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<br />le._ *r <br />4 4 t" I <br />in. <br />_ i <br />lc .. <br />at., <br />er, <br />?s <br />le <br />s <br />d. <br />CS <br />r<:; <br />fe - <br />it <br />in . <br />a : 'r <br />f- <br />Ie .e <br />is 1E <br />to <br />is <br />m <br />in <br />Ie <br />at <br />ct = 3 <br />ro <br />in <br />rs <br />1e <br />S_ <br />at <br />1- <br />is <br />'o <br />d <br />X <br />is <br />A <br />)n <br />a- <br />1 <br />ld 1 <br />ld <br />I- " <br />W <br />1t <br />estimates put the total population size of the northern sported owl at <br />2500 pairs (48). In western Oregon and Washington, the remaining <br />old-growth forest is restricted mainly to 12 national forests that arc <br />largely contiguous. To comply with the National Forest .Manage- <br />ment Act of 1976, which requires that native verrcbrate species be <br />maintained well distributed throughout their range on federal land, <br />the U.S. Forest Service developed a plan to preserve the northern <br />sported owl. Originally, this was based on the supposition that <br />protection from logging of territories for about 500 pairs distribut- <br />ed throughout the region would maintain enough genetic variability <br />for the population ro survive (23). However, models of stochastic <br />demography and habitat occupancy indicate that the plan is likely to <br />cause extinction of the owl because suitable habitat in the region will <br />be too sparsely distributed to support a population (49). An <br />independent assessment by the Forest Service also predicts extinc- <br />tion on demographic grounds, but essentially the same management <br />strategy remains in effect (50). <br />The red-cockaded woodpecker, Picoides borealis, ranges across the <br />southeastern United States, inhabiting pine forests more than about <br />80 years old, most of which currently exist on federal lands. Their <br />preferred habitat has substantial openings, and is maintained by <br />recurring fires that prevent succession to hardwoods. Thew birds <br />live in colonies, composed of one breeding pair and up to several <br />offspring serving as helpers, that occupy an annual home range <br />averaging about 215 acres in which they forage for insects. Nesting <br />occurs in cavities that may take a year for the birds to hollow out of <br />living, mature longleaf pines (80 to 120 years old) that have had <br />their heartwood destroyed by a fungus. The total size of the <br />breeding population was recently estimated to be 6000 individuals <br />(51). The recovery plan for the red-cockaded woodpecker, based on <br />genetic considerations, has as its goal the establishment of local <br />populations of 250 clans (500 breeding individuals) (24). The <br />species has been listed as federally endangered since the passage of <br />the Endangered Species Act in 1973, but subsequently has declined <br />rapidly throughout its range u a result of fire prevention and <br />logging of suitable, unoccupiet: habitat, which has severclv frag- <br />mentcd the remaining suitable habitat (51). For example, under <br />management by the U.S. Forest Service, populations on national <br />forests in Texas have been declining by about 10% per year (52). <br />Unless management practices arc drastically altered, it seems that the <br />red-cockaded woodpecker will soon be etrirla. <br />Scientific advisory panels reviewed the inadequacies of the man- <br />agement plans for the northern spotted owl and the red-cockaded <br />woodpecker and in both cases recommended abandoning simple <br />genetic rules for establishing minimum viable population sizes (48, <br />51). Management of particular species should incorporate details of <br />the species ecology, especially its life history and demography, <br />which may require larger populations than has been suggested on <br />genetic grounds alone. <br />Since conservation of the northern sported owl and the red- <br />cockaded woodpecker involves preserving habitats worth billions of <br />dollars to the lumber and paper industries, in principle there should <br />be no difficulty in funding long-term scientific studies to obtain the <br />information necessary for sound management decisions. Already a <br />great deal is known about both species. Why then does it appear that <br />the conservation plans for these species are unsuccessful? Short-term <br />economic and political interests often dominate scientific consider- <br />ations in the development and implementation of management <br />plans for threatened or endangered species. Whether economics and <br />politics continue to produce scientificallv deficient conservation <br />plans will be decided in manv cases only by extended litigation. <br />Future conservation plans should incorporate both deco aphv <br />T- genetics inIDassessing etheth regwrenrnts for species <br />sill 'rely" , ra)?ecognizing that tocKw[ld po ulaaons" do o _ap`(uc taaors <br /> <br />G <br />may usuallv be of more immediate im trance than genetic factors. <br />Alva lsnc mtegntuion o demography and pop on genetics, <br />applicable to species in natural environments, is a formidable task <br />that has enticed but largely eluded ecologists and evolutionary <br />biologists. The immediate practical need in biological conservation <br />for understanding the interaction of demographic and genetic <br />factors in the extinction of small populations therefore may provide <br />a focus for fundamental advances at the interface of ecology and <br />evolution. <br />REFERENCES AND NOTES <br />1. N. Myers, in Conservation Biology, the Science of Searoty and Diversity, M. E. Stink, <br />Ed. (Sinauer, Sunderland. M& 1996). pp. 411 161; D. Simberlotf, in Dynamio of <br />E:tinaion, D. K. Elliot, Ed. (Wiley, New York 1986), pp. 165-180; D. labkxnski, <br />Snenu 231, 129 (1986). <br />2. E. C. Wolf; in State ofthe World 1988, L R. Brown et al., Ede. (Norma. New York <br />1988), pp. 101-117 and 210-213. <br />3. M. L Oldfield, The Value of Conserving Genetic Resancer (U.S. National Park <br />Service, Washingam, DC, 1984). <br />4. C, H. Southwick Ed., Global Ecology (Sinauer, Sunderland, MA. 19831. <br />S. R. Lando and G. F. Banowdough, in Viable Populatwtu Jut Conservation, M. E. <br />Soule, Ed. (Cambridge Univ. Press, New York 1987), pp. 87-123; R. Lunde, <br />Am. Nat. 116,463 (1980). <br />6. R. H. MacArthur and E. O. Wilson, The Theory of Island Biagro. apky (Princeton <br />Univ. Press, Princeton, 1967); D. K. Elliot, Ed.. Dynania of Extinction (Wife%-, <br />New York 1986). <br />7. M. E. Souii and D. Simberloff, Biol. Conserv. 35, 19 (1986). <br />8. M. E. Souk and B. A. Wilcox, Eds., Conservation Biology, in Evolutionary-Fealogiml <br />Perspective (Sinauer, Sunderland. AMA, 19801; O. H. Frarnkel and M. E. SoultiConservation and Evolution (Cambridge Univ. Press, New York 19811; C. M. <br />Schonewald-Cox. S. M. Chambers, B. MacBryde, L Thomas. Eds., Generics and <br />Conservation: A Reformer for Untoging Wild Animal and Plans Populations i Benjamrn- <br />Cummin8s, London, 1983); M. E. SoWl _ Ed.. Conservation Biology, the Science of <br />Starnty and Diversity (Sinauer, Sunderland, MA, 1986). <br />9. D. S. Falconer, Introduction to Quantitative Generics (Longman, London, ed. 2, <br />1981). <br />10. S. Wright, Evolution and the Genetics of Populations, Experimental Results and Evolution- <br />ary Dedumonu (Univ. of Chicago Press, Chicago. 1977), vol. 3. <br />11. M. J. Simmons and 1. F. Crow, Amtu. Rev. Genet. 11, 49 (1977). <br />12. R. Lando and D. W. Schemske- Evolution 39, 24 (1985); D. Charlesworth and B. <br />Charlesworth, Amu. Rev. Flol. Syst. 18, 237 (1987). <br />13. K. Ralls and 1. Ballou. in, Genetics and Conservation: A Ref re= l6? :If&%ggmq Wild <br />Animal and Plant Populations. C. M. Schcxwald-Cox, S. M. Chambers, B. Mac- <br />Bryde, L Thomas, Eds. (Benjamin-Cumrritngs, London, 1983). pp. 164-184. <br />14. K. Ralls.atnd J. D. Ballots, Eds. Zoo Biol. 5 (no. 2), pp. 81-238 (enure issue), <br />(1986). <br />15. O. A. Schwartz, V. C. Bleieh, S. A. Holl, Biol. Conserv. 37, 179 (1986). <br />16. J. W. Senor, in Conservation Biology, an Evolutionary-Ecologual Perspective, M. E. <br />Sould and B. A. Wilma, Eds. (Sinauer, Sunderland. .AM, 1980), pp. 209-224. <br />17. R. C. Lewontin, The Genetic Basis of Evolutionary Change (Columbu Univ. Press, <br />New York, 1974), p. 91. <br />18. S. Wright, Evolution and the Genetics of Populations. the Theory of Gene Frequenntier <br />(Univ. of Chicago Press, Chicago, 1969), vol. 2, <br />19. S. Wright, Ann. Eugert. 15, 323 (1951); A. Robertson, Genetics 37,189 (1952); C. <br />1. Goodnight, Evolution 41.80 (1987); E. H. Brvam S. A. McCommas, L. M. <br />Combs, Genetics 114, 1191 (1987). <br />20. 1L Londe, Genet. Res. 26,221 (1975); see also W. G. Hill ibid. 40,225 (1982); M. <br />Lynch, Evolution 39, 804 (1985); M. Lynch, Genet. Res. 51, 137 (1988). <br />21. 1. R. Franklin, in Conservation Biology, in Evolutionary-Ecological Perspective, M. E. <br />SoulE and B. A. Wilcox, Eds. (Sinauer, Sunderland. MA, 1980), pp. 135-149. <br />22. U. S. Seal and T. Foose, J..t/init. Acad. Sn. 49, 3 (1983/84). <br />23. Final Regional Guide and Final Environmental Impact Statement fir the Pon_ht Northwest <br />Region (U.S. Forest Service, Portland, OR. 1984). <br />24. Red-cockaded Woodpecker Recovery Plan (U.S. Fish and Wildlife Service, Atlanta, GA, <br />1985). <br />25. F. W. ABendorf et al., Heredirat 91, 19 (1979); S. O'Brien et al., Sciew 221, 459 <br />(1983). <br />26. G. G. Simpson. The .Naior Features of Evolution (Columbia Univ. Press, New York, <br />19531, pp. 77-80 and 129-132. <br />27. M. SLidun and R. Linde, Am. Nat. 110, 31 (1976); B. Chadesworth, Paleobiology <br />10. 319 (1984). <br />28. P. R. Ehrlich, in Genetics and Conservation: A Refrrerue for Iblamogmng V4 old .Animal and <br />Plain Populations, C. M. Schonewald-Cox, S. M. Chambers. B. MacBrvdc, L <br />Thomas, Eds. (Benjamin-Cummings, London, 1983), pp. 152-163. <br />29. T. Foose, Ito. Zoo Yearb. 20, 154 (1980); D. G xxlff t. in Conservation Biology, in <br />Evolutionary-Ecology Perspective, M. E. SouM and B. A. Wilcox, Eds. (Sinauer, <br />Sunderland, MA, 1980), pp. 171-195. <br />30. W. C. Allee et al., Principles of Animal Ecology (Saunders, London, 1949); H. G. <br />Andrewartha and L. C. Birch. The Distribution and Abundance of Animals (Univ. of <br />Chicago Press. Chicago, 1954). <br />31. E. G. Leigh, Jr., J. Theor. Biol. 90, 213 (1981). <br />32. D. Goodman, in Viable Povulations for Conservation, M. E. Soul& Ed. !Cambridge <br />p. 11-34. <br />Univ. Press, New York 1987), pp.- <br />ARTICLES <br />4-1 16 SEPTEMBER 1988 [459