273 Biodiversity

Traditionally, ecologists have measured biodiversity, a general term for the variety present in the biosphere, by taking into account both the number of species and their commonness. Biodiversity can be estimated at a number of levels of organization of living things. These estimation indexes, which came from information theory, are most useful as a first step in quantifying biodiversity between and within ecosystems; they are less useful when the main concern among conservation biologists is simply the loss of biodiversity. However, biologists recognize that measures of biodiversity, in terms of species diversity, may help focus efforts to preserve the biologically or technologically important elements of biodiversity.

The Lake Victoria cichlids (Figure 1) provide an example through which we can begin to understand biodiversity. The biologists studying cichlids in the 1980s discovered hundreds of cichlid species representing a variety of specializations to particular habitat types and specific feeding strategies: eating plankton floating in the water, scraping and then eating algae from rocks, eating insect larvae from the bottom, and eating the eggs of other species of cichlid. The cichlids of Lake Victoria are the product of an adaptive radiation. An adaptive radiation is a rapid (less than three million years in the case of the Lake Victoria cichlids) branching through speciation of a phylogenetic tree into many closely related species; typically, the species “radiate” into different habitats and niches. The Galápagos finches are an example of a modest adaptive radiation with 15 species. The cichlids of Lake Victoria are an example of a spectacular adaptive radiation that includes about 500 species.

At the time biologists were making this discovery, some species began to quickly disappear. A culprit in these declines was a species of large fish that was introduced to Lake Victoria by fisheries to feed the people living around the lake. The Nile perch was introduced in 1963, but lay low until the 1980s when its populations began to surge. The Nile perch population grew by consuming cichlids, driving species after species to the point of extinction (the disappearance of a species). In fact, there were several factors that played a role in the extinction of perhaps 200 cichlid species in Lake Victoria: the Nile perch, declining lake water quality due to agriculture and land clearing on the shores of Lake Victoria, and increased fishing pressure. Scientists had not even catalogued all of the species present—so many were lost that were never named. The diversity is now a shadow of what it once was.

The cichlids of Lake Victoria are a thumbnail sketch of contemporary rapid species loss that occurs all over Earth and is caused by human activity. Extinction is a natural process of macroevolution that occurs at the rate of about one out of 1 million species becoming extinct per year. The fossil record reveals that there have been five periods of mass extinction in history with much higher rates of species loss, and the rate of species loss today is comparable to those periods of mass extinction. However, there is a major difference between the previous mass extinctions and the current extinction we are experiencing: human activity. Specifically, three human activities have a major impact: destruction of habitat, introduction of exotic species, and over-harvesting. Predictions of species loss within the next century, a tiny amount of time on geological timescales, range from 10 percent to 50 percent. Extinctions on this scale have only happened five other times in the history of the planet, and they have been caused by cataclysmic events that changed the course of the history of life in each instance. Earth is now in one of those times.

Types of Biodiversity

Scientists generally accept that the term biodiversity describes the number and kinds of species in a location or on the planet. Species can be difficult to define, but most biologists still feel comfortable with the concept and are able to identify and count eukaryotic species in most contexts. Biologists have also identified alternate measures of biodiversity, some of which are important for planning how to preserve biodiversity.

Genetic diversity is one of those alternate concepts. Genetic diversity or variation is the raw material for adaptation in a species. A species’ future potential for adaptation depends on the genetic diversity held in the genomes of the individuals in populations that make up the species. The same is true for higher taxonomic categories. A genus with very different types of species will have more genetic diversity than a genus with species that look alike and have similar ecologies. If there were a choice between one of these genera of species being preserved, the one with the greatest potential for subsequent evolution is the most genetically diverse one. It would be ideal not to have to make such choices, but increasingly this may be the norm.

Many genes code for proteins, which in turn carry out the metabolic processes that keep organisms alive and reproducing. Genetic diversity can be measured as chemical diversity in that different species produce a variety of chemicals in their cells, both the proteins as well as the products and byproducts of metabolism. This chemical diversity has potential benefit for humans as a source of pharmaceuticals, so it provides one way to measure diversity that is important to human health and welfare.

Humans have generated diversity in domestic animals, plants, and fungi. This diversity is also suffering losses because of migration, market forces, and increasing globalism in agriculture, especially in heavily populated regions such as China, India, and Japan. The human population directly depends on this diversity as a stable food source, and its decline is troubling biologists and agricultural scientists.

It is also useful to define ecosystem diversity, meaning the number of different ecosystems on the planet or in a given geographic area (Figure 2). Whole ecosystems can disappear even if some of the species might survive by adapting to other ecosystems. The loss of an ecosystem means the loss of interactions between species, the loss of unique features of coadaptation, and the loss of biological productivity that an ecosystem is able to create. An example of a largely extinct ecosystem in North America is the prairie ecosystem. Prairies once spanned central North America from the boreal forest in northern Canada down into Mexico. They are now all but gone, replaced by crop fields, pasture lands, and suburban sprawl. Many of the species survive, but the hugely productive ecosystem that was responsible for creating the most productive agricultural soils is now gone. As a consequence, soils are disappearing or must be maintained at greater expense.

Patterns of Biodiversity

Biodiversity is not evenly distributed on Earth. Lake Victoria contained almost 500 species of cichlids alone, ignoring the other fish families present in the lake. All of these species were found only in Lake Victoria; therefore, the 500 species of cichlids were endemic. Endemic species are found in only one location. Endemics with highly restricted distributions are particularly vulnerable to extinction. Higher taxonomic levels, such as genera and families, can also be endemic. Lake Huron contains about 79 species of fish, all of which are found in many other lakes in North America. What accounts for the difference in fish diversity in these two lakes? Lake Victoria is a tropical lake, while Lake Huron is a temperate lake. Lake Huron in its present form is only about 7,000 years old, while Lake Victoria in its present form is about 15,000 years old. Biogeographers have suggested these two factors, latitude and age, are two of several hypotheses to explain biodiversity patterns on the planet.

Biogeographer

Biogeography is the study of the distribution of the world’s species—both in the past and in the present. The work of biogeographers is critical to understanding our physical environment, how the environment affects species, and how environmental changes impact the distribution of a species; it has also been critical to developing evolutionary theory. Biogeographers need to understand both biology and ecology. They also need to be well-versed in evolutionary studies, soil science, and climatology.

There are three main fields of study under the heading of biogeography: ecological biogeography, historical biogeography (called paleobiogeography), and conservation biogeography. Ecological biogeography studies the current factors affecting the distribution of plants and animals. Historical biogeography, as the name implies, studies the past distribution of species. Conservation biogeography, on the other hand, is focused on the protection and restoration of species based upon known historical and current ecological information. Each of these fields considers both zoogeography and phytogeography—the past and present distribution of animals and plants.

One of the oldest observed patterns in ecology is that species biodiversity in almost every taxonomic group increases as latitude declines. In other words, biodiversity increases closer to the equator (Figure 3).

It is not yet clear why biodiversity increases closer to the equator, but hypotheses include the greater age of the ecosystems in the tropics versus temperate regions that were largely devoid of life or drastically impoverished during the last glaciation. The idea is that greater age provides more time for speciation. Another possible explanation is the increased energy the tropics receive from the sun versus the decreased energy that temperate and polar regions receive. It is not entirely clear how greater energy input could translate into more species. The complexity of tropical ecosystems may promote speciation by increasing the heterogeneity, or number of ecological niches, in the tropics relative to higher latitudes. The greater heterogeneity provides more opportunities for coevolution, specialization, and perhaps greater selection pressures leading to population differentiation. However, this hypothesis suffers from some circularity—ecosystems with more species encourage speciation, but how did they get more species to begin with? The tropics have been perceived as being more stable than temperate regions, which have a pronounced climate and day-length seasonality. The tropics have their own forms of seasonality, such as rainfall, but they are generally assumed to be more stable environments and this stability might promote speciation.

Regardless of the mechanisms, it is certainly true that all levels of biodiversity are greatest in the tropics. Additionally, the rate of endemism is highest, and there are more biodiversity hotspots. However, this richness of diversity also means that knowledge of species is lowest, and there is a high potential for biodiversity loss.