The Meaning of Biodiversity

William Silvert
Instituto Nacional de Investigao Agrria e das Pescas IPIMAR, Avenida de Braslia, s/n 1449-006 Lisboa, Portugal silvert@ipimar.pt

Keywords: Biodiversity, extinction, conservation, society, politics

Abstract
The concept of biodiversity is widely used but poorly defined. Although there exist formal definitions, they are so broad and vague that their long-term effectiveness in environmental management is questionable. Some countries interpret conservation of biodiversity so rigorously that all development can be blocked by the prospect of the extinction or loss of critical habitat of any single species it seems unlikely that this policy will prevail, at least not until the far greater problems of human population growth and industrial expansion are resolved. Furthermore, the tendency to define biodiversity in terms of species richness is a poor political strategy in marine ecosystems, since global (as opposed to local) extinction is virtually unknown in marine populations, with the exception of mammals and birds. In order to defend biodiversity in an effective way we need to present the concept in a coherent fashion that pragmatic politicians and members of the general public can understand and evaluate from the perspective of their own values; this will provide the basis for negotiated rather than adversarial settlement of conflicts between biodiversity and human activities. In order to achieve this we need to move away from the traditional definitions of biodiversity, often expressed in terms of obscure taxonomic distinctions, and identify socially meaningful effects of species and habitat loss. For example, agronomists have long promoted biodiversity by arguing that monocultures are prone to disease, and pharmaceutical companies have publicised the number of new drugs that they have discovered in obscure species. However, it is unlikely that we will be able to conserve all existing species, and we must be prepared to demonstrate the probable consequences of all activities that are likely to have environmental impacts. This paper describes a strategy for analysing, presenting and defending biodiversity.

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William Silvert, The Meaning of Biodiversity. Talk at EMBS38.

Introduction
Biodiversity has emerged as a scientific topic with a high degree of social prominence and consequently of political importance. To the extent that scientists wish to see programs to conserve biodiversity implemented, they must be aware of the political aspects of the problem and be prepared to make the compromises that are necessary when one enters into the political arena. The need to compromise with public opinion is well recognised by NGOs (NonGovernmental Organisations) and other groups which practice environmental activism, and often the result is a political agenda that has only a tenuous connection with scientific reality. An example is the issue of avoiding unnecessary suffering to animals although most scientists believe that clubbing animals to death is actually less painful than most other forms of slaughter, there is widespread public opposition to the clubbing of cute white harp seals but relatively little concern about how other types of seals, which are usually ugly and grey, and domestic cattle are killed. Scientists are familiar with the pressure to treat experimental animals, such as pretty white experimental mice, in a humane fashion, and yet few voices are raised against the incredibly cruel methods used to exterminate rodents when they are seen as pests. Conservationists face similar issues just as campaigns against cruelty to animals often deal more with the human appeal of the animal than with its actual suffering, so species loss is often seen more in terms of the attractiveness of the species to humans than any biological factors. This is a practical reality that we have to address, and if we hope to accomplish the kinds of conservation goals that many scientists support, we need to understand how to integrate these with the often irrational processes that go into the formulation of public policy.

Why focus on biodiversity?

The current public emphasis on biodiversity is fairly new, and it is interesting to consider how the concept developed. During the past half-century or so there has been a growing awareness of the importance of natural ecosystems and a desire to conserve rather than simply exploit our environment. The concept probably goes back to foresters and game managers like Aldo Leopold (1933) but came to the public attention with popular books like Fairfield Osborns Our Plundered Planet (1948). During the 1960s there was an explosion of literature about the kinds of damage we were doing to the living world, such as Rachel Carsons Silent Spring (1962) and Wesley Marx The Frail Ocean (1967), which led both to public awareness of the value of conservation and to pressure for public action (Mitchell and Stallings 1970). The focus at the time was naturally limited to issues that were easily understood, such as conservation of easily recognised species this had been a matter of public concern at least since the founding of the Audubon Society in 1886 (Graham 1990). Over time our appreciation of ecology has developed and most people today appreciate that the lowly earthworm has as much importance (actually more) as beautiful egrets and cuddly pandas. So the next step in public understanding was that all species play a role in the global ecosystem and should be conserved. Even for those species whose merit was not clear, the terrible finality of extinction is forever is so alarming that steps were taken to conserve all species, such as the U. S. Endangered Species Act of 1973 which states that the United States has pledged

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William Silvert, The Meaning of Biodiversity. Talk at EMBS38. itself as a sovereign state in the international community to conserve to the extent practicable the various species of fish or wildlife and plants facing extinction. The problem with all this is that extinction is a natural phenomenon which occurs even in the absence of human pressure, and evolutionary history is a record of species being wiped out by natural occurrences or being superseded by better adapted life forms (for example, the replacement of dinosaurs by warm-blooded mammals). Trying to conserve all species increasingly appears to be a futile task, and there is growing awareness that we need to be more judgemental in our approach to conservation. Rather than simply seeking to conserve everything, we have to decide when to fight and when to let go. One option is to reduce the emphasis on individual species and focus on biodiversity, a vague concept but one which captures the idea that we dont want a lot of species to go extinct. Because of the transition from conserving species to conserving biodiversity, much attention has been directed to tropical gulfs and forests and other hotspots where many different species are to be found. However, this too has not proved to be a perfect target for political action, since few people would be prepared to downgrade the limited but exciting fauna of the polar regions relative to the species-rich understory of a tropical rain forest. The transition from conserving species to conserving biodiversity is a confusing one, and even in the scientific literature it is not always clear what is meant. However, given the social importance and prominence of biodiversity, it is vital that scientists be able to communicate with the public about what is happening and what the consequences of our actions or inaction in this field are likely to be. To accomplish this we need to develop concepts of biodiversity that are not only scientifically valid, but also meaningful to non-specialists.

Why conserve biodiversity?

Although the question may seem like heresy to an ecologist, we cannot take it for granted that all sectors of society see value in biodiversity. Quite the contrary, humans often deliberately reduce biodiversity to achieve their goals. In many parts of the globe fields of richly varied plant types have been replaced by vast uniform fields of maize, wheat, and other valued crops. Some of these commercial monocultures are even monoclonal, the ultimate in low diversity. Programs of pest control, both agricultural and residential, strive to eliminate unwanted creatures with no concern for the resulting impact on biodiversity. Although it is difficult to exterminate pests in the sea, it has certainly been tried many countries have sought to cull seals and other marine predators that compete with man. Closer to home, few people want a diverse ecosystem flourishing where they live, and those that do keep it in flower pots and behind glass walls. Modern mankind is not always comfortable with nature, and prefers to visit it during holidays rather than share space with it. This has had an impact on biodiversity, since the result has been a drastic reduction in habitat for plants and animals that could quite happily coexist with humans, but are excluded from recreational areas like beaches and golf courses as well as residential areas, garbage dumps, and highways. Even national parks are sued for containing dangerous wild animals, as in a recent case where a woman was killed by a bear (Mansfield 2001). The excessive use of antibacterial products is evidence of our desire to live in a biologically antiseptic environment, even though the end result

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William Silvert, The Meaning of Biodiversity. Talk at EMBS38. is faster evolution of resistant strains. It is hard to avoid the conclusion that many of the people who support biodiversity only support it somewhere else. In almost every case where undesirable species are removed or suppressed no effort is made to consider the impact on biodiversity. Healthy crops or abundant fish are considered far more important than maintaining the diversity of insect fauna or marine mammal populations. The unappealing nature of many vital organisms, such as the vast variety of marine worms, does not contribute to public appreciation of their roles. So it cannot be taken for granted that everyone supports biodiversity, and conservation may at times encounter surprising resistance.

Types of biodiversity
There are many different kinds of definition that have been devised for biodiversity, but only three main categories will be considered here. One is genetic diversity, which can refer to the diversity of genes within a single species as well as between species. Another is taxonomic diversity, based of course on the different taxa contained within an ecosystem. The third is functional diversity, which recognises the variety of roles that different organisms including the separate life stages of individual species play in the ecosystem. The importance of genetic diversity has long been recognised in agriculture, where the danger of disease wiping out a single strain of organism is a constant concern. Many nations maintain seed banks where different plant varieties are maintained, and many examples, such as the importation of Phylloxia-resistant grape vines from California into Europe, show the necessity of maintaining this kind of diversity. Genetic diversity protects ecosystems against other forms of environmental change, not only against disease. Diversity within a species is the driver behind evolutionary adaptation. If we had a species that was optimally suited for a temperature of 17 and the temperature changed to 18 it might be seriously threatened, but because typically the species would include genotypes with a range of temperature optima, we would instead expect to see rapid dominance of the 18 genotype and a decline in the abundance of the lower temperature varieties. Within a few generations the population could be well adapted to the changed environment. Over the long term there might also be favourable mutations which could further enhance the survival of the species, but maintenance of a diverse gene pool is an essential mechanism for adapting to change. The concept of genetic diversity also applies between populations, since species replacement in response to environmental change usually indicates that the replacement species is genetically better suited to the changed conditions. We tend to think of these changes as different, especially when the replacement is a previously rare or exotic species, but the underlying idea that under changed conditions there is likely to be a shift to a population better suited to the changed conditions is fundamentally the same whether they come from the same or different species. Genetic diversity becomes especially important in the context of climate change and other local or global environmental shifts, since it plays a critical role in determining how communities will adapt to stress. If the components of an ecological community have sufficient genetic breadth to adapt, then the community structure will be relatively robust and will probably survive with little change. However, if the community is stressed then it is likely that genetic breadth will be reduced and only

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William Silvert, The Meaning of Biodiversity. Talk at EMBS38. those individuals optimally suited to current conditions will be present, and these are the ones that will be most affected by environmental changes. Taxonomic diversity is probably the most widely recognised form of biodiversity, but it may also be the least meaningful. It can be defined in many ways, but basically it involves identifying the number of different taxa (usually at the species level) and possibly weighting them by the abundance of individuals. Calculations of taxonomic biodiversity tend to be limited by the taxonomic expertise available, especially at lower trophic levels whales are easy to classify, but nematode taxonomists are in short supply. There is a tendency to lump some of the more confusing taxa together at the genus or higher level, which introduces a degree of arbitrariness to the calculations. An additional problem is that the degree of separation between species varies, and it is not always convincing to argue that the difference between two species of isopods is just as significant as that between a shark and a sea anemone. Warwick (2004) considers a type of diversity measure in which the taxonomic distance between species is taken into account, so that an ecosystem in which different families or classes are well represented would have a higher biodiversity than one in which there were many species from a single genus. This resolves some of the concerns about taxonomic differences, although it still seems a bit artificial for example, carnivorous zooplankton turn out to be an incredibly diverse group with many members from several different phyla (such as Cnidaria, Ctenophora, Chaetognatha and Crustacea, to say nothing of various larval stages) even though they do not seem that different from each other in terms of their functioning with the ecosystem. Perhaps the most important form of biodiversity is functional diversity, the kind of diversity that ensures that every task that needs doing within an ecosystem gets done. It doesnt help to have thousands of species of herbivores in a system if there are no primary producers to feed them and no detritivores to clean up after them. But there are major difficulties in defining functional diversity, not least of which is defining what the various functions are and who is capable of carrying them out. In some cases the answer is obvious, as when there is a single top predator or dominant bioturbator in the system the role of the polar bear in the arctic ecosystem, Arenicola in the Wadden sea, Calanus in the Atlantic Ocean or alligators in the Florida Everglades is so clear that these species and genera are easily seen to have clearly defined functional roles. For other taxa, such as the myriad number of benthic worms, it is not clear at all what the differences are between their ecological functions, and for others, such as benign parasites, it is difficult to find any role at all they just seem to be passive participants.

Generalised niche theory

The niche is classically associated with existing populations (Hutchinson 1957) and strict Hutchinsonians reject the concept of an empty niche, which by definition is not associated with any existing organisms. This is however a valuable concept in understanding biodiversity, so it is worth exploring. The potential niche of a population is the range of environmental conditions in which it can persist, the actual niche is the range in which it is currently found (often called the realised niche), and an empty niche represents an environment which could in principle support a population of some sort, but currently does not. The reason that this concept fits in with biodiversity is that it helps to be able to describe an ecosystem where some

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William Silvert, The Meaning of Biodiversity. Talk at EMBS38. species have been removed as containing empty niches, which can represent either ecological roles which are no longer being carried out, or opportunities for invasive species. This can be visualised by analogy to Soviet-era housing, with several families sharing apartments designed for one. The potential niche of each family is the apartment to which it has the key, its actual niche is the room in the apartment to which it has been assigned, and if a room or apartment is vacant it represents an empty niche. A diverse ecosystem is usually one in which the niches are closely packed, in which every room is occupied by a different family to stretch the analogy (Christiansen and Fenchel 1977). Invasion by a super-competitor which can expel many of the native species from their niches, in effect taking over an entire apartment, greatly reduces the diversity of the community. But what are the consequences of this for ecosystem function, and is the loss of diversity always bad? While we cannot make totally sweeping assertions about loss of diversity, and in particular about the replacement of several species occupying small closely-packed niches by a single species with a broad niche, there are certainly grounds for concern. First of all, the success of an invading species is often due to the lack of a predator adapted to remove it, but if its predators eventually manage to follow it the result is destabilisation of the ecosystem. Second, an invader with a broad actual niche which is almost as large as its potential niche is unlikely to exploit it as fully as native species whose actual niches are compressed through competitive pressure. In particular, if environmental conditions shift, an exotic species with a broad niche may have the ability be squeezed into a smaller actual niche leaving some of the environmental hyperspace (to use Hutchinsons phrase) unoccupied in other words, an empty niche might be created. While it might be stressing the housing analogy too far to argue that full occupancy of all the niches in an ecosystem is desirable (to say nothing of appearing to be an endorsement of Soviet-era housing policy!), it is at least a plausible viewpoint. The more species there are competing to utilise all available resources, the greater the potential for full exploitation of the resources and for maximisation of system throughput and productivity. While the idea of defining biodiversity in terms of niches rather than species is speculative, and it will certainly not sit well with some of the classical interpretations of the niche, it offers a potential formalism for investigating biodiversity under conditions of environmental change that may have some advantages over functional biodiversity, since it may be easier to identify changes in the possible niche structure of a changed environment than changes in functional requirements.

Triage
No matter how we choose to define biodiversity, our ability to conserve it is limited. Aside from the inevitability of natural extinction, there are forces beyond the control of the scientific community or environmental agencies which force us to admit that some battles cannot be won. The chances of restraining the exploding human population, industrialisation and consequent loss of natural habitat are nil (Ehrlich 1971). The Horsemen of the Apocalypse, especially war and famine, are as deadly to natural ecosystems as they are to humans. If we commit ourselves to saving every

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William Silvert, The Meaning of Biodiversity. Talk at EMBS38. species on earth we are likely to end up saving none, but by tempering idealism with pragmatism we may achieve at least some success. One pragmatic approach in cases where the challenges are too great is triage, a gruesome but often necessary means of compromise. The concept of triage was developed by Baron Dominique-Jean Larrey (1832), Napoleon's chief surgeon, and is still used today in medical situations where the need for medical attention exceeds the resources available. Basically it consists of directing care to those patients who are seriously at risk but are likely to be saved by medical attention. The walking wounded who are likely to recover in any case and those who are too seriously injured to have a good chance of recovery constitute the other two branches of the triage system and receive minimal treatment. The same approach can be considered in dealing with species at risk. Many species are not endangered and need not be the targets of conservation campaigns. Others are so severely affected that there is almost nothing that can be done to save them, such as those which are restricted to vanishing habitats. Conservation efforts can better be directed at species which are endangered, but can plausibly be saved to pick an early example, the success of the Audubon Society in saving the snowy egret and other bird species is in part due to the fact that the greatest threat came from illegal hunters who killed the birds for their plumes, and the effective response was enforcement of antipoaching legislation coupled with public campaigns to change the fashion of womens hats (Graham 1990). When species are threatened by population growth, famine, loss of critical habitat and war it is far more difficult to save them. In many cases the only way to preserve a rare species is through captive breeding programs, and while these are sometimes effective, maintaining a species in a zoo does little for biodiversity, although these programs, like seed banks, do preserve a measure of genetic diversity. The concept of triage is similar to the use of cost-effectiveness criteria in economics. A battlefield physician confronting three casualties and evaluating the probable outcomes of treating them might conclude that one patient needs treatment to save his eyesight another will probably die anyway, but with a small chance of recovery with treatment and the third is bleeding to death, but medical care would probably save his life. All merit treatment, but if there is only time to treat one, the physician will probably chose saving life over saving eyes, and will pick the patient with the best chance of recovery. This would be the best use of his time. Similarly, the costs of conservation efforts have to measured against the net benefits, which have at times threatened to be negative if the preservation of an endangered species generates so much antagonism that the entire conservation movement suffers. Evidence of this is apparent in the hostility generated when major developments are blocked by concern for species of little public appeal, such as sand flies (Kresge 2003). There is however a factor that can interfere with application of the triage principle, and it applies to conservation of biodiversity as much as to medical practice. If a physician recognises that the wounded soldier who comes for treatment is his son, or a close friend, then he is almost certain to receive treatment no matter what the likely outcome. The same is true of some endangered species no matter how close to extinction they may be, or how great the cost of saving them, they must be saved. The whooping crane (Grus Americana) was at one point down to fewer than two dozen individuals, but millions of dollars have been spent bringing it back from the brink. The emotional factor cannot be ignored.

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William Silvert, The Meaning of Biodiversity. Talk at EMBS38.

Setting value on species

One of the hardest parts of conservation is trying to decide where to direct ones limited effort and resources, and this is especially hard when the outcome is likely to be extinction for the species we choose not to protect. The terrible finality of extinction is a factor that upsets both scientists and laymen, and for the scientific community the prospect of losing a species before we learn about it is doubly distressing. Even so, we have to balance our priorities carefully to make wise choices. Above all, if we want to be politically effective and gain public support for biodiversity conservation, we have to recognise the social factors which can generate support and earn political attention. These may not always tally with scientific priorities, but in the political arena we must learn to compromise. At the top of any list of factors that make a species a good candidate for public support is charisma. Some animals capture the public attention and will always lead the campaign for conservation, regardless of biological concerns. For example, the giant panda (Ailuropoda melanoleuca) is a rare bamboo-eating bear found only in southwest China, and yet is has captured the imagination of the world and has been adopted as the symbol of the WWF (originally the World Wildlife Fund), one of the leading advocacy groups in the conservation movement. Baby harp seals (Phoca groenlandica) have far more charisma than their ugly grey cousins and receive vast sympathy and support. The great whales, elephants, apes, Siberian tigers and many other impressive species are cherished and generate tremendous popular interest. A second factor that again may not correspond to the priorities of scientists, nor of many environmentalists, is the value of a species to man. There has been tremendous concern about the depletion of stocks of Atlantic cod (Gadus morhua) in recent years, but the endangered status of the barndoor skate (Raja laevis), a species of virtually no commercial importance, passed virtually unnoticed (Casey and Myers 1998). Fortunately there is a place for ecological concerns in conservation, and the role of species, their function within the ecosystem, is a matter of growing public awareness. There is for example awareness of the importance of top predators, with the result that even frightening creatures like sharks are now the object of publicly funded conservation programs. Alligators (Alligator mississippiensis) are also protected but public appreciation is probably based more on their fearsome appearance and their predatory status than on their essential role in maintaining habitat through the creation of gator holes, ecologically vital depressions where water remains during dry periods. However, the role of less spectacular creatures is not as well appreciated. The various Calanoid copepods which support the Atlantic food web are virtually unknown outside the scientific community, even though they garner enormous interest within it. Detritivores like amphipods and hagfishes (Myxinidae) are important but repulsive, which makes it hard to garner public support for their conservation. One can probably find few examples of essential species less charismatic than the dung beetles (Scarabaeidae), without which the dry terrains of India and many other countries would be covered with semi-fossilized cattle faeces, and yet the ancient Egyptians appreciated and even revered these extraordinary insects. Some highly unattractive species can penetrate social consciousness, most notably the lowly earthworm. Generating popular support for repulsive animals is not easy, but can be done. However, there are more technical issues about the ecological roles that plants and animals play which probably have to be worked out within the scientific community. -8-

William Silvert, The Meaning of Biodiversity. Talk at EMBS38. Some species truly have unique and essential ecological functions, ranging from the common earthworm up to the mighty polar bear, and it is relatively easy to make a case for their conservation. Many other species though have close competitors which could possibly fill the same ecological role, and thus could be replaced if they went extinct. We know very little about the degree of redundancy within ecosystems, and yet without this information it is almost impossible to predict the effect of the loss of one of a number of closely related species. Gray (2004) points out a lack of correspondence between species richness and marine productivity when the North Sea and the Baltic are compared, but whether we can safely assume that this means that most of the species in the North Sea are redundant is questionable. This is certainly one of the areas of biodiversity research that needs attention in future years. All of these concerns ignore a factor that does not necessarily fall under the heading of biodiversity, at least as it is sometimes defined, but which seems critical to any discussion of species conservation uniqueness. Some species are simply unique. There is nothing like them, and regardless of whether they are charismatic or play some essential role, their loss would greatly reduce the variety of nature. Some, like the coelacanth, are living fossils, antiquities from millions of years ago. Others, like the placozoan Trichoplax adhaerens, the loriceferan Nanoloricus mysticus and the cycliophoran Symbion pandora, are the only members of their respective phyla and appear to have virtually no relation to anything else in the animal kingdom. Surely any meaningful measure of biodiversity would recognise the importance to conserve these cryptic creatures. The factors described above are quite different from some of the considerations raised in official biodiversity documents. For example, the Texel-Faial criteria (OSPAR 2003) do not provide a great deal of guidance on which species should receive priority in conservation (they mostly deal with identifying whether a species is endangered), aside from criterion 5 which defines Keystone species: a species which has a controlling influence on a community. Control can be good or bad, and it is not always clear how this criterion should be applied. Clearly the definition applies to top predators like sharks and polar bears, but it also applies to lampreys and zebra mussels in the Great Lakes. One can reasonably ask to what extent a global definition of desirable species is even possible, or whether conservation should be considered on a case-by-case basis. Given the complexity of the criteria it seems unlikely that a universal rating system could be developed, but clearly this is an objective to which many scientists are committed.

Obstacles to conservation
There are many obstacles that make the life of a conservationist hard, and it is important to understand the rationale behind resistance to conservation efforts. Although scientists tend to view all species and habitats with the enthusiasm that a thirst for knowledge generates, this attitude is not widely shared, and often the environments that most intrigue the scientific community, such as marsh lands, generate little excitement among the population at large. In a recent story about the possible extinction of a sand fly (Kresge 2003) the interviews with entomologists are summarised as, Scientists believe the fly spends most of its life buried a meter or two deep in the sand, and no one knows for sure what it does there. Although the insect was first collected around 1890 and described as a new species in 1941, much of its life history remains a mystery. While they go

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William Silvert, The Meaning of Biodiversity. Talk at EMBS38. on to suggest that it is a key species in a unique habitat, it can easily be seen that the conservation effort is not going down well with the local community: Frustrated city officials tick off the projects the fly has slowed, changed or derailed entirely. A manufacturing project. The county hospital. A cemetery expansion. An interchange. The result, they say, is lost opportunity, lost sales tax and lost property rights. While the case of the endangered fly in California may seem extreme, there are many cases where conservation efforts engender enormous secondary costs, either by lessening support in general for conservation or sometimes by the need for financial support to affected communities. The governments of the United States and California do not offer compensation to developers whose projects are blocked by habitat concerns, but in countries like India, where protected elephant species can destroy crops, villages, and human lives, the costs and liabilities must be viewed as a governmental responsibility (Bist 2002). Other endangered species, such as some of the great apes and whales, are traditional food sources for native communities.

Exotic species
In speaking of biodiversity we tend to focus uncritically on conservation of species, but there are some species that might arguably be eliminated from the biosphere. There is general agreement that certain disease organisms should be eliminated, such as those responsible for smallpox and poliomyelitis, with only a few reservoirs maintained so that if they reappear we can produce vaccines. There would probably be few opponents to a campaign to rid the world of other noxious pests, such as malarial mosquitoes, but there are many species offensive to some which play useful ecological roles or are even prized by others. House flies are obnoxious pests and it is disgusting to see their maggots in our trash, but it is still a blessing that something is willing to digest and recycle garbage. Lampreys are considered a terrible parasite in the Great Lakes and are exterminated at great cost, but in parts of Europe they are considered a delicacy fit for the finest gourmets. And even the possibility of eradicating some of the 30 deadly mosquito species that carry the malaria parasite (out of about 2,500 known mosquito species) is controversial and subject to criticism (Judson 2003). We tend to think of non-native species as irrelevant to calculations of biodiversity, but this raises the question of how long an invasive species must be present before it is considered a native. No species has been present forever, so we need to ask how long a species has been present, and how well integrated it is, in order to evaluate its place in the biodiversity debate. The mustang is a wild horse introduced to the Americas by the Spanish conquistadors within the past 500+ years, and there are grounds for debating whether the feral stock should now be considered an indigenous species worth protecting, or an exotic to be harvested and possibly eliminated. Some marine species like Macoma balthica were presumably transported to North America on the hulls of early ships and are now well established in the Gulf of Maine and other sites. There is a common sense that non-native species always reduce biodiversity, but this probably reflects the fact that it is the invaders which do the greatest harm that attract the most attention rabbits in Australia, goats in the Galapagos, water hyacinth in American waterways, mosquitoes in Hawaii, zebra mussels in the Great Lakes are only a few examples. There are probably many exotic species which fit in well to new environments and may not even be identified as foreign. Should we take these species

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William Silvert, The Meaning of Biodiversity. Talk at EMBS38. into account when calculating biodiversity? The answer is not clear, and this underscores the fact that we have to consider the actual consequences of any change in community composition rather than focussing on measures of biodiversity, no matter how well thought out they may be. In some cases the non-native species are a natural replacement for native species which have become extinct. It is likely that many indigenous species of earthworm and similar invertebrates in Canada and the northern United States became extinct during glaciation periods and that some of the existing species originated in Europe, but certainly no one would argue that earthworms should be exterminated because they are not part of the native fauna. The line between native and introduced species is rarely clear, and adds a further challenge to any attempt to quantify biodiversity. Even for recent introductions we need to ask whether the process is reversible even if it is possible to turn back the clock by removing an invasive species, we need to determine whether the system will return to its previous state. If native species have gone extinct or been severely depleted, they may never recover.

Speciation factories
Some of the attention in biodiversity studies has focussed on biodiversity hotspots, regions where enormous numbers of distinct species can be found. If our objective is to save many species, then certainly we should focus our attention on areas like tropical rainforests and the Gulf of Thailand where there are far more species to save than, say, in polar and temperate regions. This viewpoint should be looked at critically, and it is not always clear what the presence of many different species means in terms of any reasonable measure of ecosystem performance. Consider for example an archipelago or any other system of separated islands. In almost every case we find that each island has several unique species, usually of organisms like snails which diffuse slowly. Although unique, these species are often closely related and functionally similar. Suppose that one of these species becomes extinct, and a member of the same genus is brought from a neighbouring island will the ecosystem collapse? Will its ecological niche remain vacant? Or will the imported species happily adapt and perhaps after a few generations evolve into yet another unique species? Although this is an open matter for conjecture (if the experiment has been done, this author has not seen the work), it seems plausible. We do not really understand what determines the number of species in an ecosystem, or why some systems have many more than others (Hutchinson 1959). Speciation can occur through many mechanisms island biogeography is just one of many ways in which a species can split into two, as separated subpopulations evolve slightly differently, especially in the presence of any environmental gradient. Even in the absence of any divisive effect, species can split into two through bifurcation for example, in the presence of abundant food some organisms can either search actively for food or simply become ambush predators and let the prey come to them, and both turn out to be locally optimal strategies (Silvert 2002). This ability of organisms to develop a variety of new tactics for coping with environmental challenges leads to a proliferation of life history strategies, and our inability to predict precisely which choice an organism will make has been the basis for attacks on ecological theory for example, Slobodkin (1965) argued that the failure of mathematical theory to predict that certain rotifers protect themselves with spines against predation reflected a fundamental weakness of theoretical ecology. This line of reasoning implies that - 11 -

William Silvert, The Meaning of Biodiversity. Talk at EMBS38. evolution is deterministic and thus it cannot account for speciation, but an alternative argument is that rotifers have many options for avoiding predators not only spines, but also shells, speed, transparency, hiding in refugia, and so on all of which are equally optimal (of equal fitness), and over time the population would probably explore all of them. The result, of course, would be a proliferation of species as different subpopulations develop separate defensive strategies. This leads us into the Pandoras box of evolutionary theory, and it is difficult to see how we can judge biodiversity without understanding how it arises. The concept of speciation occurring at periods when organisms reach a stage where they can choose between alternate life history patterns, the idea that there can be a bifurcation in the optimal strategy, is akin to the theory of punctuated equilibrium (Gould 1980) which remains controversial and will not be further explored here. However, the underlying concept that the high biodiversity of certain environments is due to some mechanism which gives rise to a speciation factory, and the implication that perhaps the high biodiversity of a tropical rain forest should not necessarily take precedence over the low biodiversity of the polar icecaps remains one worth considering.

Regime shifts
Underlying some of the questions about ecosystem dynamics and the ways in which biodiversity changes are the possibilities of major alterations, either natural or anthropogenic, in the structure of communities. Such changes have been documented in the fossil record and by studies of fish scales in sediments (Soutar & Isaacs 1969, 1974), and there is evidence of recent dramatic changes in the composition of the North Sea community (Lindeboom et al. 1995). Such regime shifts are not well understood and although some mechanisms have been proposed, such as a cyclic interaction between the pelagic and benthic subsystems (Silvert 1991), it is hard to find any conclusive evidence or explanation for them. This is particularly true in exploited marine systems, since any changes in community structure are likely to be masked by population shifts to due fishing and other human activities. However, if regime shifts occur they are likely to be accompanied by large changes in biodiversity since they involve many species, and any policy on biodiversity should take this factor into account.

Risk of species loss

One of the most frightening aspects of dealing with biodiversity is the realisation that if we fail to conserve some species and it goes extinct, it will be gone forever there is no chance of recovery. This element of finality is probably one of the reasons why conservation has achieved the social impact it has, and the awareness that mankind has wiped out such impressive creatures as the woolly mammoth (Mammuthus primigenius), the Stellers sea cow (Hydrodamalis gigas), the dodo (Raphus cucillatus) and the passenger pigeon (Ectopistes migratorius) is alarming evidence that the permanent loss of prized species is possible. There is however a growing awareness that there are many less obvious candidates for extinction, and if we accept the thesis that some battles will be lost, then we have to confront the risk that we may let some essential species fade away without appreciating its value until it is too late.

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William Silvert, The Meaning of Biodiversity. Talk at EMBS38. Unfortunately we do not always know what the biological cost of extinction is. We can assess the aesthetic loss of mammoths and dodos, but we have little idea of the ecological impact even of the loss of an estimated 5 billion passenger pigeons, undoubtedly a large fraction of the total bird population of the 19th century (Schorger 1955). Unless we know what will happen if a species goes extinct, we have to rely on conjecture. For example, the venom of some little-known snail species may turn out to have medicinal value (Sreenivasan 2002), or a vanished species might be replaced by one which is the vector for some terrible disease. Gray (2004) has documented some truly extraordinary benefits that can spring from the most unlikely species. There is clearly scope for valuable research to support policy decisions on biodiversity conservation, but in the absence of such research we still have to provide advice, and the less we know, the greater the risks but they cannot be avoided. Even when we fully understand the ecological role of a species, we may not be able to judge the consequences of its loss. Surely the earthworm must be one of the most essential members of the terrestrial community, but even though almost all the native earthworms in Canada were apparently eradicated by glaciation 10,000 years ago or more and were only replaced within the last few centuries by European species, the impact of this loss seems to have been far less than we might expect (Hogner 1953). Perhaps we know far less about the robustness of ecological communities than we think we do. This is not meant to suggest that the risk of extinction should be downplayed. Aside from the loss in the richness of our own lives when something that has taken millions of years to evolve vanishes from the earth because of our negligence, there can be no doubt that without certain species the world will be a very different, and possibly worse, place to live in, for us and for other creatures. The problem is that we are not always able to tell which these key species are.

Summary
Biodiversity is a huge field and this article has touched only a few of its aspects. The underlying message however can be summarised in a few simple statements. Biodiversity is an important concern of scientists, environmental activists, society as a whole, and politicians. Conservation of biodiversity requires communication and cooperation between all of these parties. Concepts of biodiversity must therefore be meaningful to all parties.

This means that the scientific community must develop definitions of biodiversity which they can explain and defend to the general public, and they must also be sensitive to public concerns which may not always mesh with their scientific judgments. Politics is based on compromise, and compromise usually means giving up something which you value. It is unlikely that we will ever achieve a perfect policy on conservation of biodiversity, but a good policy is better than none.

Acknowledgement
This work was prepared under the auspices of the European Fisheries Ecosystem Plan, funded by the EU Quality of Life and Management of Living Resources programme. Contract no: QLRT-CT-2001-01685.

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William Silvert, The Meaning of Biodiversity. Talk at EMBS38.

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