The World of Northern Evergreens by E. C. Pielou by E. C. Pielou - Read Online

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The World of Northern Evergreens - E. C. Pielou

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Predictions

Preface to the Second Edition

The world has changed since 1988 (the year when the first edition of this book appeared). At last it is dawning on governments that forests are more than just a source of timber. They provide, as well, indispensable ecological services. Were they to disappear, climate change would speed up because the world would lose its greatest carbon sink.

Ways to estimate the monetary worth of ecological services have recently been devised. So far, they have been carried out in detail in only a few places in the world. For example,* a closed-canopy forest in Kenya (East Africa) was found to supply $320 million in services, every year, from 1600 square miles (about 4100 km²).

As the true worth of forests comes to be appreciated, naturalists’ knowledge is regarded with more respect than it was in the days when their activities were looked on as no more than an enjoyable hobby. Their expertise has become useful and widely appreciated. The purpose of this new edition is to introduce new material on the evergreens in northern North America and to bring the earlier book up-to-date.

Some particulars: I have described the contrast between conifers and broadleafs (formerly, and less precisely, known as hardwoods) in much more detail. The enormous gap between these two kinds of plants is obscured by labeling them all simply as trees. It conceals their great dissimilarity. They have been evolving divergently from each other for more than a hundred million years (see chapter 6). New chapters are devoted to the forest floor (chapter 7) and to the geographical extents of different forest ecosystems (chapter 12). The effects of logging are discussed in chapter 11, and how global warming is affecting the forests and vice versa, in chapter 13.

Some paragraphs have been added on animals whose connections with their special habitats are unusually close, for example, caribou, some grizzly bears, and beavers.

And much else besides. No branch of science, and that includes natural history, ever remains static.

E. C. PIELOU

Comox, British Columbia

* Jen Fela, Reforestation Key to Economic Growth in Kenya, in Frontiers in Ecology and the Environment, vol. 8, no. 2, 2010, p. 63.

Preface to the First Edition

For many people, certainly for the majority of North Americans with homes in the northern half of the continent, coniferous trees constitute a large fraction of all the living material they will see in a lifetime. Naturalists, hikers, backpackers, canoeists, cross-country skiers—in fact all those whose work or recreation takes them outdoors in northern North America—are accustomed to seeing coniferous trees by the tens of millions, whether they consciously notice them or not.

Outdoor people have a wide spectrum of interests. Specialists tend to specialize in interesting items: a birder is more likely to concentrate on owls than on starlings, and the average plant-hunter finds orchids more fascinating than crabgrass. Because of this preoccupation with the hard-to-find, the beautiful, and the unusual, most of the commonest objects in nature are apt to be ignored. They are simply there, part of the background. But to assume that because a thing is common it is therefore uninteresting is a mistake.

For most outdoor people the fact that they will encounter rank upon rank of coniferous trees in excursion after excursion in the future neither pleases nor displeases them. They don’t even think about it. If there are innumerable coniferous trees in your future, why not take advantage of the fact, look at the trees more closely, and learn something about them? Knowledge cannot fail to bring interest and appreciation.

Learning to identify the different species of coniferous trees is only a beginning. Once you know the trees, many things can be observed if you know what to look for. There is, however, a world of difference between seeing and interpreting. The ability to interpret is the hallmark of the true naturalist, and developing that ability is one of the pleasures of being a naturalist. The well-informed naturalist understands and enjoys a thousand things that the uninformed one doesn’t even notice; and the more people who understand and enjoy the woods, the more there will be to protect them.

E. C. PIELOU

Denman Island, British Columbia

Chapter 1

Origin of the Evergreen Forests

Conifers and the Ice Age

Of all the people who enter a northern forest, only a handful ever ask themselves these two questions: Where has the forest come from? And why are the great majority of its trees conifers rather than deciduous, broad-leafed trees—broadleafs for short.

The answers are by no means obvious. The questions have engaged the interests of ecologists and motivated years of research. What has been investigated and measured is not, for the most part, observable on a hike in the woods, but the hike would certainly be more interesting for somebody knowing about the questions, and the answers that have been discovered so far.

Consider the first question, where have the trees come from? The only certain answer is that the trees in the area we are concerned with, the area that was ice-covered at the end of the last ice age, must have descended from ancestors that lived elsewhere.

About 18,000 years ago, when the ice sheets of the most recent ice age had reached their maximum extent (figure 1.1), they covered nearly all of northern North America. The glaciated areas must have been like Greenland and Antarctica were until recently—barren expanses of ice, devoid of plant life. Then, as now, the ice was on the verge of disappearing. Conditions in the unglaciated regions near the margin of the ice sheets must have been bleak. The contrast with conditions now is worth contemplating.

Figure 1.1. The area in North America covered by ice sheets at the end of the last ice age.

Once the climate began to warm up and the ice sheets started shrinking, newly exposed land became available for small plants, and hardy evergreen forests gradually invaded the area. Seeds were blown in from the south. At the time of maximum glaciation, evergreen forests stretched across the continent south of the ice margin; even the Great Plains were forested. East of the continental divide the most abundant trees were spruces and jack pines (the evidence that allows us to visualize the forests of the distant past is described in chapter 3). The forests south of the ice on the west coast had a richer mix of tree species and may have been much like they are now.

The northward march of the forests into the newly ice-free land was inevitably slow. There was no soil to start with—only lifeless mixtures of boulders, gravel, sand, and clay, laid bare by the melting ice. The development of soil adequate for trees must have taken a considerable time. Different species of trees arrived to occupy their present geographic ranges at different times. Those that survived the ice age farther south than the spruces and jack pines had farther to come.

And those that require shade had to wait until forests of sun-loving trees were casting enough shade for the shade-growers to invade. For example, hemlock is believed to have reached what is now northern Michigan about 3000 years later than white pine.¹

The climate continued to change as it has done throughout time, independently of the current global warming (the topic of chapter 13). It was thought to have warmed fairly steadily between 18,000 and 10,000 years ago, except for a temporary cold interval around 12,000 years ago. In any case, warming resumed and temperatures reached a maximum about 8000 years ago, even before the ice sheets had completely melted. Because of the very long time required to melt huge masses of ice at the then-prevailing temperatures, probably around 2° Celsius higher than now (2010), two remnants of the original ice sheets persisted, one on each side of Hudson Bay, until 6500 years ago. They finally disappeared after the time of maximum warmth. Because of the slight downward trend of temperatures until very recently, some species of evergreen have lost ground and do not, nowadays, grow as far north as they did 8000 years ago.

The Advantages of Being Evergreen

The second of the two questions posed at the beginning of this chapter was, why are the great majority of the trees in the northern forests evergreen conifers? Why conifers rather than broadleafs? To say that conifers are better adapted to the environment is no answer. The question then simply becomes, in what way are they better adapted? The answer cannot be simple, because conifers do better than broadleafs in two contrasting environments. They obviously thrive in the mild, moist climate of the Pacific coast, and they also prosper, if less luxuriantly, in the harsh cold, dry climate of the subarctic.

Consider the Pacific coast rain forest first. The mild winters and abundant rain create an ideal climate for many trees, not only for conifers. Poplars, aspens, and alders (about which more in chapter 5) flourish in rain forest and grow much larger than do inland representatives of their species. But the magnificent conifers, the enormous trees that arouse the wonder of everyone who sees them, greatly outnumber the broadleafs.

It is believed that it is their evergreenness that makes conifers superior to broadleafs in the Pacific Northwest. Because they have green leaves all through the year, conifers can perform photosynthesis whenever the weather is warm enough. They can profit from warm spells in late fall, early spring, and even in winter, when the broadleafs are leafless. Therefore, they are actively growing during a much larger fraction of the year than is possible for broadleafs.

For rain forest, the most stressful time of the year comes during the summer drought. This is unusual, for everywhere else in our area winter is more stressful than summer. Water shortage affects both conifers and broadleafs, but the broadleafs are much less able to cope with this adversity because their big, thin leaves dry out more quickly than do the small, needlelike leaves of the conifers (more on this in chapter 4). Thus the annual drought comes at a time when the broadleafs are most vulnerable, the time when they are in leaf.

In the north country, on the other hand, the ability to photosynthesize on any warm day of the year is no use to a tree growing where the only warm days are in summer. There must be some other explanation for the success of conifers in the vastly different conditions of the far north.

The Advantages of Long-Lived Leaves

The success of evergreen conifers vis-à-vis broadleafs in the north probably has less to do with the effect of cold on the trees, and more to do with its effects on the soil. In a cold climate, dead vegetation takes a long time to decay because the bacteria that bring about decay act slowly in the cold. In a word, humus is slow to form. And if the ground is dry, dead vegetation simply dries out, without forming humus. Some dry soils are little more than dry sand, and others are merely thin, dusty coatings over hard bedrock. (For more on forest soil, see chapter 7.)

The question that now arises is, why should conifers do better than broadleafs on poor soils? Once again, evergreen leaves probably confer an advantage. In this case the advantage is that evergreen leaves last for several years. Therefore an evergreen conifer, unlike a deciduous broadleaf, does not have to grow a new set of leaves every spring, and its demands for nourishment are correspondingly less. Moreover, evergreen leaves can begin photosynthesis earlier in the year than leaves on deciduous trees can, because the latter have to grow before they can function.

The deciduous conifers in our area (tamarack and two other larch species, or collectively, larches) are an awkward exception to this generalization. How can they afford to grow new sets of leaves each year? A strong contributing cause is this: Larch leaves are small and widely spaced so that on a given tree, they shade one another to a lesser degree than do those on both evergreen trees and broadleafs. All larch leaves are more or less equally well illuminated and photosynthesize sugars with maximum efficiency.² This accords with the subjective impression of lightness and brightness that larches give, and it is also an objective, scientifically measurable fact. The conclusion is that larches are as well adapted as evergreen conifers to the evergreen forest habitat; they will be treated as honorary evergreens throughout this book.

Compared with broadleafs, conifers lead more frugal lives. Their life processes take place more slowly (details in chapter 6). In short, conifers live on a waste-not-want-not system; broadleafs, on an easy-come-easy-go system. This enables conifers to make do with inferior soils.

Enduring the Cold

If there is one adaptation that is more important than any other to a very far northern tree, it is cold hardiness. Trees and tall shrubs have