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The Black Hills forest, the prairie riparian forest, and other forests of the western United States can be separated by the distinctly different combinations of species they comprise. It is easy to distinguish between prairie riparian forest and Black Hills forest—one is a broad-leaved forest of ash and cottonwood trees, the other is a coniferous forest of ponderosa pine and white spruce trees. One has kingbirds; the other juncos (birds with white outer tail feathers). The fact that ecological communities are indeed, recognizable clusters of species led some early ecologists, particularly those living in the beginning of the twentieth century, to claim that communities are highly integrated, precisely balanced assemblages. This claim harkens back to even earlier arguments about the existence of a balance of nature, where every species is there for a specific purpose, like a vital part in a complex machine. Such a belief would suggest that to remove any species, whether it be plant, bird, or insect, would somehow disrupt the balance, and the habitat would begin to deteriorate. Likewise, to add a species may be equally disruptive.

One of these pioneer ecologists was Frederick Clements, who studied ecology extensively throughout the Midwest and other areas in North America. He held that within any given region of climate, ecological communities tended to slowly converge toward a single endpoint, which he called the "climatic climax". This "climax" community was, in Clements's mind, the most well-balanced, integrated grouping of species that could occur within that particular region. Clements even thought that the process of ecological succession—the replacement of some species by others over time—was somewhat akin to the development of an organism, from embryo to adult. Clements thought that succession represented discrete stages in the development of the community (rather like infancy, childhood, and adolescence), terminating in the climatic "adult" stage, when the community became self-reproducing and succession ceased. Clements's view of the ecological community reflected the notion of a precise balance of nature.

Clements was challenged by another pioneer ecologist, Henry Gleason, who took the opposite view. Gleason viewed the community as largely a group of species with similar tolerances to the stresses imposed by climate and other factors typical of the region. Gleason saw the element of chance as important in influencing where species occurred. His concept of the community suggests that nature is not highly integrated. Gleason thought succession could take numerous directions, depending upon local circumstances.

[■]Who was right? [■] Many ecologists have made precise measurements, designed to test the assumptions of both the Clements and Gleason models. [■] For instance, along mountain slopes, does one life zone, or habitat type, grade sharply or gradually into another? [■] If the divisions are sharp, perhaps the reason is that the community is so well integrated, so holistic, so like Clements viewed it, that whole clusters of species must remain together. If the divisions are gradual, perhaps, as Gleason suggested, each species is responding individually to its environment, and clusters of species are not so integrated that they must always occur together.

It now appears that Gleason was far closer to the truth than Clements. The ecological community is largely an accidental assemblage of species with similar responses to a particular climate. Green ash trees are found in association with plains cottonwood trees because both can survive well on floodplains and the competition between them is not so strong that only one can persevere. One ecological community often flows into another so gradually that it is next to impossible to say where one leaves off and the other begins. Communities are individualistic.

This is not to say that precise harmonies are not present within communities. Most flowering plants could not exist were it not for their pollinators—and vice versa. Predators, disease organisms, and competitors all influence the abundance and distribution of everything from oak trees to field mice. But if we see a precise balance of nature, it is largely an artifact of our perception, due to the illusion that nature, especially a complex system like a forest, seems so unchanging from one day to the next.

EXTRA1 Look at the four squares [▇] that show where the following sentence could be inserted in the passage. Where could the sentence best be added? Click on a square [▇] to insert the sentence in the passage.

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