparing the major work
he had contemplated:

My appointment of Director of the Bureau of Sci-
ence in 1919 clearly indicated to me that I could
scarcely hope to consummate my plan of prepar-
ing and publishing a general descriptive flora of
the Philippines, as I soon realized that most of
my botanical work would of necessity have to be
done outside of office hours. I accordingly com-
promised with myself and . . . commenced the
actual preparation of my 'Enumeration of Philip-
pine Flowering Plants.'’

The four-volume Enumeration was issued be-
tween 1922 and 1926. In it Merrill attempted to:

account for all binomials accredited to the
Philippine flora, adjust the synonymy, cite all
important literature references, illustrative
[specimens] when desirable, detennine the Phil-
ippine and extra-Philippine distribution of each
species and record native names.®

While it was not the complete, descriptive
work that he had hoped to produce, it was a
valuable summation of all that he and his col-
leagues had accomplished. The Enumeration
allowed Merrill to outline his conclusions on
the relationship of the Philippines' climate, geo-
logic history, and plant life to those of adjacent
regions. Also included were discussions of the
original settlement of the islands; their peoples
and languages; and the history of botanical
study in the Philippines. Unexpectedly, the

E. D. Menill 19

Enumeration served as a kind of closure to
Merrill's years in the Philippines, for as it
turned out, he left Manila in the fall of 1923
never to return.

The Scientist-Administrator Moves On

Merrill's departure tvas almost as abrupt as his
arrival: he was given only a week to decide
whether to accept a position as dean of the Col-
lege of Agriculture at the University of Califor-
nia. Had there been no family dependent on
him, he would undoubtedly have remained in
the Philippines. But in 1907 he had married
Mary Augusta Sperry of Illinois. After the wed-
ding in Manila, the couple spent a year traveling
to China and Japan, followed by a several-month
stay in Washington, D.C., and visits to London,
Leiden, Berlin, and Florence, where Merrill
studied in herbaria. Once settled back in
Manila, Mary gave birth to three children over
the next seven years. When the third child died
in infancy, the Merrills concluded that "Manila
was not the proper place in which to bring up a
family." In 1915, at the end of another visit to
Washington, Mrs. Merrill stayed on with the
two children. Elmer returned to Manila and did
not see his fourth child, born in 1916, until she
was nearly five years old.

It was not easy to leave the scene of so many

years of work, the city in which I made such

reputation I bear as a botanist.^

As he left Manila in 1923 Merrill took some
comfort in the good will of his American and
Filipino colleagues in the Bureau of Science and
in the resources that he left behind for the ongo-
ing work of inventorying the Philippine flora: a
fine library and herbarium, and an exhaustive
body of research. Through the field collecting of
Merrill and his coworkers, the list of known
Philippine species had been extended from
2,500 plants of all types in 1900 to 8,120 species
of flowering plants, 1,000 species of ferns, and
3,000 species of cryptogams by 1926, when the
final volume of the Enumeration was published.
Perhaps the greatest of all the ironies in
Merrill's life would come during World War 11,
when the collections of the Bureau of Science
were destroyed by Japanese bombs.

By that time Merrill was at the Arnold Arbo-
retum and in a position to help rebuild the col-

lections. As soon as the fighting ended, he ral-
lied curators at Harvard and other major her-
baria to send duplicate specimens and library
materials to the Philippines. Work on the com-
plete flora of the Islands has been carried
forward in recent years by Philippine and
American botanists at the Philippine National
Herbarium, the Bishop Museum, and the
Botanical Research Institute of Texas, using
Merrill's meticulous scholarship as a starting
point. Tragically, many of the plants to be
included may no longer exist by the time the
flora is published, since rainforest is being
destroyed in the Philippines at a rate second
only to Madagascar's. Of the extensive primary
forests that once covered the mountainous
archipelago, current estimates are that less than
three percent remain intact.

Endnotes

* E. D. Merrill (hereafter EDM.), "Autobiographical:
Early years, the Philippines, California," Asa Gray
Bulletin n.s. (1953) 2(4): 338.

^ Ibid.

^ EDM, "The Ascent of Mount Halcon, Mindoro,"
Philippine Journal of Science, section A (1907) 2(3):
200.

Ibid., 195.

5 EDM, "An interpretation of Rumphius's Herbarium
Amboinense," Bureau of Science Publication. Manila
(1917) 9: 25-26.

^ EDM, "A bibliographic enumeration of Bornean
plants," Journal of the Straits Branch of the Royal
Asiatic Society (1921) Special number: 27-28.

^ EDM, "Autobiographical," 357.

8 Ibid.

9 Ibid, 359.

Eor Eurther Reading

Merrill, E. D. 1945. Plant Life of the Pacific World. NY:
Macmillan.

. 1955. Real values. Asa Gray Bulletin n.s. 3(1):

27-32.

Editors. 1946. Merrilleana, a selection from the general
writings of Elmer Drew Merrill. Chronica
Botanica 10: 127-394.

Ida Hay, a member of the Arnold Arboretum staff for
over twenty years, is the author of Science in the
Pleasure Ground, a history of the Arnold Arboretum
published in 1995 by Northeastern University Press. She
now lives in Northampton, Massachusetts.

Light in a Bottle: Plant-Collecting in

the Philippines

Rob Nicholson

There has been no better time to be a field botanist. It is from the world of
plants that cures are again being sought, and compounds isolated from plants
are being tested for their anti-viral, anti-cancer, and anti-fungal activity. A
small corps of botanists are journeying to the field, gathering samples of bark,
leaf, and root, and trundling them back to the biochemistry labs of the world.

The status of botanical compounds has risen in
recent years with the success of the taxane
group of compounds in fighting cancer. How-
ever, funding for research on medicinal plants is
notorious for its boom-and-bust cycles, and if
the current well-funded efforts bring no major
leads, a new downcycle in support may he trig-
gered. At the same time, there is an urgent sense
among botanists returning from the field that
the work must be done now or never. The
relentless spread of human population means
that forests continue to he turned into fields and
pastures and that individual plant species are
harvested into extinction by the multitudes
of people who value their timber, flowers, or
aphrodisiac bark.

Botanical field collecting for medicinal leads
usually proceeds in two phases. Initially a broad
assortment of plants is collected from a certain
region or from a specified set of plant families.
Extracts are made and laboratory-tested for
effectiveness against a variety of diseases. If a
particular species shows promise — the San
Pedro cactus, let us say — the second phase
begins with a search for different populations
of the same species or for other species within
the genus — perhaps the San Roberto cactus^ —
in hopes of achieving a still higher level of effec-
tiveness. Essentially, then, it is find a needle in
the haystack, then find a better needle.

Over the last twelve years most efforts have
focused on cancer and the HIV virus, and scores
of botanical compounds have been tested on cell

cultures of these diseases. The work is coordi-
nated by the Developmental Therapeutics Pro-
gram of the National Cancer Institute (NCI)
under the directorship of Dr. Gordon Cragg.
Since 1985 over 30,000 plant extracts have been
tested for in-vitro effectiveness against sixty
types of cancerous tumors, and over 52,000 have
been tested against HIV since 1987. Once an
effective compound has been isolated and pat-
ented, the NCI licenses it to a pharmaceutical
firm for further development.

Homolanthus as a Potential Anti-HIV
Therapy

Dr. Cragg reported in 1994 that "four novel
plant-derived agents with in-vitro anti-HIV
activity have been isolated and selected for
preclinical development." Among these are
extracts from a Conospermum species of Aus-
tralia, Ancistrocladus korupensis of Cameroon,
and Calophyllum of southeast Asia, which was
collected and identified by researchers at
the Arnold Arboretum and is now in Phase II
human clinical trials. A fourth genus of interest
to NCI is Homalanthus.

There are about 35 species of Homalanthus,
ranging throughout Indomalaysia and Poly-
nesia. Because none of these is very appealing
aesthetically, little had been written about them
since 1914, when Elmer Merrill (later director of
the Arnold Arboretum) described a number of
Homalanthus species from the Philippines.
Then in the late 1980s, after a decade of work-

Light in a Bottle 21

In lush and humid lowland lainfoiest. the plant-collecting party stops to catch their breath before a final
push up Mt. Apo. the highest peak in the Philippines.

Amid razor-sharp sawgrass and burnt trunks, expedition members were successful in locating a handful
of young Taxus plants to be tested for anti-cancer activity.

ME! VIN SIIEMLUCK

22 Arntildia 1998 Spring

ing with native healers in Samoa, botanist Paul
Cox ot Brigham Young University returned to
the United States with a number of samples for
laboratory analysis, together with documenta-
tion of their local medicinal uses. One of Cox's
specimens was Homalanthus nutans. As mem-
bers of the euphorbia family, Homalanthus are
related to such plants as crotons, poinsettia, cas-
sava, and castor beans. Some members of the
family contain milky latexes that cause gas-
trointestinal poisoning, dermititis, or tumors,
but a number of them have been used medici-
nally worldwide to relieve toothaches and oral
infections, or as emetics and laxatives.

Plants of the genus Homalanthus are small
trees, weedy colonizers that thrive along the
edges of roads and fields or in newly opened
spaces in the forest canopy. Samoans know
them as mamala and use their bark, leaves,
stems, and roots to treat a variety of ailments.

one being yellow fever. In Western laboratories
a compound extracted from the wood,
prostratin, was found to strongly inhibit the
killing of human host cells in-vitro by the HIV
virus. This first flash of promise set off a cycle
of activity. Botanists began collecting and study-
ing the plant, and chemists initiated studies to
decipher its mode of activity, thus giving it sta-
tus as a candidate for clinical trials.

Botanists Take To the Field
At the same time that studies were revealing
the medicinal potential of Homalanthus, my
research partner. Dr. Melvin Shemluck, and I
secured a grant from the United States Depart-
ment of Agriculture to continue our previous
research on wild populations of yew, this time
collecting live material in the Philippines.
Taxus, the yew genus, had been the subject of
intense research for over a decade thanks to the

The cabbage fields of Mt. Pulog represent a classic dilemma of developing countries: increased food production vs.
biodiversity conservation.

Light in a Bottle 23

discovery of the compound taxol, an anti-cancer
agent found in its needles and bark. We were
focusing on disjunct yew populations in order
to learn which species or populations produce
the most taxol, a crucial piece of information
for selecting the best ones for biotechnological
applications or plantations. We also hoped to
determine the conservation status of wild
stands of yews in the Philippines, as well as to
procure samples for research groups that are
studying the genetic and ecological aspects of
yew biology. Since the intriguing results of tests
against the HIV virus using prostratin were
becoming well known by this time, the USDA
extended our mandate to enable us to search for
new species or populations of Homolanthus
while we looked for yew.

Our pretrip research led us to an obscure book
on the shelf of the Smith College library. The
Vegetation of the Philippine Mountains, written
in 1919 by William Brown, an associate of Elmer
Merrill. His meticulous accounts were rich in
all the kinds of data that help in planning a col-
lecting trip, including the timing of monsoons
and the altitude of various plant habitats. Brown
also described the broad categories of forests
found in the Philippines. At the lowest altitudes
is the lowland Dipterocarp forest, a tropical
rainforest with three stories of trees and a shrub
and herb layer at the base. Some of these
Dipterocarp trees can reach 130 feet in height
and have been the source of Philippine
mahogany lumber for hundreds of years. The
lower montane, or mid-mountain, forest is
found at altitudes of 3,300 to 8,200 feet. Ever-
green oaks and laurels are the major component
of this two-storied forest, with southern hemi-
sphere conifers such as Podocarpus and Agathis
found in this association. Finally, the lower
montane mist forest, or mossy forest, is a
high-altitude, single-layer assemblage of low-
growing, gnarled, mossy trees bathed by daily
mists or rain.

We acquired visual familiarity with
Homalanthus by studying dried, pressed speci-
mens in herbaria, noting the botanical charac-
teristics of species that would help us
distinguish them in the field. Field notes on her-
barium labels give invaluable clues to location
and often include altitude, associated species.

local names of plants, and sometimes, more
colorful information, as with the sample of
Homalanthus nutans. Collected in the Solomon
Islands by S. F. Kajewski in 1931, the notes
included, "When a man has been infected by an
evil spirit the sap of this tree is drunk to get rid
of this spirit."

On the Ground in the Philippines

Our first stop in the Philippines was the
National Museum in Manila, where we con-
sulted with Philippine flora expert Dr. Domingo
Madulid and put together our team. Our next
destination was Mt. Pulog; at 9,607 feet it is the
highest point on the northern island of Luzon.
We drove north from Manila through a snarl of
jeepneys, buses, cars, and motorbikes, at one
point passing a 40-foot deep lahar flow six miles
from its source, Mt. Pinatubo. Our party of four
botanists and driver stayed that night in the
mountain resort of Baguio, a city much damaged
by a recent earthquake. The next day we began
our ascent of Mt. Pulog on a road whose quality
declined drastically as we hit steeper terrain. As
if to further emphasize the power of nature in
the Philippines, a front of thunderheads began
to drop its moisture on us, making the last
twenty miles a battle up the deeply rutted
road, its clay soil slippery from the rain. Each
time our efforts seemed doomed, we somehow
cajoled our vehicle forward.

After reaching the entrance of Mt. Pulog
National Park, we moved our gear into the cabin
that serves as the park's headquarters and the
camp for its solitary caretaker. While the torrent
continued into dusk, we dried out under the tin
roof, peering out at a Homalanthus plant across
the stream that had once been a road. The clear
light of morning showed the magnitude of
settlement on the surrounding slopes: the for-
ests of Benguet pine, Pinus insularis, had been
intensively cut right up to the park's boundary,
and the fields of the local Ifugao tribesmen sur-
rounded our cabin.

Ironically, this degraded forest was ideal habi-
tat for Homalanthus, and we had no problem
finding several specimens of a large-leaf species,
H. megaphyllus, a small tree with thick
branches, large minaret-shaped buds, and a dis-
tinctive, rounded leaf with a red stem. We soon

MELVIN SHEMLUCK

24 Arnoldia 1998 Spring

A robust plant of Homalanthus megaphyllus
headquarters of Mt. Pulog National Park.

had a collection of a half-dozen samples of wood
with corresponding cuttings for propagating
back at the Smith Botanic Garden.

The cabin was sited at 7,500 feet, the point
where the pine forest gave way to the moist,
low, and dense mossy forest. Rather than carve
out a trail through the heavy undergrowth, we
stayed on the path to the summit of Mt. Pulog
as we searched for yew trees and other
Homalanthus species. Some of the species here
were recognizable as common farther north in
more temperate zones — spicehush (Clethra
luzonica), Berberis barandana, Deutzia
pulchra, Ilex crenata f. luzonica, and dense
shrubs of Rhododendron subsessile. The
most unusual collection of the day was a white-
flowered epiphytic rhododendron unique for its
thin needle-like leaves.

As we climbed higher the effects of the colder
climate became evident in the stunted shruh-
like forms of species that grew as trees lower
down. At 8,700 feet, the woody flora disap-
peared altogether, giving way to a tussock
grassland interspersed with a dwarf bamboo,
Arundinaria niitakayaniensis. At this point we
were still 900 feet from the summit and the
temptation to see the ocean from the top of the
island beckoned, hut our Philippine colleagues
counseled that the rains would come again by
noon. Since no Homalanthus or Taxus would be
found above treeline, we retreated to search
along other paths lower down. The rest of the

morning produced nothing, and
as predicted, the rain began
again at noon, reducing visibil-
ity to 75 feet. Melvin and I
spent the afternoon walking
along the edges of cabbage
fields and up and down the
crude paths that crisscrossed
the slopes. It was evident that
the forest had been severely
depleted since yew was last col-
lected on Mt. Pulog. Each hour
of trudging past three-foot-wide
pine stumps made us more
depressed, and we began to talk
of an epitaph for this popula-
tion of yew.

We returned to the cabin
dejected and prepared for defeat, but our gloom
changed to joy when our Philippine colleague
Ephrain greeted us with a fold of newspaper con-
taining a sprig of yew. Not a thousand yards
from the cabin our Ifugao guide had found five
adult yew trees (up to 80 feet tall) and a trio of
saplings on the edge of a cabbage patch. Once
the guide confirmed that this was the plant we
sought, he remembered seeing more of them far-
ther north on the other side of the mountain.

We realized, however, that the day of the yew
on Mt. Pulog is almost past: because it germi-
nates and grows under a solid canopy, yew
is found only in forests of long standing.
Homalanthus, by contrast, rapidly establishes
itself in recently disturbed areas — for instance,
in clearcuts or canopy gaps. With farmers clear-
ing the forests higher and higher up the moun-
tains on Luzon, Homalanthus is enjoying a
newfound prosperity.

To ensure broad genetic diversity, we wanted
to collect from widely separated populations.
After returning to Manila, we flew to Mindanao,
the largest of the Philippines' southern islands,
to seek out the yew and Homalanthus popula-
tions previously documented on Mt. Apo.
Mindanao is also home to some rare endemic
species of Homalanthus, and we hoped to locate
some of these.

Mt. Apo is the highest peak in the Philip-
pines; the only access is by foot. Our party of
four botanists and three porters began the ascent

found near the .

Light in a Bottle 25

at the village of Kitapowan and walked upward
through lush lowland forest. Towering above us,
one spectacular tree, damar, Agathis dammara,
had an eight-foot-wide trunk oozing a sticky,
milky resin, which is the source of a resin used
in varnishes.

Clumps of epiphytic orchids, some of them
three feet across, cluttered the path, brought
down by their own weight from the branches
above. Some of the trees were so tall that the
first branch was too high to identify, although
tentative identification could be made from
fragments of floral matter fallen on the
path. Other plants, like the bizarrely primitive-
looking screwpine (Pandanus) or the lush tree
ferns, grew in the lower canopy layer and were
unmistakable. We found one species of Homa-
lanthus on the uphill climb — H. populneus — a
solitary specimen of 45 feet that had sprouted
near a treefall. After a grueling five-hour climb,
we set up camp at 7,200 feet on the shores of
Lake Venado. Again we found ourselves in a
lush, mossy forest belt of southern conifers like
Daciydium and Falcatifolium, with the vivid
scarlet blossoms of epiphytic vireya rhododen-
drons punctuating the green curtain.

Rain was pouring down again as we rested
in camp and botanized around the lakeshore.
Here we found trees
more closely related to
the southern flora:

Tasmannia, Leptosper-
mum, and Pittosporum,
but no yew. From our
camp we could see the
north-facing slope of
Mt. Apo. A fire had
burned a considerable
portion of its forest in
1986, and silvery dead
trunks now rose out of a
sea of ten-foot sawgrass.

The lone path traversing
this thicket seemed our
only practical route for
the next day's climb.

We set off early, aware
now that the workday
would be cut short by
afternoon rains. The only

prior collection of Taxus on Mt. Apo had been
by the botanist Robbins in 1965, at an altitude
of 7,500 feet. We moved upward nervously, fear-
ing that Robbins had approached the peak from
another direction, or that the fire had eradicated
what may have been the only Taxus population
on Mt. Apo. But at 7,300 feet we spotted a yew
seedling, an eight-inch sprig in the moss on the
edge of the path, and within the next 400 feet of
altitude, we found an additional fifteen seed-
lings and saplings. These few plants had prob-
ably sprouted from seed deposited in the soil
prior to the fire, and it seems unlikely that they
will survive competition from the sawgrass or
withstand the intensity of the full sun should
they rise above it.

Although the specimens were too small to
yield a sample for laboratory testing, we col-
lected a few cuttings from each to root back at
the Smith College greenhouses and to provide
material for researchers working on other
aspects of the yew. We tried one additional
foray off-path in the direction of a promising
ravine. But when clouds pumped up and pos-
tured threateningly, we turned back. Suddenly
the rainy season began in earnest, with a down-
pour that dwarfed all previous storms. On the
way back down the mountain we found one

While curious townspeople looked on. the author and other expedition team
members prepared samples for the trip down the mountain.

26 Ainoldia 1998 Spring

more species of Homolanthus, H. rotundifolius,
bringing our total to four.

Conclusion

Upon our return to the U.S. our Taxus samples
were analysed for relevant medicinal com-
pounds, and data concerning individual tagged
trees was sent back to the Philippines. Should
plantation culture become an option, these tags
can identify elite trees. Cuttings were rooted at
Smith College, and from these plants, material
was supplied to other researchers working on the
yew's taxonomy, biochemistry, and genetics.

Samples from our Homalanthus collection
were sent to the Natural Products Branch of the
National Cancer Institute to begin the long pro-
cess of analysis and trial. Initial extracts showed
significant activity against HIV cell lines, but
further development has stalled for a variety of
reasons; other plant compounds have shown
more promise, as have certain non-plant-derived
compounds. Although prostratin, unlike other
compounds in its class (phorhol-esters), does not
induce tumors, taint by association has damp-
ened interest on the part of pharmaceutical
firms. Dr. Paul Cox, the botanist who brought
prostratin out of the rainforest, has suggested
some plants may produce "gray pharmaceuti-
cals," drugs of proven safety and efficacy that
are not marketable in the Western world. Possi-
bly prostratin may fall into this category, a low-
cost, plantation-grown treatment option that
offers an alternative to high-priced Western drug
regimes. The colonizing nature of Homalanthus
may make it an ideal subject for plantations in
forested areas.

In the daily grind of plant-collecting, it is easy to
fixate on the immediate goal, the plants them-
selves, and to forget that each collection, long
shot though it is, may be the basis of a cure for
thousands or millions of people. The renewed
interest in botanical compounds makes collect-

ing far afield possible, yet with each trip comes
the realization that we may be chasing and bot-
tling the last rays of light before an eclipse of
uncertain dimension and duration.

Bibliography

Balick, M. 1., and P. A. Cox. 1996. Plants. People and
Culture: The Science of Ethnobotany. NY:
Scientific American Library.

Brown, W. H. 1919. Vegetation of the Philippine
Mountains. Manila: Bureau of Science.

Cox, P. A. 1993. Saving the Ethnopharmacological
Heritage of Samoa. Journal of Ethno-
pharmacology 38{2-3]-. 181-188.

. 1994. The Ethnobotanical Approach to Drug

Discovery: Strengths and Limitations. In
Ethnobotany and the Search for New Drugs.
ed. D. L Chadwick and f. Marsh (CIBA
Foundation Symposium 185). NY: J. Wiley.

Cragg, G. M., et al. 1994. Ethnobotany and Drug
Discovery: The Experience of the U.S.
National Cancer Institute. In Ethnobotany and
the Search for New Drugs, ed. D. J. Chadwick
and J. Marsh (CIBA Foundation Symposium
185). NY: I. Wiley.

Gulakowski, R. L, et al. 1997. Antireplicative and
anticytopathic activities of prostratin, a non-
tumor-promoting phorbol ester, against human
immunodeficiency virus (HIV). Antiviral
Research 33(2): 87-97.

Gustafson, K. R., et al. 1992. A nonpromoting phorbol
from the Samoan medicinal plant
Homalanthus nutans inhibits cell killing
by HIV-1. Journal of Medicinal Chemistry
35: 1978-1986.

Merrill, E. D., and M. L. Merritt. 1910. The Flora of
Mount Pulog. Philippine Journal of Science C.
Botany V(4).

Merrill, E. D, 1914. Notes on Philippine Euphorbiaceae,
11. Philippine Journal of Science C. Botany
IX(5).

Rob Nicholson manages the conservatories of the Smith
College Botanic Garden, Northampton, Massachusetts.
His article on cutleaf maples appeared in the summer
1997 issue of Arnoldia.

The Ecology and Economics of Elm Replacement in
Harvard Yard

Peter Del Tredici

Two new Dutch elm disease-tolerant American elms have rekindled interest
in restoring the species to the landscapes it once dominated.

These new selections of Uhnus americana —
'New Harmony' and 'Valley Forge', recently
introduced by the United States National Arbo-
retum— represent an important horticultural
breakthrough,^ but some basic biological issues
should be considered before any new plantings
of the American elm are undertaken. The pur-
pose of this article is to articulate these ques-
tions in economic as well as ecological terms in
order to facilitate the decision-making process
that many landscape architects, designers, and
town managers now face concerning the use of
American elms in historic landscapes.

The American Elm in History

The first question that should be asked is why
this tree came to be so widely planted across
eastern and central North America in the first
place. The explanation can be found in the hor-
ticultural literature of the 1800s, particularly in
a beautiful book by Lorin Dame and Henry
Brooks that was published in 1890, Typical
Elms and Other Trees of Massachusetts^ Pro-
fusely illustrated and written long before Dutch
elm disease appeared in North America, it
serves as a portrait of the American elm at
the pinnacle of its landscape dominance. The
authors made many important contributions to
our knowledge of the American elm, but most
significant is their documentation of a key fact:
that the huge elms of the past reached their
great size by virtue of rapid growth rate, rather
than by great age. Other writers who described
the American elm in the nineteenth century
include F. A. Michaux (1819), A. J. Downing
(1841), D. 1. Browne (1846), F. ]. Scott (1870), G.

B. Emerson (1875), and C. S. Sargent (1890).'^
These authors make it clear that the American
elm was widely planted for a number of reasons,
only one of which was its great size and beauty.
Other, more pragmatic reasons gleaned from
the literature of the period can be summarized
as follows:

• The American elm, a native species, was
widely distributed throughout eastern and
central North America, typically growing
on moist bottomland or along disturbed
roadsides.

• It was so easy to transplant that it could lit-
erally be ripped out of swamps and planted
along roadways.

• It recovered well from the heavy pruning it
received following these careless transplant-
ing practices.

• It was highly adaptable, growing equally
well on wet or dry sites.

• It grew very rapidly, averaging about a half-
inch increase in caliper per year and reaching
two to three feet in diameter within seventy
years. Shade was in short supply in the nine-
teenth century, and the American elm pro-
vided it more quickly than any other tree.

• It shed its lower branches naturally, making
it well suited for locations along heavily traf-
ficked city streets and country roads.

Dutch Elm Disease

The Dutch elm disease fungus, along with its
dispersal agent, the European elm-bark beetle,
arrived in North America in 1930, hidden under
the bark of European elm burl-logs that had

2S Arnnhlia 19 >8 Spring

Elm Replacement 29

been shipped into the United States for making
veneer/ The disease spread so rapidly and killed
American elms so quickly that it seemed at first
that the species was headed for extinction. For-
tunately, this prediction has not come to pass.
The species still thrives as a wild tree in wet
woods and along streambanks throughout east-
ern North America.® As a landscape plant, how-
ever, the American elm is close to extinction.
The grand old specimens, four to five feet in
diameter, that once graced virtually every town
common in New England have been replaced by
trees that seldom reach more than eighteen
inches across before succumbing to DED or
some other disease.

Since the 1960s, there has been considerable
publicity about various efforts to "save" or to
"bring back" the American elm. Long-term
approaches have involved either selecting DED-
tolerant American elm cultivars or hybridizing
American elm with other elm species that
are DED-resistant. In contrast, short-term
approaches focus on preserving existing speci-
mens by spraying for the beetle, injecting
infected trees with fungicides, and removing
diseased limbs as quickly as possible. While
these treatments have saved individual trees for
up to twenty years, they are at best temporary
solutions; the inevitable infirmities of old age
are already catching up with older specimens.

Case Study: The Harvard Elms
The American elm has been the mainstay of the
Harvard Yard landscape for well over a hundred
years. The trees have faced many threats during
this time, but none has been as serious as the
introduction of Dutch elm disease.® In 1979,
when Harvard University began to implement
an integrated elm protection program, there
were 285 elms on campus. Most of them were
American elms, but a number of English and
European elms of uncertain identity were also
mixed in. Most of the trees were about seventy
or eighty years old when the protection program
began. By 1994, after fifteen years of treatment,
there remained only 165 elms, a mortality rate
of 42 percent. Detailed figures do not exist, but
the average cost of the total protection program

American elms have long been valued for their
exceptional growth rate. This ‘Princeton’ elm grew
more than two meters in a single year. Jack
Alexander, Arnold Arboretum propagator, points to
the start of the current year’s growth.

for the Harvard Yard elms over the period is
estimated at $25,000 a year, broken down
roughly as follows:

• $14,000 for two sprays each year (one dor-
mant oil and one foliage spray)

• $3,000 for fungicide injection each year

• $3,000 for fertilization every third year

• $5,000-$ 10,000 for pruning and removals

Over the fifteen-year period the total amount
spent on the elms was approximately $375,000,
or $100 per tree per year, in spite of which, mor-
tality was at 42 percent after fifteen years. By
extending these figures out, one can calculate
the cost of elm maintenance over twenty years
at roughly $500,000, with mortality approach-

Ulmus americana ‘Princeton’, a Dutch elm disease-tolerant American elm cultivar, has been growing on the
northeast slope of Bussey Hill since 1935.

PETER DEL TREDICI

30 Ainoldia 1 998 Spring

ing 50 percent. By comparison, the annual cost
for maintaining non-elm trees in the Yard is
approximately $20 a tree.

Replacement Costs

In 1994 an elm replacement program was initi-
ated.^ The cost of planting 200 new trees in the
Yard, most of them four to eight inches in cali-
per and ten to twenty feet tall, was $470,000, or
$2,350 per tree, including a one-year mainte-
nance contract and guarantee.

Essentially, the numbers show that the cost of
planting 200 new trees was roughly equal to the
cost of maintaining 285 elms for twenty years,
of which only half will still he alive and' the
other half in a state of decline at the end of
twenty years. To put it another way, twenty
years of maintaining one large elm with only a
50-percent chance of survival costs the same as
planting one new four-to-eight-inch caliper tree.

There is no absolute answer to the question of

how much one should invest in an elm protec-
tion program, and in any case, the question
should not be decided purely on an economic
basis. Elm protection programs cannot save a
tree forever, and in anticipation of the death
of the elms, such programs should always be
undertaken in conjunction with a program of
planting other species of trees. It is clear that the
high density of American elms that was seen in
many cities and towns during the first third of
the twentieth century should not be recreated.®
Indeed, it was this high density that allowed the
elm-hark beetle population to build up rapidly,
leading to the epidemic spread of DED. One sees
more elms surviving these days than in the past,
not because trees are more tolerant of Dutch
elm disease .than before, but because the
reduced elm population has resulted in lower
elm-bark beetle populations. This in turn allows
more elms to escape detection by their preda-
tors. Planting new trees of different species are

fust months after this photograph was taken, this hundred-year-old American elm. one of the last still standing
on Boston’s Commonwealth Avenue Mall, succumbed to a heavy wet snow that brought it crashing down onto
cars and into townhouse windows.

Elm Replacement 31

The remnants of an allee of American elms tower over replacement plantings of Zelkova serrata on the grounds
of Phillips-Exeter Academy in Andover, Massachusetts. The zelkova has many merits, but neither in scale nor
stature does it resemble the American elm.

an investment in the future that softens the
blow when a big elm dies, as it inevitably does.

Achieving Diversity

Unfortunately very few, if any, trees possess the
combination of graceful form and great size of
the American elm. The honey locust, Gleditsia
triacanthos, comes about as close as any tree,
but it grows much more slowly. Zelkova serrata
has roughly the same shape but is much
smaller. As Roller and Weaver point out, there
is no perfect replacement for the American
elm.® The key to successful substitution is to
choose species with the same landscape impact
or stature, regardless of whether they possess
the American elm's structure.

Within the genus Ulmus, there are several
potential candidates, but none are without some
drawback. The ubiquitous Siberian elm (U.
pumila), for example, is highly resistant to DED
but is very messy and graceless in form. The

lacebark elm (U. parviflora) is a handsome tree,
but much smaller than its American cousin.
Some of the more recent hybrid elms (involving
U. davidiana. japonica, and wilsoniana], may
eventually prove to be excellent replacements,
but they have not yet been thoroughly tested
under landscape conditions.

It must also be remembered that DED is only
one of several diseases that kill American
elms." In particular, phloem necrosis and elm
yellows can be lethal to many of the cultivars
that have been selected for their tolerance to
DED." And the elm-leaf beetle, along with a
host of other insects, had been killing elms long
before DED arrived on the scene. If the Ameri-
can elm is to make a comeback in the modern
American landscape — either as a hybrid or as a
disease-tolerant selection — it should be used on
an equal footing with other species, never as the
predominant species in the landscape.

Another approach to replacing American

32 Ainoldia 1998 Spring

elms involves working within a single genus or
family, which allows one to approach unifor-
mity and diversity simultaneously. In the Ter-
centenary Theater part of Harvard Yard, for
example, a grouping of legumes including
Cladrastis, Gleditsia, Gymnocladus, and
Styphnolobium (formerly Sophora], all share a
characteristic arching trunk and flat-topped
crown, but clearly differ in other aspects of their
habit. One can also group different oak species
to achieve a measure of uniformity amidst
diversity. In the oldest section of the Yard, a
grouping of oak species includes Quercus rubra,
palustris, phellos, coccinea, alba, bicolor, and
acutissima.

The advantages of increased species diversity
can be summarized as follows:

• It offers a measure of protection against an
epidemic spread of insects or of fungal and
bacterial diseases.

• It allows one to match different microcli-
mates on the site with the most appropriate
species.

• It provides greater variation in flower and
foliage displays, making a walk across cam-
pus a more interesting and potentially a
more educational experience.

Conclusion

The desire to restore the American elm to its
former status as the primary street tree in the
East is very strong. But if "restoring" a given
historic landscape means replanting the Ameri-
can elm — or any of its disease-tolerant selec-
tions or hybrids — at the density it occupied
historically, then it is a mistake. In the popular
literature on elms, the unspoken assumption
seems to be that if we could only conquer Dutch
elm disease, then we could easily recreate the
grand, elm-lined streets of the past. This idea is
biologically unsound. Because of the dynamic
nature of the interaction between host and
predator, "disease tolerance" is always a relative
phenomenon, not a fixed genetic trait; total
immunity is unattainable. Historical accuracy
and aesthetic tastes notwithstanding, it is in no
one's interest to bring the American elm hack to
its former position of landscape preeminence.

Endnotes

' United States National Arboretum Plant
Introduction Announcement, Ulmus americana
'Valley Forge' and 'New Harmony' (January, 1997).

^ An excerpt from Typical Elms and Other Trees
appeared in Arnoldia (1982) 42(2): 49-59.

^ F. A. Michaux, “Ulmus americana," in The North
American Sylva, vol. 3, tr. by A. L. Hillhouse (Paris
and Philadelphia, 1819); A. J. Downing, A Treatise on
the Theory and Practice of Landscape Gardening, 9th
ed. (NY: Orange Judd, 1873); D. J. Browne, Trees of
America (NY: Harper & Bros, 1846); F. J. Scott, The
Art of Beautifying Suburban Home Grounds of Small
Extent (NY: D. Appleton, 1870); G. B. Emerson, A
Report on the Trees and Shrubs Growing Naturally
in the Forests of Massachusetts. 2nd ed. (Boston:
Little Brown, 1875). C. S. Sargent, “Ulmus
americana" . in Silva of North America, vol. 7.
(Boston: Houghton Mifflin, 1890).

Berton Roueche, "Profiles: A great green cloud," The
New Yorker {]uly 15, 1961), 35-53; Donald C. Peattie,
A Natural History of Trees. 2nd ed. (Boston:
Houghton-Mifflin, 1964).

^ R. M. Burns and B. H. Honkala, eds., “Ulmus
americana, " Silvics of North America, vol. 2,
Hardwoods (USDA Forest Service Agriculture
Handbook 654, 1990).

® ]. Shaw, "Every tree doomed," Harvard Magazine
(1994) 96(4): 46-53.

' Ibid; M. Van Valkenburgh and P. Del Tredici,
"Restoring the Harvard Yard Landscape," Arnoldia
(1994) 54(1): 2-11.

® J. O. Dawson and M. A. Khawaja, "Change in street-
tree composition of two Urbana, Illinois,
neighborhoods after fifty years: 1932-1982," fournal
of Arboriculture (1985) 11(11): 344-348.

’ G. L. Roller and R. E. Weaver, Ir., "Replacing the
American Elm: Twelve Stately Trees," Arnoldia
(1982) 42(2): 88-100.

Michael Dirr, Manual of Woody Landscape Plants,
4th ed. (Champaign, IL: Stipes, 1990); F. S. Santamour,
Ir., and S. E. Bentz, "Updated checklist of elm
(Ulmus) cultivars for use in North America, fournal
of Arboriculture (1995) 21(3): 122-131; A. M.
Townsend et ah, "Variation in response of selected
American elm clones to Ophiostoma ulmi," Journal
of Environmental Hoticulture (1995) 13(3): 126-128;
G. Ware, "New elms for urban landscapes," Morton
Arboretum Quarterly (1995) 31(1): 1-9.

" Sinclair et ah. Diseases of Trees and Shrubs (Ithaca,
NY: Cornell University Press, 1987).

Santamour and Bentz, op cit.

Peter Del Tredici is Director of Living Collections at the
Arnold Arboretum.

Summer 1998,

^ The Magazine of the Arnol^Arboretum

GRAY HERBARIUM

amoldia

Volume 58 Number 2 1998

Amoldia (ISSN 004-2633; USPS 866-100) is
published quarterly by the Arnold Arboretum of
Harvard University. Second-class postage paid at
Boston, Massachusetts.

Subscriptions are S20.00 per calendar year domestic,
$25.00 foreign, payable in advance. Most single copies
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Editorial Committee
Phyllis Andersen
Robert E. Cook
Peter Del Tredici
Gary Roller
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Copyright © 1998. The President and Eellows of
Harvard College

Page

2 AN ECOLOGICAL HISTORY OF
MASSACHUSETTS FORESTS

John O’Keefe and David R. Foster

4 I. Dynamics in the Postglacial Era

8 n. Disturbance Prior to European
Settlement

11 HE Changes After European
Settlement

26 IV. Present Conditions and Future
Prospects

32 CASE STUDIES IN FOREST HISTORY
AND ECOLOGY

33 How Land Use Determines
Vegetation: Evidence from a New
England Sand Plain

Glenn Motzkin and David R. Foster

35 Forest Response to Natural
Disturbance Versus Human-
Induced Stresses

David R. Foster, John Aber, Jerry Melillo,
Richard D. Bowden, and Fakhri Bazzaz

41 Ecosystem Response to an
Introduced Pathogen:

The Hemlock Woolly Adelgid

David A. Orwig and David R. Foster

Front cover: Harvard Pond, Petersham, Massachu-
setts. Photograph by David R. Foster.

Inside front cover: Al Cline, director of the Harvard
Forest, 1939-1946, leans against a windthrown white
pine that was uprooted in the 1938 hurricane. From
the Harvard Forest Archives.

Inside back cover: Dead white pine snag on the
Pisgah tract of the Harvard Forest in southern New
Hampshire. This large snag is a remnant of the old-
growth forest that blew down in the 1938 hurricane
and is surrounded by the vigorous growth of a new,
regenerating forest. Photograph by David R. Foster.

Back cover: Aerial photograph of the Prospect Hill
tract of the Harvard Forest taken by David R. Foster
in 1995. The fire tower rises above conifer and hard-
wood forests.

2 Arnoldio 1998 Summer

Old hemlock forest in the Slab City tract of the Harvard Forest.

An Ecological History of Massachusetts Forests

John O’Keefe and David R. Foster

The forests of Massachusetts present a history
of almost continual change, albeit change that
has varied greatly in scale, rates, and causes
through time. The importance of human dis-
turbance relative to natural disturbance has
steadily increased — gradually at first, as abo-
riginal activity expanded to include agriculture,
and then rapidly since European settlement.
Just as the variety, frequency, and extent of
human disturbance have increased through
time, they can be expected to continue increas-
ing and changing into the future.

European settlement in the seventeenth and
eighteenth centuries brought a dramatic trans-
formation as much of the land was deforested
and farmed and the remainder was logged,
grazed, or burned. Since the mid-nineteenth
century, agriculture and forest use have
declined, forest area and age have increased,
and the land has become more ''natural" than
at any time postsettlement. But despite the
natural appearance of much of the modern
landscape, a legacy of intensive past use
remains in vegetation structure and compo-
sition, in landscape patterns, and ongoing
dynamies. Consequently, an understanding of
the history of human influence must be an
integral part of ecological study and a critical
component of conservation planning and
resouree management.

For millennia our forests had been evolving
into the landscape that greeted the settlers. One
reasonable start for our forests' history is the
end of the last glacial period, more than 13,000
years ago.

ICE AGE

13,000

Tundra

I 1,500

Spruce woodland and forest

10,000

Human arrival

9,500

Pine forest

8,000

Mixed deciduous forest

5,000

Hemlock decline

3,000

Arrival of chestnut trees

1,000

Native American agriculture

250-350

European settlement

150

Peak of agricultural clearing

85

Chestnut blight

60

1 938 hurricane

PRESENT

Approximate chronology of important events
in the development of Massachusetts’ forests,
given in years before present.

1. Dynamics in the Postglacial Era

Over 15,000 years ago, at the peak of the
last glaciation, most of present-day
Massachusetts was covered by ice up
to a mile thick. Cape Cod and the offshore
islands — Nantucket, Martha's Vineyard, and the
Elizabeth Islands — consist largely of what geolo-
gists call moraines, piles of debris accumulated
at the front of the advancing ice sheet and left
behind when the glaciers finally melted. The
advancing glaciers not only smoothed and
shaped the landscape by scraping and plucking
the bedrock as they advanced, they also left
behind a layer of ground-up rock, or till, which
has since evolved into soil. As the glaciers
melted, the tremendous volume of water pro-
duced seasonal streams that carried and sorted
much of this material and deposited sands and
gravels wherever they slowed.' The soils of
Massachusetts are a product of this massive
natural engineering, later supplemented by
organic material from the vegetation that even-
tually covered the landscape. Along major rivers
fine silt was deposited when the rivers over-
flowed their banks in spring; in some depres-
sions a surplus of moisture allowed thick layers
of peat or muck to develop. The resulting pat-
tern of soil types has strongly influenced the
type and distribution of trees and forests.

Of course, during the glacial period, there
were no forests in Massachusetts: The modern
tree species of New England were restricted to
favorable locations called refugia south of the
glacial zone, presumably scattered across the
southern Appalachians and the eastern coastal
plain. The huge quantities of water trapped on
land as glacial ice had once been seawater,- con-
sequently, sea level was several hundred feet
lower than at present, leaving vast areas of con-
tinental shelf off the present-day East Coast
exposed as refugia as well. The forests in these
various refugia contained species mixtures

unlike any we are familiar with today. As the
climate warmed and the glaciers melted, the
trees began their migration north at rates deter-
mined by the method of seed dispersal and the
climatic tolerance of each species.

We are fortunate to have a record of this post-
glacial migration in the form of pollen preserved
in sediments at the bottom of lakes and ponds
and below many wetlands. Each year pollen
from plants is carried away on the wind. When
it lands on the water of a pond or lake, it sinks
to the bottom, and along with other wind-borne
material is incorporated into the sediments.
These sediments form a chronological sequence
of layers, the oldest at the bottom, which scien-
tists can recover as a thin cylinder or core of
mud. Each layer in the core can be dated using
radioisotopes, and since pollen is extremely
resistant to decay — the pollen of different plants
can be identified at least to genus and in some
cases to species — the presence and relative
abundance of different types of pollen at each
layer enables paleoecologists to reconstruct the
major vegetation changes at a site.

The pollen diagram for the Harvard Forest
Black Gum Swamp (page 6) shows changes in
vegetation over time that are typical of sites
across Massachusetts. As the glaciers melted
and the climate warmed, a period of tundra is
followed by boreal (northern conifer) forest,
succeeded in turn by pine forest with rapidly
increasing amounts of several deciduous species
(birch, oak, beech) by 8,000 years ago. Although
mixed deciduous forests have been dominant for
the last 8,000 years, the mixture of species has
changed continually during that period, and
these changes can tell us much. Species have
behaved quite independently, presumably
migrating to Massachusetts from different loca-
tions at different rates, each species responding
in its unique fashion to the various combina-

The Postglacial Era

D

Alders, bearberry, and other pioneer vegetation follow a receding glacier in Alaska.

tions of climatic, soil, biotic, and historical fac-
tors found in the area. The major influences on
these changes are long-term climate change,
1} migration rates of individual species, and natu-

[i ral disturbance processes.

Population dynamics of selected species
I help us understand these processes. Hemlock

increased rapidly in importance after its arrival
‘ about 9,000 years ago. A little less than 5,000

years ago, it decreased dramatically in a very
; short time, then slowly recovered. This sudden

* decline, seen in pollen records throughout the

Northeast, has been attributed to a severe out-
I break of insects or disease that greatly reduced

hemlock populations for nearly 1,000 years.

Regional pollen analyses suggest that tem-
, peratures during the period from 8,000 to 5,000

, years ago were milder than those of the last

4,000 years. ^ It was during this warm period that
I many tree species now common in Massachu-

setts migrated into the area, with some species

extending well north of their current ranges. At
a number of sites, fire frequency, indicated by
charcoal in sediment cores, was also greater
than in the more recent period.-^ The increase
in spruce pollen over the last 2,000 to 3,000
years was probably caused by more recent cool-
ing. All these changes in forest communities
were complex; while spruce, a northern species,
increased — evidently in response to gradual
cooling — chestnut, a southern species, was also
migrating north across Massachusetts. In fact,
chestnut is the most recent arrival in the pollen
record, appearing less than 3,000 years ago,
much later than the other important deciduous
species that occur in the region today.

The Natural Environment

The distribution of forest types across Massa-
chusetts during the period preceding European
settlement was controlled by the physical geog-
raphy, or physiography, of the landscape; the

I'i; I l.U 1)11. I Ul.nK'l

6 Arnoldia 1998 Summer

Pollen diagram from the Black Gum Swamp at the Harvard Forest in central Massachusetts depicting the
major changes in the vegetation over the past 12,000 years. Tundra communities were replaced by boreal forest
dominated by spruce until approximately 9,200 years before present (B.P.) when pine and other tree species
became important.

Changes in the relative abundance of species resulted from climate change, species migrations, disease
(hemlock decline at about 5,000 B.P.), and fire until 250 to 300 years B.P., when European settlement resulted
in major deforestation and the increase in agricultural weeds, herbs, and early successional species.*

underlying geology; and the frequency of distur-
bances such as windstorm and fire. Massa-
chusetts, excluding Cape Cod, is roughly
rectangular, 125 miles (200 km) east to west
and 50 miles (80 km) north to south. Today it
receives approximately 40 inches (100 cm) of
precipitation annually, distributed fairly evenly
throughout the year, with a mean annual tem-
perature that ranges from a mean of several
degrees Fahrenheit below freezing in January to
about 70 degrees in July, and averages nearly 50
degrees over the year.

Within a relatively small, compact area, Mas-
sachusetts contains six broad physiographic
regions: the coastal lowlands, the central
uplands, the Connecticut River valley, the Berk-
shire Mountains, the Berkshire valley, and the

Taconic Mountains. The geologic substrate var-
ies across the state. Except for parts of the Con-
necticut River valley, the Taconic Mountains,
and the Berkshire valley, the soil is acidic, poor
in nutrients, and shallow with patches of
exposed bedrock. Elevation generally increases
from east to west, reaching a maximum at Mt.
Greylock in the Berkshires (3,487 ft [1060 m]).
These are the physiographic and geological con-
ditions that interact with climate to produce
vegetation zones sometimes referred to as
ecoregions.

Within Massachusetts natural vegetation
zones are largely determined by climate, which
in general varies with elevation except in areas
near the moderating influence of the ocean.
Southeastern Massachusetts, all of Cape Cod,

The Postglacial Era 7

Map of southern New England depicting the major
physiographic regions. From east to west they are
coastal lowlands, central uplands, Connecticut
River valley, Berkshire Mountains, Berkshire valley,
and Taconic Mountains.^-^

and the offshore islands fall
within the pitch pine-oak zone.

This vegetation type, occurring
on sandy and gravelly soils laid
down as glacial moraines or out-
wash deposits, is characterized
by drought-tolerant and fire-
adapted species, including pitch
pine, scrub oak, and huckle-
berry. This type also occurs
on scattered outwash deposits
in inland Massachusetts. The
remainder of the coastal low-
lands, the southern central
uplands, and the southern
Connecticut River valley fall
within the central hardwood-
hemlock-white pine zone. This
vegetation type represents the
northern extension of the oak-

hickory dominated forests of the central Appa-
lachians and the Middle Atlantic States.

North and west of the central hardwood zone
we generally find the transition hardwood zone,
which also extends up the major river valleys in
the western part of the state. This zone is char-
acterized by increasing amounts of species
found farther north, such as yellow birch, black
(or sweet) birch, sugar maple, and beech, with
less oak (especially white oak) and with paper
birch occurring on heavily disturbed sites. The
higher elevations in the Berkshire and Taconic
mountains and the extreme northern border of
central Massachusetts fall within the northern
hardwood and spruce-fir zones. The spruce-fir
zone, in which red spruce is the dominant coni-
fer, is restricted to the highest elevations, gener-
ally above 2,000 feet, while the northern
hardwood zone occurs just below the spruce-fir
zone and has hemlock and white pine as its
dominant conifers. Both zones have mixtures of
hardwoods dominated by sugar maple, yellow
birch, beech, and red maple.

Spruce-fir/
northern hardwood

Northern hardwood/
hemlock/white pine

Transition hardwood/
hemlock/white pine

Central hardwood/
hemlock/white pine

Central hardwood/
hemlock

Pitch pine/oak

The major forest vegetation zones in southern New England. Sandy
glacial deposits, found on Cape Cod and in extreme southeastern
Massachusetts, support a dry forest of pitch pine and scrub oak.
Transition hardwood forest dominates the central uplands and much of
the Connecticut River and Berkshire valleys, with northern hardwood
forest on the higher elevations in the Berkshires and Taconics and spruce-
fir forest on the highest elevations.^

11. Disturbance Prior to European Settlement

atural Disturbance

The major natural disturbances affect-
ing Massachusetts forests include
windstorms, pathogens (insect and diseases),
and fire. Evidence of past storms may persist
for as long as a thousand years in the pits
and mounds often found in our forests,*'^
but these microtopographic features offer no
information about the frequency, intensity, and
distribution of storms, which vary with chang-
ing climatic conditions. However, if, as seems
likely, storm patterns have remained stable in
Massachusetts over the postglacial period, we
can infer trends from observations over the past
few hundred years.

Two different types of windstorms cause
significant damage to Massachusetts forests:
tropical storms, or hurricanes; and downbursts,
or microbursts — sudden, straight-line winds,
often from the northwest, associated with
severe thunderstorms and occasionally accom-
panied by tornadoes. Downburst winds are
probably the dominant wind disturbance in the
Berkshires and western Massachusetts. They
continue east across central Massachusetts
and become less important in areas near the
coast under the stabilizing influence of mari-
time winds. While commonly confined to
small areas, the potential destructiveness of
these winds was demonstrated in July 1995,
when hundreds of thousands of acres of forest
from the Adirondack region of New York
into western Massachusetts were severely dam-
aged by an extremely large and long-lived
downburst front.

Tropical storms represent the most important
wind disturbance in central and eastern Massa-
chusetts. Historical evidence indicates that hur-
ricanes may affect central and eastern areas
approximately every hundred years, with the
Cape and islands affected more often. The stron-
gest winds in these large, counterclockwise-

rotating storms are on the easterly side, as
illustrated by the catastrophic storms of 1815
and 1938. These storms produced the greatest
damage on south- and east-facing hillsides
while steep north and west slopes were pro-
tected from the strongest winds." Western
exposures and ridges, on the other hand, are
prone to selective damage from the more patch-
ily distributed microburst winds associated
with severe thunderstorms.

The only strong evidence of pathogenic dis-
turbance in the paleoecological record is the
widespread hemlock decline nearly five thou-
sand years ago. The rapidity and extent of this
decline, not associated with declines in other
species or identifiable climatic change, points to
a species-specific pathogen as the cause. Hem-
lock remained at low population levels for
nearly a thousand years. The extent of its even-
tual recovery differed across the region. At
many sites, hemlock gradually approached its
predisturbance abundance between a thousand
and fifteen hundred years later." At other sites,
it never fully regained its former importance,
presumably because of competition with
recently immigrating species or slight climatic
changes over the intervening period. Of course,
this event offers many comparisons with the
human-transported pathogens (gypsy moth,
chestnut blight, hemlock woolly adelgid) that
our forests are coping with today.

Fire, like windstorms, probably differed sig-
nificantly in its impact across Massachusetts as
a result of differences in climate, fuel type and
abundance (which varies by vegetation type),
and ignition sources (lightning and aboriginal
human populations). When we look at the
impacts of fire, we encounter the earliest strong
evidence for human influence on Massachusetts
forests. Charcoal in sediment cores indicates
that fires were less frequent and therefore less
important in the Berkshires than in southeast-

Prior to European Settlement 9

Paths of the major hurricanes that have impacted New England from 1600 to the present are shown on the
left. On the right is the damage inflicted on forests in the region by the 1938 hurricane. Approximately three
billion board feet of timber were windthrown by the storm, more than six hundred lives were lost, and damage
costs exceeded a hundred million dollars.^'*'^^

ern Massachusetts and on Cape Codd^ The
droughty, sandy soils of the latter areas sup-
ported a highly fire-prone vegetation largely
dominated by pitch pine, scrub oak, other
oaks, and huckleberry. Pitch pine, like all coni-
fers, contains resins in its needles that make it
much more flammable than broadleaf, decidu-
ous trees. Huckleberry, although a broadleaf,
deciduous species, also contains resins in its
leaves and therefore creates a very flammable
understory. At the same time, this type of veg-
etation is highly fire-adapted. All the oaks, espe-
cially scrub oak, are prolific sprouters follotving
injury. Pitch pine is unique among Massachu-
setts' native conifers in possessing dormant
buds beneath the bark and near the base of its
trunk that enable the tree to sprout and survive
if the main stem is severely damaged by fire.
Moreover, the cones of pitch pine tend to be
serotinous, which means they may remain

closed with seed inside until the heat from a
fire triggers an opening mechanism to release
the seeds onto the recently burned landscape.
Although pitch pines in Massachusetts rarely
exhibit this behavior today, it is commonly
observed in the frequently burned New jersey
pinelands.

The northern hardwood species — ^sugar
maple, beech, and yellow birch — while capable
of sprouting, tend to have thinner bark that
provides less protection from understory
fires. Hemlock, a major associate in the north-
ern hardwood forest, is also thin-barked as
well as slow-growing, long-lived, and incapable
of sprouting. Therefore, where these species
were dominant, we can conclude that fires were
not as frequent or severe as in southeastern
Massachusetts. Moreover, during the growing
season broadleaf foliage normally holds enough
water to be nonflammable. This moisture tends

10 Ainoldia 1998 Summer

to limit the fire season in our broadleaf forests
to spring and fall, when the fallen dry leaves
often burn in surface or brush fires. In fact,
the combination of these factors led some to
nickname the northern hardwood forest "the
asbestos forest."

Aboriginal Human Impact

American Indian populations migrated into
Massachusetts shortly after the trees, some
ten thousand years ago, but their populations
remained quite low until four or five thousand
years ago. Some researchers have speculated
that the hemlock decline of about that time
and the subsequent increase in species such as
oak and hickory, which produce abundant large

Areas of concentrated aboriginal human populations
in southern New England during the Late Woodland
period (1000 B.P.-400 B.P.) immediately preceding
European settlement. Populations were concentrated
along major river valleys, the coast and the larger
islands of Nantucket and Martha's Vineyard, and the
lower reaches of broad upland areas.

nuts edible by both wildlife and humans, may
have contributed to the increase in aboriginal
populations.

Archaeological evidence indicates that
human populations — like fires — were more
numerous in the eastern than in the western
part of the state during the period four hundred
to a thousand years ago, with settlements also
found along the major river basins.’^ There is
little evidence that these populations cleared
extensive areas for agriculture. It is more likely
that they created a patchwork of recently
cleared areas, abandoned fields, and village sites
in a matrix of intact forest. Population density
(and presumably human impact on the forest)
gradually decreased moving away from the
coast, from a high of up to fifty people per
square mile on Nantucket and Martha's Vine-
yard, to four to ten per square mile in inland
eastern and southeastern Massachusetts and
the Connecticut River valley. It is unlikely
that permanent settlements were made in the
upland Berkshires.

Interestingly, the areas of high population
density shown here match the distribution of
the central hardwood and pitch pine-scrub oak
forest zones seen on page 7. These are the forest
types most suitable for burning. Although
extensive debate continues regarding the
frequency, extent, and broad-scale impacts of
aboriginal burning, there is general agreement
that these populations did burn forests to create
fields and to rejuvenate understory browse for
deer and other animals they hunted. No
doubt this burning was largely restricted to dry
areas where most vegetation was already
adapted to fire, but it also intensified these con-
ditions by reducing any fire-sensitive species
that may have been present.

III. Changes After European Settlement

The Colonial Period

European settlement spread through
Massachusetts at uneven rates. The
coastal counties Essex, Suffolk, Norfolk, Ply-
mouth, and Bristol were largely settled by 1675,
as was the Connecticut River valley, with set-
tlers moving northward from settlements in
the Springfield area that dated from the early
1600s. Concentrated in the coastal lowlands
and major river valleys, these early settlement
zones corresponded closely with the areas
where aboriginal practices had most affected the
forests. Settlement spread westward from the
coast into Middlesex and Worcester counties in
the late seventeenth and early eighteenth centu-
ries and into the foothills of the Connecticut
River valley during the same
period. In 1725 Massachusetts
began using land grants to pay
off debts, especially for mili-
tary service,'® which encour-
aged settlement in the central
upland areas. The last areas to
be settled, from the second half
of the eighteenth century into
the beginning of the nineteenth
century, were the Berkshire
Mountains and the northern-
most portions of the central
uplands.

Forest clearing was initially
quite slow, for reasons that
included the lack of markets
for excess production and a
town organization based on the
European model of a central-
ized settlement and common
field system.'®''® More than a hundred years
after its settlement in 1635, Concord was still
more than fifty percent forested. This rate of
deforestation, about 0.4 percent a year, was
typical of towns in the seventeenth century.'®

A shift to a town pattern of dispersed settlement
and individual ownership of private land, with
all land in the township distributed, led to
much more rapid deforestation toward the
middle of the eighteenth century. Rates of 0.8
percent to 1.0 percent per year were common
in the second half of the eighteenth century
both in older towns like Concord and new ones
like Petersham.

This acceleration in deforestation coincided
with a shift toward a market economy, partly
driven by a developing beef trade with the West
Indies. Cattle were suitable for remote hilltown
farms during this period because they could he
walked to market on the rudimentary roads that
precluded the long-distance transport of most

products. The difficulty of transport also partly
explains the methods most commonly used to
clear the forest: girdling the trees and leaving
them in place to fall apart slowly, or cutting and
burning them.'® Except where water transport

1720

1760

1800

1840 1880

Year

1920

I960

Changes in the percentage of the land area covered by forest during the
historical period in the state of Massachusetts, town of Petersham, and
the Prospect Hill tract of the Harvard ForestP°

HARVARD FOREST ARCHIVES

12 Arnoldia 1998 Summer

1

After European Settlement 13

Soil Suitability

■ Wet

^ M Rough and Stoney
I I Moist Loams

These maps of the township of Petersham. Massachusetts, depict soil suitability and stone walls, on the left,
and on the right, forest cover in 1830 and 1985. Stone walls and agricultural land are concentrated in areas of
more productive soil.'^'

was available, trees had value as lumber or fire-
wood only locally. Potash, a relatively more
compact and transportable product, was prob-
ably the major marketable product from the
trees of these early farms.

Pasture was the most suitable use for most
of Massachusetts, since the rockiness of most
areas made preparing land for tillage a long and
backbreaking chore. It has been said that it took
two generations to clear upland farms for plow-
ing, the first to remove the trees and the second
to remove the stones. The massive stone walls
surrounding abandoned fields across the state

attest to the effort required by the second task.
And yet the great number of rocks scattered
throughout the remaining pastures and second-
growth woods suggest that the majority of the
landscape was never tilled, but rather grazed or
at most mowed. The principal exceptions, of
course, were the major river valleys, where
postglacial alluvial deposits provided excellent
tillage once the trees were removed. These areas
are notable for their lack of stone walls.

In upland areas, hilltops were often selected
for village centers and initial clearing for agri-
culture because they had good drainage and

Above left, presettlement forest; below, initial clearing and subsistence farming, 1740. Most human
disturbances prior to European settlement were infrequent and scattered. Nonetheless, the forest communities
that the settlers encountered had evolved under dynamic conditions and had been in place only a relatively
short time.

These images are from the dioramas (three-dimensional models) in the Land-Use History series on
permanent display at the Fisher Museum of Forestry, Harvard Forest. Petersham, Massachusetts. They depict
typical changes in the upland Massachusetts landscape, from initial European settlement through the 1930s
(when they were constructed). Other dioramas in this remarkable series depict forest management and
conservation practices.

HARVARD FOREST ARCHIVES

14 Arnoldia 1998 Summer

relatively few stones. Except for the broad river
valleys, inland lowlands often offered poor
drainage and a shorter growing season. Colo-
nists commonly evaluated land quality on the
basis of topography and on their knowledge of
the site preferences of different tree species and
forest types.

Initially, a farmer might clear six to eight
acres over the course of several years. When
tilled, this initial clearing could support a typi-
cal family of five to seven. During this period
the dominant economic base of rural Massachu-
setts was low-intensity agriculture combined
with artisanship. Few individuals provided
for all their needs through their own labor,
but through cooperation and exchange, town-
ships could be largely self-sufficient. Towns
supported a range of artisans, shops, mills, and
tanneries. Roads provided internal circulation
but relatively poor access to external markets.
At the same time, coastal communities were
developing extensive fishing, manufacturing.

and shipping industries, exploiting local forests
for shipbuilding materials and export products.
By the mid- 1700s Salem was the most prosper-
ous port in the country and a center of world-
wide trade.

Agricultural Period

The period from the late 1700s through the first
half of the nineteenth century saw a major
transformation of the economy, social struc-
ture, and landscape of Massachusetts. The
rural economy underwent a shift from home
production and local consumption to market-
oriented intensive agriculture, enabled by the
improvement in transportation brought about
by newly constructed roads, canals, and rail-
roads. Farmers responded to the expanding mar-
kets by clearing more forested land and draining
wetlands, often on marginally productive sites.
Pasture remained the primary land use, with
beef and wool the dominant farm products
until canal and rail connections with the West

Height of intensive farming, 1830. The percentage of land cleared for tillage, pasturage, orchards, and building sites in
central Massachusetts was about 70 percent. A century later, the percentages of cleared and forested areas had been
reversed. The stone walls testify to the tremendous labor required to farm land that is better adapted to growing trees than
hay and grain.

After European Settlement 15

and relaxation of wool tariffs in the 1830s and
1840s reduced their profitability.^^ Most farm
families also engaged in home production of
some sort (shoes, hats, clothes), and many
earned some income from mills or tanneries.
Local industry thrived, and most hill towns
reached their peak levels of agricultural and
commercial activity, as well as population, dur-
ing this period.'^ However, this period also rep-
resented the start of the region's shift to
industry, a factor that together with the expand-
ing national transportation network and west-
ward settlement initiated the decline of New
England agriculture.

Many settlements literally moved downhill,
changing from ridgetop agricultural villages to
riverside industrial towns for a variety of rea-
sons. Hill towns without significant water-
power resources were unahle to participate in
the transition from an agricultural to a manu-
facturing economy. The new factories for pro-
ducing textiles, wooden products, and tools
needed water to power their machines.^® The
developing railroad network, which followed
the same watercourses that the factories used
for power, transported raw materials and fin-
ished products to and from the factories. The
new roads and railroads allowed many non-
perishable farm products to he shipped from
the Midwest more cheaply than they could be
produced in Massachusetts.

Many factors contributed to the decline of
Massachusetts agriculture, but depletion of the
fertility of the land was not a major one. In fact,
there is evidence that the quality of tilled land
in hill towns improved through the eighteenth
and nineteenth centuries. The disadvantages
of Massachusetts farmland included stony soil
and small fields divided by numerous stone
walls, which were incompatible with mechani-
zation. Industrial production and improved
transportation removed opportunities for
supplemental income by reducing the need for
local artisanship. Social factors also contributed
to the decline of Massachusetts' hill-town agri-
culture; attractiveness of urban amenities and
income, the decline of interest in agricultural
life, and the shrinking economic opportunities
in small towns.

The pattern of decline was strongly influ-
enced by regional geography. Towns adjacent to

developing industrial centers like Worcester and
Fitchburg had a ready market for fuelwood, pro-
duce, and milk, while those more distant pro-
duced butter, cheese, and hay. The farthest
distant towns declined most rapidly.^^'^' In 1810
Massachusetts was an agricultural state with a
population of 412,000 that was remarkably
evenly distributed in rural areas (79 percent),
with the exception of Boston, Salem, and a few
other coastal communities. Industrialization
brought a tremendous increase and concentra-
tion of population in urban and, more recently,
suburban centers. In 1975, 85 percent of the
population of 5.8 million was located in urban
areas. In contrast, many rural communities
have greatly declined in population over the
past hundred years.

How did our forests fare during the agricul-
tural period? By the late eighteenth century the
gradual clearing of the first half of that century
had become a rapid deforestation that continued
until the mid-nineteenth century. Forest clear-
ing was concentrated in the uplands, with the
wetter swamps and steep, rocky slopes generally
left as woodlots. The Berkshires were the last
areas to be cleared and were never developed for
agriculture to the extent that the remainder of
the state was. The statewide peak in the level of
deforestation was reached about 1860, by which
time nearly 70 percent of the land was cleared.
Many areas east of the Berkshires experienced
the same pattern as Petersham and the Prospect
Hill tract of Harvard Forest, with maximum
clearance in the 1840s to 1860s, when less than
20 percent of the forest remained. The location
and amount of forest left uncleared varied by
geography. For example, in the north-central
portion of Massachusetts from the Connecticut
River valley to eastern Worcester County, the
hills east of the valley, with many rocky ridges,
remained largely forested, as did the north-
south-trending, poorly drained valleys farther
east. Most of the rest of the region was cleared.

Of course, even areas not cleared for agricul-
ture were harvested intensively by the nine-
teenth century. The growing rural populations,
whose numbers peaked in the mid-1800s,
required large amounts of cordwood for fuel.
Petersham, for example, had a population of
nearly 1,800 people in 1840. Assuming an aver-
age household size of six, this population would

16 Arnoldia 1998 Summer

1980

ASHBURNHAM

H Forest
□ Meadow
I I Open

1830

1830

1980

After European Settlement 17

have represented three hundred households to
heat. If each household used 15 cords a year (a
conservative figure when fireplaces are used),
they would have required a total of 4,500 cords
of fuelwood a year. The 20 percent of Petersham
that remained forested in 1840 represented
about 6,000 acres. Because Massachusetts for-
ests can be expected to grow between one-half
and one cord of hardwood per acre per year, vir-
tually all the woodland growth in Petersham
could have been used for fuelwood.

These hardwoods were probably managed by
means of a coppice system, in which trees
would be harvested very young (every twenty to
forty years), left to resprout, and then harvested
again as soon as the new growth was big enough
to burn. Across upland Massachusetts most
farms could maintain woodlots to satisfy their
own fuel needs, but near cities and along the
coast where settlements had been in place
longer, the fuelwood was soon exhausted and
had to be brought great distances by ship at con-
siderable expense.

Although fuel was by far the dominant use
of wood in the early 1800s, the remaining for-
ests also faced other demands. Hemlock and
chestnut trees, especially, were cut to provide
tanbark for tanneries. Lumber was needed for
constructing houses, barns, and public build-
ings. Wood was used to make charcoal, and
fences had to be built. The scarcity of wood
by the early 1800s probably accounts for many
of the stone walls that still exist along bound-
aries and in pastures, where the stones would
not have had to be removed for plowing or mow-
ing; by then stones were more readily available
than wood.

Postagricultural and Modern Periods

The decline of agriculture in the second half of
the nineteenth century was accompanied by a
corresponding regrowth of forest. Our present
forests can be divided into secondary forest on
land formerly cleared and used for agriculture

(plowed or grazed), and primary forest on land
never actually cleared but harvested throughout
the agricultural period. As we have seen, the
major portion of upland farmland was used for
pasture, and even tilled land may have reverted
to mowing or pasture before final abandonment.
The resulting sod surface was not hospitable to
many "pioneer" tree species such as birch and
aspen, whose small, windblown seeds would
often dry out and die after germinating; the
sprouting seeds were trapped in the grass unable
to reach mineral soil. The sod did, however, pro-
vide a suitable seedbed for the windblown but
larger seeds of white pine, which colonized vast
areas of abandoned farmland. Pines were much
less likely to have been cut for fuelwood, and
those left as shade trees in a pasture or along a
fencerow could colonize many acres with dense
stands of young pine. Moreover, animals still
grazing these pastures would avoid pine seed-
lings while devouring most broadleaf species.

These new forests grew quickly, and by
the late 1800s supported renewed harvesting
for lumber and especially for shipping contain-
ers. The new portable steam sawmill, in com-
mon use by the turn of the century, permitted
logging throughout the backwoods areas. Tre-
mendous volumes of "old-field" (or abandoned-
field) white pine were harvested, peaking
in 1910-1911. During this timber boom, exten-
sive harvesting of all species across the state
resulted in large tracts of even-aged, young, low-
value stands. Many of these cut-over stands,
considered nearly worthless at the time, were
acquired by the state for overdue taxes and have
formed the basis of our state forest system. It
was at this time, the early twentieth century,
that the excesses of the timber industry
throughout the East gave rise to the conserva-
tion movement, which was strongly represented
m Massachusetts.

When the old-field pines were harvested, they
were unable to sprout from the remaining
stumps and roots (except for pitch pine, our only

Maps of three townships characteristic of different physiographic regions in central Massachusetts depicting
distinctive amounts and patterns of forest, open land, and meadow in 1830 and 1980.

Ashburnham, on rocky hills near the New Hampshire border, was least extensively cleared and today is
the most forested. Barre, on rolling terrain in the central uplands, was extensively cleared for agriculture but
has largely reverted to forest. Deerfield, in the Connecticut River valley, was extensively cleared except for a
few north-south bedrock ridges, and the fertile valley bottom remains in agriculture today.

HARVARD FOREST ARCHIVES

18 Arnoldia 1998 Summer

Agricultural abandonment and establishment of old-field white pine, 1850. Almost immediately the forest
started to reclaim the idle fields and pastures. They were quickly seeded to white pine, with hardwoods such
as red maple, white ash, red oak, chestnut, gray and paper birch forming a minor element.

First crop of old-field white pine harvested. 1910. From 1890 to 1920 portable sawmills appeared everywhere,
and many new wood-using factories were established. With yields of 25 to 50 thousand board feet per acre
and standing lumber valued at $10 per thousand, one might well envy a farmer who owned a 100-acre woodlot,
worth perhaps $30,000 — a wholly volunteer crop on which only taxes had been expended.

After European Settlement 19

Old-field white pine is followed by hardwood. 1915. Five years after logging, hardwoods are primarily growing
in the open lanes between the windrows of slash. These trees originated as stump sprouts — red maple, red and
white oak, white ash. hard maple, chestnut, black cherry, black birch — or as seedlings of light-demanding
species — gray and paper birch, pin cherry, and poplar.

Hardwood stand reaches cordwood size, 1 930. Twenty years after logging on moderately moist soil, red maple,
gray and black birch, and most other species begin to slow their growth upwards. Red oaks maintain a steady
growth in height and girth so that by sixty years they will have formed an overstory above the other trees.

20 Arnoldia 1998 Summer

native conifer with this ability) and so had to
reestablish themselves on the site from seed.
As the old-field pines were growing, however,
various hroadleaf species, including oaks, red
maple, and cherry usually established them-
selves beneath them from seeds carried in by
animals or blown in by the wind. All these hard-
woods have the ability to sprout from cut or
damaged stems. Therefore, even if they were cut
back when the pines were harvested, the hard-
woods could grow much more quickly from
their established root systems than could the
tiny pine seedlings. This succession from a first
generation of old-field white pine to a second
generation of mixed hardwoods has been typical
across most of Massachusetts.

The proliferation of old-field pine across
Massachusetts in the second half of the nine-
teenth century, before they were harvested and
replaced by hardwoods, led to problems as well
as economic benefit. The vast expanses of young
pines fed an epidemic of a native insect, the
white pine weevil. The larvae of this insect eat
the terminal buds of young pines, killing the
leader and releasing the branches in the topmost
whorl to replace it. At best, the growing trunks
develop a crook; at worst, they divide into mul-
tiple, spindly stems. In either case the economic
value of the trees is greatly reduced. White pine

blister rust, a fungal disease lethal to white pine,
also spread rapidly through the tracts of old-field
pine. This disease requires an alternate host of
the genus Rihes (currants and gooseberries) for
part of its life cycle. During the 1930s the state
and federal governments conducted a massive
eradication program for Ribes, with men march-
ing through the woods tens of feet apart pulling
up wild Rihes plants. The prevalence of white
pine weevil and blister rust also led in the 1920s
and 1930s to red pine being planted across the
state on many sites where white pine might
normally have grown, because red pine is not
affected by either pathogen. Although red pine
is at the very southeastern edge of its range in
western Massachusetts, these plantations have
generally done well. Many are now maturing
and being harvested.

The extensive old-field white pine stands also
played a major role in the most dramatic natu-
ral disturbance to affect our forests in the twen-
tieth century, the hurricane of September 21,
1938. Historically, hurricanes have been a major
force in shaping Massachusetts forests. The
1938 storm followed a track similar to that of
other historically significant storms (1788,
1815), but several factors conspired to make it
the most destructive storm in our recorded his-
tory. The week before had been very wet, satu-

The Dexter Woodlot, situated
iust south of Petersham
village. Before the hurricane
of September 21, 1938, this
was one of the most
attractive white pine groves
in the region.

After European Settlement 21

rating the soils and predisposing trees to
windthrow. The added rain from the storm pro-
duced massive property damage from flooding
along rivers, compounding the wind damage.
Large areas of central Massachusetts still sup-
ported stands of old-field pine on land aban-
doned in the late nineteenth century. Even pine
stands as young as thirty years of age suffered
severe damage if their sites were not protected
topographically from the southeast winds.
Hardwood stands on similar sites were not as
susceptible to damage unless they were at least
twice that age.^^ The prevalence of old-field
pines set the stage for the unprecedented impact
of the storm on our forests, nearly three billion
board feet of timber blown down. We had unin-
tentionally created about as vulnerable a land-
scape as possible. There is evidence that the
storms of 1788 and 1815 may have been similar
in intensity and path, but they encountered
a landscape with much less forest and their
impacts were quite different.

In 1938 the vast tracts of blown-down pine
presented a problem: the threat of fire. Fires
often follow other disturbances, especially in
conifer stands where the resinous foliage and
lack of new green sprouts contribute to flamma-
bility. With this in mind, and in an attempt to
recover some of the value of the blown-down

timber, a massive salvage operation was under-
taken that recovered much of the windthrown
timber. Logging crews were brought in from all
over the Northeast, temporary camps were set
up, and logs were salvaged and brought to the
mills. Because the volume of logs far exceeded
the capacity of all the available mills, logs were
stored in every pond in the area. As long as the
logs remained underwater, away from oxygen
in the air, they were preserved. Many ponds in
central Massachusetts were dammed and raised
to their present levels in order to accommodate
as much salvaged timber as possible. The tre-
mendous volume of lumber produced by the
hurricane salvage also drastically lowered lum-
ber values. To stabilize the price, the federal
government bought up the vast supply, stamp-
ing "U.S." at the end of each log. Mobilization
for World War II finally made use of this vast
lumber supply.

Humans have been unwitting accomplices
in several other recent forest disturbances as
well. Increased mobility of people and products
has resulted in numerous forest pests and patho-
gens being introduced from abroad. In many
instances these organisms pose special prob-
lems because native plants possess little resis-
tance to the exotic pests. Several such
"immigrants" have severely affected our forests.

STILLMAN ARCHIVES OF THE ARNOLD ARBORETUM

22 Arnoldia 1998 Summer

A1 Cline, director of Harvard Forest. 1939-1946, surveys white pine logs awaiting milling at Harvard Pond.
Following the 1938 hurricane more than half of the fallen timber across New England was salvaged, purchased
by the federal government, and stored in lakes and ponds to prevent insect damage, staining, and decay until
the material could be milled. Today, occasional logs stamped on the end with “U.S. " will be pulled from the
mud bottom of a pond and, when dried, provide perfectly intact and usable wood.

and Massachusetts has the dubious distinction
of being the introduction site of one pest that
has damaged forests on a national scale. Gypsy
moths were introduced into the United States
in 1869, when Leopold Trouvelot imported
them to Medford with the intention of using
them as silkworms to develop a local silk indus-

try. The moths quickly proved unsuited for this,
purpose and escaped into the local forests,
where they found the native deciduous species,
especially oaks and aspen, to be an ideal food
source. Since then, gypsy moths have gradually
expanded their range, and during periodic
regional outbreaks consume virtually every

Tne Arnold Artoretum

s U M M E

NEWS

The Arboretum Reaches Out to Influence the World

Gary Koller shares his ht)rticultural knowledge at an Arboretum Fall Plant Sale.

Robert E. Cook, Director

Whether it is our annual
crop of interns, our visiting
research fellows, or our
experienced staff, the
Arnold Arboretum serves as
a training ground for career
growth and achievement. As
a result, this often leads to
the loss of individuals to
other institutions as they
seek advancement in their
careers. Such losses, though
sad, are evidence of the
strength of the Arboretum
as a respected resource for
knowledge and training in
the fields of botany and hor-
ticulture. Over the past few
years, the Arboretum has
peppered its “alumni” throughout
the country.

Two years ago Dr. Kim Tripp,
who had been a Putnam fellow
from 1994 to 1996, accepted the
directorship of The Botanic
Garden of Smith College in
Northampton. Though she has
taken on a new and challenging
role, Kim retains a research
appointment at the Arboretum.

In 1996, Richard Schulhof,
director of public programs, was
named director of Descanso Gar-
dens in Los Angeles, after almost
seven years at the Arboretum.

Dr. Candace Julyan has been
appointed to replace Richard in a
re-organized education depart-
ment. Before coming to the Arbo-
retum in 1995 as project director

for our Community Science Con-
nection, Candace developed and
directed the National Geographic
Kid’s Network. Though we miss
him greatly, Richard continues to
work with Candace by participat-
ing in the Community Science
Connection.

After 17 years of service, Pat
Willoughby, our superintendent
of grounds, moved to a wonderful
opportunity in the Boston sub-
urbs, becoming superintendent
of grounds at Wellesley College
in 1997. His able assistant for
six years, Julie Coop, has been
appointed to his position at the
Arboretum. Julie was in charge of
maintenance at the Case Estates in
Weston from 19S8 to 1991. With
Julie’s promotion, we have hired

a new assistant superintendent,
Tom Akin, who comes from
Weston Nurseries.

Chris Strand, who supervised
visitor services for four years,
announced his departure from the
Arboretum in the fall of 1997.

He has been appointed manager
of Green Spring Gardens Park
in Virginia. Ellen Bennett has
recently assumed Chris’ restruc-
tured and renamed position as
manager of horticultural informa-
tion. Prior to her position at the
Arboretum, Ellen worked at the
Massachusetts Horticultural Soci-
ety for three years as director of
education and as community out-
reach coordinator.

With the new year came three
other critical changes in Arbore-

Karen Madsen

turn staff. Ciary Roller, who had
been our senior horticulturist for
over twenty years, announced that
he would retire in July to w-ork
full-time on a thriving landscape
design business. Ciary had been
serving half-time at the Arbore-
tum for the past five years while
his business was growing, and
in retirement he will retain an
appointment as horticultural fel-
low and will continue to teach in
our adult education program and
write for Arnold ia.

Stephen Spongberg, who had
been horticultural taxonomist
with us for more than a quarter
century, also retired to accept a
wonderful opportunity as director
of Polly Hill Arboretum on
Martha’s Vineyard, an entirely
new institution that will be devel-

oping its collections and programs
over the coming years under
Steve's leadership. He, too, has
retained a research appointment at
the Arboretum, and we anticipate
growing close relations with our
sister institution on the Vineyard.

Finally, Professor Peter Stevens
has announced that he will be
leaving with his wife. Professor
Toby Kellogg, a former Putnam
fellow at the Arboretum, for the
University of Missouri at St.

Louis, where they wdll fill two
tenured professorship positions.
Peter is a world expert on the
tropical flora of Southeast Asia,
and his systematic knowledge and
experience will be greatly missed.

In each of the above instances,
our colleagues have moved to an
exceptional opportunity for career

development; these changes have
given the Arboretum the chance
to recruit new staff and rebuild
programs. For instance, three job
searches are currently underway,
for a horticulturist, a botanical
taxonomist, and a researcher
experienced in the biology of
tropical forests.

The Arnold Arboretum is a
strong institution dedicated to
education as well as research, and
an important part of education
involves shaping individuals for
positions of leadership and respon-
sibility. Though inevitably these
positions will often be elsewhere,
the Arboretum will continue to
serve the fields of botany and hor-
ticulture in this important way. It
is one more way we can influence
the world.

1998 Interns

The interns of 1998: top
row from left, Yong
(ihan Park, Kristin
Berry, Bethany Grasso,

Sandy Talcott, Nigel
Gurnett, Marc
Fortenberry, Loise
Cretinon, Derrik Daly;
bottom row from left,

Fritz Bottger, Matthew
Coyner, Angie Martin,

Nicki Richardson, Gwen
Newman, and the
manager of the intern
program. Assistant
Superintendent Tom
Akin. Missing was
Lise Caron.

With members that represent our local region, the
international community (France and Korea), and
many states in between, the Arboretum’s intern class
of 1998 worked enthusiastically throughout spring
and summer. They took classes taught by Arboretum
staff on a variety of horticultural subjects, among
them woody plant identification, integrated pest man-
agement (IPM), pests and diseases, and plant propaga-
tion. In addition to working on an erosion abatement

project, the interns were involved with numerous
maintenance and renovation projects throughout the
grounds. To supplement their experiences at the
Arboretum, interns participated in several educational
outings, including trips to the New York Botanical
Garden, Mount Auburn Cemetery, and the new Polly
Hill Arboretum on Martha’s Vineyard. The interns’
diligence and enthusiasm made a valuable contribu-
tion to the Arboretum that is apparent at ever)- turn.

2

SUMMER 1998

Karen Madsen

Frances Maguire Named Harvard Hero

In June, Sally Zeckhauser,
Harvard University’s vice
president for administration,
announced this year's
Harc'ard Heroes — employees
in Ms. Zeckhauser’s unit
whose job performance has
been outstanding. Frances
Maguire, director of finance
and administration, has
been named the Arnold
Arboretum's 1998 Harvard
Hero. Director Bob Cook
noted that Frances has done a
superb job in managing the
financial affairs of the insti-
tution, saving many thou-
sands of dollars over the past
decade due to her careful
consideration of income and expenditures. Her deep knowledge of
the difficult Harvard accounting system has been invaluable to
the Arboretum. Recently, Frances has taken responsibility for
helping the Arboretum’s transition to a completely revamped
accounting system being adopted by Harvard University. For
these and numerous other reasons, Frances deserves great recogni-
tion and thanks from the Arboretum and from Harvard.

On June 10, all the Harvard Heroes
were recognized in a ceremony in
Sanders Theater, Memorial Hall.
Harvard president Neil Rudenstine
and his senior staff from central
administration were in attendance
to express their appreciation of the
Harvard Heroes.

Fundraising Reaches
New Heights

Lisa Hastings
Director of Development

The Arnold Arboretum completed
a record-bteaking year with
respect to the Campaign for the
Arboretum. Total gifts and com-
mitments for the fiscal year end-
ing June 30, 1998, reached
$2,016,834, the largest amount
raised from philanthropy in the
Arboretum’s history. The total
represents an increase of 191 per-
cent over the $692,1 1 1 raised in
fiscal year 1992.

MAJOR GIFTS: New records were
established both in total dollars
raised and total number of donors.
Gifts over $10,000 reached a total
of $1,704,373, an unprecedented
number that includes the
Arboretum’s first million-dollar
gift. The number of donors of
more than $10,000 increased 170
percent, from 10 in 1997 to 27 in
1998. Overall, the Arboretum

• continued on page 4

1997 Intern Is 1998 Apprentice

Tom For joined the Arboretum staff in April as
apprentice in the living collections department,
only the second person to hold this relatively new
position. Originally from St. Thomas, Ontario, he
comes to the Arboretum after graduating from the
Niagara Parks Commission Botanical Gardens and
School of Horticulture. Tom also has received a
bachelor of arts degree in history wdth honors from
the University of Toronto. The Arboretum is not
an entirely new entity for Tom, as he served as an
Arboretum intern in 1997. As a member of the living
collections department, he will be responsible for a
variety of projects on the grounds, including erosion
abatement on the esker just off Meadow Road.
During his year at the Arboretum, he will circulate
through all the units within living collections, thus
acquiring deeper knowledge of the maintenance
requirements of a scientific collection of
woody plants.

ARNOLD ARBORETUM NEWS

3

Karen Madsen

• from page 3

received 85 gifts over S 1 an
increase of 255 percent over the
22 gifts of this size received in
fiscal year 1993, the first year of
the campaign.
MEMBERSHIP/ANNUAL
APPEAL: A total of 638 new
members joined the Friends of
the Arnold Arboretum last year,
bringing the total number of
members at year's end to 3,364.
New members, coupled with a
renewal rate of 80 percent and a
healthy increase in membership
upgrades, resulted in an increase
of nearly 20 percent in member-
ship and 23 percent in member-
ship income. The largest

New Staff

In July, Ellen Bennett started as
manager of horticultural informa-
tion at the Arboretum. Ellen
comes from the Massachusetts
Horticultural Society, where she
served as director of education and
as community outreach coordina-
tor. Originally from Virginia, she
earned her master of science degree
in public horticulture administra-
tion through the Longwood
Graduate Program at the Univer-
sity of Delaware. As a member of
the education department, Ellen
will create a new docent program
and work on a variety of issues
related to the public face of the
Arboretum. She will also spend a
portion of her time working with
the living collections department.

membership increases (40 percent)
came at the level of Sustaining
iVIember (SI 00) and above, a testa-
ment to increased outreach and
the active participation of new and
reinvigorated volunteer leader-
ship. In addition, funds donated
through the Annual Appeal raised
S89,01 1 from 271 donors, a 29
percent increase in dollars and a
27 percent increase in the number
of donors over the previous year.
THE WOMEN'S MATCHING
FUND: The Women's Matching
Fund offered through Harvard
University has had a swift and
positive impact on our fund-
raising. Due to gifts and pledges
made by nine women, the Arbore-

Laurie Chidester has started work
as curriculum and web specialist
in the education department.
Laurie spent last year as a student
in the Harvard Graduate School of
Education working on her master
of education degree, focusing on
technology in education. Earlier,
during 13 years in software devel-
opment, Laurie made use of her
master's degree in computer sys-
tems engineering. Originally from
the Baltimore area of Maryland,

Laurie will be focusing on the
Community Science Connection
project funded by the National
Science Foundation. She will use
the internet to support teachers
and students in an incjuiry- based
curriculum model in the study
of trees.

turn has qualified for $391,000
since the fund was announced.

To date, $10 million of the total
fund balance of $15 million is
committed across the university;
the opportunity to qualify for
matched funds will end when the
fund balance is depleted.
SUMMARY: Total gifts and
pledges in the Campaign for the
Arboretum stand at $5,596,243,
representing 68 percent of the
Arboretum's $8.2 million goal.
With less than 18 months left
before the conclusion of the uni-
versity-wide campaign, the com-
ing year will be critical to closing
the gap and successfully complet-
ing the Arboretum's campaign.

Karen O’Connell, membership
coordinator, joined the Arboretum
development staff in July, replac-
ing Kelly Harv'ey who left the

Arboretum to begin graduate
studies in veterinary science.
Karen brings a wealth of experi-
ence in data management. She
worked for two years as data man-
ager in the marketing and devel-
opment department of the Boston
Ballet, prior to which she spent
four years with the Environmental
Defense Fund serving as office
manager and records manager.

A 1 989 graduate of Boston
College with a degree in market-
ing, she will oversee member
sers'ices and is looking forward
to meeting the Arboretum’s
growing constituency.

E

SUMMER 1998

After European Settlement 23

green leaf in the forest, leaving it in mid-July
looking nearly as barren as in midtvinter. Defo-
liation for two successive years is especially
harmful. The outbreak of 1980-1981 across the
Northeast was particularly severe, causing
extensive oak mortality. Today the gypsy moth
has spread throughout the Northeast and into
the Middle Atlantic and midwestern states
and is one of the most destructive forest pests
throughout the region.

Probably the most dramatic effect on our
forests by an introduced pathogen has been
that produced by the chestnut blight fungus.
Although the details of its introduction are not
certain, the fungus was first noticed in New
York in 1904 and rapidly spread throughout

the range of the American chestnut, passing
through Massachusetts in 1913-1914. An espe-
cially virulent pathogen, chestnut blight fungus
is the only pest that has effectively eliminated
mature individuals of its host, greatly altering
our forests in the process. Chestnut was cer-
tainly one of the most useful trees in the nine-
teenth-century forests, providing abundant
crops of edible nuts, bark for tanning, and excel-
lent wood that was beautiful, decay-resistant,
and as strong as oak but lighter. It also sprouted
vigorously and grew very quickly and therefore
increased in numbers in areas that were repeat-
edly harvested.

By the early 1920s all the large chestnuts in
the state had been killed. However, because in

20 20 20 40 60 20 20 40 20 40 60 20 20 40

Percent

Pollen diagram from the humus soil in a hemlock forest at the Harvard Forest, Petersham. Massachusetts. The
site is a primary forest that was never cleared for agriculture hut was clear-cut early in the settlement period
(apparent in the pollen diagram at a depth of approximately 17 centimeters) and then cut repeatedly for
firewood.

Each tree species responded differently to the series of human impacts. Chestnut benefitted greatly from
the cutting activity until its virtual elimination by blight in 1913 (seen at about 8 centimeters in depth). Other
major changes are the decline in several long-lived, shade-tolerant species during the agricultural period. Both
hemlock and beech are very sensitive to fire and could be largely eliminated from upland areas by repeated
fires, a rather common agricultural practice. Beech and sugar maple never recovered to presettlement levels of
abundance and have been replaced by oak, pine, and red maple, which have gradually increased: hemlock has
become the dominant species on this moist lowland following the loss of chestnut to the blight.

HARVARD FOREST ARCHIVES

24 Arnoldia 1998 Summer

Farm abandonment accidentally provided more favorable conditions for wildlife than did old forest: low-growing game
food and cover are much more plentiful and varied. Shrubs and apple trees furnish fruit and browse; valuable herbaceous
species and a wealth of insects are within reach of young birds. Deer, rabbits, woodcock, and aquatic birds are among the
wildlife that flourish in old fields, abandoned millponds, and stone walls.

effect the fungus kills by girdling the trees —
gaining access through cracks in the bark and
preventing transport of vv^ater and nutrients past
the point of infection — the roots and base are
not affected and can send out new sprouts. The
chestnut's decay resistance, especially within
the sapwood, has preserved many stumps that
testify to the former importance of this species,
and today chestnut sprouts are common in our
woods. Individual stems are usually killed by
the time they are several inches in diameter,
when the bark naturally develops cracks, only
to be replaced by new sprouts. Chestnut's place
in the forest has been taken by a mixture of spe-
cies, especially oaks, hut its wood and its nuts
cannot he replaced.

Other native tree species have also been sig-
nificantly affected by human-introduced agents,
although none so dramatically as the chestnut.
Dutch elm disease, a wilt fungus transported
by a bark beetle, completely transformed the
appearance of almost every town in the state in

the 1950s and 1960s by killing the stately shade
trees that lined most of our main streets. The
disease is passed from tree to tree by insects
above ground and through root grafts below
ground in areas where the trees grow adjacent to
each other, as in street plantings. Its effects were
somewhat less traumatic in our forests, because
elm occurred in mixed stands and exhibited a
greater range of natural resistance than did
chestnut. Nevertheless, the devastation of the
elms in our urban landscapes once again demon-
strates the susceptibility of manmade monocul-
tures to various pathogens.

More recently, many of our beeches have been
killed or disfigured by beech-bark disease. This
disease, caused hy the coincident impact of a
fungus and a scale insect working together, is
steadily spreading southward after being intro-
duced into the Canadian Maritimes. Hemlock
woolly adelgid is beginning to cause mortality
in the southern Connecticut River valley area
and has been reported in many other areas of

After European Settlement 25

the state as it slowly advances north. This
aphidlike insect, first introduced to the West
Coast and then to Maryland on nursery stock
from Japan, poses a dire threat to hemlock for-
ests because hemlocks have shown little resis-
tance to its effects and are incapable of
sprouting. Moreover, because many hemlock
stands occupy steep habitats and produce deep
shade, thereby creating unique microenviron-
ments, their loss would represent drastic
changes to many of our forests.

Over the past several decades,
however, it is through logging and
land conversion for suburban devel-
opment that humans have most
affected our forests directly. The
regrowth that followed the cutting
of old-field pine stands and other
forests early in this century and the
1938 hurricane has produced an
abundant middle-aged and maturing
forest, much of which has been and
is being harvested at varying degrees
of intensity. Limitations set for envi-
ronmental reasons on harvesting fed-
erally owned trees together with a
strong export market have resulted
in added pressures on Massachusetts'
forests. Despite these pressures,
however, the average size of our trees
has been steadily increasing. In some
instances we have even managed to
reduce the impact of suburban devel-
opment on the forest. Significant
numbers of people are now building
homes on large forested lots, clearing only the
area immediately around the houses, and in
some developments buildings are clustered
together, reserving the majority of land as forest
or open space. While both of these development
patterns alter the forest, they are much less
destructive than traditional tract development.

Wildlife species in Massachusetts have been
significantly influenced by human-induced
changes in the landscape as well as by hunting.
Information on this subject is difficult to gather
and much of it is indirect. It is believed, how-
ever, that most members of the large, broad-
ranging species, including elk, wolf, mountain
lion, and moose, were eliminated during the

initial period of forest clearance. Deer were
nearly eliminated by the mid-1800s. Because
they represent an edge species, however — using
open areas for browsing and forests for cover —
and are tolerant of human activities, deer have
responded so favorably to the return of the for-
est that they have reached densities detrimental
to the vegetation in those areas where they are
not controlled by hunting.^' Beavers were extir-
pated by the 1700s owing to the value of their
pelts. They were successfully reintroduced in

West Stockbridge in 1928 and have subse-
quently expanded their range to the point of
overutilizing existing habitat. More recently,
wild turkeys have been reintroduced very suc-
cessfully, and moose are returning on their
own as part of their growing northern popula-
tions migrate south. These three species are
responding to the expansion of our woodland
area, as have the black bear and the fisher,
which have significantly expanded their ranges
and numbers within the past seventy-five years.
Other species, most notably open-land birds
such as the hohwhite and meadowlark, have
decreased in number as the forest has regrown
and matured.

Changes in the relative abundance of selected animal species in
Massachusetts over the past 400 years. Whereas the wolf has been
eliminated and remains absent, beaver have been reintroduced and the
coyote represents a new species in the landscape.^°

PETER DEL TREDICl

IV. Present Conditions and Future Prospects

How do the forests we see today differ
from the forests the colonists found?
The process of deforestation and refor-
estation has produced landscape patterns that
vary with the distribution of natural and cul-
tural features within local areas. Today, open
agricultural land is primarily found only in
broad river valleys and on the crests of broad
ridges. Urban concentrations developed first
along the coast and major rivers and later along
the railroads, which tended to follow the rivers.
More recently, suburban development has

Old stone walls surrounded by forest bear witness to
abandoned farmland.

occurred along major highways, especially near
junctions. Forests now predominate outside
these zones, and in protected reserves and some
of the wetlands within them, and are under the
greatest pressure at the edges of these zones.

The changes have strongly favored a new
landscape of even-aged forests, with sharp
boundaries between types. Agricultural clearing
and abandonment, heavy cutting for fuelwood,
intensive harvesting of old-field pine and other
species early in this century, the hurricane of
1938 with its subsequent salvage harvesting —
all these phenomena have combined to create
the even-aged forests we find throughout Massa-
chusetts. Land-use regulations and land owner-
ship boundaries create visible differences in
vegetation that tend to be perpetuated through
time and through changes in ownership. Trends
in the size of fields and farms and in regional
timber harvesting practices have imposed a
repetitive patchwork of forest types that has
replaced the natural vegetation patterns. These
even-aged forests and imposed patterns increase
the possibility that disturbances will in the
future cause far more damage than might be
expected in a more diverse forest. Moreover,
the relative scarcity of very young forests inhib-
its the growth of species that require this type
of habitat.

Although they tend to be even-aged, over the
past one hundred-plus years, the forests in
Petersham have continually increased m area
and size, a trend that has been repeated m for-
ests throughout the state. How has the compo-
sition of our forests been affected? The most
dramatic change has been the increase in white
pine following agricultural clearing and subse-
quent abandonment. Prior to European settle-
ment white pine was confined mainly to sandy
outwash soils that had undergone natural dis-
turbance or to sites heavily burned by Indians;
or it appeared as scattered, emergent individuals

Present Conditions and Future Prospects U

Forest cover (shown in green) for north-central Massachusetts in 1830, at
the approximate peak of agricultural clearance, and in 1980. Major
physiographic regions include the Connecticut River valley, the rough
Pelham Hills to its east, and the undulating central upland regions farther
east. ND indicates no data.^^

in old stands of mixed species.

Following agricultural abandon-
ment, especially of pastures,
white pine proliferated through-
out most of the state on sites
it would never have occupied
without prior clearing and graz-
ing. Despite intensive harvesting
and the 1938 hurricane, white
pine remains much more domi-
nant and widely distributed today
than It was before European
settlement.

On woodlots, repeated cutting
for fuelwood and burning for agri-
culture m the nineteenth century
favored an increase in invasive,
shade-intolerant pioneer species —
gray and paper birch, aspen, pin
and black cherry — as well as
species that sprout prolifically —
chestnut, oak, red maple, hickory,
and birch. Chestnut is probably
the species that responded most
favorably to these nineteenth-
century disturbances because it
sprouts prolifically from dormant
basal buds and is capable of phe-
nomenal rates of growth both in
height and diameter when repro-
ducing vegetatively.^^'^^

As our forests have reappeared
following the period of extensive
cutting at the turn of the century,
the loss of chestnut in the teens,
the 1938 hurricane, and the more
recent initiation of fire-suppression policies,
long-lived, shade-tolerant species have gradu-
ally increased in number — hemlock, sugar
maple, and, to a lesser extent, the highly distur-
bance-intolerant beech. However, early survey
records and pollen analyses suggest that these
species, especially beech, remain well below
their presettlement distributions. On the other
hand, oak, which requires at least moderate dis-
turbance for successful regeneration, may be
more common than before settlement.

Significant new harvesting has followed on
the recent maturation of considerable areas of
Massachusetts forest. At the same time, the

environmental awareness that began to develop
in the 1960s has led to new regulations on
forest-cutting practices, these in turn have
raised the extent and quality of professional for-
est management across the state. The growing
understanding that forests do not function in
isolation has highlighted the need for regionally
mapped information for use in designing man-
agement practices within the context of a region
or ecosystem. Fortunately, the requirement to
file a cutting plan for all harvests greater than
25,000 board feet has made it possible to map
current harvesting patterns in much finer detail
than was possible in the past.

HARVARD FOREST ARCHIVES

28 Arnoldici 1998 Summer

The Pisgah old-growth tract in Westminster, New Hampshire. This forest of 300-year-old white pine, hemlock,
and hardwoods was purchased by the Harvard Forest in 1922 in order to protect it from logging and to provide
a study area for investigating natural forest processes. In the hurricane of September 22, 1938, it was
completely blown down, and the area has formed an important long-term study in forest dynamics and
recovery from disturbance.

Growing environmental interest has also
led to the discovery of remnant patches of
old-growth forest, once assumed to have been
entirely eliminated by the extensive clearing
and harvesting of the nineteenth and early
twentieth centuries. Although exact definitions
of "old-growth" vary considerably, these rem-
nants typically include dominant trees well
over two hundred years old and show minimal
evidence of human disturbance. At present
between 500 and 1,000 acres of old-growth
forest are recognized in Massachusetts,^* and
the number continues to rise as more areas are
investigated by scientists with a better under-
standing of what they are looking for, which is
not necessarily huge old trees.

Many of these remnants are small patches of
barely ten acres — which, according to one cur-
rent working definition, is the minimum size

necessary to prevent significant edge impacts —
but some remnants are considerably larger.
Most are located on steep, rocky slopes, often on
headwater streams, where they were inacces-
sible for harvesting from either the stream
valley or the broad ridges and were somewhat
protected from natural disturbances as well.

Not even these sites offer protection from
recent human disturbances, which are subtle
but pervasive. These include atmospheric pollu-
tion and rising carbon dioxide (CO,) levels, both
of which may he implicated in the global warm-
ing predicted by many scientists. While neither
of these forces has yet had serious, measurable
impacts on our forests, both have the potential
to significantly alter them in the future.

Because the effects of pollutants are
extremely complex, the eventual impact of
long-term exposure is still largely unknown.

Present Conditions and Future Prospects 29

Map of currently known old-growth stands in
Massachusetts.

The amount of nitrogen (NOx) — an important
component of atmospheric pollution — increases
as one moves farther west and to higher eleva-
tions in Massachusetts.^^ As the major limiting
nutrient for plant growth in our soils, nitrogen
works initially as a fertilizer, but at higher con-
centrations it may lead to nutrient loss through
leaching.^® Ozone found at low atmospheric
elevations is another pollutant with potentially
serious forest impacts.

Elevated CO2 levels affect the quality of
organic matter in the soil through their influ-
ence on plant growth, competitive interactions,
and leaf chemistry, and have the potential to
change global climate. We do not yet understand
how forest communities and ecosystem pro-
cesses might ultimately be changed by elevated
CO2 levels, nor do we know the local effects of
global warming. Massachusetts' forests do have
some impact on the global CO2 level: because
they are still relatively young and growing, our
forests take up and store significant amounts
of CO2, slightly offsetting the increases from
fossil-fuel burning and deforestation. CO2 levels
and pollution are both international issues that
will require unprecedented levels of cooperation
if they are to be controlled.

Our forests have changed constantly through-
out geologic and historical time, but human-
induced changes over the past three hundred
years have been much more frequent, varied, and
far-reaching. These changes have been superim-
posed upon natural disturbances, and where the
two processes have acted in concert, as in the
1938 hurricane, the impact has been substantial.

The forests across the state today are quite
different from those the colonists and Indians
saw — indeed, they are quite different from those
of sixty years ago. They will continue to change,
but recent trends in carbon storage and increas-
ing volume of wood will undoubtedly continue.
Changing ownership patterns will increasingly
affect forest development. Over the past fifty
years both the term of ownership and the aver-
age size of forest properties have continued to
shrink as our population has become less and

The tower of the Harvard Forest’s Environmental
Measurement Station measures exchanges of gases,
energy, and moisture between the atmosphere and the
forest. These measurements document the surprisingly
high rate of carbon uptake by temperate forests that are
recovering from earlier land use. It also documents
seasonal changes in forest activity due to meteorological
changes and ozone stress.

JOHN O'KEEFE

30 Arnoldia 1998 Summer

less agrarian and more and more mobile, and —
as suburbs encroach on rural areas — as more and
more people have built their homes in wooded
areas. These trends will influence our forests
and their management into the next century.
At the same time, demands for forest conserva-
tion and preservation will no doubt increase —
especially on public lands — as the value of our
forests for recreation, amenity, and watershed-
protection increases even more rapidly than
their value for development or other economic
uses. Humans — directly, indirectly, and in con-
junction with natural processes — will continue
to be the dominant force acting on our forests.

The recovery of Massachusetts' forests is
testimony to the resilience of our landscape in
the face of centuries of natural disturbance and
unthinking human activity. However, it is
critical that decisions regarding conservation.

Endnotes

* A. N. Strahler, A Geologist’s View of Cape Cod
(Garden City, NY: Natural History Press, 1966).

^ M. B. Davis, Three pollen diagrams from central
Massachusetts. American Journal of Science (1958)
256: 540-570.

^ W. A. Patterson and A. E. Backman, Fire and disease
history of forests. In Vegetation History, ed.

B. Huntley and T. Webb (Dordrecht: Kluwer, 1988),
603-632.

D. R. Foster and T. M. Zebryk, Long-term vegetation
dynamics and disturbance history of a Tsuga-
dominated forest in central New England. Ecology
(1993) 74: 982-998.

^ I. K. Wright, Regions and landscapes of New England.
In New England's Prospect: 1933 (New York:
American Geographical Society, 1933), 14-49.

^ N. Jorgensen, A Guide to New England's Landscape
(Chester, CT: Globe Pequot Press, 1977).

^ M. V. Westveld, Natural forest vegetation zones of
New England. Journal of Forestry (1956) 54: 332-338.

* E. P. Stephens, The historical-developmental method
of determining forest trends. Ph.D. thesis
(Cambridge: Harvard University, 1955).

® W. H. Lyford and D. W, MacLean, Mound and pit
relief in relation to soil disturbance and tree
distribution in New Brunswick, Canada. Harvard
Forest Paper (1966) 15.

J. Jenkins, Notes on the Adirondack blowdown of July
15, 1995: Scientific background, observation,

responses and policy issues. Wildlife Conservation
Society Draft Report, 1995.

forestry, wildlife management — in fact, all
environmental decision-making — begin with
knowledge and an appreciation of the history
and dynamic nature of our landscape. Without
these, any plan will almost certainly produce
surprises, if not failure. The forests have
reclaimed abandoned farmland and now cover
nearly two-thirds of Massachusetts. As our
population expands onto this land, the new
suburban forest owners, largely unaware of the
history of our forests and only slightly more
informed about changes now occurring, must
become more knowledgeable about the dynamic
nature of their backyard forests. We are blessed
with a landscape and a climate that are ideally
suited for growing trees and forests, but with-
out an understanding of the past we may
unwittingly lose many of the values these
forests can provide.

'* D. R. Foster and E. R. Boose, Patterns of forest
damage resulting from catastrophic wind in central
New England, U.S.A. Journal of Ecology (1992) 80:
79-98.

T. D. Allison, R. E. Moeller, and M. B. Davis, Pollen
in laminated sediments provides evidence for a mid-
Holocene forest pathogen outbreak. Ecology (1986)
67: 1101-1105.

W. A. Patterson and K. E. Sassaman, Indian fires in
the prehistory of New England. In Holocene Human
Ecology in Northeastern North America, ed. G. P.
Nicholas (New York: Plenum Publishing, 1988),
107-135.

’■* D. M. Smith, Storm damage in New England forests.
M.S. thesis (New Haven, CT: Yale University, 1946).

D. R. Foster, Species and stand response to
catastrophic wind in central New England, U.S.A.
Journal of Ecology (1988) 6: 135-151.

G. C. Whitney, From Coastal Wilderness to Fruited
Plain: A History of Environmental Change in
Temperate North America from 1500 to the Present
(Cambridge, UK: Cambridge University Press, 1994).

W. Cronon, Changes in the Land: Indians, Colonists
and the Ecology of New England (New York: Hill
and Wang).

C. Clark, The Eastern Frontier: The Settlement of
Northern New England 1610-1763 (New York:
Knopf, 1983).

D. R. Foster, Land-use history and four hundred
years of vegetation change in New England, U.S.A.
In Global Land Use Change: A Perspective from
the Columbian Encounter, ed. B. L. Turner et al.
SCOPE Publication (Madrid: Consejo Superior de
Investifaciones Cientificas, 1995), 253-321.

Present Conditions and Future Prospects 31

Information is derived from the following sources:
For Massachusetts, D. R. Dickson and C. L. McAfee,
Forest Statistics for Massachusetts. 1972 and 1985,
Resource Bulletin NE-106 (USDA Forest Service,
1988); W. R MacConnell, Remote sensing 20 years
of change in Massachusetts, Massachusetts
Agricultural Experiment Station Bulletin 630 (1975);
F. W. Rane, Fourth Annual Report of the State
Forester of Massachusetts for the Year 1907 (Boston:
Wright and Potter, 1908); and Fi. I. Baldwin, Forestry
in New England (Boston: National Resources
Planning Board Publication 70, 1942).

For Petersham, FI. M. Raup and R. E. Carlson, The
history of land-use in the Fiarvard Forest, Harvard
Forest Bulletin (1941) 20: 1-64; W. P MacConnell
and W. Niedzwiedz, Remote sensing 20 years of
change in Worcester County, Massachusetts, 1951-
1971 (Amherst, MA: Massachusetts Agricultural
Experiment Station, University Massachusetts,
1974); Fi. O. Cook, The Forests of Worcester County.
The Results of a Forest Survey of the Fifty-Nine
Towns in the County and a Study of Their Lumber
Industry (Boston: State Printing Office, 1917); and
Rane, Fourth Annual Report of the State Forester.

For Prospect FFill, D. R. Foster, Land-use history
(1730-1990) and vegetation dynamics in central New
England, U.S.A., fournal of Ecology (1992) 80: 753-
772; and S. H. Spurr, Stand composition in the
Harvard Forest as influenced by site and forest
management, Ph.D. thesis (New Haven, CT: Yale
University, 1950).

Maps are compiled from the atlas of Worcester
County (1830, unpublished) and analysis of aerial
photographs for 1985.

M. R. Pabst, Agricultural trends in the Connecticut
valley region of Massachusetts, 1800-1900. Smith
College Studies in History (1941) 26: 1-135.

^ A. H. Baker and H. V. Izard, Production Changes and
Marketing Strategies of Worcester County Farmers,
1780-1865: A Case Study of the Wards of Shrewsbury
(Dublin, NH: Society for History of Early American
Republic, 1987).

C. Merchant, Ecological Revolutions. Nature, gender
and science in New England (Chapel Hill: University
of North Carolina, 1989).

A. K. Botts, Northbridge, Massachusetts, A Town
That Moved Down Hill, fournal of Geography {193A]
33: 249-260.

N. lones. An ecological history of agricultural land-
use in two Massachusetts towns. Senior thesis
(Amherst, MA: Hampshire College, 1991).

A. H. Baker and H. I. Patterson, Farmers' adaptations
to markets in early nineteenth-century Massa-
chusetts. Proceedings of the Dublin Seminar
(London: Antiquities Society, 1986), 95-108.

Data from U.S. Census with maps modified
from R. W. Wilkie and J. Tager, Historical Atlas

of Massachusetts (Amherst: University of
Massachusetts Press, 1991).

D. R. Foster, T. M. Zebryk, P. K. Schoonmaker, and A.
Lezberg, Post-settlement history of human land-use
history and vegetation dynamics of a hemlock
(Tsuga) woodlot in central New England, fournal of
Ecology [1991] 80: 773-786.

Modified from W. E. Bickford and U. J. Dyman,
An Atlas of Massachusetts River Systems:
Environmental Designs for the Future. (Amherst:
University of Massachusetts Press, 1990).

T. Kyker-Snowman, 1989 Quabbin forest
regeneration study (Boston: Metropolitan District
Commission (MDC) Publication, 1989).

R. Zon, Chestnut in southern Maryland, U.S.D.A.
Bureau of Forestry Bulletin 53 (1904).

F. L. Paillet and P. A. Rutter, Replacement of native
oak and hickory tree species by the introduced
American chestnut (Castanea dentata) in
southwestern Wisconsin. Canadian fournal of
Botany [1989] 7: 3457-3469.

D. R. Foster, G. Motzkin, and B. Slater, Land-use
history as long-term broad-scale disturbance: regional
forest dynamics in central New England. Ecosystems
(1998) 1: 96-119.

Massachusetts General Laws, chapter 132, sections
40-46.

P. W. Dunwiddie and R. T. Leverett, Survey of Old-
Growth Forest in Massachusetts. Rhodora (1996) 98:
419-444.

37 S. G. McNulty, J. D. Aber, T. M. McLellan, and S. M.
Katt, Nitrogen cycling in high elevation forests of the
northeastern U.S. in relation to nitrogen deposition.
Ambio (1990) 9: 38-40.

33 L D. Aber, K. J. Nadelhoffer, P. Steudler, and L M.
Melillo, Nitrogen saturation in northern forest
ecosystems. BioScience (1989) 9: 376-386.

The authors are ecologists on the staff of the Harvard
Forest, a research and educational department of
Harvard University located in Petersham, Massa-
chusetts. David Foster is director of the Harvard Forest
and teaches forest ecology and conservation in the
Department of Organismic and Evolutionary Biology
and the Kennedy School of Government. John O'Keefe
is coordinator of the Fisher Museum, a small museum
devoted to the land-use history, ecology, and
management of New England's forests.

This article is adapted from "An Ecological History
of Massachusetts Forests," in Stepping Back to Look
Forward: A History of the Massachusetts Forest, edited
by Charles H. W. Foster, published in 1998 by the
Harvard Forest, and distributed by the Harvard
University Press. The book can also be obtained postpaid
from Harvard Forest, Petersham, MA 01366, where
proceeds from the $28 price go to support undergraduate
research and education in forest ecology.

.^2 Arnoldia 1998 Summer

oc

<

K IQH

'.ifj

m

m

Old-growth forest in New England before the hurricane of 1938.

CASE STUDIES IN FOREST HISTORY AND ECOLOGY

How Land Use Determines Vegetation:
Evidence from a New England Sand Plain

Glenn Motzkin and David R. Foster

Human activity exerts long-lasting impacts on
natural ecosystems, and because today's forest
composition reflects past land uses as well as
natural phenomena, knowledge of land-use
history and an understanding of its effects are
integral to ecological study and critical for con-
servation planning. However, attempts to deter-
mine the relative contribution of land use vs.
physical factors in controlling vegetation pat-
terns have consistently been confounded by the
strong correlation between past land use and
original site conditions.

To evaluate the effects of historical land use,
researchers at the Harvard Forest studied pitch
pine-scrub oak communities on a broad sand
plain in the Connecticut River valley of Massa-
chusetts, a site that is unusually homogenous
in topography, drainage, and soil texture. Such
homogeneity enabled the researchers to detect
the effects of differing land-use on the structure
and composition of the vegetation. Also motivat-
ing the study was the rarity of pitch pine-scrub
oak communities. They support several rare
plant and animal species but have been substan-
tially degraded by industrial, commercial, and
residential development, and are therefore priori-
ties for conservation throughout the Northeast.

The paleoecological record of the study area —
1,900 acres on a flat outwash delta composed
primarily of sand and gravel — suggests that pre-
European fires were common, some of them,
perhaps, ignited by a large regional Indian popu-
lation. Like other sand plains in the region, the
area was used for wood products from the eigh-
teenth to the mid-nineteenth century. In the
early nineteenth century, the Reverend Timo-
thy Dwight, author of Travels in New England
and New York (1821), described the Montague
Plain and surrounding areas as "an extensive

yellow pine plain covered with a lean, miserable
soil." Nonetheless, 82 percent of it was subse-
quently plowed for crops before being abandoned
m the early twentieth century. Agriculture had
a major impact on vegetation but a relatively
minor long-term effect on physical and chemi-
cal soil properties on this site. For instance,
although visually striking, the difference be-
tween plowed and unplowed soil horizons is pri-
marily one of color, the result of organic matter
being redistributed within the soil by plowing.

Agricultural fields on Montague Plain were
abandoned and allowed to reforest at least fifty
years ago — some more than one hundred years
ago — and yet the plant composition remains
very different on areas that were once plowed
compared to those that were never plowed.
Some species are just as common today on
formerly plowed areas as on unplowed sites.
However, several species, most notably pitch
pine (Finns rigida) and gray birch (Betula
populifolia), are much more common on plowed
sites. Of particular interest is a group of species
that is characteristic of sand plain habitats and
is common on unplowed portions of Montague
Plain, but that has not successfully re-colonized
former agricultural lands even though they were
abandoned more than half a century ago. This
group includes some familiar species, such as
black huckleberry (Gaylussacia baccata), win-
tergreen (Gaultheria procumbens), blueberries
(Vaccinium spp.j. Intensive soil and population
analyses suggest that the failure of these species
to re-colonize these sites does not result from
plowing; instead, it appears that successful
sexual reproduction is rather infrequent in these
species. Their rates of vegetative spread are so
slow that even one hundred years is not long
enough for re-colonization.

34 Arnoldia 1998 Summer

Cross-sectional diagram across the major land-use, vegetation, and soils
boundary at the Montague sand plain. An unplowed site is on the left, a
plowed site on the right. Even after a century of forest growth, the soils on
plowed sites exhibit a deep and light-colored A horizon, sharply separated
from the underlying B horizon.

Fire has been important on the Montague
Plain: high charcoal-to-pollen ratios in the
paleoecological record, documentation of many
large fires during this century, and field evi-
dence of fire in 83 percent of the research plots
testify to that. How^ever, fire does not appear to
be the primary determinant of vegetation pat-
terns on Montague Plain. Rather, fire is appar-
ently important within a pattern of species
associations that is largely controlled by prior
land use. For instance, pitch pine requires
exposed mineral soil and open canopy condi-
tions for successful establishment, conditions
that may be met through physical disturbance
such as plowing or severe fire. Because few
recent fires have been severe enough to create
these conditions, nearly all extant pitch pine
stands are located on abandoned plowed fields.
After pitch pine became established, fire con-
tributed to its dominance on old fields.

The high frequency of extensive fires early in
this century (when most stands were becoming
established) probably favored pitch pine, which
produces seed at a much younger age than hard-
woods or white pine. Young white pine is more
susceptible to fire than pitch pine and lacks the
ability to resprout, an ability that is shared by
pitch pine and associated hardwood species. In
the absence of fire, hardwoods and especially
white pine increase in number, whereas fre-
quent light fires limit white pine and increase
the understory of hardwood sprouts without

disturbing the pitch pine over-
story.

On unplowed sites, on the
other hand, the vegetation
structure varies from closed
canopy hardwood-huckleberry
stands to dense scrub oak
thickets. We suspect that on
these sites repeated cutting and
burning converted formerly
forested areas into shrublands:
fire promotes the stability of
scrub oak stands by encourag-
ing vigorous sprouting and by
removing developing tree cano-
pies. In the absence of fire,
however, hardwood trees slowly
become reestablished in scrub
oak stands to form an open forest canopy. Since
scrub oak is intolerant of shade, this canopy
eventually reduces its numbers. The continuum
from scrub oak thickets to hardwood forests on
unplowed sites is therefore believed to result
both from fire and cutting history.

In all of these examples, a distinction remains
between the vegetation of plowed sites and that
of unplowed sites. We have therefore concluded
that modern vegetation patterns on Montague
Plain are the result of complex disturbance his-
tories, with fire serving to modify the species
assemblages that originally developed after land
use was discontinued. In other words, the inter-
actions of human, physical, and biotic factors
have resulted in a remarkably heterogeneous
landscape on a site with homogeneous soils.

In view of the heterogeneity of vegetation and
the equally variable natural disturbance pat-
terns in the area, conservation efforts might
best focus on long-term protection of the entire
landscape, rather than simply protecting the
portions that currently support uncommon spe-
cies. Only in this way will long-term protection
of this complex, dynamic system be ensured.

Glenn Motzkin is a plant ecologist at the Harvard Forest,
where David Foster is the director. Their research is
reported in full in "Controlling Site to Evaluate Histor>’:
Vegetation Patterns of a New England Sand Plain" by
Glenn Motzkin, David Foster, Arthur Allen, lonathan
Harrod, and Richard Boone, and published in Ecological
Monographs (1996) 66(3): 345-365.

CASE STUDIES IN FOREST HISTORY AND ECOLOGY

Forest Response to Natural Disturbance Versus
Human-Induced Stresses

David R. Foster, John Aber, Jerry Melillo, Richard D. Bowden, and
Fakhri Bazzaz

Retrospective studies, which employ
historical, archaeological, paleoeco-
logical, and dendrochronological tech-
niques to unravel past changes in
landscapes and the environment, pro-
vide one of the few opportunities for
comparing forest disturbance and vege-
tation dynamics across major cultural
and temporal boundaries. By extending
our perspective from decades to mil-
lennia, historical forest reconstruc-
tions permit us to assess the effects of
infrequent events, long-term trends,
and gradual environmental change.
They also enable us to document the
successional or developmental changes
in forest ecosystems that are often
comprised of tree species that can live
many centuries. Therefore, these assess-
ments provide important insights into
fundamental ecological processes and
the provenance of modern conditions
and can also serve as the basis for
informed management decisions.

One important source of long-term
information on forest history and
dynamics in response to natural envi-
ronmental change and disturbance are
old-growth forests and other sites on
which human impact has been mini-
mal. Recent surveys have discovered
surprisingly large numbers of old-
growth stands — even in the densely
populated eastern U.S. — and these for-
ests have become an important focus of
reconstructive studies on both basic
and applied issues. The presence of old
trees with lengthy tree-ring records of

Saplings grow between moss-covered logs of old-growth white pine
and hemlock downed in the hurricane of 1938.

36 Ainoldia 1998 Summer

growth, and undisturbed sediments and soils that
contain stratigraphic records of pollen, charcoal,
and other semi-fossilized plant and animal
materials, enable us to develop long-term site
histories of disturbance and forest change.

Reconstructive studies of old-growth and
other sites have revealed the remarkable resil-
iency of the temperate forests in the Northeast
to a wide range of physical disturbances, includ-
ing windthrow, fire, ice damage, and pathogens.
However, a question of great interest to ecolo-
gists and with major implications for policy-
makers is whether these forests will be equally
resilient in the face of chronic chemical and cli-
matic stress brought on by changes in the global
earth-atmosphere system. To address this ques-
tion, the Harvard Forest initiated its Long Term
Ecological Research (LTER) program in 1988 to
analyze the effects on ecosystem structure and
function of historically important natural dis-
turbances in comparison with those of recent
and projected chemical and climatic stresses.

Three disturbance and stress processes are
being investigated in considerable detail: hurri-
cane blowdowns, the chronic additions of nitro-
gen that occur presently as a consequence of
acid rain and the burning of fossil fuels, and the
warming of the soil environment that will occur
as global temperatures rise with predicted cli-
mate change. To develop meaningful compara-
tive results, we designed experimental
treatments that closely simulate
the major impacts of the relevant
disturbances and stresses, and
we chose sites m typical second-
growth forests on widespread
upland soils with similar forest
and environmental conditions.

The Experimental Blowdown

Hurricane blowdowns cause
catastrophic damage in New
England forests every fifty to one
hundred and fifty years. The
LTER studies undertook to
evaluate the effects of a storm
similar to that of the 1938
hurricane on an eighty-year-old
hardwood forest. To simulate a
hurricane on an experimental
plot, researchers used a power-

driven winch to pull over in one direction, as
would occur m a natural windstorm, a subset of
trees comparable to those blown down in 1938.

The experimental blowdown had an immedi-
ate and dramatic impact on forest structure:
the basal area (cross-sectional area of wood)
and density of trees declined more than 70 per-
cent; the average distance between canopy trees
increased from 3.3 to 7.6 meters, and the maxi-
mum distance increased from 7.8 to 20.9
meters; and more than 260 new mounds and
pits resulting from the uprooting of trees cov-
ered 8 percent of the soil surface.

Nonetheless, as a result of releafing and
sprouting, more than 75 percent of uprooted and
broken trees survived the first year and more
than 40 percent continued growing after four
years. Survival varied considerably according to
species and extent of damage: 60 percent of yel-
low and black birch iBetula alleghaniensis and
B. lentaj survived, as did approximately 50
percent of white pine (Pinus strobus) and red
maple (Acer rubrum), and approximately 30
percent of red oak IQuercus rubra), white birch
(B. papyrifera), and white ash (Fraxinus
americana). The standing and relatively undam-
aged trees remaining after the blowdown
showed the same low mortality rate (4 percent)
as the control site for the first four years.

The extent of tree regeneration in the
"blowdown" understory was high. Saplings and

A skidder's winch and cable were used at the Harvard Forest to pull dowm '
trees across a two-acre area to simulate the effects of a hurricane in 1990. .

lOHN () Kl.LI L

Natural Disturbance Versus Human-Induced Stresses 37

Researchers attach markers to a root mound upended in a simulated
hurricane, 1992. The many microsites created by the uprooting of trees
provide important habitat and microenvironmental diversity in the forest.

sprouts increased from fewer than 6,000 per
hectare before the blowdown to nearly 8,000 per
hectare one year later and to over 25,000
per hectare after three years, with only slight
compositional change (birches increased,
black cherry [Prunus serotina] and white pine
declined). Increased light levels one to three
meters above the ground resulted in greater
growth in diameter and height of existing sap-
lings and new sprouts on the blowdown site
than on the control site.

Net ecosystem productivity declined follow-
ing the experimental blowdown: in the second
year, litterfall in the blowdown was only 59
percent of that found in the control site; by
the fourth year, however, that number had
increased to 71 percent. In general, the blow-
down site underwent major reorganization of
forest structure and a subtle compositional
change along with rapid redevelopment of
canopy cover. In contrast, and contrary to gen-
eral expectation, the soil environment showed
no major changes. Temperature and moisture
remained the same, as did the nitrogen cycling
pattern and net exchanges of important green-
house gases between soils and the atmosphere.

The hurricane blowdown caused a marked
change in the appearance of the forest, but the
long-term impact on important processes was
far less dramatic: it was limited to a vertical
reorganization of the canopy and foliage from
the top of the canopy to one to five meters above

the ground. The combination
of high survival rates and pro-
lific sprouting by broken and
uprooted trees, coupled with
rapid growth by understory
plants, resulted in rapid recovery
of total leaf area and little change
in the soil microenvironment,
including temperature and mois-
ture: as damaged trees died off,
the growth of saplings and
understory plants has provided
additional leaf area. By ensuring
the continuity of shade on the
forest floor, floristic composition
has changed less than might be
expected based on successional
theory and on previous studies of
the 1938 storm.

From an ecosystem perspective, which seeks
to examine the structure and function of forests,
the blowdown experiment is highly instructive.
Despite massive structural alteration of the for-
est, net energy and nutrient processes remained
largely intact. Productivity, as measured by
litterfall, declined immediately following the
disturbance, but recovered rapidly within four
years. The similarity of nitrogen cycling pat-
terns, soil respiration rates, and gaseous effluxes
in the control and blowdown plots indicates
that changes in nutrient availability were mini-
mal. Importantly, the maintenance of efficient
cycling of nutrients suggests that the forest was
able to retain these constituents despite the
major physical change. Continuous vegetation
production and cover provided a high degree of
control by the vegetation over critical microcli-
matic conditions and ecosystem processes.

The Experimental Nitrogen Increases

In industrialized parts of North America and
Europe, the atmospheric deposition of nitrogen
has vastly increased since the 1940s as a conse-
quence of fossil fuel combustion. In the high
elevation forests of New York and New England
and throughout central Europe, nitrogen deposi-
tion has been implicated in tree mortality and
forest dieback. This conclusion is somewhat
paradoxical because nitrogen is limiting in most
terrestrial ecosystems and an increase might be
expected to simply increase growth rates. How-

)()HN I KEtFL

38 Arnoldia 1998 Summer

After the experimental blowdown, gridded frames were used to map fine-
scale microtopography and to release and track tree seed as part of a study of
seed dispersal. The root mound stands nearly two meters high.

ever, as nitrogen availability through acid rain
exceeds demand, the ecosystem may become
"saturated," basic processes may change in del-
eterious ways, and the excess nitrogen may leak
through soils into streams and lakes. There is
strong concern that this increased nitrogen will
damage water quality and alter critical soil pro-
cesses, including nutrient cycling and trace gas
fluxes, throughout the temperate zone.

The chronic nitrogen, or nitrogen saturation,
experiment was designed to examine the effects
of continuous, low-level additions of nitrogen
caused by acid deposition or forest ecosystem
structure and function. Two stands were selected
for study: a second-growth hardwood stand and
a mature red pine (Pinus resinosa) stand. Impor-
tantly, the eighteenth- and nineteenth-century
land-use history of the hardwood forest involved
repeated cutting, which probably depleted nutri-
ents on the site, whereas the red pine stand had
been planted in the 1920s on an old agricultural
field that had been plowed and fertilized,
thereby maintaining or enhancing its natural
abundance of nitrogen.

Three 30-by-30-meter plots — control (no nitro-
gen), low nitrogen-addition, and high nitrogen-
addition — were established in each stand; nitro-
gen was applied in the form of ammonium
nitrate in six equal monthly doses from May to
October beginning in 1988. The high nitrogen-
addition plots receive doses similar to levels
occurring in central European countries today.

In marked contrast to the blow-
down experiment, the chronic
nitrogen experiment has so far
produced only minimal structural
changes. Tree mortality has not
been affected, and species com-
position and canopy structure
remain similar to those in control
plots. However, dramatic changes
in ecosystem function have
occurred that may foreshadow
future changes in structure.

For example, whereas the con-
trol plots retain essentially all of
their nitrogen and lose none
through soil leaching, the high-
nitrogen pine plot has shown
accelerated nitrate loss, indicative
of nitrogen saturation, since year
three. In the sixth year of treatment, the low-
nitrogen pine plot also began showing nitrate
losses. By contrast, the more nitrogen-limited
hardwood stand has shown nearly total reten-
tion of nitrogen; detectable nitrate losses were
not measured until year six, and then only in the
high-nitrogen plot. Changes in internal carbon-
and nitrogen-cycling rates and in net exchanges of
methane between soils and the atmosphere have
also been substantial. This result is important
as it demonstrates that nitrogen excess and
saturation triggers very fundamental changes in
a range of ecosystem processes.

Ecophysiological theory predicts that higher
concentrations of a limiting nutrient like nitro-
gen will lead, in turn, to higher rates of net pho-
tosynthesis and tree growth through a fertilizer
effect. Indeed, more tree growth has occurred
in the hardwood stand (originally nitrogen-
limited), with a 45-percent increase in wood
production. However, in the pine stand — where
high losses from leaching suggested that nitro-
gen saturation occurred very rapidly — wood pro-
duction over six years has been 28 percent lower
than that of the control plot. Combined with
even more dramatic growth declines and mor-
tality increases in conifer stands located in high
nitrogen-deposition regions of the world, these
results suggest a general decrease in the vigor
and wood production of conifer stands, and per-
haps even the onset of serious forest decline, in
response to nitrogen saturation.

Natural Disturbance Versus Human-Induced Stresses 39

The Soil-Warming Experiment

The rate of increases in global temperature that
are currently projected far exceed the changes
that occur naturally through glacial-interglacial
cycles. One consequence of global warming may
be changes in the rates of critical temperature-
dependent ecosystem processes, which in turn
control forest growth and health. Important soil
processes that may be expected to change with
increasing global temperatures include decom-
position, methane production, nitrogen miner-
alization and nitrification, and phosphorus
availability. Since many of these changes may
result in the release of greenhouse gases, soil
warming may itself exacerbate soil warming.

The soil-warming experiment at the Harvard
Forest was designed to assess the response of a
second-growth hardwood forest to elevated soil
temperatures. We paid particular attention to
the effects of warming on soil processes that
may alter ecosystem function, atmospheric
chemistry, and global climate. Eighteen 6-by-6-
meter plots were established in April 1991; each
was randomly assigned to one of three treat-
ments: heated plots, in which the average soil
temperature was raised 5 degrees Centigrade
above ambient using buried heating cables; dis-
turbance control plots, in which buried cables
were installed but received no electrical power,-
and undisturbed control plots.

Initial results from the first
growing season of the study indi-
cated that heating increased
emissions of carbon dioxide by
40 percent. Nitrogen cycling was
also affected dramatically, with a
doubling of mineralization rates
in the upper soil layers. During
the second and third growing
seasons, warming had a much
less dramatic effect on carbon
dioxide emissions but a sus-
tained dramatic effect was seen
in nitrogen mineralization,
which again doubled in rate.

The following scenario pro-
vides one interpretation of why
the rate of carbon dioxide emis-
sions in the heated plots relative
to that of control plots decreased
between the first and second

years relative to disturbance control plots,
whereas the rate of nitrogen mineralization con-
tinued to increase. There are two major pools of
organic matter in soils, a "fast" pool and a
"slow" pool. The fast pool contains material
with a high ratio of carbon to nitrogen (for
instance, recently fallen litter), whose decay
results in relatively large losses of carbon diox-
ide but a small net loss of nitrogen. In contrast,
the slow pool contains material with a low ratio
of carbon to nitrogen (such as partially decom-
posed and relatively stable humus), whose decay
results in a smaller loss of carbon dioxide and a
larger net loss of nitrogen.

Because elevated soil temperature increases
the rate of decay in the slow pool, increased
amounts of both carbon and nitrogen are
released. The nitrogen then becomes available
for uptake by plants. However, the carbon-to-
nitrogen ratio of living plant material is substan-
tially higher than that of organic matter in the
soil of the slow pool, and because of that, warm-
ing may lead to increased carbon storage in the
ecosystem without increased nitrogen storage.

The study highlights the importance of long-
term experiments while underscoring the
potential for complex changes resulting from
global climate change. Although the immediate
result of soil warming is carbon release — and

One of the eighteen 6-by-6-meter plots used in the soil-warming experiment
at the Harvard Forest. Heating coils placed below the soil surface raised the
average temperature by five degrees Centigrade and caused subtle changes
in the environment.

lOHN O'KEEFE

40 Arnoldia 1998 Summer

feedback on global change — this effect is tran-
sient. Over the longer term, the net release of
nitrogen might be a much more important effect
and might signal fundmental though subtle
shifts in ecosystem processes.

Comparison Between the Effects of Natural
Disturbances and Those of Novel Stresses

Which of these changes — "natural" disturbance
or climatic and chemical stress — is actually
most disruptive to the integrity of the commu-
nity and most likely to lead to long-term
changes in ecosystem function? Comparison of
results from the three experiments led to the
surprising conclusion that structural integrity —
that is, the actual appearance of a forest — is not
a good indicator of forest ecosystem function.
Whereas the blowdown site appeared severely
disturbed, fundamental internal processes were
not altered significantly, and the stand is on a
path to recovery of structure and function in
keeping with the cyclic pattern of disturbance
and regeneration in this forest type.

By contrast, the nitrogen-addition and soil-
warming plots are visually intact and apparently
healthy, yet the subtler measures of ecosystem
function suggest serious imbalances with
possible future implications for community
structure, internal ecosystem processes, and
exchanges with the global atmosphere.

In the nitrogen-addition plots, nitrogen losses
into the deeper soil are being induced and major
changes in the amounts of trace gases (carbon
dioxide, methane, nitrous oxide) released from
the soil to the atmosphere have occurred. In the
soil-warming plots, carbon dioxide exchanges
have become negative, nitrogen cycling has
increased dramatically, and nitrogen losses (as
nitrate) are increasing. In time, alterations in
chemical or physical environments caused by
these novel stresses will create changes in nitro-
gen concentrations and in the rates of carbon
and nitrogen cycling, which in turn will alter
ecosystem productivity.

We cannot predict what the ultimate trajec-
tory of these changes will be because there is no
historical analog for these experiments and
none of our present-day species evolved in an
environment that included these stresses. It is
reasonable to believe, however, that in the long

run system function will continue to be dis-
rupted more by these novel disturbances than
by natural disturbances. The plant-response
mechanisms seen in the hurricane experiment,
which have presumably evolved as a conse-
quence of natural selection for recovery from
natural, physical disturbance, may not exist for
situations where large quantities of nitrogen are
added to the soil because of human activity, or
soil becomes warmer very rapidly because of
climate change.

Several conclusions emerge from studies of
natural ecosystems and from our experimental
study of "natural" physical disturbance and
novel climatic and chemical stresses: (1) all eco-
systems are dynamic as a consequence of natu-
ral disturbance, natural environmental change,
and human impacts,- (2) as a consequence of
natural changes there are no static baseline con-
ditions for assessing current ecosystems or for
establishng correct goals for environmental
management; (3) ecosystems are incredibly
resilient, but the rate and novel character of
modern human disturbances raises the question
of whether they exceed this threshold of resil-
iency,- and (4) a comparison of hurricane distur-
bance, nitrogen deposition, and global change
(soil warming) suggests that physical appear-
ance is a poor indicator of ecosystem integrity
and that major and deleterious changes in eco-
system function may occur as a consequence of
these novel stresses.

David Foster is director of the Harvard Forest of Harvard
University. ]ohn Aber is director of the Complex
Systems Research Center, Institute for Study of Earth,
Oceans, & Space at the University of New Hampshire.
Jerry Melillo is co-director of the Ecosystem Center at
the Marine Biological Laboratory of Woods Hole.
Richard Bowden is assistant professor in the Department
of Environmental Science, Allegheny College. Fakhri
Bazzaz is H. H. Timken Professor of Science in the
Department of Organismic and Evolutionary- Biology of
Harvard University.

This research is reported in full in "Forest
Response to Disturbance and Anthropogenic Stress,"
published in BioScience (1997) 47(7): 437-445. This
summary also draws on "Ecological and conservation
insights from reconstructive studies of temperate
old-growth forests" by D. R. Foster, D. A. Orwig, and
I. S. McLachlan, published in TREE: Trends in Ecology
and Evolution (1996) 11(10): 419-424.

CASE STUDIES IN FOREST HISTORY AND ECOLOGY

Ecosystem Response to an Imported Pathogen:
The Hemlock Woolly Adelgid

David A. Orwig and David R. Foster

One of the provocative issues that have emerged
in the field of ecology over the past few decades
has concerned the question of whether indi-
vidual plant or animal species may play a criti-
cal role in controlling ecosystem processes and
determining major community characteris-
tics— that is, are there really "keystone" spe-
cies’ In an era when species are being driven
either locally or globally extinct, the possibility
that individual taxa may be crucial for main-
taining the integrity of ecosystems has become
a major impetus for the study of biodiversity.

This issue has been the focus of considerable
research in the northeastern United States in
particular, in large part because the region has
experienced a series of species eliminations or
reductions in numbers primarily as a conse-
quence of human activities. The decline of hem-
lock across its range 4,800 years ago due to an
insect pathogen, followed by a very gradual
recovery; the elimination of chestnut in this
century by a fungal blight introduced from east-
ern Asia; the outbreak of gypsy moth on oak,
aspen, and birch forests; and the purposeful
elimination or drastic reduction of animal popu-
lations— wolves, passenger pigeons, deer, bea-
vers, to name only a few — all have undoubtedly
had major consequences for forest, aquatic, and
wetland ecosystems across the Northeast,
although they remain largely unquantified.

Widespread, abundant, long-lived, and extraor-
dinarily shade-tolerant, eastern hemlock (Tsuga
canadensis) has always been classified as a
preeminent part of the "climax" forest that
existed before European settlement. Thanks to
the extremely dense shade and deep, acidic lit-
ter it produces, hemlock is able to determine
forest composition and create distinctive wild-

life habitats. Although it is currently found in
greatest abundance on rocky ridges or talus
slopes and in moist, cool ravines, hemlock
grows on a broad range of sites, and, in fact,
hemlock plays an important role in controlling
ecosystems all across New England, from the
rocky highlands through the mesic woodlands
and riparian areas, and into the wetlands.

Now, nearly 5,000 years after a pathogen deci-
mated hemlock populations across its entire
range, an introduced inseet, hemloek woolly
adelgid, or HWA (Adelges tsiigae), threatens
another catastrophe for hemlock, and at the same
time offers an opportunity to examine the eon-
sequences for the ecosystems it occupies. In con-
trast to the case of the chestnut blight, whose
effects we understand only partially, we hope to
learn from the case of HWA what happens to a
forest when its dominant species is lost.

It is important to point out that the HWA-
hemloek system behaves very differently from
other pathogen and pest systems. The damage
caused by HWA is unlike that of chestnut
blight, gypsy moth, dutch elm disease, or the
related balsam woolly adelgid (Adelges piceae)
in that HWA attacks overstory trees, saplings,
and seedlings alike, and therefore has the poten-
tial to eliminate hemlock from a site within a
few years. Moreover, hemlock trees lack the
ability to sprout or refoliate after defoliation;
consequently reestablishment can only occur
from the seedbank or from seed transported
from surviving trees. Hemlock seed typically
remains viable only one growing season,
although a few researchers have reported seed
viability for up to four years. It is believed that
on most sites, episodic establishment of hem-
lock occurs only infrequently, under unusual

42 Arnnldia 1998 Summer

conditions related to the nature of the seedhed,
moisture, and seed availability.

White-tailed deer often represent another
obstacle to regeneration; many studies have
shown that deer browse can severely reduce
hemlock seedling densities. Deer herd density
in Connecticut has more than doubled since
1980, resulting in high local population densi-
ties that change understory composition and
structure. All these problems raise concerns
about the long-term viability, stability, and
composition of hemlock ecosystems. Many
hemlock stands are scattered across the land-
scape on sites that have been protected from
intensive human and natural disturbance. If hem-

lock is eliminated from even a portion of these
sites, seed production will be eliminated or dras-
tically reduced across broad geographic areas. It
should he noted that it took eastern hemlock
one to two thousand years to recover following
the mid-Holocene decline 4,800 years ago.

In 1995 the Harvard Forest initiated a major
research effort consisting of stand, landscape,
and ecosystem components in central Connecti-
cut along a 370-mile transect that extends from
Long Island Sound to the Massachusetts border.
To document the damage already caused by
HWA since it arrived in 1985, we used aerial
photographs to map each hemlock stand greater
than three hectares (seven and a half acres). We
have compiled extensive data on forest
composition from 115 stands, which we
have overlain on a CIS (Geographic Infor-
mation Systems) map that shows other
biological features as well as soil types and
topographic characteristics. This map will
facilitate our analysis of the way HWA
spreads and help us determine whether the
damage patterns observed in the present
study are consistent across various types
of landscape.

We also began to examine in detail the
response of eight hemlock stands with dif-
fering levels of HWA infestation; the range
of damage found in this sample was in gen-
eral representative of conditions observed
in dozens of stands throughout south-
central Connecticut. Results after the sec-
ond year of analysis of the eight sample
stands showed continued deterioration of
hemlock, with annual mortality levels
ranging from 5 to 15 percent in stands that
had already lost 20 to 95 percent of all
overstory hemlock. All surviving hem-
locks except those in one isolated stand
are infested with HWA and have suffered
moderate to severe canopy damage. Conse-
quently, many of our study sites currently
have a high proportion of standing dead
snags or trees whose only remaining foli-
age is in the outer ends of upper branches.

In severely damaged stands, many dead
treetops and boles have snapped off during
recent windstorms, and deterioration in
the surviving trees has continued due to

This Tsuga canadensis branch is infested with HWA ovisacs,
located along the twigs at the base of the needles.

The Hemlock Woolly Adelgid 43

the presence of secondary organisms
such as hemlock borer, shoestring
fungi, and conk fungi. The conse-
quences of this continued mortality
will be substantial pulses of woody
debris, dramatic changes in the age,
structure, and composition of for-
ests, and altered wildlife habitat.

Light availability as well as above-
and below-ground space should
increase as HWA spreads across the
landscape. We have recently begun to
study how these substantial changes
in forest structure will affect the
timing, magnitude, and duration of
nitrogen cycling.

Our analysis of the effects of HWA
damage shows dramatic changes in
the understory microenvironment
and in the response of vegetation to
these changes. Particularly in stands
that previously had a high percentage
of hemlock in the overstory, black
birch (Betula lenta) seedlings have
established themselves prolifically,
encouraged by the increased amount
of light reaching the previously
shaded forest floor. The organic layer
in the soil under hemlock canopies
appears to be an ideal substrate for
germination of this wind-dispersed
species. Several researchers have
reported similar increases following
partial cutting or tree mortality due
to chestnut blight or windthrow. An
increased abundance of black birch is
also expected in less severely dam-
aged stands in the future, since it is
already present in the overstory and
produces copious numbers of seeds.

Seedlings of red maple and of several oak
species were also present in low densities in
most of our study sites; they may become more
numerous in the future. The overall scarcity of
hemlock seedlings in these forests suggests that
neither they nor seedbanks will be of much help
in hemlock reestablishment. Very few shrub
species have colonized these stands; exceptions
are bird-dispersed grape species (Vitus) and
Virginia creeper (Parthenocissus quinque folia).

Severe HWA damage on the Guilford, Connecticut, study site.
Note the dead hemlocks in the foreground and background,
adjacent to trees that retain sparse canopies. Hemlocks typically
die with their branch structures intact and gradually fall apart
over three to ten years.

Several opportunistic herbaceous species have
also become established in low abundance
throughout the study area, and invasion has
already begun of several exotic tree, shrub, and
herbaceous species that may increase and fur-
ther affect revegetation processes in the future.
Many factors, including pre-HWA site and soil
characteristics, hemlock mortality rates, herbi-
vore pressure, and the percentage of overstory
space occupied by hardwood species, will also

44 Arnoldia 1998 Summer

Black birch seedlings have established themselves prolifically under HWA-damaged hemlocks, encouraged by
the increased amount of light reaching the previously shaded forest floor.

play a role in determining the eventual compo-
sition of vegetation.

By documenting damage patterns and changes
in forest structure and composition resulting
from an introduced forest pest as they develop,
we have gained valuahle insight into the initial
recovery processes of forests when a dominant
species is removed. The results of this study and
direct observations in dozen of stands through-
out the state indicate that Connecticut forests
are being severely impacted by HWA. To date,
the rate and intensity of infestation is not attrib-
utable to any site factor or stand characteristic,
and there is no apparent impediment to the
widespread expansion of HWA and devastation
of eastern hemlock across its range.

Varying degrees of HWA infestation at the
stand level resulted in high hemlock mortality
rates, pulses of downed woody debris, and dra-
matic changes in microenvironment character-
istics due to overstory canopy gaps. These

structural changes have initiated rapid vegeta-
tion responses in understories typically devoid
of vegetation. In addition, our results suggest
that some sites may experience a complete
change in cover type from hemlock to forests
dominated by birch, oak, and maple species.
Under this scenario, forest composition and
structure at the landscape level would become
increasingly more homogenous. The outlook for
hemlock persistence in southern New England
forests is bleak. If HWA dispersal continues
unimpeded, dramatic reductions of hemlock
across broad geographical areas appear immi-
nent unless natural or introduced predators of
HWA are found.

David A. Orwig is a forest ecologist at the Harvard
Forest, where David R. Foster is the director. Their
research has heen published in full in "Forest Response
to the Introduced Hemlock Woolly Adelgid in Southern
New England, USA" in Journal of the Torrey Botanical
Society (1998) 125(1): 60-73.

V

- - Xi.’-

arnoldia

The Magazine of the Arnold Arboretum

U s.

DEC 1 7 1998

GRAY herbarium

amoldia

Volume 58 Number 3 1998

Amoldia (ISSN 004-2633; USPS 866-100) is
published quarterly by the Arnold Arboretum of
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Karen Madsen, Editor
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Editorial Committee
Phyllis Andersen
Robert E. Cook
Peter Del Tredici
Gary Roller
Stephen A. Spongherg
Kim E. Tripp

Copyright © 1998. The President and Fellows of
Harvard College

Page

2 In the Shadow of Red Cedar
Wade Davis

1 1 The First and Final Flowering of
Muriel's Bamboo
Peter Del Tredici

18 Nature Study Moves into the
Twenty-First Century
Candace L. Julyan

25 Native vs. Nonnative: A Reprise
Letters to the Editor

30 Native Plants: Another View
Harrison L. Flint

Front cover: Bigleaf maples (Acer macrophyllum)
on Washington State's Olympic Peninsula photo-
graphed by Graham Osborne.

Inside front cover: Branchlets and cones of
western red cedar (Thuja plicata). Photograph by
Peter Del Tredici.

Inside back cover: Bunchberry (Cornus canadensis)
and ferns in Skagit Valley, Washington. Photograph
by Graham Osborne.

Back cover: Western red cedar and hemlocks
(Tsuga heterophylla) in Mt. Rainier National Park,
Washington. Photograph by Graham Osborne.

;RAHAM DiBORNf

In the Shadow of Red Cedar

Wade Davis

In the shadow of red cedar, along a stream col-
ored hy salmon, in a place where plants draw
food from the air and small creatures living on
dew never touch the forest floor, it is difficult to
imagine a time when the coastal temperate
rainforests of North America did not exist.
Today, these immense and mysterious forests,
which in scale and wonder dwarf anything to be
found in the tropics, extend in a vast arc from
northern California 2,000 miles north and west
to the Copper River and the Gulf of Alaska.
Home to myriad species of plants and animals,
a constellation of life unique on earth, they
spread between sea and mountain peak, reach-
ing across and defying national boundaries as
they envelop all who live within their influence
in an unrivaled frontier of the spirit.

It is a world anchored in the south by giant
sequoias (Sequoiadendion giganteum), the
most massive of living beings, and coast red-
woods (Sequoia sempervirens) that soar 300 feet
above the fogbanks of Mendocino. In the north,
two trees flourish: western hemlock (Tsuga
heterophylla), with its delicate foliage and
finely furrowed bark; and Sitka spruce (Picea
sitchensis), most majestic of all, a stunningly
beautiful species with blue-green needles that
are salt tolerant and capable of extracting min-
erals and nutrients from sea spray. In between,
along the silent reaches of the midcoast of Brit-
ish Columbia, behind a protective veil of Sitka
spruce, rise enormous stands of Douglas fir
(Pseudotsuga menziesii). Intermingled with
hemlock and fir, growing wherever the land is
moist and the rains abundant, is perhaps the
most important denizen of the Pacific slope, the
western red cedar (Thuja plicata), the tree that
made possible the florescence of the great and
ancient cultures of the coast.

To walk through these forests in the depths of
winter, when the rain turns to mist and settles

softly on the moss, is to step back in time. Two
hundred million years ago vast coniferous for-
ests formed a mantle across the entire planet.
Dinosaurs evolved long supple necks to browse
high among their branches. Then evolution
took a great leap, and flowers were born. What
made them remarkable was a mechanism of
pollination and fertilization that changed the
course of life on earth. In the more primitive
conifers, the plant must produce the basic food
for the seed with no certainty that it will be fer-
tilized. In the flowering plants, by contrast, fer-
tilization itself sparks the creation of the seed's
food reserves. In other words, unlike the coni-
fers, the flowering plants make no investment
without the assurance that a viable seed will be
produced. As a result of this and other evolu-
tionary advances, the flowering plants came to
dominate the earth in an astonishingly short
time. Most conifers went extinct, and those that
survived retreated to the margins of the world,
where a small number of species maintained a
foothold by adapting to particularly harsh con-
ditions. Today, at a conservative estimate, there
are over 250,000 species of flowering plants. The
conifers have been reduced to a mere 700 spe-
cies, and in the tropics, the hotbed of evolution,
they have been almost completely displaced.

On all the earth, there is only one region of
any size and significance where, because of par-
ticular climatic conditions, the conifers retain
their former glory. Along the northwest coast of
North America the summers are hot and dry,
the winters cold and wet. Plants need water and
light to create food. Here in the summer there is
ample light for photosynthesis but not enough
water for most deciduous trees, except in low-
lying areas where broadleafed species such as
red alder (Alnus lubia), cottonwood (Populus
balsamifera ssp. thchocarpa), and vine maple
(Acer circinatum) flourish. In the winter, when

Western red cedar and hemlock in Stoltmann Wilderness, British Columbia.

4 Arnoldia 1998 Foil

both water and light arc sufficient, the low tem-
peratures cause the flowering plants to lose
their leaves and become dormant. The ever-
green conifers, by contrast, are able to grow
throughout the long winters, and since they use
water more efficiently than broadleafed plants,
they also thrive during the dry summer months.
The result is an ecosystem so rich and so pro-
ductive that the hiomass in the best sites is eas-
ily four times as great as that of any comparable
area of the tropics.

Indeed it is the scale and ahundance of the
coastal rainforests that overwhelm the visitor.
White pine (Finns strobus), the tallest tree of
the eastern deciduous forests, barely reaches
two hundred feet; in the coastal rainforests
there are thirteen species that grow higher, with
the redwoods reaching nearly four hundred
feet, taller than a twenty-five-story building.
Red cedars can be twenty feet or more across at
the base. The footprint of a Douglas fir would
crush a small cabin. The trunk of a western
hemlock, a miracle of biological engineering.

stores thousands of gallons of water and sup-
ports branches festooned with as many as 70
million needles, all capturing the light of the
sun. Spread out on the ground, the needles of a
single tree would create a photosynthetic sur-
face ten times the size of a football field.

These giant trees delight, but the real wonder
of the forest lies in the details, in the astonish-
ingly complex relationships; a pileated wood-
pecker living in the hollow of a snag, tiny
seabirds laying their eggs among the roots of an
ancient cedar, marbled murrelets nesting in a
depression in the moss in the fork of a canopy
tree, rufous hummingbirds returning each
spring, their migrations timed to coincide with
the flowering of salmonberries iRubus
spectabilis). In forest streams dwell frogs with
tails and lungless salamanders that live by
absorbing oxygen through their skin. Strange
amphibians, they lay their eggs not in water but
on land, in moist debris and fallen logs.

Invertebrate life is remarkably diverse. The
first survey to explore systematically the forest

C'.RAHAM OSBORNE

Red Cedar 5

Mossy branches of bigleaf maple, Acer macrophyllum, Olympic
Peninsula, Washington.

canopy in the Carmanah Valley of Vancouver
Island yielded 15,000 species, a third of the
invertebrates know^n to exist in all of Canada.

Among the survey's collections were 500 spe-
cies previously unknown to science. Life is
equally rich and abundant on the forest floor.

There are 12 species of slugs, slimy herbivores
that in some areas account for as much as sev-
enty percent of the animal biomass. A square
meter of soil may support 2,000 earthworms,

40,000 insects, 120,000 mites, 120,000,000
nematodes, and millions upon millions of proto-
zoa and bacteria, all alive, moving through the

earth, feeding, digesting, repro-
ducing, and dying.

None of these creatures, of
course, lives in isolation. In
nature, no event stands alone.
Every biological process, each
chemical reaction, leads to the
unfolding of other possibilities
for life. Tracking these strands
through an ecosystem is as com-
plex as untangling the distant
threads of memory from a myth.
For years, even as industrial log-
ging created clearcuts the size
of small nations, the coastal
rainforests were among the least
studied ecosystems on the planet.
Only within the last decade or
two have biologists begun to
understand and chart the dynamic
forces and complex ecological
relationships that allow these
magnificent forests to exist.

One begins with wind and rain,
the open expanse of the Pacific
and the steep escarpment of
mountains that makes possible
the constant cycling of water
between land and sea. Autumn
rains last until those of spring,
and months pass without a sign of
the sun. Sometimes the rain falls
as mist, and moisture is raked
from the air by the canopy of the
forest. At other times the storms
are torrential, and daily precipita-
tion is measured in inches. The
rains draw nutrients from the soil, carrying
vital food into rivers and streams that fall away
to the sea and support the greatest coastal
marine diversity on earth. In the estuaries and
tidal flats of British Columbia, in shallows that
merge with the wetlands, are six hundred types
of seaweed, seventy species of sea stars. Farther
offshore, vast, underwater kelp forests shelter
hundreds of forms of life, which in turn support
a food chain that reaches into the sky to nour-
ish dozens of species of seabirds.

The land provides for life in the sea, but the
sea in turn nurtures the land. Birds deposit

6 Arnoldia 1998 Fall

excrement in the moss, yielding tons of nitrogen
and phosphorus that are washed into the soil by
winter rains. Salmon return by the millions to
their native streams, providing food for eagles
and ravens, grizzly and black bears, killer
whales, river otters, and more than twenty other
mammals of the sea and forest. Their journey
complete, the sockeye and coho, chinooks,
chums, and pinks drift downstream in death and
are slowly absorbed back into the nutrient cycle
of life. In the end there is no separation between
forest and ocean, between the creatures of the
land and those of the sea. Every living thing on
the raincoast ultimately responds to the same
ecological rhythm. All are interdependent..

The plants that dwell on land nevertheless
face particular challenges, especially that of
securing nutrients from thin soils leached by
rain throughout much of the year. The tangle of
ecological adaptations that has evolved in
response is nothing short of miraculous. As
much as a fifth of the biomass in the foliage of
an old-growth Douglas fir, for example, is an
epiphytic lichen, Lobaiia oiegana, which fixes
nitrogen directly from the air and passes it into
the ecosystem. The needles of Sitka spruce
absorb phosphorus, calcium, and magnesium,
and their high rate of transpiration releases
moisture to the canopy, allowing the lichens
to flourish.

On the forest floor thick mats of sphagnum
and other mosses filter rainwater and protect
the mycelia of hundreds of species of fungi;
these elements form one of the richest mush-
room floras on earth. Mycelia are the vegetative
phase of a fungus, small hairlike filaments that
spread through the organic layer at the surface
of the soil, absorbing food and precipitating
decay. A mushroom is simply the fruiting
structure, the reproductive body. As the myce-
lia grow, they constantly encounter tree roots.
If the species combination is the right one, a
remarkable biological event unfolds. Fungus
and tree come together to form mycorrhizae, a
symbiotic partnership that allows both to ben-
efit. The tree provides the fungus with sugars
created from sunlight. The mycelia in turn
enhance the tree's ability to absorb nutrients
and water from the soil. They also produce
growth-regulating chemicals that promote the

production of new roots and enhance the
immune system. Without this union, no tree
could thrive. Western hemlocks are so depen-
dent on mycorrhizal fungi that their roots barely
pierce the surface of the earth, even as their
trunks soar into the canopy.

The story only gets better. All life requires
nitrogen for the creation of proteins. Nitrates, a
basic source, are virtually absent from the
acidic, heavily leached soils of the rainforest.
The mycorrhizae, however, contain not only
nitrogen-fixing bacteria that produce this vital
raw material but also a yeast culture that pro-
motes the growth of both the bacteria and the
fungus. There are scores of different mycor-
rhizae— the roots of a single Douglas fir may
have as many as forty types — and, like any other
form of life, the fungus must compete, repro-
duce, and find a means to disperse its spore. The
fruiting body in many cases is an underground
mushroom or a truffle. When mature, it emits
a pungent odor that seeps through the soil to
attract rodents, flying squirrels, and red-backed
voles, delicate creatures that live exclusively on
a refined diet of truffles. As the voles move
about the forest, they scatter droppings, neat
little bundles of feces that contain yeast culture,
fungal spores, and nitrogen-fixing bacteria — in
short, all that is required to inoculate roots and
prompt the creation of new mycorrhizae.

Fungi bring life to the forest both by their
ability to draw nutrients to the living and by
their capacity to transform the dead. In old-
growth forests twenty percent of the biomass —
as much as six hundred tons per hectare — is
retained in fallen debris and snags. There is as
much nutrition on the ground as there is within
it. The moss on the forest floor is so dense that
virtually all seedlings sprout from the surface of
rotting stumps and logs, which may take several
hundred years to decay.

When a tree falls in the forest, it is immedi-
ately attacked by fungi and a multitude of
insects. The wood provides a solid diet of carbo-
hydrates. To secure proteins and other nutri-
ents, the fungi deploy natural antibiotics to kill
nitrogen-fixing bacteria. Chemical attractants
emitted by the fungi draw in other prey, such as
nematode worms, which are dispatched with
exploding poison sacs and an astonishing arse-

Red Cedar 1

Western red cedar near Port Angeles. Washington.

8 Arnoldia 1998 Fall

Douglas firs at sunrise.

nal of microscopic weapons. The assault on the
log comes from many quarters. Certain insects,
incapable of digesting wood directly, exploit
fungi to do the work. Ambrosia beetles, for
example, deposit fungal spores in tunnels bored
into the wood. After the spores germinate, the
tiny insects cultivate the mushrooms on minia-
ture farms that flourish in the dark.

In time other creatures appear — mites and
termites, carpenter ants that chew long galleries
in the wood and establish captive colonies of
aphids that produce honeydew from the sap of
plants. Eventually, as the log progresses through
various stages of decay, other scavengers join

the fray, including those that consume
white cellulose, turning wood blood-
red and reducing the heartwood to dust.
An inch of soil may take a thousand
years to accumulate. Organic debris
may persist for centuries. Dead trees are
the life of the forest, but their potential
is realized only slowly and with great
patience.

This observation leads to perhaps the
most extraordinary mystery of all. Lush
and astonishingly prolific, the coastal
temperate rainforests are richer in their
capacity to produce the raw material of
life than any other terrestrial ecosystem
on earth. The generation of this
immense natural wealth is made pos-
sible by a vast array of biological inter-
actions so complex and sophisticated
as to suggest an evolutionary lineage
drifting back to the dawn of time. Yet
all evidence indicates that these forests
emerged only within the last few thou-
sand years. In aspect and species com-
position they may invoke the great
coniferous forests of the distant geo-
logic past, but as a discrete and evolving
ecosystem the coastal temperate
rainforests are still wet with the inno-
cence of birth.

Some twenty thousand years ago,
what is today British Columbia was a
place of turmoil and ice. The land was
young, unstable, given to explosive
eruptions that burst over the shore. A
glacial sheet more than 6,000 feet deep
covered the interior of the province, forging
mountains and grinding away valleys as it
moved over the land, determining for all time
the fate of rivers. On the coast, giant tongues of
ice carved deep fjords beneath the sea. The sea
levels fell by 300 feet, and the sheer weight of
ice depressed the shoreline to some 750 feet
below its current level. Fourteen thousand years
ago, an instant in geologic time, the ice began to
melt, and the glaciers retreated for the last time.
The ocean invaded the shore, inundating coastal
valleys and islands. But the land, freed at last of
the weight of eons, literally sprang up. Within a
mere one thousand years, the water drained

Red Cedar 9

back into the sea, and the coastline became
established more or less as it is today.

Only in the wake of these staggering geologi-
cal events did the forests come into being. At
first the land was dry and cold, an open land-
scape of aspen and lodgepole pine (Pinus
contOTta). Around ten thousand years ago, even
as the first humans appeared on the coast, the
air became more moist and Douglas fir slowly
began to displace the pine. Sitka spruce flour-
ished, though hemlock and red cedar remained
rare. Gradually the climate became warmer,
with long seasons without frost. As more and
more rain fell, endless banks of clouds sheltered
the trees from the radiant sun. Western hem-
lock and red cedar expanded their hold on the
south coast, working their way north at the
expense of both fir and Sitka spruce.

For the first people of the raincoast, this eco-
logical transition became an image from the
dawn of time, a memory of an era when Raven
slipped from the shadow of cedar to steal sun-
light and cast the moon and stars into the heav-
ens. Mythology enshrined natural history, for
it was the diffusion of red cedar that allowed
the great cultures of the Pacific Northwest to
emerge. The nomadic hunters and gatherers
who for centuries had drifted with the seas
along the western shores of North America
were highly adaptive, capable of taking advan-
tage of every new opportunity for life. Although
humans had inhabited the coast for at least five
thousand years, specialized tools first appear in
the archaeological record around 3000 B.C.,
roughly the period when red cedar came into its
present dominance in the forests. Over the next
millennium, a dramatic shift in technology and
culture occurred. Large cedar structures were
in use a thousand years before the Christian era.
A highly distinct art form developed by 500 B.C.
Stone mauls and wooden wedges, obsidian
blades and shells honed to a razor's edge allowed
the highly durable wood to be worked into
an astonishing array of objects, which in turn
expanded the potential of the environment.

^ ^

In Oregon and Washington only ten percent of
the original coastal rainforest remains. In Cali-
fornia only four percent of the redwoods have
been set aside. In British Columbia, roughly

sixty percent has been logged, largely since
1950. In the last two decades, over half of all
timber ever extracted from the public forests of
British Columbia has been taken. At current
rates of harvest, roughly 1.5 square miles of old
growth per day, the next twenty years will see
the destruction of every unprotected valley of
ancient rainforest in the province.

In truth, no one really knows what will happen
to these lands once they are logged. Forests are
extraordinarily complex ecosystems. Biologists
have yet to identify all of the species, let alone
understand the relationships among them.
Although we speak with unbridled confidence
of our ability to reproduce the ecological condi-
tions of a forest and to grow wood indefinitely,
there is no place on earth that is currently cutting
a fourth generation of timber on an industrial
scale. The more imprecise a science, the more
dogmatically its proponents cling to their ability
to anticipate and predict phenomena.

Forestry as traditionally practiced in the
Pacific Northwest is less a science than an ide-
ology, a set of ideas reflecting not empirical
truths, but the social needs and aspirations of
a closed group of professionals with a vested
interest in validating its practices and existence.
The very language of the discipline is disingenu-
ous, as if conceived to mislead. The "annual
allowable cut" is not a limit never to be
exceeded but a quota to be met. The "fall down
effect," the planned decline in timber produc-
tion as the old growth is depleted, is promoted
as if it were a natural phenomenon when it is
in fact a stunning admission that the forests
have been drastically overcut every year since
modern forestry was implemented in the 1940s.
"Multiple-use forestry" — which implies that
the forests are managed for a variety of purposes,
including recreation, tourism, and wildlife —
begins with a clearcut. Old growth is "har-
vested," though it was never planted and no one
expects it to grow back. Ancient forests are
"decadent" and "overmature," when by any
ecological definition they are at their richest
and most biologically diverse state.

The most misleading of these terms is "sus-
tained yield," for it has led the public to believe
that the trees are growing back as fast as they
are being cut. But they are not. In British

10 Arnoldia 1998 Fall

Columbia alone there are 8.7 million acres of
insufficiently restocked lands. We continue to
cut at a rate of 650,000 acres per year. Every year
2.5 million logging-truck loads roll down the
highways of the province. Lined up bumper to
bumper, they would encircle the earth twice. In
practice, sustained-yield forestry remains an
untested hypothesis: after three generations we
are still cutting into our biological capital, the
irreplaceable old-growth forests. As a scientific
concept, sustained yield loses all relevance
when applied to an ecological situation the
basic parameters of which remain unknown. At
best, sustained yield is a theoretical possibility;
at worst, a semantic sleight of hand, intended
only to deceive.

Anyone who has flown over Vancouver
Island, or seen the endless clearcuts of the inte-
rior of the province, grows wary of the rhetoric
and empty promises of the forest industry. Fish-
ermen and women become skeptical when they
learn that logging has driven 142 salmon stocks
to extinction and left 624 others on the brink.
Timber for British Columbia mills now comes
from Manitoba. Truck drivers from Quesnel, a
pulp-and-paper town in the center of the prov-
ince, haul loads hundreds of miles south from
Yukon. Just one of the clearcuts southeast of
Prince George covers five hundred square kilo-
meters, five times the area of the city of
Toronto. This, after sixty years of official com-
mitment to sustained-yield forestry. The lament
of the old-time foresters — that if only the public
understood, it would appreciate what we do —
falls flat. The public understands but does not
like what it sees.

Fortunately, this orthodoxy is now being
challenged. Many in the Pacific Northwest,
including the best and brightest of professional
foresters, recognize the need to move beyond, to
an era in which resource decisions are truly
based on ecological imperatives, in which the
goal of economic sustainability is transformed
from a cliche into an article of faith. To make
this transition will not he easy, and it will
involve much more than tinkering with the
edges of an industry that generates $15.9 billion
a year in the province of British Columbia alone.
Dispatching delegations to Europe to reassure
customers, or devising new regulations that if

implemented may mitigate some of the worst
ecological impacts, will neither restore the
public's confidence and trust nor address
the underlying challenge of transforming
the economy.

Any worker who has wielded a saw or ripped
logs from a setting knows that in the end it all
comes down to production. The enormous
wealth generated over the last fifty years has
been possible only because we have been willing
to indulge egregious practices in the woods that
have little to do with the actual promise of for-
estry. Spreading clearcuts ever deeper into the
hinterland is a policy of the past, crude and
anachronistic, certain to lead to a dramatic
decline in the forestry sector and to bitterness
and disappointment in the communities that
rely upon the forests for both spiritual and
material well-being. Revitalizing cutover lands
with vibrant tree plantations, implementing
intensive silviculture to increase yields, estab-
lishing the finest model of forest management
on a finite land base — these are initiatives that
will both allow communities to prosper and
enable them to fulfill a moral obligation to leave
to the future as healthy an environment as the
one they inherited.

There is no better place to pursue a new way
of thinking than in the temperate rainforests of
the coast. At the moment, less than six percent
has been protected; the remainder is slated to be
logged. If anything, this ratio should be reversed.
We live at the edge of the clearcut; our hands
will determine the fate of these forests. If we do
nothing, they will be lost within our lifetimes,
and we will he left to explain our inaction. If we
preserve these ancient forests, they will stand
for all generations and for all time as symbols of
the geography of hope. They are called old
growth not because they are frail but because
they shelter all of our history and embrace all of
our dreams.

Wade Davis is an ethnobotanist and prolific writer.
This article is excerpted from his most recent book,
Shadows in the Sun: Travels to Landscapes of Spirit and
Desire, published by Island Press/Shearwater Books
(1.800.828.1302 or www.islandpress.org).

Photographer Graham Osborne specializes in alpine
and coast subjects. The photographs in this article and
on the covers have been published in his book Rainforest.
published by Chelsea Green, Vermont.

The First and Final Flowering of Muriel's Bamboo

Peter Del Tredici

Regular readers of Ainoldia can appre-
ciate the many satisfactions that come
from working at the Arnold Arbore-
tum, with its endless opportunities
for studying plants. Even after twenty
years of daily contact, there's always
something new and exciting. Some
days it is the first flowers on a recently
planted specimen,- on others, it is
stumbling, sometimes quite literally,
across an amazing old plant never
before noticed. The highlight of the
1998 season was definitely the discov-
ery of flowers on Muriel's bamboo,
Fargesia muiielae, which appeared at
the Arnold Arboretum for the first —
and last — time.

Muriel’s bamboo, Fargesia murielae, in the full flush of its spring growth.

The Flowering

Fargesia murielae is native to the
mountains of central China, where it
grows at elevations between two and
three thousand meters. The species is
one of the principal foods of the giant
panda bear and arguably one of the
most ornamental of the hardy species
of bamboo. Its graceful, arching stems
reach two to three meters in height
and add a measure of exotic elegance
to any garden. As a clump-forming
species it expands slowly, in stark con-
trast to bamboos that spread by long,
underground stems — the "running
bamboos" — which are often the bane of unwary
gardeners. Experienced bamboo growers are uni-
versal in their praise of Fargesia murielae, not
only for the above-mentioned traits, hut also
because Muriel's bamboo is among the hardiest
of the entire family, growing well in USDA zone
5 and, with protection, into zone 4.

For all of its attractiveness, however, the
most interesting feature of Muriel's bamboo is

its monocarpic life cycle — it flowers once in its
life and then dies. Gardeners are used to seeing
sunflowers germinate, grow, and die in a single
season, and foxgloves die after two years, but
the idea of a plant flowering after eighty to one
hundred years and then dying seems more than
a little strange. And strange indeed it is, being
found only among the "woody" monocots, such
as the well-known century plant [Agave sp.).

12 Arnoldia 1998 Fall

a few genera of palms (most notably in the
genus Corypha], and an Andean bromeliad of
tree-sized proportions {Puya raymondii], which
tend to come into flower when they reach a
critical size.'

Monocarpic bamboos are unique even among
this unusual group because they do not flower
according to their size, but according to a prede-
termined maturation cycle, the length of which
appears to he genetically fixed for each species.^
The eighty-to-one-hundred-year flowering cycle
of Muriel's bamboo, while certainly not the
longest on record, is among the most widely
known and well documented. Indeed, it was the
widespread flowering and subsequent death of
the umbrella bamboo in China in 1971, along
with that of several closely related species, that
created worldwide concern about the survival
of the giant panda. The panda population in
central China, it was found, had become overly
dependent on the high-elevation species of
Fargesia after bamhoo species growing at lower
elevations were eliminated by land clearance for
agriculture.^

Even more remarkable than their long flower-
ing cycle, many hamhoos are also synchronous
in their flowering behavior. This term refers to
the tendency of most or all of the individuals of
a given species to come into flower at more or
less the same time. This unusual behavior has
led some authors to postulate that flowering in
these bamboos is controlled not by climatic fac-
tors but by some sort of internal clock. In real-
ity, however, the synchronicity is less precise
than generally believed, particularly when
plants in their native habitat are compared with
same-aged cohorts in cultivation that have been
repeatedly propagated by division. ■* It may be
propagation by subdivision that affects culti-
vated hamhoos, but in any case their flowering
cycle occurs as much as twenty years later.

While many authors have speculated on the
possible evolutionary and ecological signifi-
cance of the monocarpic habit in bamboos,
nothing has been proved. One theory, proposed
by Daniel Janzen,® is that the long delay in flow-

ering is a strategy for preventing a buildup of
predators that would feed on the highly nutri-
tious seeds if they were produced on a predict-
able schedule. However, this idea does not
explain why the flowering intervals of many
bamboos greatly exceed the lifespans of most
animals that would feed on their seeds. More
likely, the real reason is inextricably embedded
in a complex matrix of physiological, ecological,
and evolutionary factors.

The Introduction

The history of the plant's introduction into cul-
tivation in the West, like that of so many other
plants, is cloaked in mystery and confusion. It
was first collected hy the Arnold Arboretum's
most famous plant collector, E. H. Wilson, who
assigned it number 1462. The Arboretum has
most of Wilson's field hooks in its archives, and
those for his first Arboretum expedition, from
February 1907 through April 1909, contains the
following entry: "1462. Bamhoo, 12 ft., stems

golden, thickets, 7000-9000 ft. Fang. Plants,

Unfortunately, the last three words of

the passage are unintelligible, but one impor-
tant piece of information is unequivocal: living
plants, along with the usual herbarium speci-
mens, were part of this collection.

With the help of Alfred Rehder, Wilson
reworked his field notes and published them in
Plantae WilsonianaeF a work in three volumes
that appeared in sequence in 1913, 1916, and
1917. The reference to the umbrella bamboo
occurs on page 64 of volume II:

Aiundinaiia sp. Western Hupeh: Fang Hsien,
uplands, alt. 2000-3000 m., April 17, 1907
(No. 1462; 2-4 m. tall, stems golden). Without
flowers. This plant is in cultivation. It forms on
the mountains of north-western Hupeh dense
thickets and with its clear golden slender stems
is one of the most beautiful of Chinese Bamboos.

A picture will be found under No. 0111 of the
collection of my photographs. E.H.W."*

The photograph that Wilson referred to is
found in a bound volume entitled "Arnold Arbo-
retum Second Expedition to China: 1910-191 1.

\ ^ ' r

f .

ft * I _ I ,

ARCHIVES OF THE ARNOLD ARBORETUM

Muriel’s Bamboo 13

Photographs by E. H. Wilson." Photograph
#0111 clearly shows a clump-forming bamboo
growing below a group of fir trees (Abies
fargesiij. According to the notes on the accom-
panying label, the photograph was taken on June
19, 1910, and the plant, Wilson's #1462, is seen
growing "behind Fang Hsien" at an altitude of
8,000 feet. Wilson's diary for this day includes
the following entry:

The rain had ceased when we woke at 5 am ik
though dark mist obscured everything from a
hundred yards above St around us I prophesied a
fine day. It remained fine for about two hours St

then commenced to rain steadily. It increased as
the day advanced St we had a line soake. All were
soon drenched to the skin St everything became
sodden. We hurried on as fast as possible St
reached the head of the pass at 10 am . . . Much
of the Bamboo has been burned and cut away
from the path which is considerable improved
since our last visit . . . This bamboo is the hand-
somest I know with Its bright golden yellow
culms some 10-15 ft high shrubs and with arch-
ing plume. It must be very hardy lor the climate
here is very rigorous. Patches of original forest
remain here and there St especially near water-
course silver fir S. many Birch with willows
and Rhod. are practically the
sole constituents.

The final reference to bam-
boo #1462 in the Arboretum
archives comes from an un-
dated notebook in Wilson's
own handwriting entitled:
"Numerical list of seeds [no.
1-1474, 4000-4462], collected
on his Arnold Arboretum expe-
ditions to eastern Asia, 1907-
09, 1910, which were planted
in the arboretum nurseries."®
Under #1462, a single bamboo
plant is listed as being located
in the "Greenhouse St Frames"
area of the nurseries. Unfortu-
nately, the Arboretum's perma-
nent records of plants growing
on the grounds do not contain
any mention of #1462, strongly
suggesting that the plant was
never cultivated out-of-doors.

Fargesia murielae photographed by
E. H. Wilson in its native habitat,
Fang Hsien. China, at 8,000 feet.
The plants are ten to fifteen feet
high with yellow culms. Abies
fargesia stand in the background.
Below is Wilson’s field book entry
for collection #1462.

(T ^ 'O'

f ' J I.' ^ 1 ^ I- - / D f f I /I * »

1 4 Arnoldia 1 998 Fall

The first scientific description of Wilson's
#1462 did not appear until 1920, in an article in
Kew Bulletin of Miscellaneous Information^^
under the name Arundinaria murielae Gamble.
In the notes following 1. S. Gamble's Latin de-
scription, W. 1. Bean, Kew horticulturist, noted
that, "By Mr. Wilson's special wish the species
is dedicated to his daughter, Muriel Wilson."
Bean went on to detail the plant's history:

This Bamboo was presented to Kew from the
Arnold Arboretum in the autumn of 1913. A
single plant came in a pot, and this was divided
up into about half a dozen pieces, which were
repotted and grown for a few months in a green-
house. They were then planted out in the collec-
tion of Bamboos near the Rhododendron Dell
where they have grown luxuriantly and promise
to be as ornamental as any hardy species. They
are at present (October 1920| about 8 ft. high
forming dense masses of culms, the outer ones of
which arch outwards towards the top and give
the plants a very graceful appearance . . . On the
whole A. murielae is a distinct and most attrac-
tive addition to hardy bamboos.

At the Royal Botanic Garden, Kew, the only
record of Wilson's #1462 is in the accession
books, which noted its arrival on December
12, 1913.

Wilson's only other reference to #1462 is in A
Naturalist in Western China, published in Lon-
don in 1913 and New York in 1914. On page 49
he describes the vegetation behind Fang Xian by
paraphrasing his journal entry of June 19, 1910:

The summit is of hard limestone with rare
outcroppings of red sandstone. Stunted wind-
swept Silver Fir and various kinds of Currant
extend to the summit. Rhododendron and a
dwarf Juniper (J. squamata) are also common.
The descent was through woods of Birch and
Bamboo to an open, grassy, scrub-clad, sloping
moorland, through which a considerable torrent
flows. The Bamhoo, so common hereabouts, is
very beautiful, forming clumps 3 to 10 feet
through. The culms are 5 to 12 feet tall, golden
yellow, with dark, feathery foliage; the young
culms have broad sheathing bracts protecting
the branchlets. Taken all in all, this is the hand-
somest Bamboo 1 have seen.*

The footnote at the bottom of the page reads:
"In 1910 1 successfully introduced it into culti-
vation." In the revised edition of the book, pub-

lished in 1929 under a new title, China, Mother
of Gardens, Wilson makes clear that the name-
less bamboo mentioned in the 1913 edition was
collection #1462 by removing the footnote and
adding the following to the end of the above-
quoted paragraph: "In 1910, 1 successfully intro-
duced It into cultivation. It has been named
Arundinaria Murielae in compliment to my
daughter.""

From all this information, it appears that only
one plant of Wilson's #1462, collected on May
17, 1907, survived the long journey from Fang
Xian in China to the Arnold Arboretum, where
it was observed growing in the greenhouse in
1910. In December 1913, without ever being
cultivated out-of-doors here, the plant was sent
to Kew Gardens where it was divided — it must
have been quite large — and planted out in the
bamboo collection. Although Fargesia murielae
was widely distributed throughout Europe dur-
ing the first part of the century, the Arnold
Arboretum did not get another plant until
1960, when the U.S. National Arboretum in
Washington, D.C., sent one (under the name
Sinarundinaria murielae] that had been
imported in 1959 from the Royal Moerheim
Nurseries in Dedemsvaart, Holland."

Flowers at Last

The first flowers of Fargesia murielae in the
West appeared in Denmark in 1975." While
these plants were clearly representative of the
species, it is not certain that they were part of
Wilson's #1462 clone. The plants were said to be
smaller than Wilson's, and they came into
bloom several years earlier than plants known
to be divisions of #1462.

While the origin of the Danish plants will
never be determined with certainty, the fact
remains that in 1998 the flowering of known
clones of Wilson's Fargesia murielae appears to
be virtually complete, more than ninety years
after it was collected from the wild. Some of the
plants of #1462 have produced seed, but it is
important to remember that they are the result
of self-pollination, and as such they are likely
to suffer from the deleterious effects of inbreed-
ing depression. Only by re-collecting the species
in central China — from seedlings that germi-
nated following the widespread flowering that

Muriel’s Bamboo 15

History of Fargesia murielae in the West

1892: The French missionary P. Farges collects a herbarium specimen of an unknown
flowering bamboo in Szechuan Province, China. In 1893, the French taxonomist A. Franchet
assigns it to a new genus, Fargesia, with the specific name spathaceaF'^

17 May 1907: On his first expedition to China for the Arnold Arboretum, E. H. Wilson collects
plants and three sterile herbarium specimens of an unknown bamboo at Fang Xian, Hubei,
under collection #1462.

[1910]: Wilson makes note of a single plant from his collection #1462 growing in the "green-
houses and frames" area of the Arnold Arboretum.

10 June 1910: On his second Arboretum expedition to China, Wilson revisits Fang Xian and
photographs #1462, labelling the photograph #0111.

12 December 1913: One plant of Wilson's #1462 is received by Kew Gardens from the Arnold
Arboretum. The plant is divided into six pieces that are planted out in the bamboo area.

1916: Wilson labels #1462 as Arundinaria sp. in volume II of Plantae Wilsonianae, but lists
the wrong collection date.

1920: Wilson's #1462 is given the name Arundinaria murielae by }. S. Gamble.

1935: T. Nakai of Japan reclassifies Arundinaria murielae as Sinarundinaria murielae.

23 December 1959: U.S. National Arboretum botanist F. Meyer arranges for the importation
of plants of Sinarundinaria murielae (PI #262266) from the Royal Moerheim Nurseries,
Dedemsvaart, Holland. The plants are probably divisions of Wilson's #1462. One of them is
received by the Arnold Arboretum on 8 November 1960, under accession #1239-60.

1975: Plants of Sinarundinaria murielae in Denmark, possibly divisions of Wilson's #1462,
come into flower.

1979: Based on the flowering specimens of the Danish plants, T. Soderstrom proposes the name
Thamnocalamus spathaceus, for the umbrella bamboo. Based on the same specimens, other
botanists argue that the species should be classified as either Fargesia murielae (Gamble) or
F. spathacea (Franchet).

1988: At Kew Gardens, the original plants of Wilson's #1462 come into flower for the first time.

1995: C. Stapleton makes the case for preserving the name Fargesia murielae, but proposes
correcting the spelling of the specific to murieliae.^^

May 1998: Arnold Arboretum plants of Fargesia murielae, received from the U.S. National
Arboretum in 1960, come into flower for the first time.

PETER DEL TREDICI

16 Arnoldici 1998 Fall

The flowers of Fargesia murielae are inconspicuous

occurred there during the 1970s— can we hope
to obtain material comparable in quality to the
original Wilson #1462.

The story of the introduction of Muriel's bam-
boo is typical of the interplay between meticu-
lousness and confusion that often surrounds the
introduction of a new plant. That we can follow
the Fargesia story as well as we can hears wit-
ness to the care and effort that the Arnold Arbo-
retum in general, and Wilson in particular, put
into the process of collection and documenta-
tion. The story illustrates another point as well:
the importance of sharing plants among botani-
cal gardens. Kew Gardens, and especially its
horticulturist W. I. Bean, deserve credit for
propagating and eventually distributing the
plant throughout Europe. Distributing rare
plants is an act both of generosity and of self-
preservation; if you have two plants and give
one away, you can get it hack when you lose the
one you kept. Such losses happen frequently.

but the tradition of sharing plants provides an
important safety net that greatly increases the
chances of successful introduction. Given the
rate at which the forests of the world are disap-
pearing, failure to thoroughly document collec-
tions— and to share them — can represent the
loss of a resource that can never be recaptured.

Endnotes

’ S. A. Renvoize, Thamnocalamus spathaceus and its
hundred-year flowering cycle. Kew Magazine (1991)
8(4): 185-194.

^ E. I. Fortanier and R. H. Jonkers, Juvenility and
maturity of plants as influenced by the ontogenetical
and physiological aging. Acta Horticulturae (1976)
56: 37-44.

^ G. B. Shaller, ). Hu, W. Pan, and J. Zhu, The Giant
Pandas of Wolong (Chicago: University of Chicago
Press, 1985).

* Renvoize, op cit.

^ D. lansen. Why bamboos wait so long to flower. Ann.
Rev. Ecol. Syst. (1976) 7: 347-391.

^ E. H. Wilson, AA Manuscript #39526: First
expedition for Arnold Arboretum; Feb. 1907-Sept.
1909; collecting numbers 1-1474 (undated).

^ C. S. Sargent, ed., Plantae Wilsonianae 11 (Arnold
Arboretum, 1916), 64.

® Wilson's diary entry for April 17, 1907, makes no
mention of any bamboo, but when 1 checked the
herbarium specimens that document #1462, I found
all of them dated "17/5/07" in Wilson's handwriting.
In the absence of any journal for the month of May
1907, this discrepancy in dates was resolved by
checking Wilson's other herbarium specimens
collected at Fang Xian. According to former
Arboretum director R. A. Howard, in his 1980 article
"E. H. Wilson as Botanist" (part 1, Ainoldia 40(3):
102-138; part 11, 40(4): 154-193), the Fang Xian
material all had collection dates in May 1907. This
clearly suggests that the date of April 17 published m
Plantae Wilsonianae is an error, and that May 17,
1907, noted on the specimen, was the actual date for
the collection of Fargesia murielae.

’ E. H. Wilson, AA Manuscript #3961 1: Numerical list
of seeds |no. 1-1474, 4000-4462], collected on his
Arnold Arboretum expeditions to eastern Asia, 1907-
08, 1910, which were planted in the arboretum
nurseries (undated, probably 1910-1911).

J. S. Gamble, in Anon., Decades Kewenses:
Plantarum novarum in Herbaria Horti Regii

Tne Arnola ArLoretum

L L

NEWS

Field Studies Are Inspired by the Work of Volunteers

Diane Syversou
Manager of School Programs

A fourth-grade teacher from the
Baker School brought her class to
the Arboretum this October for a
field study called Plants in Autumn:
How Seeds Travel. Her response to
the question, “What did you like
most about your experience at the
Arboretum?” was:

I really admire the fact that
this program is sraffed by vol-
unteers. I think it’s important
for kids to see people donating
their time and energy because
they want to. Additionally the
atmosphere was inviting,
which made the experience
that much better.

Volunteer guides who are
personally invested in their work
create an invigorating learning
environment for school classes thar
come to the Arboretum for Field
Study Experiences. Visiting
schoolchildren find themselves in
a group facilitated by any one of
the 25 men and women who guide
children on these fall and spring
programs. Each guide is trained to
support the children’s science
learning, as together they examine
the plants and habitats within the
Arboretum landscape.

As volunteers, rhe school
program guides are dedicated to
enriching children’s connection
with science, nature, and the
Arboretum through the Field
Study Experience. These volun-
teers are men and women whose
commitment might originate

from a personal interest in
children’s education: they include
former teachers, a school librarian,
an education graduate student,
grandparents, and a person consid-
ering a career change to education.
Other volunteers come with per-
sonal experience and interesr in
life science: as do a part-time
science teacher who saves a day per
week to “teach ” at the Arboretum,
an ex-biology instructor, a retired
chemist, a self-employed horticul-
turist, and many impassioned gar-
deners. Many of our volunteers
know and love the Arboretum

from the perspective of neighbor
and supporter; it is from this per-
spective that they invest in shar-
ing its richness with others.

School program guides make
a one-year commitment to their
job that includes thirty hours of
Field Study Experience training;
weekly guiding of elementary age
children, fall and spring; and
attendance at education meetings
during the winter months. For
more information or to observe a
field study program, phone Diane
Syverson, manager of school pro-
grams, at 617/524-1718 xl63-

No Complaints Here

Peter Del T redid, Director of Living Collections

Gardeners are notorious for their
ability to complain endlessly
about the weather. If it’s not too
wet, then it’s certainly too dry; if
it’s not too hot, it’s certainly too

cold. The right amount of snow is
great, but too little or too much
is always a problem. And so on
down the line. This tendency in

• continued on page 3

Kirstin Behn

Karen Madsen

Former Intern
Returns as Putnam
Fellow

I MU ft! lirogna, Putnum l-elloif

I 've been fortunate, as a child of a
foreign service family, to travel in
Asia, Europe, and the U.S. and to
live in very different kinds of
places, including suburban north-
ern Virginia, downtown Tokyo,
and the rural Northeast Kingdom
of Vermont. Somewhere along the
way, noticing my surroundings, I

became a student of landscapes
and landscape history.

I consider my Putnam fellow-
ship an opportunity to continue
my investigations into the work-
ings of New England landscapes,
which I began officially as a
graduate student in landscape
architecture at Harvard’s Graduate
School of Design (GSD). One of
my projects here will be the study
of planning and land management
issues, including development and
tourism pressures on working
farms and forests. 1 also will inves-
tigate how the Arboretum func-
tions within its three increasingly
urbanized watersheds in order to
prepare a stormwater maintenance
plan for the site. Finally, I will
research the land-use history of
the area proposed for a new sun-
loving shrub and vine collection.

Farewell to Peter Stevens

The imminent departure of Peter Stevens, professor of biology and
a curator of the Arnold Arboretum and Gray 1 lerbaria, represents
a serious loss to the Arboretum’s group of specialists in Asian
botany. Peter will be joining his wife. Dr. Elizabeth Kellogg —
known to us all as Toby — on the faculty of the University of
Missouri at St. Louis. Toby will hold the E. Desmond Lee Chair
in Botanical Studies, Peter will be a professor of biology, and both
will also hold adjunct positions at the Missouri Botanical Garden.

Peter joined the staff of the Arnold Arboretum as an assistant
curator in 1973 after three years as a botanist in the Papua New
Guinea Forest Service, and worked his way up through the ranks:
quite a feat at Harvard! He has pursued two groups of interests
here. One has been in theoretical aspects of the history and
practice, of systematics, and particularly how botanists use the
characters of plants in classification and to interpret evolution.
But Peter may well be remembered most for his elegantly crafted
systematic treatments, in the St.John’s wort family, Clusiaceae,
and especially its large and subtly varying tropical tree genus
Calophyllunr. in the rhododendron family, Ericaceae; and in vari-
ous other taxa that have presented interesting problems to him.

Peter has played a seminal part in the teaching of plant
systematics at Harvard. His undergraduate course Bio 103,
Evolution and Diversification of Flowering Plants, and his gradu-
ate course Bio 218, The Families of Flowering Plants, have
attracted a growing number of students who found them dense
and therefore difficult but, thanks to Peter’s ebullient enthusiasm
for his subject, immensely stimulating.

We wish Peter and Toby good success in this new phase in their
careers and will welcome their future visits here.

Peter Ashton. Director. 1978—1987

During my tenure as Putnam
Fellow, I am dividing my time
between the Institute for Cultural
Landscape Studies (ICLS) and the
Department of Living Collections.
By straddling departments, I am
allowed a wonderful balance in
my work. I may spend one day
devoted to ICLS in the Widener
Library stacks at Harvard, track-
ing down references to farmland
conservation or cultural geogra-
phy. The next day (after studying
USGS topographical maps and

poring over city wastewater
flow diagrams), I m out following
the course of our own Bussey
Brook.

1 first joined the Arboretum
as a horticultural intern in the
summer of 1993. After graduating
from the GSD in 1997, 1 spent
one year working in a landscape
architecture firm before returning
to the Arboretum in September of
this year. I’m thrilled to be back
and feel happily at home here
among the trees.

2

FALL 1998

Karen Madsen

• from page 1

gardeners has only been exacer-
bated over the course of the last
ten or twenty years, as weather
extremes become the norm and
the so-called hundred-year flood
seems to happen once a decade.

All ot which takes me to the
point of this article, namely, that
the living collections department,
which has certainly done its share
of complaining about the weather
in the past, doesn't have anything
to complain about this year. The
winter was mild and the spring
was cool and moist. Remarkably,
the summer, which was consid-
ered very dry over most of the
East Coast, was no problem in
Boston where we enjoyed adequate
rainfall through the treacherous
months of July and August.
Indeed, nearly every time we
talked about watering our newly
installed plants, it started raining.
And the same is true for the fall.
Two weeks never passed without
substantial rain.

Taking full advantage of this
“anomalous" weather pattern, the
grounds crew planted more than
120 conifers m the collections
from mid-September through
mid-October. It was particularly
gratifying to plant these trees,
given that the Pinetum area was
badly damaged by the blizzard of
April 1, 1997. "While it requires a
certain amount of imagination on
the part of the visitor, it is now

possible to envision the appear-
ance of the Arboretum twenty
years into the future, when the
new plantings reach adulthood.

This fall’s planting list was
heavily laden with arborvitae
(Thuja Occident alls and T. plicata),
but we also accessioned many
pine, fir, larch, and spruce. As it
happened, during the long Colum-
bus Day weekend, and just a few
days after we planted our last tree.

On October 15, the Massachusetts
Horticultural Society (MHS)
awarded Peter Ashton the Thomas
Roland Medal of Honor during
the MHS Annual Awards Cer-
emony at the Boston Harbor
Hotel. Dr. Ashton served as direc-
tor of the Arnold Arboretum from
1978 to 1987 and currently is the
Charles Bullard Professor of For-
estry at Harvard University.

First bestowed in 1927, the
Thomas Roland Medal was
awarded in recognition of “excep-
tional skill in horticulture.” In
presenting the award. Dr. John C.
Peterson, president of MHS,
lauded Dr. Ashton “for his exten-
sive work that has ensured a won-
derful public treasure in Boston’s
Arnold Arboretum, and for the
demonstration of what is without
question exemplary skill in the
field of horticulture.”

it rained more than four inches in
three days, saturating the ground
in a way that no amount of hand
watering or irrigation ever could.
Remarkably, the universal law of
compensation seems to have
worked its mysterious magic at
the Arboretum, making the prob-
lems of the past few years seem
like distant memories. Unfortu-
nately, I’m sure that next year will
be a completely different story.

Dr. Peter Ashton, Arnold
Arboretum director, 1978-1987

Dr. Ashton’s successor as
director of the Arnold Arboretum,
Dr. Robert Cook, extended the
congratulations of the entire staff.
“Peter is also the world’s foremost
authority on the tropical forests
of Asia,” Cook noted. “We thank
Peter for bringing the Arboretum
to a position of leadership for
conservation of Asian tropical
forests.”

Dr. Peter Ashton Receives Honorary Medal

Arnold Arboretum Council members Wendy
Pearson, Sarah Jolliffe, and Bob Bartlett prepare to
embark on a tour of the living collections follow ing
the fall Arboretum Council meeting. Council
members serve as advocates for the Arboretum,
advise the director in their specialized areas of
expertise, and support the institution in a variety of
ways. Events of the day included presentations on
new initiatives, ongoing projects, and a panel
presentation of landscape maintenance issues.

ARNOLD ARBORETUM NEWS

John Burley

Two Collaborative Projects of the AA/NPS Win ASIA Awards

lioth I- airs ted: A Cult aval Landscape Report for the I'rederick Law Olmsted
National Historic Site. Volume 1 : Site History and Landscape Explorers:

Uncovering the Power of Place won 199H Merit Awards from the Amer-
ican Society ot Landscape Architects (ASLA). Both publications are
the result of collaborations between the Arnold Arboretum and the
National Park Service that began in the early 1990s.

Fairsted, the Frederick Law Olmsted National Historic Site in
Brookline, iMassachusetts, was the home and professional office of
Frederick Law Olmsted and the subsequent firms headed by his sons
and others. The National Park Service acquired the site in 19S0. The
Fairsted Report, produced jointly by the Olmsted Center for Landscape
Preservation of the National Park Service and the Arnold Arboretum,
includes a detailed history of the landscape of Fairsted by the noted
Olmsted scholar Cynthia Zaitzevsky and an afterword that describes
the horticultural and cultural context of the Olmsted’s work by the
garden historian Mac Griswold. Peter Del Tredici, director of living
collections of the Arnold Arboretum, participated in the evaluation of historic documentation of the site
and provided valuable expertise in plant identification from historic photographs. This report is an integral
part of the restoration process for the Fairsted landscape, which began in 1991.

Although the report documents a site of only 1 .76 acres, it is (to quote the ASLA) “a fascinating look at
Olmsted’s most intimate work: the design, literally, of the master’s own backyard.” Copies of the report have

been distributed to libraries nationwide. Individual copies can be purchased
through the Eastern National Bookstore at the Frederick Law Olmsted
National Historic Site, 99 Warren Street, Brookline, MA 02446. For mail
orders, contact Alan Banks at 617/566-1689 x221.

The ASLA calls Landscape Explorers “the first — and thus far — the only
curriculum designed to teach elementary-school students about the impor-
tance of landscape and place in everyone’s lives.” This unit of study invites
students to explore the landscape from the perspective of an artist, a historian,
or a naturalist. The stated hope that drives the unit is that “children who
understand the role of ‘place’ in their evolving sense of self tend to become
adults with a commitment to conserving and enhancing their immediate
neighborhoods and the larger landscapes of which they are a part.” The
authors of this work are Diane Syverson, manager of school programs at the
Arboretum, and Liza Stearns, education specialist for the Frederick Law
Olmsted National Historic Site. Participating students begin their exploration of place by examining their
own schoolyard and learning what it means to “read” a landscape. They then apply those newly learned skills
in a visit to the Arboretum, exploring this landscape in one of the three distinct ways described above. For
further information about Landscape Explorers, contact Diane Syverson at 6l7/524-l'’18 xl63.

Give the Gift of Membership

Membership in the Vrieiids of the Arnold Arhoretiim makes a unique gift for family and
friends at any time of year. The special recipient of your gift w ill enjoy a year’s worth of

exciting benefits.

Please help support the Arnold Arboretum hy purchasing a gift inenihership today!

Call the Membership Office at 6 17/524-1 ''18 xl65 for more information.

FALL 1998

Muriel’s Bamboo 17

Fargesia murielae at Kew Gardens, lifeless at the conclusion of its flowering in 1997.

M. A. Franchet, Fargesia. nouveau genre de
Bambusees dela Chine. Bull. Mens. Soc. Linn. (Paris,
1893) 2: 1067-1069.

C. Stapleton, Muriel Wilson's Bamboo. Newsletter of
the Bamboo Society {European Bamboo Society,
Great Britain, January 1995), 21.

Acknowledgments

The author would like to express his thanks to Dr. Chris
Stapleton, consulting taxonomist at Kew Gardens, for
his help in sorting out the complex history of the
introduction of Fargesia murielae. and to Keiko Satoh,
Putnam Research Fellow at the Arnold Arboretum, for
help in sifting through the Wilson Archives, housed at
the Arnold Arboretum.

Peter Del Tredici is Director of Living Collections at the
Arnold Arboretum.

conservatarum. Kew Bulletin of Miscellaneous
Information (1920) 10: 344-345.

" Perhaps Wilson used 1910 as the date for
"successfully" introducing Fargesia murielae
because it was then that he inventoried his
collections for those that were actually alive "in the
arboretum nurseries." An alternative, and rather
unlikely, interpretation is that Wilson recollected the
hamboo in 1910 and simply recycled #1462 from
the 1907 expedition. Of course, one can not discount
the possibility that Wilson just made a mistake
in giving 1910 as the date for the introduction of
F. murielae.

At the National Arboretum the plant was given
inventory number 262266; at the Arnold Arboretum,
It became accession number 1239-60.

T. R. Soderstrom, The Bamboozling Thamno-
calamus. Garden (1979)3(4): 22-27; Renvoize, op. cit.

Nature Study Moves into the Twenty-First Century

Candace L. July an

The veining of the leaves and the construction of the stalks . . . are as interesting to me
as the construction of a locomotive is to an engineer. When you get to know the plants,
you feel as though you ought to have a garden where you can take care of real plants
and study them.

Plants move, though many people do not know it. It is true that they do not move with
a jerk, but they move very slowly. When the corn gets beaten down by a heavy rain or
hail storm, it gradually works itself up again, although it never gets perfectly straight
as before. When we move, we bend our joints. That is the way also with the corn. It
bends at the nodes.

Reports from fourth-grade students at the Francis Parker School, Chicago, 1915.^

A rri^ A 5 r»m An

In many respects these reports
could be more readily attributed to
students today than to those at the
beginning of this century. The
study of plants is now considered a
routine part of the elementary cur-
riculum, and reports are a standard
form of communication between
teacher and student. However,
classroom practice that encourages
students' observations of nature,
considered laudable today, was
much more controversial at the
beginning of the century. At the
Arnold Arboretum, education for
children has been shaped by our
strong belief that the most power-
ful learning happens out in the
landscape, a belief that was articu-
lated at the turn of the century by
participants in the "nature-study
movement." The fourth-graders
quoted above, students at a school
founded on the principles of this
movement, had studied plants by
observing corn growing in their
schoolyard, rather than by reading
about it in a textbook. A closer

Nature Study 19

look at the tenets of nature-study serves
to identify the roots of our beliefs and to
illuminate new ways to approach the
study of nature.

The nature-study movement, which
peaked between 1890 to 1920, was part
of a progressive education philosophy
that proposed a child-centered approach
to learning by encouraging engagement
and play in contrast to more traditional,
text-driven practices. Nature-study edu-
cators (who used the hyphen to signify
that their nature study included a peda-
gogical approach) proposed that learning
about the natural world was as impor-
tant as studies of reading, writing, arith-
metic, and grammar. The key precepts of
the nature-study movement can be sum-
marized briefly;

• The objects of study can be ordinary,
seasonal phenomena.

• Direct observation is central to learn-
ing; drawing can be a useful, comple-
mentary tool.

• The teacher guides the students'
exploration; fostering discussions is
considered more critical than memori-
zation.

• Truly significant learning about nature takes
place outdoors, "in nature."

• Education should instill a love of nature in
the child. ^

Much of the impetus for this movement came
from a concern that the rigid approach to teach-
ing was not resulting in significant learning
by students. Samuel Jackson, an important
spokesman for the movement, summarized the
dissatisfaction of many with traditional book-
centered study:

Instead of providing the child with proper condi-
tions which cause him to grow out of the old
into the new, usually, the teacher merely smites
him with a definition. The child is finally bela-
bored into saying, "The earth is round like a
globe or a ball," and the matter is dropped; but
most of his geography forever conforms to his
picture of the old flat earth of his childhood.^

Such misgivings were certainly not new.
Over two centuries earlier, the Moravian monk

Tree. XIII. .\rbor.

A Plamt, i. growelh PlanUt, i. procrescil

from a Sftd. e Stmimt,

A plant waxeth to a

t.

Frutex in Arbcrtm, 3.
Radix, 4.

Sustentat arborem.

Sitirfs (Siemma) 5.
Surgit c radicc.

Stirfi se diridit
in Ramai, 6.

9t Frandet, 7.
factais e Faliis, 8.

John Amos Comenius (1592-1670) wrote a cri-
tique of the approach to children's education
at that time:

Hitherto the schools have done nothing with the
view of developing children, like young trees,
from the growing impulse of their own roots, but
only with that of hanging them over with twigs
broken off elsewhere. They teach youth to adorn
themselves with others' feathers, like the crow
in Aesop's Fables. They do not show them things
as they are, but tell them what one and another,
and a third and a tenth, had thought and written
about them, so that it is considered a mark of
great wisdom for a man to know a great many
opinions which contradict each other.'*

Comenius developed his ideas in the first
illustrated children's book, Orbis Pictus, pub-
lished in 1658 and focused on topics familiar to
young people. The book's small woodcut graph-
ics are accompanied by short texts that deal
with a wide range of topics drawn from both
nature and ethics — from clouds, trees, and ani-
mals, to honesty, respect, and love.

PUata abit

SJkaa/, 3. in Frutuem,

A ShKd to a Tret, 3.

The Root, 4.
bearetb up the Tree.

The Body or 5item, 5.
riscth from the Roou
The Stem divideth it self
into BamgAs, 6.
and green BrameAes, 7.
outde of Leaves, 8.

D

SPECIAL COLLECTIONS, MUNROE C. GUTMAN LIBRARY OF HARVARD UNIVERSITY GRADUATE SCHOCU OF EDUCATION

20 Arnoldia 1 998 Fall

Another writer influential in the develop-
ment of the nature-study movement was Jean
Jacques Rousseau (1712-1778). Many of his
ideas were incorporated into the movement's
philosophy: the principles of science are discov-
ered by the child, not learned as facts,- learning
should begin with observation of common
phenomena; the order of learning should be
determined by the learner's interests and expe-
riences, not by the organization of science; and
the objective should be enthusiasm for the dis-
cipline and methods of science, rather than a
body of memorized facts. ^

As the nature-study movement gathered
momentum in the late nineteenth century, its
leaders built upon these ideas to create an
approach to education with careful study of the
outdoor environment as its centerpiece. While a
growing number of teachers found these ideas
exciting and in line with their own thinking,
many others were baffled by the idea of teaching
without books and using natural objects and
phenomena to help children understand the
world around them. Ultimately the movement
lost strength as educators turned away from the
ideas of progressive education in favor of more
traditional approaches.

The Relevance of Nature-Study Today

While the philosophy of the nature-study move-
ment could be found in small pockets of schools
throughout this century, the ideas gained favor
again in the 1960s and 1970s with the growth of
environmental education and of science educa-
tion that focused on experience and, more
recently, in the 1980s and 1990s, with a renewed
focus on science education. The notion of a
compatibility between science and nature-
study was not prevalent at the turn of the cen-
tury. Although exceptions existed, such as Louis
Agassiz, a nineteenth-century scientist whose
credo was "Study nature, not books," generally,
nature-study educators and professors of science
held significantly different ideas, as suggested
in these passages written by Anna Comstock
m 1911:

For a long time botanical science, in the popular
mind, consisted chiefly of pulling flowers to
pieces and finding their Latin names by the use

of the analytical key. All the careful descriptions
of the habits of plants in the classic books were
viewed solely as conducive to accuracy in plac-
ing the proper label on herbarium specimens.
Long after the study of botany in the universities
had become biological rather than purely sys-
tematic, the old regime held sway in our second-
ary schools; and perhaps some of us today know
of high schools still working in the first ray that
pierced primeval darkness. . . .

To-day nature-study and science, while they
may deal with the same objects, view them from
opposite standpoints. . . . The child, through
nature-study, learns to know the life history' of
the violet growing in his own dooryard, and the
fascinating story of the robin nesting in the cor-
nice of his own porch.®

Comstock explained that nature-study "does
not start out with the classification given in
books, but in the end it builds up a classification
in the child's mind which is based on fundamen-
tal knowledge; it is a classification like that
evolved by the first naturalists, it is built
on careful personal observations of both form
and life."’

She would, no doubt, be surprised to learn
how the teaching of science has shifted in the
intervening years. In 1994, the National Acad-
emy of Science convened a large group of scien-
tists and educators to consider how and what
children should learn about science and the
environment. The conclusions of this group,
published in 1996 as the National Science Edu-
cation Standards (NSES), suggest certain "big
ideas" to be addressed at each grade level and
propose an approach to teaching that in many
ways resembles the one endorsed by the nature-
study authors at the turn of the century:

Learning science is something students do, not
something that is done to them. In learning
science, students describe objects and events,
ask questions, acquire knowledge, construct
explanations of natural phenomena, test those
explanations in many different ways, and com-
municate their ideas to others.®

The Arboretum's work with children employs
a combination of the nature-study philosophy
and scientific practice. Begun in 1984, the
Arboretum's Field Studies Experiences are
designed for small groups of elementary stu-

Nature Study 21

dents who come to the Arboretum to observe
closely and make sense of what they see. In the
fall, students look for seeds and determine their
mode of travel; in the spring, they discover the
stages of transformation from flower to fruit. In
both of these activities, careful observation is
supplemented by conversations with the guides,
who help students make sense of what they see.
This program is based on a belief that children
learn best through experiences in the landscape,
guided by attentive adults.

A decade later, we explored ways to add data
collection to these observation-based activities.
In 1995, with funding from the National Science
Foundation (NSF), the Arboretum began the
development of a program that could serve as a
model for partnerships between elementary
schools and institutions involved in science.
While based on many of the principles of nature-
study, this new project, called Seasonal Investi-
gations, also includes an emphasis on keeping
systematic records of observations and sharing
those data with others using a computer
web site.

A Design for Nature Study in the Twenty-
First Century

Before I investigated a twig in winter, I just
thought that the leaves fell off a tree and gradu-
ally grew back. But boy, did I learn a lot about
trees from just one little twig!

Maybe I should tell you about some things I
learned. ... I learned the names of the different
parts of a twig, like the Terminal Bud, which is
the bud at the tip, and the Lateral Buds, the little
buds on the sides. I, myself, liked the names our
class made up better. Like the name I gave to the
Terminal Bud, Kiss-End Tail (an off-spring from
the expression "Kiss and Tell").

Another thing I learned from my twig is that
the different colors along the twig signal yearly
growth. We also determined the yearly growth
for 1995-96 by looking at the first ring from the
top. Then we measured from that ring to the
very tip of the twig. Get this, my twig grows one
centimeter less each year! So next year, if my
twig only grew one centimeter since 1995, my
twig will probably stop growing. Or maybe it
will start a whole new growth. I think that
the reason my twig's health has been declining is
because of the harsh winters we've been having.
Well it'll sure he a big surprise [this spring]!

My twig was a
informative soui
learned more i
trees than I couh
fit into one report. So I
better go before I start an-
other paragraph telling you
about how great trees are!"

— Report from a fourth-grade
student at the Murphy School,

Boston, 1996.

Like the student reports from
1915 quoted at the beginning, this
one displays an enthusiastic, obser-
vation-based consideration of the sub-
ject. The author is a student in one of
the classrooms participating in the
Arboretum's Seasonal Investigations
program (originally called the Commu-
nity Science Connection), designed to
help elementary teachers strengthen
their science curricula by replacing the
usual one- to two-week unit based on
textbook explanations with a year-
long study of trees in the schoolyard.

To date, we have worked with fifty
teachers from the Boston, Newton, and
Brookline school districts; all attended
summer institutes at the Arboretum
before guiding their students through
the study. The goals of the program are
very similar to those of the nature-study
movement:

• to encourage use of the outdoors as an ex-
tension of the classroom, whether in the
schoolyard, the Arboretum, or
the students' neighborhood;

• to provide the structure of a
year-long study of individual trees
that incorporates close observations, sys-
tematic data collection, and guidance in
making sense of the data;

• to give the students opportunities to com-
municate on the web with others studying
the same topic.

The program proceeds through three seasons.
The fall investigation focuses on the general
characteristics of trees and on the ways species
differ, such as in the dates that leaves change

22 Arnoldia 1998 Fall

>>

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Location jhttp 7 /arboretum harvard edu/csc /seasonal /current /SPQTDIR /leaves /types .Mm*

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Patterns A Changing Leaf Color

What pattern of
leaf color
change do you
notice on the
leaves of your
tree?

yt ■'/<>/

/•?//'■ \ Some students

who visited the

Arboretum last year noticed that leaves changed color In a number
of different patterns,

♦ Some found leaves that changed color at the edge r •
orthetips. Click on the example image

• Some found that the color changed (or It stayed ,
green) through the veins. Click on the example -4^
image,

• Others found leaves with blocks of color change

Click on the example image,

What sorts of patterns did you find"? Look over your field notes or
visit your tree to see if any of these patterns describe how the
leaves on your tree changed Did you find another pattern
altogether?

Share your discoveries and your questions about leaf color
change!

[What Other* Fojtdl

What I Fotjnd

Return io Spotlighi | Whei I Found | Wh&t Others Pound

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Document Done.

color and fall from the tree. In the winter inves-
tigation, students learn to "read" a twig and use
their new knowledge to determine which was
the best recent growing year for the schoolyard
trees. The spring investigation revisits the
features examined in the winter to learn
whether and how those features change in the
spring and to determine when the flowers are
"open for business."

The student report quoted
above was written as part of the
winter twig investigation. The
twigs, initially viewed by students
as a bag of sticks, constitute the
major focus of the class investiga-
tion. Each twig soon becomes a
treasured resource. Students begin
by making careful drawings and
identifying features of the twig,
later naming the features. These
names are often revealing. For
example, one student named the
annual growth-ring marks "grow-
ing up lines." Many students pre-
ferred their own names to those
of scientists, hut they were fluent
in both.

The Role of the Web Site

Now in our final year of NSF fund-
ing, we are designing a web site
for Seasonal Investigations that we
believe will support both the class-
room and outdoor work and allow
a greater number of teachers to
take part in the project. While the
program can be (and sometimes
is) completed successfully using
only the classroom and school-
yard, the on-line environment pro-
vides an important support for the
four activities central to the proj-
ect— observation, data collection,
communication, and publication —
with a web site feature dedicated to
each of these activities.

The Spotlight feature changes
weekly throughout each seasonal
investigation; the topic of each
entry is chosen to encourage
closer observation. In the fall investigation, stu-
dents were invited to consider patterns of leaf
change, to view other students' drawings of pat-
terns they found, and to share their observations
about leaf patterns with others. Another Spot-
light entry asked them to consider how bark ac-
commodates the expansion of a tree's girth.
Three possibilities — fissures, plates, and peels —
were illustrated with photos; students were

Nature Study 23

asked to look at their school-
yard trees and report their
findings.

The Tree Talk feature facili-
tates communication among
classes, from initial letters of
introduction to later conversa-
tions about questions or find-
ings. Contributions to most of
these "conversations" can be
made and viewed at any time,-
in addition, there is an option
for a live, scheduled chat with
either Arboretum staff or other
classrooms.

The Activities feature pro-
vides the structure for sharing
data among classes. Students
are asked to provide specific
data about their schoolyard
trees, changing or adding to the
data as the study progresses.

The combined data provide
opportunities for discussion in the classroom or
with other students.

The Publication feature is intended to elicit a
creative activity at the end of each investiga-
tion, perhaps a report or drawing, that brings
together the ideas, surprises, and discoveries
from the investigation.

The first three years of the project were spent
perfecting the model and developing a set of
investigations that could be completed in the

schoolyard, with supporting visits to the Arbo-
retum. During this, the last year of the project,
the focus is on perfecting the web site to ensure
that the program will continue after NSF fund-
ing ends.

Even before the project's completion, the
framework of Seasonal Investigations has been
adopted as a model by other institutions
engaged in science education. Descanso Gar-
dens in Los Angeles is replicating the entire pro-
gram as a pilot project with the Los Angeles
Unified School District. The Garden's director,
Richard Schulhof, had first-hand experience
with the project as a member of the Arboretum
staff at the time it began, and is enthusiastic
about using the program as a new approach to
science teaching for his own staff as well as for
the Los Angeles teachers. In addition, the Mas-
sachusetts Audubon Society is using the Sea-
sonal Investigations framework to develop both
teacher institutes and investigations of vernal
pools in three locations across Massachusetts.

Future Directions

Many of the ideas of the nature-study move-
ment are alive and in practice in today's
programs at the Arboretum, but new issues

24 Arnoldia 1 998 Fall

are also being raised. What role can the web
play, not as an end in itself but as a springboard
to investigations outdoors? How might it pro-
vide an avenue for sharing our educational
ideas, many of which have century-old roots,
with interested educators around the glohe?

In many ways, the words of Anna Comstock
have as much relevance at the end of this cen-
tury as they did at the beginning:

When the child has become acquainted with the
conditions and necessities of plant life, how dif-
ferent will the world seem to him! Every glance
at forest or field will tell him a new story. Every
square foot of sod will be revealed to him as a
battlefield in which he himself may count the
victories in the struggle for existence, and he
will walk henceforth in a world of miracle and of
beauty, — the miracle of adjustment to circum-
stances, and the beauty of obedience to law."’

The young author who wrote about her twig
is one of a growing number of students whose
science experiences have been shaped, either
directly or indirectly, through a connection with
the Arboretum and its staff. As we enter the
twenty-first century, we continue to seek oppor-
tunities for sharing our ideas about the compat-
ibility of nature, science, and technology with
teachers and students eager to learn about trees
and plants. Our hope is that ideas about
children's education, developed and nurtured at
the Arboretum, can grow into viable "seeds"
locally and around the country.

Endnotes

' From the Francis Parker School Year Book, vol. IV,
lune 1915 (Archives of Gutman Library, Harvard
University).

^ Culled from W. S. Jackman, Nature-Study and
Related Subjects for Common School. Part 11. (New
York: Henry Holt, 1891); A. C. Boyden, Nature Study
by Months (Boston: New England Publishing, 1898);
G. L. Clapp, "Real and sham observation by pupils,"
Education, January 1892; C. B. Scott, Nature-Study
and the Child (Boston: D. C. Heath, 1901); A. B.
Comstock, Handbook of Nature-Study (Ithaca, NY:
Comstock Publishing, 1911).

^ Quoted in Jackman, op. cit., pp. 9-10.

* Quoted in N. A. CaJings, "History of Qbject
Teaching," Barnard’s American fournal of Education
(December, 1962) 12: 637.

^ Quoted in T. Minton, "The History of the Nature-
Study Movement and Its Role in the Development
of Environmental Education" (Unpublished dis-
sertation, University of Massachusetts, Amherst,
1979), pp. 30-31.

* Quoted in G. F. Atkinson, First Studies of Plant Life
(Boston: Ginn Co., 1901), p. iii.

^ Comstock, op. cit., p.5.

* National Research Council, National Science
Education Standards (Washington, D.C.: National
Academy Press, 1996), p. 20.

’ Atkinson, op. cit., p. v.

Candace Julyan is Director of Education at the Arnold
Arboretum.

Native vs. Nonnative: A Reprise

Letters to the Editor

To the Editor;

I thoroughly enjoyed the article "An Evolutionary Perspective on Strengths, Fallacies,
and Confusions in the Concept of Native Plants" hy Stephen Jay Gould [Arnoldia
Spring 1998). Gould uses the native plant issue to clarify some of the common mis-
conceptions surrounding the basic theory of natural selection. He also reasserts him-
self as one of Darwin' prevailing "hulldogs." I do not take issue with anything that
Gould includes in his analysis, but rather what he fails to include. Little effort is
made to address the evolutionary ecology perspective in the concept of native plants.

Gould touches lightly on the merits of native plants by indicating that they "have
generally been present for a long time and have therefore stabilized and adapted" to
local conditions. This is for me a key reason, from an evolutionary perspective, for
promoting native plants, since natives have presumably coevolved with other local
organisms and the chemical and physical environment. An ecological balance has,
therefore, generally been struck that prevents the unbridled increase in any one
species' numbers. Exotic plant species (i.e., nonnative plants), on the other hand,
have a greater propensity for rampant population growth. Darwin even comments on
the ecological consequences of invasive exotic plant species and points out in chap-
ter three of On the Origin of Species that;

. . . cases could be given of introduced plants which have become common throughout
whole islands in a period of less than ten years. Several of the plants now most numerous
over the wide plains of La Plata, clothing square leagues of surface almost to the exclu-
sion of all other plants, have been introduced from Europe. ... In such cases the geometri-
cal ratio of increase, the result of which never fails to be surprising, simply explains the
extraordinarily rapid increase and wide diffusion of naturalized productions in their new
homes. . . . when a plant or animal is placed in a new country amongst new competitors,
though the climate may be exactly the same as in its former home, yet the conditions of
its life will generally be changed in an essential manner.

Irruptions of invasive exotic plant and animal species typically occur, as Darwin
implies, because they do not generally have the competitor, predator, or pathogen
load typically associated with native plants. Exotic species are new players in an
environment and do not adhere to the "rules" that govern native species. Admittedly,
the vast majority of exotic species are not invasive since they do not seem to com-
pete well with native plants. Those that are invasive, however, have wrought havoc
on local and regional ecosystems. Many native plants are maligned as invasive
because of their weedy nature, but there is a distinct difference; native weeds do not
disrupt natural communities nor do they tend to form monocultures. (I would like to

PETER DEL TREDICI

Native vs. Nonnative 11

see the term "invasive" restricted to ecologically disruptive exotic plants and the
term "aggressive" adapted for native weeds.)

Gould also gives little mention to the ecologically disruptive consequences of
invasive exotics to biodiversity other than saying that he "treasures nature's bounte-
ous diversity of species," and that "cherishing native plants does allow us to defend
and preserve a maximal amount of local variety." This is precisely why native plants
should be the first choice for landscaping among ecologically sensitive individuals.
Second choice should be exotic species that are not invasive or those that have a
very low potential for becoming invasive. I am not suggesting that we adopt the
"Naziesque" approach to plant material choice. I too am awed by our "bounteous"
species diversity but it is only diminished by invasive exotic species. And I hope
that Gould, by pointing out that the argument for using native plants is evolution-
arily fallacious, has not encouraged what he so stridently abhors: a misconstrued
Darwinian alibi for depraved behavior — in this case, using ecologically disruptive,
invasive plant species.

Organisms, native or otherwise, respond to their environment through the adaptive
creativity of natural selection. Theirs is a life without intent. There is no desire
among plants to become a garden pest or to disrupt natural communities. Humans,
by purposefully homogenizing the world's flora, have forced the occurrence of
unlikely species interactions, some of which we greet with delight (culinary herbs,
vegetable crops, and the majority of ornamentals) and some with dread (kudzu, privet,
and water hyacinth). Gould weakly dissuades the introduction of invasive exotic
plant species by maintaining that there should be "sensitive and respectful mixing of
natives and exotics." From this I read: proceed with caution. I feel that a stronger
position needs to be taken on this issue. Invasive exotics are a major threat to
biodiversity and the genetic diversity contained within. I therefore challenge botani-
cal gardens and arboreta, plant nurseries, and private gardeners to promote the use of
ecologically judicious plant choices in our public and private gardens.

John Randall

Conservation Curator, North Carolina Botanical Garden of
the University of North Carolina at Chapel Hill

To the Editor:

Stephen lay Gould ("An Evolutionary Perspective on Strengths, Fallacies, and Con-
fusions in the Concept of Native Plants," Arnoldia Spring 1998) offers an excellent
argument for his characterization of the concept of native plants as "a remarkable
mixture of sound biology, invalid ideas, false extensions, ethical implications, and

Grapevines (Vitus sp.) in northeastern Connecticut.

28 Ainoldici 1998 Fall

political usages." It seems worth adding to his commentary, written from an evolu-
tionary perspective, an argument from the perspective of practical horticulture.

But first, I hasten to point out that in 1998, we in the arenas of botanic gardens and
horticulture are already working to promote the use of environmentally appropriate
plants specific to the requirements and use of the planted site. Plantsmen are not rec-
ommending aggressive exotics as landscape plants of preference regardless of environ-
mental consequences. That would not only he irresponsible hut would ultimately
destroy the green industry and its important contribution to the U.S. economy.

The primary criteria for plant selection in managed environments today is whether
a plant is reasonably well adapted to the site and, hence, will survive and thrive with-
out requiring regular use of pesticides, and within that context, whether the plant
satisfies the ornamental, agricultural, and/or functional demands of the site and its
constituents. Given these criteria, there are many instances when exotic plants are
the clear choice for a given landscape — especially when we recall that not all exotics
are invasive or aggressive (in fact only a small minority have proven to he so), and that
not all natives are nonaggressive (for instance, our native staghorn sumac). The real
challenge, of course, is to determine with intelligence and sensitivity to site con-
straints what are the environmentally appropriate species for a given site that are
likely to succeed there. And, I might add, what may be the appropriate cultivars,
which are capable of great phenotypic and physiological divergence within one
species — in some cases, even greater divergence than the wild-type species can offer
within a genus.

We cannot ignore the reason that invasive exotics have been used in the first place,
which is: Managed environments (cities, residential neighborhoods, parks, disturbed
wetlands, timber production lands, and so on) are already drastically altered and have
already been interfered with, resulting in significantly inhibited natural selection and
the ability of the prior extant site natives to thrive. For this very reason, a managed
environment often requires conscious choice of potentially aggressive plants if there
are to be any plants at all that live there.

One of the reasons that botanic gardens, arboreta, and many types of public gardens
maintain living collections of plants is to allow for evaluation and comparison of
plant growth and development, and landscape performance long-term in real time in
a given regional landscape. The ability to carry out these evaluations allows us to
select well-adapted plants for an area. The broader the palette of well-adapted plants
available, the more effectively an environmentally sound landscape can be built.

All sites, whether managed or wild, including severely disturbed and altered stress-
ful environments (such as urban parks), require plants adapted to the conditions on
that site. A well-adapted plant for a managed site may or may not he a regional
native, depending on the specific stresses associated with the given managed environ-
ment. We cannot effectively plant many of the natives of the humid Northeast United
States in, for example, parking lot beds, or even in many new suburban garden sites
that have been stripped of topsoil (excepting only such broadly adapted and aggres-
sive natives as, for instance, poison ivy).

Native vs. Nonnative 29

Many Asian natives serve as good landscape plants in the Northeast precisely
because they are well-adapted to our most common types of "disturbed" landscapes.
The climatic and soil similarities between eastern North America and eastern Asia
are well documented and widely understood and accepted. Should we, in spite of this
natural botanical gift, restrict the plants grown on these sites to a few U.S. natives
that will thrive there because they will basically thrive anywhere? Do we want our
managed outdoor stressful environments to be planted with only a limited palette of
regionally native aggressive plants? Would the residents of Washington, D.C., really
want us to replace the flowering Asian cherries with native pin cherries (which are
actually more susceptible to tent caterpillars)?

How do we define "regionally native," in any case? — plants found growing within
a 100-mile radius of the site now, 100 years ago, 1000 years ago? Plants found grow-
ing within the state now, 50 years ago, 100 years ago? Plants found growing within
the region now, 50 years ago, 100 years ago, 1000 years ago?

Much as we may wish to, we cannot turn back the clock and erase the huge distur-
bances that we have thoughtlessly imposed throughout most of our native habitat.
This makes it even more critical that we preserve and protect what small acreages of
undisturbed habitat remain, as much as is possible.

Unfortunately, with increasing population and urbanization, the likelihood is that
over the next 100 years, these small acreages of undisturbed, or little-disturbed, or
restored habitat will become even more fragmented, pressured, and fragile. It is
imperative that we learn to manage our expanding areas of managed environments
wisely, using a diversity of plants that result in environmentally sound as well as
beautiful, productive, and functional landscapes. We cannot achieve that goal by
relying solely on "regionally native" plants for every single managed landscape no
matter its location or purpose.

Clearly, a reasonable, moderate, thoughtful, site-specific, and non-arbitrary
approach to plant selection is required for each individual landscape. Known rampant
invasives, regardless of provenance, should not be planted. The decision process for
what plants to include in a native wetland restoration project should clearly be dras-
tically different from that of choosing plants for an urban pocket park. In all cases,
effort to use plants suited to the region and the site must be made.

I write this response to remind us of what we all as plantsmen are already working
to achieve, that is, to bring reason, responsibility, knowledge, and moderation to bear
on the process of how we choose plants for managed environments, and what choices
we make. In the face of the next hundred years of increasing pressure on the land, the
future of our flora and the quality of our lives depends on this.

Kim Tripp

Director of the Botanic Garden of Smith College
and Arnold Arboretum Associate

Native Plants: Another View

Harrison L. Flint

To close this circle, for the time being, we reprint very nearly verbatim an
article from Arnoldia, Winter 1982-1983. When we asked Professor Flint to
update it to serve as a companion to the letters to the editor, he found very
little, and nothing at all of substance, that he wished to change.

Following the tradition of such great
midwestern naturalists as Jens Jensen, Aldo
Leopold, and May Theilgaard Watts, contempo-
rary landscape planners have grown in aware-
ness of native plants and their usefulness in
designed landscapes. The movement toward
landscaping with native plants now has spread
widely and has not yet reached its full potential.
Its ultimate expression is found in re-creating
natural plant communities, a stepwise and
time-consuming process now being carried out
by relatively few landscape planners. Such plan-
ners usually are sophisticated horticulturists
and landscape architects who have elected to
specialize in this particular area.

Yet, while many landscape planners have
developed close familiarity with a great range of
plants, carefully selecting those most appropri-
ate for the situation at hand, less-sophisticated
members of their profession have eschewed all
forms of vegetation that are not "native." For
some this position is taken with a sense of mis-
sionary zeal; for others it may simply offer con-
venience in requiring knowledge of a smaller
number of landscape plants.

To select landscape plants on the basis of
whether or not they are native, one must first
determine which species are "native." In New
England, for instance, is it permissible to select
black locust (Robinia pseudoacacia), a common
wild tree in much of the area, yet native only
farther south and west? Must redbud (Cercis
canadensis) be excluded in southwestern Wis-
consin, since it is an exotic species in that state,
even though it grows naturally a dozen miles
away in northwestern Illinois? In Indiana, must
another tree legume, American yellowwood

(Cladrastis kentukea, formerly C. lutea), be
restricted in use to only those few counties
where it is indigenous?

Any question about species eligibility for use
in re-creating'or preserving a natural plant asso-
ciation finds its answer in the planner's knowl-
edge of the association. Clearly, only certain
plants "belong." But in other areas of landscape
planning, divisions between native and nonna-
tive species blur — and perhaps are best left
blurred, allowing selection decisions to be made
according to criteria relating to function.

Exclusion of nonnative plants on principle is
based upon several generalized claims, all of
which hold at least a grain of truth:

(1) Nonnative plants look out of place in
the landscape.

If one's objective is to preserve a natural land-
scape, ample justification exists for removing
nonnative species as weeds. The same is true in
re-creating a "natural" landscape, but in other
cases the question is not so easily answered.
Must a woodland gardener in New England
be asked to plant no other species of wild
ginger (Asarum) than the native A. canadensel
Must sweetshrub (Calycanthus floridus),
galax (Galax urceolata), box huckleberry
(Gaylussacia brachyceia), and yellowroot
(Xanthorhiza simplicissima) be left to their
more southerly native haunts? And must the
New England gardener be sure to omit lily-of-
the-valley (Convallaris majalis) and English ivy
iHedera helix) as European natives? Perhaps,
but only as a matter of taste.

(2) Plant species are better adapted to the
region in which they are native than

Native vs. Nonnative 31

elsewhere, because this region has “made”
them, through distinctive selection pressures.

As logical as this view may seem at first, it has
two flaws. First, it excludes the possibility of
preadaptation. For example, the climate of
northeastern Asia so closely parallels that
of similar latitudes in northeastern North
America that many Asian species have been pre-
adapted to our climate long before they have
seen it, and turn out to be some of our most use-
ful landscape plants.

A second flaw is the tacit presumption that
the soil and climate of a particular landscape
site are similar to those of the natural region in
which it is located. Landscape designers and
contractors know that this is not true. Most
landscape sites, especially urban ones, are
exposed to soil and climatic stresses that sel-
dom exist in wild areas nearby. Soils may be
greatly modified by construction and subse-
quent restoration. Patterns of wind, solar radia-
tion, and temperature fluctuation are modified
in developed sites. Perhaps most important of
all, patterns of rainfall, runoff, and absorption of
water into the soil are drastically altered. In
short, developed sites are so greatly changed
that they may differ much more from nearby
natural areas than do certain natural areas on
the other side of the earth.

(3) Nonnative plants are weedy, reproducing
freely and invading areas where they are not
wanted.

This is a valid criticism of several nonnative
species, such as buckthorns [Rhamnus sp.),
certain Asian honeysuckles [Lonicera sp.),
kudzu vine (Pueraria lobata), some species of
Elaeagnus, Euonymiis, and others. But it is not
a fair generalization. In fact, it seems a contra-
diction to generalize that nonnative species are
not well adapted yet reproduce to the point of
being a nuisance. Again, it is necessary to know
which species, both native and exotic, are
weedy and exclude them in situations in which
they might get out of control.

(4) Native plants are less susceptible to insect
and disease problems than nonnatives and so
need less maintenance.

We as often hear the counterclaim; that nonna-
tive plants separated from their ecosystems are.

at least for a time, free of many of their natural
enemies, and examples of native species with
major problems are easily found. American elm
(Ulmus americana) has been decimated in
many areas by Dutch elm disease and phloem
necrosis. The most promising sources of resis-
tance to Dutch elm disease are Asian species
and their hybrids. The majestic American
chestnut (Castanea dentata), nearly wiped out
by blight in its native habitat decades ago, is
finding its closest replacement in the disease-
resistant Chinese chestnut (C. mollissimaj and
its hybrids.

Crabapples native to eastern North America
(e.g., Malus angustifolia, M. coronaria, and
M. ioensis) are susceptible to cedar-apple rust, a
serious enough problem to rule them out as
landscape plants in most localities where red
cedar (Juniperus virginiana), the alternate host
for the disease organism, is present. Asian
crabapples are relatively free of this problem. In
areas where red cedar does not grow wild, the
disease can be largely controlled by substituting
junipers of Asian origin for red cedar.

Resistance to insect and disease problems is
too important a consideration in selecting land-
scape plants to be left to generalization. It is bet-
ter dealt with directly by selecting troublefree
plants than indirectly by selecting only native
or nonnative plants, in the expectation that they
will tend to be more resistant to problems than
their opposite numbers.

(5) We need to make better use of the
tremendous pool of genetic diversity inherent
in native plant species, a pool that has been
barely sampled thus far.

Amen! And the same can be said for nonnative
species. How often is our knowledge of an Asian
species, for instance, limited to a few clones or
at best a narrow slice of the germplasm that
exists in the natural range? Intrepid plant
explorers have introduced to us many new spe-
cies from remote corners of the world. Notwith-
standing the many collections made over the
past decade, we have largely failed to follow up
on their discoveries by assembling larger
samples of those species for evaluation, just as
surely as we have neglected to observe fully the
variation that exists in native species. As a

32 Arnoldia 1998 Fall

result, our narrow knowledge of diversity in
plant species confounds the issue of their
nativeness.

The U.S. Department of Agriculture has
taken an important step to improve this situa-
tion with regard to crop species through its net-
work of plant germplasm repositories. It is up
to other institutions, including botanical gar-
dens and arboreta, to develop stronger programs
relating to preservation and development of
germplasm of value to landscape improvement.

There are, of course, landscape situations
where nonnative plants are clearly inappropri-
ate and so to be avoided. This includes preserva-
tion, restoration, and re-creation of natural areas
and plant associations. In many other situations
the constraint of using only native plants,
intended to produce a natural effect, itself

becomes artifact. In such situations it is more
sensible to return to the basics of plant selec-
tion, considering adaptability and intended
function first, then maintenance requirements
and seasonal interest. When a pool of plants
having the desired requirements has been
assembled, final selections can be made on the
basis of individual taste.

The search for a broad range of prospective
landscape plants, and their thoughtful use, has
made our landscape increasingly functional and
interesting. Continuing the search will enrich
our lives in the process.

Harrison Flint is professor emeritus of horticulture
at Purdue University in Lafayette, Indiana. He is
the author of The Country Journal Book of Trees and
Shrubs and Landscape Plants for Eastern North
America, 2nd edition.

U.S. POSTAL SERVICE

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(Required by 39 U.S.C. 3685)

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County, MA 02130-3500, publisher,- Karen Madsen, Arnold Arboretum, 125 Arborway, Jamaica Plain, MA 02130-
3500, editor. 10. Owner: The Arnold Arboretum of Harvard University, 125 Arborway, Jamaica Plain, Suffolk County,
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4

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