February 2002

Insects make up the vast majority of animal species. Photo: Oksana Hlodan.

How is a biological species defined?

Wilson: The biological species concept is a classical concept. It applies, however, to species that reproduce sexually and is as follows: a species is a population or a series of populations of individuals that interbreed freely with one another. In other words, a species is a more-or-less genetically isolated element, evolving on its own.

This concept implies a closed gene pool. Is this universal?

Wilson: It is not universal. It can be applied to the vast majority of animals and a great many flowering plants. It may be applicable to many kinds of microorganisms. But there are many other organisms, especially plants and microorganisms, to which this concept cannot be applied, or at best applied with difficulty. Obviously, it cannot be applied if there is no sexual reproduction.

In the case of asexual species, it’s possible to classify them arbitrarily based on genetic differences. For example, a standard used for bacteria is when two strains of bacteria differ by as much as 30% of their DNA, they can be treated as different species. This is arbitrary, of course, but it’s also reasonably workable.

Can a subspecies be defined as clearly as a species?

Wilson: Not to the same degree. In theory, a subspecies is a geographical race. A species may be divided into geographical races that differ from one another consistently, in a set of diagnostic traits, such as color, length of wing, breeding habits. Overall, subspecies have many difficulties that make them far more an arbitrary category than species.

One of the difficulties is that traits tend to vary separately from one another. Take for example a butterfly species of Eastern Europe. You might find that populations grow progressively larger going from north to south but are darker in color going from east to west. A closer study may reveal that they have more spots on the wing going from southeast to southwest. So, what are you to do? Will you select one or two characteristics and define the subspecies that way? Or do you try to combine many characteristics and, if so, you get a jumble of many subspecies?

Discordance of traits is probably the most difficult aspect of applying the subspecies concept. However, subspecies is still used and it has some validity, especially in the case of island races.

What about chrono, or ancestral, species?

Wilson: A chronospecies is simply a necessity of science because we have to distinguish between populations that live today and populations that are ancestral to them. They lived so long ago that they are very different. For example, we are the species Homo sapiens — and it is fairly certain that we descended directly from another species, Homo erectus — and it would be very confusing, somehow intuitively incorrect, to try to put them together. Even though Homo erectus evolved through stages into Homo sapiens, we can’t apply the biological species concept to this obviously. Nonetheless we want to make a distinction between them, even though it is arbitrary.

What are the origins of biodiversity?

Wilson: First is the apparent cycle that species go through in populating a new area and diversifying. When an island or an archipelago is formed, for example, or an area is cleared by glaciation or other major physical event of its original biodiversity, there is first a flood of immigrant species. They interact with one another and form a community that we call an ecosystem. If the new area is left undisturbed, then typically there is an episode of rapid evolution — an adaptation of new species to the environment. If there is enough area and enough geographically isolated parts in that area to sustain populations that have little contact with one another, then we also have rapid species formation.

A typical example would be Hawaii. Evidence shows that the archipelago filled up with a small number of species and that they evolved and diversified into many species over a relatively short period of time, by which I mean, centuries or thousands of years. That’s short compared to the evolution we see in many other parts of the world.

Diversification finally reaches a level where the fauna and flora become stabilized, with coadaptation occurring. That means that one species is dependent on another, such as one species specialized on feeding on another. At this point, speciation slows down. It’s similar to the growth of an organism - rapid at first with assembly of new parts that interact and finally a level of maturity can be sustained for a long period of time.

The other set of principles of biodiversity have to do with amounts, or what determines the ultimate amounts. We know, for example, that tropical rainforests have many more species per unit area than do the tundra or coniferous forests of the northern hemisphere. The number of species in a given area, i.e., in a square mile or a square hundred miles, increases steadily as one approaches the equator. The number also increases as you go from small islands to large islands, as in the Caribbean.

What determines this increase of species?

Wilson: There appear to be three factors, which I like to refer to as ESA. E is for energy, S for stability, and A for area.

The more energy that is available to the evolving community of species, the more species there are. That maximizes as you go towards the equator. The more stable a region, as in a constant-climate area, the more species accumulate because they have more time to adapt and fit together. The larger the area, the larger the population and the more diverse it is. For example, South America has more species than the West Indies.

These three driving forces working together appear to account for the large amount of variation of species in the world.

What is the most prevalent form of speciation?

Wilson: If sympatric speciation occurs so widely in insects — the most diverse organisms on earth — as our studies suggest it might, it could be the most prevalent form of speciation. Sympatric speciation is a more difficult phenomenon to study than the easily observed and clearly observed allopatric speciation but it could be very prevalent, we don’t know.

Sympatric refers to similar organisms in close proximity that don’t interbreed because of differences in behavior, even though they theoretically could. Allopatric refers to similar organisms that don’t interbreed, even though they theoretically could, because they are geographically separated.

How quickly can these and other mechanisms produce new species?

Wilson: Instantly. Well, in a few generations. First of all, in plants, there’s a mechanism — called polyploidy — which, in one step, can create a strain that can’t breed with the original stock from which it came. In fact, it’s called instantaneous speciation.

Sympatric speciation can sometimes also take only a few steps to produce new species in a short period time. For example, in certain kinds of fruitflies one strain can prefer to breed on a different kind of plant, and that can happen over several mutations. Another strain comes to breed at a different season, and that can happen by several mutations. Or incompatibility can exist between two strains that differ by one gene or a very small number of genes. This can occur by mutation or recombination of existing genes in a short period of time.

So, it is theoretically possible to create new species within a few years and probably some species formation does occur that rapidly but it is difficult to observe. We are not certain how widespread very rapid speciation really is.

In 1966, you and Dr. Daniel Simberloff conducted an experiment to determine the Minimum Critical Size of Ecosystems (MCS). What did you discover?

Wilson: We were looking to determine how fast species invade an empty island and whether or not they would be affected by area or distance. What we did was to miniaturize the system in the Florida Keys by selection of mangrove islets and then remove all the organisms except trees. We then observed the return of the insects. We demonstrated that

1. Small creatures, including insects and spiders, came back very rapidly.
2. The island filled up with species until it reached approximately the number of species it had before.
3. The more distant the area for immigrant species, the longer it took to reach equilibrium in the area.
4. Although the number of species came back to original level, the composition of the community differed from the original.

This is consistent with our theoretical conception of the turnover of species, that is, extinction followed by colonization (and in the long term, speciation).

Is there is a threshold below which populations are in imminent danger of extinction?

Wilson: Yes. Population size is critical to survival. Generally speaking, when populations get below about 100 individuals, then inbreeding depression happens. And, if there are deleterious, lethal genes in the populations, e.g., cystic fibrosis in humans, then you will get a higher incidence of this genetic trait, which leads to death or sterility. In large populations, chances are that a lethal gene will occur less often.

Conservation biologists offer a 50-500 rule for the genetic health of populations. As I explain this in my 1992 book, The Diversity of Life: “a population of 50 or more is adequate for the short term only, and one of 500 is needed to keep the species alive and healthy into the distant future.”

If speciation can happen quickly, why should we worry about species extinction?

Wilson: It can occur rapidly theoretically but the product would relatively “cheapen” species differences. For example, if two species of fish differed only by one or a few genes out of the tens of thousands that they have, chances are they would differ very little from one another. Compare this to two species that diverged a million years ago and have been evolving genetically in many genes and traits. If these were wiped out, we would lose a great deal more than losing two species that differ only a little.

In any case, whether species are slightly different or markedly different, they are now disappearing at the rate somewhere of a thousand times faster than they are born due to human activity. At this rate, in one human lifetime, we can easily eliminate half the species of the world. Many of these have developed over thousands or millions of years. The clearly demarcated species that can be tracked in the fossil record, before humanity originated, appeared roughly on the order of about one species per million per year. Any rapid process of speciation that would let them begin again can’t duplicate that.

With extinction happening globally, where should we focus conservation efforts?

Wilson: On the hot spots, such as tropical forests. Hot spots are the habitats that are most endangered and have the largest number of species found nowhere else but in them. These include the forests of Hawaii and Madagascar and the rich scrub lands of southwestern Australia and southern Africa. Tropical wildernesses, such as the Amazon and the Congo, have the last of the great frontier forests able to support a mega fauna, i.e., large mammals and birds. The preservation of these places is critical.

Another area is the freshwater systems of the world, which are generally neglected. They deserve special attention because they are under the heaviest assault everywhere, as for example from pollution or drainage. These systems have the largest concentration of endangered species per unit area of any ecosystem in the world. Their equivalent in oceans is the coral reefs, a large percentage of which are now being destroyed or severely degraded. They also have a high percentage per unit of area of endangered species.

In your new book Future of Life you deflate the myth that environmental policy is hostile to economic growth. Can you elaborate?

Wilson: The living resources of the world — ecosystems and its species — are still largely unexplored, much less studied for the benefits they might hold for humans, for example, new pharmaceuticals or water purification. Some ecologists and economists have estimated that the total value of these natural ecosystems, that’s the total amount of services they provide to humanity, is in the vicinity of 30 trillion dollars a year. That’s more than the total of the gross national products of all nations combined. And it’s free!

To save and make fuller use of them in a non-obtrusive way is economically valuable to us. To destroy them is to force humanity into an artificial world in which we have to personally manage our water systems, our food supply, and our atmosphere by prosthetic devices day by day instead of relying on powerful organisms to do the work for us. Do we want to turn Earth literally into a spaceship that requires constant tinkering?

Editor’s Note (11/02): The first World Atlas of Biodiversity: Earth’s Living Resources for the 21st Century was released by the United Nations Environment Programme World Conservation Monitoring Centre (UNEP-WCMC) in August 2002 (University of California Press). Its writers estimate that:

• during the past 150 years, humans have directly impacted and altered close to 47% of the global land area
• under one bleak scenario, biodiversity will be threatened on almost 72% of Earth’s land area by 2032
• 48% of South East Asia, the Congo Basin, and parts of the Amazon will likely be converted to agricultural land, plantations and urban areas — compared with 22% today, suggesting wide depletions of biodiversity
• starting some 45,000 years ago a high proportion of larger land animals became extinct in North America, Australia, the Caribbean, and elsewhere, coinciding with human arrival

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