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Colloquium The biotic crisis and the future of evolution Norman Myers*t and Andrew H. Knoll: *Green College, University of Oxford, Oxford OX2 6HG, and Upper Meadow, Old Road, Oxford OX3 8SZ, United Kingdom; and "Department of Organismic and Evolutionary Biology, Harvard University, Cambridge, MA 02138 The biotic crisis overtaking our planet is likely to precipitate a major extinction of species. That much is well known. Not so well known but probably more significant in the long term is that the crisis will surely disrupt and deplete certain basic processes of evolution, with consequences likely to persist for millions of years. Distinctive features of future evolution could include a homogenization of biotas, a proliferation of opportunistic species, a pest-and-weed ecology, an outburst of speciation among taxa that prosper in human-dom~nated ecosystems, a decline of biodisparity, an end to the speciation of large vertebrates' the depletion of "evolutionary powerhouses" in the tropics, and unpredictable emergent novel- ties. Despite this likelihood, we have only a rudimentary under- standing of how we are altering the evolutionary future. As a result of our ignorance, conservation policies fail to reflect long- term evolutionary aspects of biodiversity loss. Human activities have brought the Earth to the brink of biotic crisis. Many biologists (e.g., refs. 1-5) consider that coming decades will see the loss of large numbers of species. Fewer scientists- witness the lack of professional papers addressing the issue appear to have recognized that, in the longer term, these extinctions will alter not only biological diversity but also the evolutionary processes by which diversity is generated. Thus, current and predicted environmental perturbations form a double-edged sword that will slice into both the legacy and fu- ture of evolution. A simple consideration of time underscores the magnitude of the challenge to scientists and public alike (ct ref. 6~. Episodes of mass extinction documented in the geological record were followed by protracted intervals of Diversification and ecolog- ical reorganization; five million years can be considered a broadly representative recovery time, although durations varied from one extinction to another (7~. Suppose, too, that the average number of people on Earth during the recovery period is 2.5 billion (by contrast with the 6 billion today). Under these conditions, the total number of people affected by what we do (or do not do) during the next few decades will be in the order of 500 trillion—10,000 times more people than have existed until now. We are thus engaged in by far the largest "decision" ever taken by one human community on the unconsulted behalf of future societies. The question of how current threats to biological diversity will affect the future of evolution was first raised by one of us in the mid-1980s (8~. It attracted virtually zero interest from fellow biologists. Thirteen years later, he revisited the question, this time with more detailed analysis, although still in exploratory form (9~. This latter publication elicited attention from the National Academy of Sciences, which undertook to sponsor a Colloquium in March 20Q0. As a "scene setter" for Colloquium participants, we drafted an overview account of topics to be tackled, and that draft makes up the bulk of this paper. We hope that it may serve the same purpose for readers of this special section of PNAS. The Core Concept One of the first truisms absorbed by biologists is that evolution is not predictable. We can no more predict the future compo- www.pnas.org/cgi/doi j10. 1 073/pnas.091092498 sition of communities than some Ordovician ecologist could have foreseen the Great Barrier Reef. However, despite our inability to predict the products of evolution the trajectories of future morphologies or the innovations of future physiolo- gies we can make meaningful estimates about evolutionary processes as they will be affected by the depletion of biological diversity. We may have little basis for predicting what large mammals might look like two million years from now, but much better reason to suppose that there will be very few of them. The evolutionary dimension to the current biotic crisis has been vividly expressed by Michael Soule (10~: "Death is one thing, an end to birth is something else." In other words, impending extinctions will be far from the full final outcome of current environmental disruption. At least as important will be the alteration of evolutionary process, and for a period that is difficult to estimate but must surely measure in millions of years. First-Order Effects. There will be several first-order effects stem- miIlg from the biotic crisis: (i) a major extinction of species within the foreseeable future, estimated by some to remove between one-third and two-thirds of all species now extant (1, 2, 5, 11~; (i') a mega-mass extinction of populations, proportionately greater than the mass extinction of species, within the foresee- able future (12); (
ecological types that thrive in human-dominated ecosystems (37, 38~? Proliferation of opportunistic species. reselected and generalist species, often appearing as opportunistic species, may prolifer- ate, especially if there is preferential elimination of K-selected species that include natural controls of reselected populations (32, 38~. Could this proliferation lead to what has been charac- terized as a "pest and weed" ecology (39, 40~? Depletion of "evolutionary powerhouses" in the tropics. Virtu- ally every major group of vertebrates and many large categories of invertebrates and plants originated in spacious zones with warm, equable climates (41, 42~. In addition, tropical species appear to have persisted for relatively brief periods of geologic time, implying high rates of evolutionary turnover and episodes of explosive speciation (21, 43, 44~. According to Jablonski (22), the tropics have been "the engine of biodiversity" for at least 250 million years. Today, we face the prospect of severe depletion if not virtual elimination of tropical forests, wetlands, estuaries, coral reefs, and other biomes, with their exceptional biodiversity and ecological complexity. Because some of these blames ap- pear, in some senses at least, to have served in the past as preeminent "powerhouses" of evolution (45, 46), their decline could entail severe consequences for rediversification as the biosphere emerges from environmental crisis. Decline of biodisparity. Elimination of species is not the only measure of an extinction event. There can be declines, as well, in biodisparity, the biota's manifest morphological and physio- logical variety (47-49~. Biodisparity impoverishment can be assessed through the surrogate measure of loss of higher taxa or guilds, and, over the past 2000 years, the preferential elimination of species-poor genera has reduced biodisparity at rates even greater than those of species loss (48~. Will the same pattern of non-random culling persist in the future? An end to speciation of large vertebrates. Even our largest protected areas will prove far too small for further speciation of elephants, rhinoceroses, apes, bears, and big cats, among other large vertebrates (30, 50, 51~. What knock-on consequences and ripple effects could there be for smaller species, indeed for biotas as a whole given, for example, the depauperizing impacts of the present-day decline of elephants (52~? Emergent novelties. There may be many emergent novelties, although these are especially difficult to predict. For instance, there could be an explosive radiation within certain higher taxa, notably small mammals and insects able to thrive in human- dominated ecosystems. The question is not whether persistent lineages can evolve in unexpected ways, but rather to what extent the environmental constraints humans place on surviving pop- ulations will channel innovations toward properties we associate with pests. Lessons from the Past? The geological record is replete with extinction events, their intensity ranging from the small and local to global mass extinctions that shattered Earth's biological order. Inevitably, extinctions were followed by rediversification, directed in the case of the largest events by ecological reorganization. What can we learn from paleobiology, other than the oft-quoted observa- tion that recovery proceeds slowly in the wake of grand scale biotic disruption (40, 53, 54~? Can we find generalities among extinction episodes that can guide thinking about our own future? Or, is it the differences among extinction events that should command our attention? As David Jablonski (63) asks in these proceedings, should we even focus on the five great mass extinctions that capture most attention, or do the more numer- ous, smaller events scattered throughout the geological record provide closer analogs for the present? The geologic record contains much evidence of bounce-back processes (49, 54-59), but how far will these serve as analytic 5390 1 www.pnas.org/cgi/doi/10.1073/pnas.091092498 blueprints for what lies ahead? How can we estimate time frames at issue? Should we anticipate a minimum period of several million years "perhaps as much as 10 million (56~] before evolution can reestablish anywhere near the biological configu- rations and ecological circuitry existing before the current crisis? Will some recovery processes operate in some sectors of the biosphere, others in others, and with widely varying rates (55, 58, 60~? In some major extinctions, for example the Cretaceous- Tertiary boundary event, environmental perturbation was swift and sure, but also short-lived. Recovery began soon after disruption. In the present biotic crisis, it is hard to envision a scenario under which the factors that are driving the biosphere toward grand scale biodiversity loss will be mitigated in the wake of such loss. On the contrary, on any time scale we can envisage (and any scenario that does not involve early mass mortality for humankind), the situation becomes bad and then stays bad for some time to come. Thus, on the time scale of the human species, environmental disruption (or at least aspects of it) is permanent. Under these circumstances (which may, to some degree, be approximated by the persistent environmental discord after the Permian-Triassic mass extinction), the prospects for rediversifi- cation are limited. Recovery Processes How will ecosystems function in a world of diminished biodi- versity? Does ecosystem function necessarily decay as diversity declines, and if so, by how much and in what manner? Can biodiversity and humans alike prosper in a world where most biological diversity will be confined to relatively small parks and reserves? If biodiversity is indeed critical to ecosystem function, do we know enough about the principles of evolution to intervene in the recovery processes? To the extent that the answer to the first part of this question is probably "yes" and the answer to the second part is almost certainly "no," what would we need to learn to attempt evolutionary interventions that will do more good than harm? More realistically, do we know enough to mitigate the loss of biological diversity? As David Western writes in his colloquium contribution, mitigating strategies will likely be carried out predominantly in ecosystems dominated or influenced by hu- mans and other species that thrive when humans are present. How we think about our evolutionary future depends directly on how successful we can hope to be in preserving biodiversity and biodisparity. Which taxa are likely to play prominent parts in recovery processes? What "survivorship" traits (ecological, biogeo- graphic, evolutionary) can we use to define those taxa that may prove more successful in surviving current events? At the same time, which taxa might tto cite Erwin's graphic phrase (55~] "win the extinction but lose the recovery?" Might certain biotas already be "stressed" by Pleistocene climatic oscillations, making them more vulnerable to depletion (61, 62~? Or are they "hardened" purged of their most vulnerable members by Pleis- tocene events (63~? Should we in fact speak of "recovery"? What is it that is supposed to be recovering (the dinosaurs didn't)? Should we not view the recovery phase as more like a transition to new and novel departures of multiple sorts (55~? Plainly there is much scope for pioneering research in response to the many questions raised (54~. We need to consider planning priorities. What research is most pressing? What is readily achievable? What is already underway? What deserves most financial or institutional support? What potential is there for interdisciplinary research, for instance that which combines genetics and restoration ecol- ogy, or paleontology and conservation biology? Myers and Knoll
Conservation Responses Should we be content simply to safeguard as much as we can of the planetary stock of species? Or should we pay equal if not greater attention to safeguarding evolutionary processes at risk (cf. refs. 64-66~? Consider, for instance, biodisparity: to cite Jablonski (49), "If we are concerned with avoiding the loss of particular functional groups, or with maximizing the potential source pool for evolutionary recovery, then biodisparity mea- sures may provide a more appropriate assessment, beyond sheer numbers of taxa, of how priorities should be set." Following on from these considerations is the question of whether we should seek to maintain the evolutionary status quo by preserving precise phenotypes of particular species, or whether we should prefer to maintain phylogenetic lines that will enable evolutionary adaptations to persist, thereby leading to new species (67, 68~. Is it sufficient for us to maintain, for example, just the two elephant species we already have, or should we try to keep open the evolutionary option of further elephant- like species in the distant future? This is an unusually significant question, with unusually sig- nificant implications for conservation strategies. Elephants, along with many other large mammals, are inclined to move around a good deal, a trait that enables them to maintain gene flow across large areas. As a result, their gene pools often tend to be fairly uniform tan elephant in East Africa may not be so different from one 4,000 km away in South Africa (68~. Re- grettably the remaining populations of elephants, substantial and extensive as they are, albeit fragmented and declining fast, are probably already below the minimum numbers to keep open the possibility of speciation (69~. In marked contrast to elephants, with their slow breeding rates, many insect species have immense breeding capacities and rapid turnover rates. These latter attributes offer quick adapt- ability to environmental shifts, whereupon genetic changes are passed along promptly. These attributes not only leave many insect species well suited to survive the environmental upheavals of human activities, but they offer exceptional scope for specia- 1. Ehrlich, P. R. & Wilson, E. O. (1991) Science 253, 758-762. 2. Myers, N. (1993) Biodiversity and Conserv. 2, 2-17. 3. Pimm, S. L., Russell, G. J., Gittleman, J. L. & Brooks, T. M. (1995) Science 269, 347-354. 4. Raven, P. H. (1999) Plants in Peril: What Should We Do? (Missouri Botanical Garden, St. Louis, MO). 5. Wilson, E. O. (1992) The Diversity of Life (Harvard Univ. Press, Cambridge, MA). 6. Ehrlich, P. R. (2000) Human Natures: Genes, Cultures and the Human Prospect (Island Press, Washington, DC). 7. Erwin, D. H. (2001) Proc. Natl. Acad. Sci. USA 98, 5399-5403. 8. Myers, N. (1985) Nat. Hist. 94, 2, 6, 12. 9. Myers, N. (1996) Environmentalist 16, 37-47. 10. Soule, M. E. (1980) in Conservation Biology: An Evolutionary-Ecological Perspective, eds. Soule, M.E. & Wilcox, B.A. (Sinauer, Sunderland, MA), pp. 151-170. 11. Pimm, S. & Raven, P. (2000) Nature (London) 403, 843-845. 12. Hughes, J. B., Daily, G. C. & Ehrlich, P. R. (1997) Science 278, 689-692. 13. Drake, J. A., Mooney, H. A., diCastri, F., Groves, R., Kruger, F., Rejmanek, M. & Williamson, M., eds. (1989) Biological Invasions: A Global Perspective. (Wiley, New York). 14. Mooney, H. A. & Hobbs, R. J., eds. (2000) Invasive Species in a Changing World (Island Press, Washington, DC). 15. Vermeij, G. J. (1991) Science 253, 1099-1103. 16. Mooney, H. A. & Cleland, E. E. (2001) Proc. Natl. Acad. Sci. USA 98, 5446-5451. 17. Walliser, O. (1995) Global Events and Event Stratigraphy (Springer, New York). 18. Woodwell, G. M., ed. (1990) The Earth in Transition: Patterns and Processes of Biotic Impoverishment (Cambridge Univ. Press, Cambridge, U.K.). 19. Hsu, K. J. (1989) Hist. Biol. 2, 1-4. 20. McLaren, D. J. (1989) Hist. Biol. 2, 5-15. 21. Briggs, J. C. (1996) Conserv. Biol. 10, 713-718. 22. Jablonski, D. (1993) Nature (London) 364, 142-144. 23. Raup, D. M. (1991) Extinction: Bad Genes or Bad Luck? (Norton, New York). Myers and Knoll tion in comparatively short order. By contrast, elephants, to- gether with other large-bodied species that reproduce slowly and hence possess restricted capacity for genetic adaptation, will be at an extreme evolutionary disadvantage. Does this factor imply that they should therefore receive all of the greater attention from conservationists or that, in a triage situation, they should rank lower in our priorities? Although this is a fundamental question, it has hardly been addressed. An even more important consideration arises concerning those origination centers and radiation lineages that serve as "evolutionary fronts" (67~. From the standpoint of future evo- lution, it is surely more appropriate to safeguard the main potential for diversity generation than to emphasize the primary focus of many current conservation programs, viz. individual taxa and, especially, endemic taxa (70, 71~. Much the same applies with respect to those functional groups that increase the potential for evolutionary recovery (49~. All in all, the prospect is that, in the wake of the present biodiversity crisis, we shall find that many evolutionary processes that have persisted throughout the Phanerozoic Eon will be slowed if not depauperized for an extended period. This is not to say, of course, that evolution will come to a halt, or even that speciation will be suspended (except for the large vertebrates). 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