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<br /> <br />evidence that increased body size and metabolic requirements result in <br />higher extinction rates in vertebrates and Terborgh and Winter in Chap- <br />ter 7 argue that population size or density is the best predictor of extinc- <br />tion in birds; thus on small islands, the probability of extinction for large <br />organisms is very high. Assuming that their generalizations hold up, it <br />means that the apparent thresholds for autochthonous speciation could, <br />at least in part, be attributable to thresholds for speciation-extinction <br />equilibria for numbers of species greater than one per genus.) <br />In practice it makes little difference whether the area effect on specia- <br />tion in Figure 4 is a reflection of taxon-specific thresholds for autochtho- <br />nous speciation, or is a reflection of thresholds for speciation-extinction <br />equilibria. Even if it is the latter, which I think is unlikely, it means that <br />existing nature reserves are still too small to generate more species than <br />they now contain. In fact, they will contain fewer and fewer species with <br />the passage of time because most reserves are now supersaturated and <br />will lose species rapidly over the next few centuries (Chapter 6). It there- <br />fore appears inescapable that, for the first time in hundreds of millions of <br />years, significant evolutionary change in most higher organisms is coming <br />to a screeching halt in the tropics. <br />One might object that this entire argument is misleading on the <br />grounds that reserves can support speciational processes because of the <br />isolation between them (that is, on the model of archipelagos). The mini- <br />mum requirements for this kind of speciation is that a species occur in <br />two or more reserves, and that the reserves be isolated long enough for <br />the evolution of intrinsic isolating mechanisms such as differences in <br />habitat preferences or post-mating reproductive barriers. Superficially, <br />this model seems reasonable, assuming that reserves are continuously via- <br />ble for thousands of years. <br />The flaw in this line of thought is that it ignores the high rate of <br />extinction of large organisms on islands. Figure 7 in Chapter 6 shows that <br />we cannot count on the species surviving long enough to speciate, even in <br />the largest tropical reserves. In fact, the extinction rate is so high that <br />half of the original large mammals species in such a reserve are expected <br />to go extinet in from 500 to 2000 years unless, that is, man intercedes. <br />The irony is that such intercession itself will prevent speciation of this <br />type from occurring. The prevention of extinction of such animals will <br />require the exchange of individuals between the populations in the differ- <br />ent reserves, this being the only available means of preventing genetic <br />deterioration of the largest or least dense specieS. Large doses of gene <br />flow, of course, are inimical to incipient speciation and the development <br />of locally adapted races because it turns the subdivided species into a <br />genetic omelette. But even if genetic deterioration were not a factor and <br />gene flow was not a necessary preventative therapy, artificial migration <br />would still be necessary in most instances in order to prevent stochastic <br /> <br />166 <br />