BIODIVERSITY



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BioDiversity BIODIVERSITY

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BioDiversity CHAPTER 1 THE CURRENT STATE OF BIOLOGICAL DIVERSITY E.O.WILSON Frank B.Baird, Jr. Professor of Science, Harvard University, Museum of Comparative Zoology, Cambridge, Massachusetts Biological diversity must be treated more seriously as a global resource, to be indexed, used, and above all, preserved. Three circumstances conspire to give this matter an unprecedented urgency. First, exploding human populations are degrading the environment at an accelerating rate, especially in tropical countries. Second, science is discovering new uses for biological diversity in ways that can relieve both human suffering and environmental destruction. Third, much of the diversity is being irreversibly lost through extinction caused by the destruction of natural habitats, again especially in the tropics. Overall, we are locked into a race. We must hurry to acquire the knowledge on which a wise policy of conservation and development can be based for centuries to come. To summarize the problem in this chapter, I review some current information on the magnitude of global diversity and the rate at which we are losing it. I concentrate on the tropical moist forests, because of all the major habitats, they are richest in species and because they are in greatest danger. THE AMOUNT OF BIOLOGICAL DIVERSITY Many recently published sources, especially the multiauthor volume Synopsis and Classification of Living Organisms, indicate that about 1.4 million living species of all kinds of organisms have been described (Parker, 1982; see also the numerical breakdown according to major taxonomic category of the world insect fauna prepared by Arnett, 1985). Approximately 750,000 are insects, 41,000 are vertebrates, and 250,000 are plants (that is, vascular plants and bryophytes). The remainder consists of a complex array of invertebrates, fungi, algae, and microorganisms (see Table 1–1). Most systematists agree that this picture is still very incomplete except

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BioDiversity TABLE 1–1 Numbers of Described Species of Living Organismsa Kingdom and Major Subdivision Common Name No. of Described Species Totals Virus   Viruses 1,000 (order of magnitude only) 1,000 Monera Bacteria Bacteria 3,000   Myxoplasma Bacteria 60   Cyanophycota Blue-green algae 1,700 4,760 Fungi Zygomycota Zygomycete fungi 665   Ascomycota (including 18,000 lichen fungi) Cup fungi 28,650 Basidiomycota Basidiomycete fungi 16,000 Oomycota Water molds 580 Chytridiomycota Chytrids 575 Acrasiomycota Cellular slime molds 13 Myxomycota Plasmodial slime molds 500 46,983 Algae Chlorophyta Green algae 7,000   Phaeophyta Brown algae 1,500 Rhodophyta Red algae 4,000 Chrysophyta Chrysophyte algae 12,500 Pyrrophyta Dinoflagellates 1,100 Euglenophyta Euglenoids 800 26,900 Plantae Bryophyta Mosses, liverworts, hornworts 16,600   Psilophyta Psilopsids 9 Lycopodiophyta Lycophytes 1,275 Equisetophyta Horsetails 15 Filicophyta Ferns 10,000 Gymnosperma Gymnosperms 529 Dicotolydonae Dicots 170,000 Monocotolydonae Monocots 50,000 248,428 Protozoa   Protozoans: 30,800     Sarcomastigophorans, ciliates, and smaller groups   30,800 Animalia Porifera Sponges 5,000   Cnidaria, Ctenophora Jellyfish, corals, comb jellies 9,000 Platyhelminthes Flatworms 12,200 Nematoda Nematodes (roundworms) 12,000 Annelida Annelids (earthworms and relatives) 12,000

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BioDiversity Kingdom and Major Subdivision Common Name No. of Described Species Totals Mollusca Mollusks 50,000   Echinodermata Echinoderms (starfish and relatives) 6,100 Arthropoda Arthropods   Insecta Insects 751,000 Other arthropods   123,161 Minor invertebrate phyla   9,300 989,761 Chordata Tunicata Tunicates 1,250   Cephalochordata Acorn worms 23 Vertebrata Vertebrates   Agnatha Lampreys and other jawless fishes 63 Chrondrichthyes Sharks and other cartilaginous fishes 843 Osteichthyes Bony fishes 18,150 Amphibia Amphibians 4,184 Reptilia Reptiles 6,300 Aves Birds 9,040 Mammalia Mammals 4,000 43,853 TOTAL, all organisms     1,392,485 aCompiled from multiple sources. in a few well-studied groups such as the vertebrates and flowering plants. If insects, the most species-rich of all major groups, are included, I believe that the absolute number is likely to exceed 5 million. Recent intensive collections made by Terry L.Erwin and his associates in the canopy of the Peruvian Amazon rain forest have moved the plausible upper limit much higher. Previously unknown insects proved to be so numerous in these samples that when estimates of local diversity were extrapolated to include all rain forests in the world, a figure of 30 million species was obtained (Erwin, 1983). In an even earlier stage is research on the epiphytic plants, lichens, fungi, roundworms, mites, protozoans, bacteria, and other mostly small organisms that abound in the treetops. Other major habitats that remain poorly explored include the coral reefs, the floor of the deep sea, and the soil of tropical forests and savannas. Thus, remarkably, we do not know the true number of species on Earth, even to the nearest order of magnitude (Wilson, 1985a). My own guess, based on the described fauna and flora and many discussions with entomologists and other specialists, is that the absolute number falls somewhere between 5 and 30 million. A brief word is needed on the meaning of species as a category of classification. In modern biology, species are regarded conceptually as a population or series of populations within which free gene flow occurs under natural conditions. This means that all the normal, physiologically competent individuals at a given time are capable of breeding with all the other individuals of the opposite sex belonging

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BioDiversity to the same species or at least that they are capable of being linked genetically to them through chains of other breeding individuals. By definition they do not breed freely with members of other species. This biological concept of species is the best ever devised, but it remains less than ideal. It works very well for most animals and some kinds of plants, but for some plant and a few animal populations in which intermediate amounts of hybridization occur, or ordinary sexual reproduction has been replaced by self-fertilization or parthenogenesis, it must be replaced with arbitrary divisions. New species are usually created in one or the other of two ways. A large minority of plant species came into existence in essentially one step, through the process of polyploidy. This is a simple multiplication in the number of gene-bearing chromosomes—sometimes within a preexisting species and sometimes in hybrids between two species. Polyploids are typically not able to form fertile hybrids with the parent species. A second major process is geographic speciation and takes much longer. It starts when a single population (or series of populations) is divided by some barrier extrinsic to the organisms, such as a river, a mountain range, or an arm of the sea. The isolated populations then diverge from each other in evolution because of the inevitable differences of the environments in which they find themselves. Since all populations evolve when given enough time, divergence between all extrinsically isolated populations must eventually occur. By this process alone the populations can acquire enough differences to reduce interbreeding between them should the extrinsic barrier between them be removed and the populations again come into contact. If sufficient differences have accumulated, the populations can coexist as newly formed species. If those differences have not yet occurred, the populations will resume the exchange of genes when the contact is renewed. Species diversity has been maintained at an approximately even level or at most a slowly increasing rate, although punctuated by brief periods of accelerated extinction every few tens of millions of years. The more similar the species under consideration, the more consistent the balance. Thus within clusters of islands, the numbers of species of birds (or reptiles, or ants, or other equivalent groups) found on each island in turn increases approximately as the fourth root of the area of the island. In other words, the number of species can be predicted as a constant X (island area)0.25, where the exponent can deviate according to circumstances, but in most cases it falls between 0.15 and 0.35. According to this theory of island biogeography, in a typical case (where the exponent is at or near 0.25) the rule of thumb is that a 10-fold increase in area results in a doubling of a number of species (MacArthur and Wilson, 1967). In a recent study of the ants of Hispaniola, I found fossils of 37 genera (clusters of species related to each other but distinct from other such clusters) in amber from the Miocene age—about 20 million years old. Exactly 37 genera exist on the island today. However, 15 of the original 37 have become extinct, while 15 others not present in the Miocene deposits have invaded to replace them, thus sustaining the original diversity (Wilson, 1985b). On a grander scale, families—clusters of genera—have also maintained a balance within the faunas of entire continents. For example, a reciprocal and apparently symmetrical exchange of land mammals between North and South America began

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BioDiversity 3 million years ago, after the rise of the Panamanian land bridge. The number of families in South America first rose from 32 to 39 and then subsided to the 35 that exist there today. A comparable adjustment occurred in North America. At the generic level, North American elements dominated those from South America: 24 genera invaded to the south whereas only 12 invaded to the north. Hence, although equilibrium was roughly preserved, it resulted in a major shift in the composition of the previously isolated South American fauna (Marshall et al., 1982). Each species is the repository of an immense amount of genetic information. The number of genes range from about 1,000 in bacteria and 10,000 in some fungi to 400,000 or more in many flowering plants and a few animals (Hinegardner, 1976). A typical mammal such as the house mouse (Mus musculus) has about 100,000 genes. This full complement is found in each of its myriad cells, organized from four strings of DNA, each of which comprises about a billion nucleotide pairs (George D.Snell, Jackson Laboratory, Maine, personal communication, 1987). (Human beings have genetic information closer in quantity to the mouse than to the more abundantly endowed salamanders and flowering plants; the difference, of course, lies in what is encoded.) If stretched out fully, the DNA would be roughly 1-meter long. But this molecule is invisible to the naked eye because it is only 20 angstroms in diameter. If we magnified it until its width equalled that of wrapping string, the fully extended molecule would be 960 kilometers long. As we traveled along its length, we would encounter some 20 nucleotide pairs or “letters” of genetic code per inch, or about 50 per centimeter. The full information contained therein, if translated into ordinary-size letters of printed text, would just about fill all 15 editions of the Encyclopaedia Britannica published since 1768 (Wilson, 1985a). The number of species and the amount of genetic information in a representative organism constitute only part of the biological diversity on Earth. Each species is made up of many organisms. For example, the 10,000 or so ant species have been estimated to comprise 1015 living individuals at each moment of time (Wilson, 1971). Except for cases of parthenogenesis and identical twinning, virtually no two members of the same species are genetically identical, due to the high levels of genetic polymorphism across many of the gene loci (Selander, 1976). At still another level, wide-ranging species consist of multiple breeding populations that display complex patterns of geographic variation in genetic polymorphism. Thus, even if an endangered species is saved from extinction, it will probably have lost much of its internal diversity. When the populations are allowed to expand again, they will be more nearly genetically uniform than the ancestral populations. The bison herds of today are biologically not quite the same—not so interesting—as the bison herds of the early nineteenth century. THE NATURAL LONGEVITY OF SPECIES Within particular higher groups of organisms, such as ammonites or fishes, species have a remarkably consistent longevity. As a result, the probability that a given species will become extinct in a given interval of time after it splits off from other species can be approximated as a constant, so that the frequency of species surviving

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BioDiversity through time falls off as an exponential decay function; in other words, the percentage (but not the absolute number) of species going extinct in each period of time stays the same (Van Valen, 1973).1 These regularities, such as they are, have been interrupted during the past 250 million years by major episodes of extinction that have been recently estimated to occur regularly at intervals of 26 million years (Raup and Sepkoski, 1984). Because of the relative richness of fossils in shallow marine deposits, the longevity of fish and invertebrate species living there can often be determined with a modest degree of confidence. During Paleozoic and Mesozoic times, the average persistence of most fell between 1 and 10 million years: that is, 6 million for echinoderms, 1.9 million for graptolites, 1.2 to 2 million for ammonites, and so on (Raup, 1981, 1984). These estimates are extremely interesting and useful but, as paleontologists have generally been careful to point out, they also suffer from some important limitations. First, terrestrial organisms are far less well known, few estimates have been attempted, and thus different survivorship patterns might have occurred (although Cenozoic flowering plants, at least, appear to fall within the 1- to 10-million-year range). More importantly, a great many organisms on islands and other restricted habitats, such as lakes, streams, and mountain crests, are so rare or local that they could appear and vanish within a short time without leaving any fossils. An equally great difficulty is the existence of sibling species—populations that are reproductively isolated but so similar to closely related species as to be difficult or impossible to distinguish through conventional anatomical traits. Such entities could rarely be diagnosed in fossil form. Together, all these considerations suggest that estimates of the longevity of natural species should be extended only with great caution to groups for which there is a poor fossil record. RAIN FORESTS AS CENTERS OF DIVERSITY In recent years, evolutionary biologists and conservationists have focused increasing attention on tropical rain forests, for two principal reasons. First, although these habitats cover only 7% of the Earth’s land surface, they contain more than half the species in the entire world biota. Second, the forests are being destroyed so rapidly that they will mostly disappear within the next century, taking with them hundreds of thousands of species into extinction. Other species-rich biomes are in danger, most notably the tropical coral reefs, geologically ancient lakes, and coastal wetlands. Each deserves special attention on its own, but for the moment the rain forests serve as the ideal paradigm of the larger global crisis. Tropical rain forests, or more precisely closed tropical forests, are defined as habitats with a relatively tight canopy of mostly broad-leaved evergreen trees 1   Van Valen’s original formulation, whose difficulties and implications are revealed by more recent research, has been discussed by Raup (1975) and by Lewin (1985). These studies deal with the clade, or set of populations descending through time after having split off as a distinct species from other such populations. They do not refer to the chronospecies, which is just a set of generations of the same species that is subjectively different from sets of generations.

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BioDiversity sustained by 100 centimeters or more of annual rainfall. Typically two or more other layers of trees and shrubs occur beneath the upper canopy. Because relatively little sunlight reaches the forest floor, the undergrowth is sparse and human beings can walk through it with relative ease. The species diversity of rain forests borders on the legendary. Every tropical biologist has a favorite example to offer. From a single leguminous tree in the Tambopata Reserve of Peru, I recently recovered 43 species of ants belonging to 26 genera, about equal to the entire ant fauna of the British Isles (Wilson, 1987). Peter Ashton found 700 species of trees in 10 selected 1-hectare plots in Borneo, the same as in all of North America (Ashton, Arnold Arboretum, personal communication, 1987). It is not unusual for a square kilometer of forest in Central or South America to contain several hundred species of birds and many thousands of species of butterflies, beetles, and other insects. Despite their extraordinary richness, tropical rain forests are among the most fragile of all habitats. They grow on so-called wet deserts—an unpromising soil base washed by heavy rains. Two-thirds of the area of the forest surface consists of tropical red and yellow earths, which are typically acidic and poor in nutrients. High concentrations of iron and aluminum form insoluble compounds with phosphorus, thereby decreasing the availability of phosphorus to plants. Calcium and potassium are leached from the soil soon after their compounds are dissolved from the rain. As little as 0.1% of the nutrients filter deeper than 5 centimeters beneath the soil surface (NRC, 1982). An excellent popular account of rain forest ecology is given by Forsyth and Miyata (1984). During the 150 million years since its origin, the principally dicotyledonous flora has nevertheless evolved to grow thick and tall. At any given time, most of the nonatmospheric carbon and vital nutrients are locked up in the tissue of the vegetation. As a consequence, the litter and humus on the ground are thin compared to the thick mats of northern temperate forests. Here and there, patches of bare earth show through. At every turn one can see evidence of rapid decomposition by dense populations of termites and fungi. When the forest is cut and burned, the ash and decomposing vegetation release a flush of nutrients adequate to support new herbaceous and shrubby growth for 2 or 3 years. Then these materials decline to levels lower than those needed to support a healthy growth of agricultural crops without artificial supplements. The regeneration of rain forests is also limited by the fragility of the seeds of the constituent woody species. The seeds of most species begin to germinate within a few days or weeks, severely limiting their ability to disperse across the stripped land into sites favorable for growth. As a result, most sprout and die in the hot, sterile soil of the clearings (Gomez-Pompa et al., 1972). The monitoring of logged sites indicates that regeneration of a mature forest might take centuries. The forest at Angkor (to cite an anecdotal example) dates back to the abandonment of the Khmer capital in 1431, yet is still structurally different from a climax forest today, 556 years later. The process of rain forest regeneration is in fact so generally slow that few extrapolations have been possible; in some zones of greatest combined damage and sterility, restoration might never occur naturally (Caufield, 1985; Gomez-Pompa et al., 1972).

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BioDiversity Approximately 40% of the land that can support tropical closed forest now lacks it, primarily because of human action. By the late 1970s, according to estimates from the Food and Agricultural Organization and United Nations Environmental Programme, 7.6 million hectares or nearly 1% of the total cover is being permanently cleared or converted into the shifting-cultivation cycle. The absolute amount is 76,000 square kilometers (27,000 square miles) a year, greater than the area of West Virginia or the entire country of Costa Rica. In effect, most of this land is being permanently cleared, that is, reduced to a state in which natural reforestation will be very difficult if not impossible to achieve (Mellilo et al., 1985). This estimated loss of forest cover is close to that advanced by the tropical biologist Norman Myers in the mid-1970s, an assessment that was often challenged by scientists and conservationists as exaggerated and alarmist. The vindication of this early view should serve as a reminder always to take such doomsday scenarios seriously, even when they are based on incomplete information. A straight-line extrapolation from the first of these figures, with identically absolute annual increments of forest-cover removal, leads to 2135 A.D. as the year in which all the remaining rain forest will be either clear-cut or seriously disturbed, mostly the former. By coincidence, this is close to the date (2150) that the World Bank has estimated the human population will plateau at 11 billion people (The World Bank, 1984). In fact, the continuing rise in human population indicates that a straight line estimate is much too conservative. Population pressures in the Third World will certainly continue to accelerate deforestation during the coming decades unless heroic measures are taken in conservation and resource management. There is another reason to believe that the figures for forest cover removal present too sanguine a picture of the threat to biological diversity. In many local areas with high levels of endemicity, deforestation has proceeded very much faster than the overall average. Madagascar, possessor of one of the most distinctive floras and faunas in the world, has already lost 93% of its forest cover. The Atlantic coastal forest of Brazil, which so enchanted the young Darwin upon his arrival in 1832 (“wonder, astonishment & sublime devotion, fill & elevate the mind”), is 99% gone. In still poorer condition—in fact, essentially lost—are the forests of many of the smaller islands of Polynesia and the Caribbean. HOW MUCH DIVERSITY IS BEING LOST? No precise estimate can be made of the numbers of species being extinguished in the rain forests or in other major habitats, for the simple reason that we do not know the numbers of species originally present. However, there can be no doubt that extinction is proceeding far faster than it did prior to 1800. The basis for this statement is not the direct observation of extinction. To witness the death of the last member of a parrot or orchid species is a near impossibility. With the exception of the showiest birds, mammals, or flowering plants, biologists are reluctant to say with finality when a species has finally come to an end. There is always the chance (and hope) that a few more individuals will turn up in some remote forest remnant or other. But the vast majority of species are not monitored at all. Like the dead

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BioDiversity of Gray’s “Elegy Written in a Country Churchyard,” they pass from the Earth without notice. Instead, extinction rates are usually estimated indirectly from principles of biogeography. As I mentioned above, the number of species of a particular group of organisms in island systems increases approximately as the fourth root of the land area. This has been found to hold true not just on real islands but also on habitat islands, such as lakes in a “sea” of land, alpine meadows or mountaintops surrounded by evergreen forests, and even in clumps of trees in the midst of a grassland (MacArthur and Wilson, 1967). Using the area-species relationship, Simberloff (1984) has projected ultimate losses due to the destruction of rain forests in the New World tropical mainland. If present levels of forest removal continue, the stage will be set within a century for the inevitable loss of 12% of the 704 bird species in the Amazon basin and 15% of the 92,000 plant species in South and Central America. As severe as these regional losses may be, they are far from the worst, because the Amazon and Orinoco basins contain the largest continuous rain forest tracts in the world. Less extensive habitats are far more threatened. An extreme example is the western forest of Ecuador. This habitat was largely undisturbed until after 1960, when a newly constructed road network led to the swift incursion of settlers and clear-cutting of most of the area. Now only patches remain, such as the 0.8-square-kilometer tract at the Rio Palenque Biological Station. This tiny reserve contains 1,033 plant species, perhaps one-quarter of which are known only to occur in coastal Ecuador. Many are known at the present time only from a single living individual (Gentry, 1982). In general, the tropical world is clearly headed toward an extreme reduction and fragmentation of tropical forests, which will be accompanied by a massive extinction of species. At the present time, less than 5% of the forests are protected within parks and reserves, and even these are vulnerable to political and economic pressures. For example, 4% of the forests are protected in Africa, 2% in Latin America, and 6% in Asia (Brown, 1985). Thus in a simple system as envisioned by the basic models of island biogeography, the number of species of all kinds of organisms can be expected to be reduced by at least one-half—in other words, by hundreds of thousands or even (if the insects are as diverse as the canopy studies suggest) by millions of species. In fact, the island-biogeographic projections appear to be conservative for two reasons. First, tropical species are far more localized than those in the temperate zones. Consequently, a reduction of 90% of a tropical forest does not just reduce all the species living therein to 10% of their original population sizes, rendering them more vulnerable to future extinction. That happens in a few cases, but in many others, entire species are eliminated because they happened to be restricted to the portion of the forest that was cut over. Second, even when a portion of the species survives, it will probably have suffered significant reduction in genetic variation among its members due to the loss of genes that existed only in the outer portions. The current reduction of diversity seems destined to approach that of the great natural catastrophes at the end of the Paleozoic and Mesozoic eras—in other words,

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BioDiversity the most extreme in the past 65 million years. In at least one important respect, the modern episode exceeds anything in the geological past. In the earlier mass extinctions, which some scientists believe were caused by large meteorite strikes, most of the plants survived even though animal diversity was severely reduced. Now, for the first time, plant diversity is declining sharply (Knoll, 1984). HOW FAST IS DIVERSITY DECLINING? The area-species curves of island systems, that is, the quantitative relationship between the area of islands and the number of species that can persist on the islands, provide minimal estimates of the reduction of species diversity that will eventually occur in the rain forests. But how long is “eventually”? This is a difficult question that biogeographers have attacked with considerable ingenuity. When a forest is reduced from, say, 100 square kilometers to 10 square kilometers by clearing, some immediate extinction is likely. However, the new equilibrium will not be reached all at once. Some species will hang on for a while in dangerously reduced populations. Elementary mathematical models of the process predict that the number of species in the 10-square-kilometer plot will decline at a steadily decelerating rate, i.e., they will decay exponentially to the lower level. Studies by Jared Diamond and John Terborgh have led to the estimation of the decay constants for the bird faunas on naturally occurring islands (Diamond, 1972, 1984; Terborgh, 1974). These investigators took advantage of the fact that rising sea levels 10,000 years ago cut off small land masses that had previously been connected to South America, New Guinea, and the main islands of Indonesia. For example, Tobago, Margarita, Coiba, and Trinidad were originally part of the South American mainland and shared the rich bird fauna of that continent. Thus they are called land-bridge islands. In a similar manner, Yapen, Aru, and Misol were connected to New Guinea. In the study of the South American land-bridge islands, Terborgh found that the smaller the island, the higher the estimated decay constant and hence extinction rate. Terborgh then turned to Barro Colorado Island, which was isolated for the first time by the rise of Gatun Lake during the construction of the Panama Canal. Applying the natural land-bridge extinction curve to an island of this size (17 square kilometers) and fitting the derived decay constant to the actual period of isolation (50 years), Terborgh predicted an extinction of 17 bird species. The actual number known to have vanished as a probable result of insularization is 13, or 12% of the 108 breeding species originally present. The extinction rates of bird species on Barro Colorado Island were based on careful studies by E.O.Willis and J.R.Karr and have been recently reviewed by Diamond (1984). Several other studies of recently created islands of both tropical and temperate-zone woodland have produced similar results, which can be crudely summarized as follows: when the islands range from 1 to 25 square kilometers—the size of many smaller parks and reserves—the rate of extinction of bird species during the first 100 years is 10 to 50%. Also as predicted, the extinction rate is highest in the smaller patches, and it rises steeply when the area drops below 1 square kilometer. To take one example provided by Willis (1979), three patches of subtropical forest

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BioDiversity isolated (by agricultural clearing) in Brazil for about a hundred years varied from 0.2 to 14 square kilometers, and, in reverse order, their resident bird species suffered 14 to 62% extinction rates. What do these first measurements tell us about the rate at which diversity is being reduced? No precise estimate can be made for three reasons. First, the number of species of organisms is not known, even to the nearest order of magnitude. Second, because even in a simple island-biogeographic system, diversity reduction depends on the size of the island fragments and their distance from each other—factors that vary enormously from one country to the next. Third, the ranges of even the known species have not been worked out in most cases, so that we cannot say which ones will be eliminated when the tropical forests are partially cleared. However, scenarios of reduction can be constructed to give at least first approximations if certain courses of action are followed. Let us suppose, for example, that half the species in tropical forests are very localized in distribution, so that the rate at which species are being eliminated immediately is approximately this fraction multiplied by the rate-percentage of the forests being destroyed. Let us conservatively estimate that 5 million species of organisms are confined to the tropical rain forests, a figure well justified by the recent upward adjustment of insect diversity alone. The annual rate of reduction would then be 0.5×5× 106×0.007 species, or 17,500 species per year. Given 10 million species in the fauna and flora of all the habitats of the world, the loss is roughly one out of every thousand species per year. How does this compare with extinction rates prior to human intervention? The estimates of extinction rates in Paleozoic and Mesozoic marine faunas cited earlier (Raup, 1981, 1984; Raup and Sepkoski, 1984; Van Valen, 1973) ranged according to taxonomic group (e.g., echinoderms versus cephalopods) from one out of every million to one out of every 10 million per year. Let us assume that on the order of 10 million species existed then, in view of the evidence that diversity has not fluctuated through most of the Phanerozoic time by a factor of more than three (Raup and Sepkoski, 1984). It follows that both the per-species rate and absolute loss in number of species due to the current destruction of rain forests (setting aside for the moment extinction due to the disturbance of other habitats) would be about 1,000 to 10,000 times that before human intervention. I have constructed other simple models incorporating the quick loss of local species and the slower loss of widespread species due to the insularization effect, and these all lead to comparable or higher extinction rates. It seems difficult if not impossible to combine what is known empirically of the extinction process with the ongoing deforestation process without arriving at extremely high rates of species loss in the near future. Curiously, however, the study of extinction remains one of the most neglected in ecology. There is a pressing need for a more sophisticated body of theories and carefully planned field studies based on it than now exist. WHAT CAN BE DONE? The biological diversity most threatened is also the least explored, and there is no prospect at the moment that the scientific task will be completed before a large

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BioDiversity fraction of the species vanish. Probably no more than 1,500 professional systematists in the world are competent to deal with the millions of species found in the humid tropic forests. Their number may be dropping, due to decreased professional opportunities, reduced funding for research, and the assignment of a higher priority to other disciplines. Data concerning the number of taxonomists, as well as detailed arguments for the need to improve research in tropical countries, are given by NRC (1980). The decline has been accompanied by a more than 50% decrease in the number of publications in tropical ecology from 1979 to 1983 (Cole, 1984). The problem of tropical conservation is thus exacerbated by the lack of knowledge and the paucity of ongoing research. In order to make precise assessments and recommendations, it is necessary to know which species are present (recall that the great majority have not even received a scientific name) as well as their geographical ranges, biological properties, and possible vulnerability to environmental change. It would be a great advantage, in my opinion, to seek such knowledge for the entire biota of the world. Each species is unique and intrinsically valuable. We cannot expect to answer the important questions of ecology and other branches of evolutionary biology, much less preserve diversity with any efficiency, by studying only a subset of the extant species. I will go further: the magnitude and control of biological diversity is not just a central problem of evolutionary biology; it is one of the key problems of science as a whole. At present, there is no way of knowing whether there are 5, 10, or 30 million species on Earth. There is no theory that can predict what this number might turn out to be. With reference to conservation and practical applications, it also matters why a certain subset of species exists in each region of the Earth, and what is happening to each one year by year. Unless an effort is made to understand all of diversity, we will fall far short of understanding life in these important respects, and due to the accelerating extinction of species, much of our opportunity will slip away forever. Lest this exploration be viewed as an expensive Manhattan Project unattainable in today’s political climate, let me cite estimates I recently made of the maximum investment required for a full taxonomic accounting of all species: 25,000 professional lifetimes (4,000 systematists are at work full or part time in North America today); their final catalog would fill 60 meters of library shelving for each million species (Wilson, 1985a). Computer-aided techniques could be expected to cut the effort and cost substantially. In fact, systematics has one of the lowest cost-to-benefit ratios of all scientific disciplines. It is equally true that knowledge of biological diversity will mean little to the vast bulk of humanity unless the motivation exists to use it. Fortunately, both scientists and environmental policy makers have established a solid linkage between economic development and conservation. The problems of human beings in the tropics are primarily biological in origin: overpopulation, habitat destruction, soil deterioration, malnutrition, disease, and even, for hundreds of millions, the uncertainty of food and shelter from one day to the next. These problems can be solved in part by making biological diversity a source of economic wealth. Wild species are in fact both one of the Earth’s most important resources and the least

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BioDiversity utilized. We have come to depend completely on less than 1% of living species for our existence, the remainder waiting untested and fallow. In the course of history, according to estimates made by Myers (1984), people have utilized about 7,000 kinds of plants for food; predominant among these are wheat, rye, maize, and about a dozen other highly domesticated species. Yet there are at least 75,000 edible plants in existence, and many of these are superior to the crop plants in widest use. Others are potential sources of new pharmaceuticals, fibers, and petroleum substitutes. In addition, among the insects are large numbers of species that are potentially superior as crop pollinators, control agents for weeds, and parasites and predators of insect pests. Bacteria, yeasts, and other microorganisms are likely to continue yielding new medicines, food, and procedures of soil restoration. Biologists have begun to fill volumes with concrete proposals for the further exploration and better use of diversity, with increasing emphasis on the still unexplored portions of the tropical biota. Some of the most recent and useful works on this subject include those by Myers (1984), NRC (1975), Office of Technology Assessment (1984), Oldfield (1984), and the U.S. Department of State (1982). In addition, an excellent series of specialized publications on practical uses of wild species have been produced during the past 10 years by authors and panels commissioned by the Board on Science and Technology for International Development (BOSTID) of the National Research Council. In response to the crisis of tropical deforestation and its special threat to biological diversity, proposals are regularly being advanced at the levels of policy and research. For example, Nicholas Guppy (1984), noting the resemblance of the lumbering of rain forests to petroleum extraction as the mining of a nonrenewable resource for short-term profit, has recommended the creation of a cartel, the Organization of Timber-Exporting Countries (OTEC). By controlling production and prices of lumber, the organization could slow production while encouraging member states to “protect the forest environment in general and gene stocks and special habitats in particular, create plantations to supply industrial and fuel wood, benefit indigenous tribal forest peoples, settle encroachers, and much else.” In another approach, Thomas Lovejoy (1984) has recommended that debtor nations with forest resources and other valuable habitats be given discounts or credits for undertaking conservation programs. Even a small amount of forgiveness would elevate the sustained value of the natural habitats while providing hard currency for alternatives to their exploitation. Another opportunity for innovation lies in altering somewhat the mode of direct economic assistance to developing countries. A large part of the damage to tropical forests, especially in the New World, has resulted from the poor planning of road systems and dams. For example, the recent settlement of the state of Rondonia and construction of the Tucurui Dam, both in Brazil, are now widely perceived by ecologists and economists alike as ill-conceived (Caufield, 1985). Much of the responsibility of minimizing environmental damage falls upon the international agencies that have the power to approve or disapprove particular projects. The U.S. Congress addressed this problem with amendments to the Foreign Assistance Act in 1980, 1983, and 1986, which call for the development of a strategy for conserving biological diversity. They also mandate that programs funded

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BioDiversity through the U.S. Agency for International Development (USAID) include an assessment of environmental impact. In implementing this new policy, USAID has recognized that “the destruction of humid tropical forests is one of the most important environmental issues for the remainder of this century and, perhaps, well into the next,” in part because they are “essential to the survival of vast numbers of species of plants and animals” (U.S. Department of State, 1985). In another sphere, The World Bank and other multinational lending agencies have come under increasing pressure to take a more active role in assessing the environmental impact of the large-scale projects they underwrite (Anonymous, 1984). In addition to recommendations for international policy initiatives, there has recently been a spate of publications on the linkage of conservation and economic use of tropical forests. Notable among them are Research Priorities in Tropical Biology (NRC, 1980), based on a study of the National Research Council; Technologies to Sustain Tropical Forest Resources (OTA, 1984), prepared by the Office of Technology Assessment for the U.S. Congress; and the U.S. Strategy on the Conservation of Biological Diversity (USAID, 1985), a report to Congress by an interagency task force. Most comprehensive of all—and in my opinion the most encouraging in its implications—is the three-part series Tropical Forests: A Call for Action, released by the World Resources Institute, The World Bank, and the United Nations Development Programme (1985). The report makes an assessment of the problem worldwide and reviews case histories in which conservation or restoration have contributed to economic development. It examines the needs of every tropical country with important forest reserves. The estimated cost to make an impact on tropical deforestation over the next 5 years would be U.S. $8 billion—a large sum but surely the most cost-effective investment available to the world at the present time. In the end, I suspect it will all come down to a decision of ethics—how we value the natural worlds in which we evolved and now, increasingly, how we regard our status as individuals. We are fundamentally mammals and free spirits who reached this high a level of rationality by the perpetual creation of new options. Natural philosophy and science have brought into clear relief what might be the essential paradox of human existence. The drive toward perpetual expansion—or personal freedom—is basic to the human spirit. But to sustain it we need the most delicate, knowing stewardship of the living world that can be devised. Expansion and stewardship may appear at first to be conflicting goals, but the opposite is true. The depth of the conservation ethic will be measured by the extent to which each of the two approaches to nature is used to reshape and reinforce the other. The paradox can be resolved by changing its premises into forms more suited to ultimate survival, including protection of the human spirit. I recently wrote in synecdochic form about one place in South America to give these feelings more exact expression: To the south stretches Surinam eternal, Surinam serene, a living treasure awaiting assay. I hope that it will be kept intact, that at least enough of its million-year history will be saved for the reading. By today’s ethic its value may seem limited, well beneath the pressing concerns of daily life. But I suggest that as biological knowledge grows the ethic will shift fundamentally so that everywhere, for reasons that have to do with the very fiber of the brain, the fauna and flora of a country will be thought part of the national

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BioDiversity heritage as important as its art, its language, and that astonishing blend of achievement and farce that has always defined our species (Wilson, 1984). REFERENCES Anonymous. 1984. Critics fault World Bank for ecological neglect. Conserv. Found. News. Nov.-Dec.:1–7. Arnett, R.H. 1985. General considerations. Pp. 3–9 in American Insects: A Handbook of the Insects of America North of Mexico. Van Nostrand Reinhold, New York. Brown, R.L., ed. 1985. State of the World 1985: A Worldwatch Institute Report on Progress Toward a Sustainable Society. W.W. Norton, New York. 301 pp. Caufield, C. 1985. In the Rainforest. A.A. Knopf, New York. 283 pp. Cole, N.H.A. 1984. Tropical ecology research. Nature 309:204. Diamond, J.M. 1972. Biogeographic kinetics: Estimation of relaxation times for avifaunas of Southwest Pacific islands. Proc. Natl. Acad. Sci. USA 69:3199–3203. Diamond, J.M. 1984. “Normal” extinctions of isolated populations. Pp. 191–246 in M.H.Nitecki, ed. Extinctions. University of Chicago Press, Chicago. Erwin, T.L. 1983. Beetles and other insects of tropical forest canopies at Manaus, Brazil, sampled by insecticidal fogging. Pp. 59–75 in S.L.Sutton, T.C.Whitmore, and A.C.Chadwick, eds. Tropical Rain Forest: Ecology and Management. Blackwell, Edinburgh. Forsyth, A., and K.Miyata. 1984. Tropical Nature: Life & Death in the Rain Forests of Central & South America. Scribner’s, New York. 272 pp. Frankel, O.H., and M.E.Soulé. 1981. Conservation and Evolution. Cambridge University Press, Cambridge, Mass. 327 pp. Gentry, A.H. 1982. Patterns of Neotropical plant-species diversity. Evol. Biol. 15:1–85. Gomez-Pompa, A., C.Vazquez-Yanes, and S.Guevara. 1972. The tropical rain forest: A nonrenewable resource. Science 177:762–765. Guppy, N. 1984. Tropical deforestation: A global view. Foreign Affairs 62:928–965. Hinegardner, R. 1976. Evolution of genome size. Pp. 179–199 in F.J.Ayala, ed. Molecular Evolution. Sinauer Associates, Sunderland, Mass. Knoll, A.H. 1984. Patterns of extinction in the fossil record of vascular plants. Pp. 21–68 in M. H.Nitecki, ed. Extinctions. University of Chicago Press, Chicago. Lewin, R. 1985. Red Queen runs into trouble? Science 227:399–400. Lovejoy, T.E. 1984. Aid debtor nations’ ecology. New York Times, October 4. MacArthur, R.H., and E.O.Wilson. 1967. The Theory of Island Biogeography. Princeton University Press, Princeton, N.J. 203 pp. Marshall, L.G., S.D.Webb, J.J.Sepkoski, Jr., and D.M.Raup. 1982. Mammalian evolution and the great American interchange. Science 215:1351–1357. Melillo, J.M., C.A.Palm, R.A.Houghton, G.M.Woodwell, and N.Myers. 1985. A comparison of two recent estimates of disturbance in tropical forests. Environ. Conserv. 12:37–40. Myers, N. 1983. A Wealth of Wild Species: Storehouse for Human Welfare. Westview Press, Boulder, Colo. 300 pp. Myers, N. 1984. The Primary Source: Tropical Forests and Our Future. W.W. Norton, New York. 399 pp. NRC (National Research Council). 1975. Underexploited Tropical Plants with Promising Economic Value. Board on Science and Technology for International Development Report 16. National Academy of Sciences, Washington, D.C. 187 pp. NRC (National Research Council). 1979. Tropical Legumes: Resources for the Future. Board on Science and Technology for International Development Report 25. National Academy of Sciences, Washington, D.C. 331 pp. NRC (National Research Council). 1980. Research Priorities in Tropical Biology. National Academy of Sciences, Washington, D.C. 116 pp. NRC (National Research Council). 1982. Ecological Aspects of Development in the Humid Tropics. National Academy Press, Washington, D.C. 297 pp. Oldfield, M.L. 1984. The Value of Conserving Genetic Resources. National Park Service, U.S. Department of the Interior, Washington, D.C. 360 pp. OTA (Office of Technology Assessment). 1984. Technologies to Sustain Tropical Forest Resources.

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