Plant communities assemble themselves flexibly, and their particular structure depends on the specific history of the area. Ecologists use the term “succession” to refer to the changes that happen in plant communities and ecosystems over time. The first community in a succession is called a pioneer community, while the long-lived community at the end of succession is called a climax community. Pioneer and successional plant communities are said to change over periods from 1 to 500 years. These changes—in plant numbers and the mix of species—are cumulative. Climax communities themselves change but over periods of time greater than about 500 years.
An ecologist who studies a pond today may well find it relatively unchanged in a year’s time. Individual fish may be replaced, but the number of fish will tend to be the same from one year to the next. We can say that the properties of an ecosystem are more stable than the individual organisms that compose the ecosystem.
At one time, ecologists believed that species diversity made ecosystems stable. They believed that the greater the diversity the more stable the ecosystem. Support for this idea came from the observation that long-lasting climax communities usually have more complex food webs and more species diversity than pioneer communities. Ecologists concluded that the apparent stability of climax ecosystems depended on their complexity. To take an extreme example, farmlands dominated by a single crop are so unstable that one year of bad weather or the invasion of a single pest can destroy the entire crop. In contrast, a complex climax community, such as a temperate forest, will tolerate considerable damage from weather to pests.
The question of ecosystem stability is complicated, however. The first problem is that ecologists do not all agree what “stability” means. Stability can be defined as simply lack of change. In that case, the climax community would be considered the most stable, since, by definition, it changes the least over time. Alternatively, stability can be defined as the speed with which an ecosystem returns to a particular form following a major disturbance, such as a fire. This kind of stability is also called resilience. In that case, climax communities would be the most fragile and the least stable, since they can require hundreds of years to return to the climax state.
Even the kind of stability defined as simple lack of change is not always associated with maximum diversity. At least in temperate zones, maximum diversity is often found in mid-successional stages, not in the climax community. Once a redwood forest matures, for example, the kinds of species and the number of individuals growing on the forest floor are reduced. In general, diversity, by itself, does not ensure stability. Mathematical models of ecosystems likewise suggest that diversity does not guarantee ecosystem stability—just the opposite, in fact. A more complicated system is, in general, more likely than a simple system to break down. A fifteen-speed racing bicycle is more likely to break down than a child’s tricycle.
Ecologists are especially interested to know what factors contribute to the resilience of communities because climax communities all over the world are being severely damaged or destroyed by human activities. The destruction caused by the volcanic explosion of Mount St. Helens, in the northwestern United States, for example, pales in comparison to the destruction caused by humans. We need to know what aspects of a community are most important to the community’s resistance to destruction, as well as its recovery.
Many ecologists now think that the relative long-term stability of climax communities comes not from diversity but from the “patchiness” of the environment, an environment that varies from place to place supports more kinds of organisms than an environment that is uniform. A local population that goes extinct is quickly replaced by immigrants from an adjacent community. Even if the new population is of a different species, it can approximately fill the niche vacated by the extinct population and keep the food web intact.
植物群体可以自由地聚集，它们特殊的结构取决于聚集区域的具体历史。生态学家使用“演替”来诠释植物群落和生态系统随着时间推移所发生的变化。演替中的第一个群落被称作先锋群落，而处于演替最后那个长期生存的群落被称为顶极群落。先锋群落和紧接着的植物群落的变化周期是从1到500年不等，植物数量和混合种类数量的变化是慢慢积累的。顶极群落本身也改变，但其变化周期超过500年。 现代一个研究池塘的生态学会发现池塘在一年当中相对而言是不变的。个别鱼类可能被替换，但年复一年鱼的总数都趋于一致。也就是说，一个生态系统自身的属性要比由单一生物体组成的生态系统更稳定。 生态学家们一度认为物种的多样性使生态系统稳定，生态系统物种越多样则生态系统越稳定。通过观察得出的结论支持了这个观点，长期持久的顶极群落通常要比先锋群落具备更为复杂的食物网和更多的物种。生态学家家们得出的结论是：顶点生态系统的稳定性明显取决于他们的复杂化程度。举个极端的例子，在单一作物的农田中，一年的恶劣天气或单一害虫的入侵就可以摧毁所有作物。与此相反，在一个复杂的顶极群落里，如温带森林，他们便可以抵御来自气候和害虫的干扰和入侵。 不管怎样，生态系统稳定性的问题非常复杂。首先，不是所有的生态学家都赞同“稳定”的含义。稳定性可以简单地定义为缺乏变化。如果是这样的话，顶极群落将被视为最稳定的，因为根据定义，他们随着时间推移而变化得最少。另外，稳定性也可以界定为生态系统在经历了严重破坏之后回复原貌的速度，比如火灾。这种稳定性也被称作弹性。在这种情况下，顶极群落将是最脆弱和最不稳定的，因为他们可能需要数百年时间才能恢复到顶点状态。 即使是这种被定义为简单地缺乏变化的稳定性并非总是与最多样的物种联系起来。至少在温带地区，会经常在演替过程中发现最多物种，而不是在顶极群落中。例如，红杉树林一旦成熟，其中的物种数量以及单个物种的数量都会减少。一般来说，多样性本身并不能保证稳定性（事实上正相反），生态系统的数学模型也可以得出同样的结论。一个更复杂的系统可能比一个简单的系统更容易被破坏（一个十五速的赛车比一个孩子的三轮车更容易损坏）。 生态学家们更想弄清楚到底哪些因素有助于促成群落的恢复，因为世界各地的顶极群落都因为人类活动而遭受到严重的损坏或毁坏。就像美国西北部圣海伦火山的猛烈喷发所造成的破坏，在人类活动对环境造成的破坏面前也相形见绌。我们必须了解对群落抵抗、破坏和恢复来说哪些是最重要的。 现在的很多生态学家们认为，顶极群落相对长期的稳定性并非来自于多样性，而是来自环境的“补缀”，随处变化的环境比始终如一的环境更有利于多种有机体的生存。当地物种灭亡后，马上就会被相邻群落的移民取代。即便是另一种不同的物种，他们也可以填补那些已灭绝生物的空缺，并保持食物网的完整。