Light-gap disturbances, recruitment limitation, and tree diversity in a neotropical forest

Science

Volumn 283, Issue 5401, 1999, Pages 554-557

  Hubbell, S P ^a   Foster, R B ^b   O'Brien, S T ^a,c   Harms, K E ^b   Condit, R ^b   Wechsler, B ^a   Wright, S J ^b   Loo De Lao, S ^b  

^a PRINCETON UNIVERSITY   (United States)

^b SMITHSONIAN TROPICAL RESEARCH INSTITUTE   (Panama)

^c W Alton Jones Foundation   (United States)

Author keywords

[No Author keywords available]

Indexed keywords

FOREST; SPECIES DIVERSITY; TREE;

ARTICLE; BIODIVERSITY; FOREST; PLANT SEED; PRIORITY JOURNAL; RECEPTOR OCCUPANCY; SPECIES DIFFERENTIATION; TREE; TROPIC CLIMATE;

EID: 0033593581     PISSN: 00368075     EISSN: None     Source Type: Journal    
DOI: 10.1126/science.283.5401.554     Document Type: Article

References (45)

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5. D. R. Strong, J. Biogeogr. 4, 215 (1977); R. E. Ricklefs, Am. Nat. 1111, 376 (1977); G. S. Hartshorn, in Tropical Trees as Living Systems, P. B. Tomlinson and M. H. Zimmerman, Eds. (Cambridge Univ. Press, New York, 1978), pp. 617-638; F. A. Bazzaz and S. T. A. Pickett, Annu. Rev. Ecol. Syst. 11, 287 (1980); J. S. Denslow, Biotropica (Suppl.) 12, 47 (1980); G. H. Orians, Trop. Ecol. 23, 255 (1982); _, in The Ecology of Natural Disturbance and Patch Dynamics, S. T. A. Pickett and P. S. White, Eds. (Academic Press, New York, 1985), pp. 307-323; Annu. Rev. Ecol. Syst. 18, 431 (1987); T. C. Whitmore, Ecology 70, 536 (1989); M. D. Swaine and T. C. Whitmore, Vegetatio 75, 81 (1988); A. Brandani et al., J. Trop Ecol. 4, 99 (1988); P. M. Vitousek and J. S. Denslow, J. Ecol. 74, 1167 (1986); J. S. Denslow et al., Oecologia 74, 370 (1987); Ecology 71, 165 (1990); Ecology 66, 682 (1985); E. W. Schupp et al., Ecology 70, 562 (1989);
6. D. R. Strong, J. Biogeogr. 4, 215 (1977); R. E. Ricklefs, Am. Nat. 1111, 376 (1977); G. S. Hartshorn, in Tropical Trees as Living Systems, P. B. Tomlinson and M. H. Zimmerman, Eds. (Cambridge Univ. Press, New York, 1978), pp. 617-638; F. A. Bazzaz and S. T. A. Pickett, Annu. Rev. Ecol. Syst. 11, 287 (1980); J. S. Denslow, Biotropica (Suppl.) 12, 47 (1980); G. H. Orians, Trop. Ecol. 23, 255 (1982); _, in The Ecology of Natural Disturbance and Patch Dynamics, S. T. A. Pickett and P. S. White, Eds. (Academic Press, New York, 1985), pp. 307-323; Annu. Rev. Ecol. Syst. 18, 431 (1987); T. C. Whitmore, Ecology 70, 536 (1989); M. D. Swaine and T. C. Whitmore, Vegetatio 75, 81 (1988); A. Brandani et al., J. Trop Ecol. 4, 99 (1988); P. M. Vitousek and J. S. Denslow, J. Ecol. 74, 1167 (1986); J. S. Denslow et al., Oecologia 74, 370 (1987); Ecology 71, 165 (1990); Ecology 66, 682 (1985); E. W. Schupp et al., Ecology 70, 562 (1989);
7. D. R. Strong, J. Biogeogr. 4, 215 (1977); R. E. Ricklefs, Am. Nat. 1111, 376 (1977); G. S. Hartshorn, in Tropical Trees as Living Systems, P. B. Tomlinson and M. H. Zimmerman, Eds. (Cambridge Univ. Press, New York, 1978), pp. 617-638; F. A. Bazzaz and S. T. A. Pickett, Annu. Rev. Ecol. Syst. 11, 287 (1980); J. S. Denslow, Biotropica (Suppl.) 12, 47 (1980); G. H. Orians, Trop. Ecol. 23, 255 (1982); _, in The Ecology of Natural Disturbance and Patch Dynamics, S. T. A. Pickett and P. S. White, Eds. (Academic Press, New York, 1985), pp. 307-323; Annu. Rev. Ecol. Syst. 18, 431 (1987); T. C. Whitmore, Ecology 70, 536 (1989); M. D. Swaine and T. C. Whitmore, Vegetatio 75, 81 (1988); A. Brandani et al., J. Trop Ecol. 4, 99 (1988); P. M. Vitousek and J. S. Denslow, J. Ecol. 74, 1167 (1986); J. S. Denslow et al., Oecologia 74, 370 (1987); Ecology 71, 165 (1990); Ecology 66, 682 (1985); E. W. Schupp et al., Ecology 70, 562 (1989);
8. D. R. Strong, J. Biogeogr. 4, 215 (1977); R. E. Ricklefs, Am. Nat. 1111, 376 (1977); G. S. Hartshorn, in Tropical Trees as Living Systems, P. B. Tomlinson and M. H. Zimmerman, Eds. (Cambridge Univ. Press, New York, 1978), pp. 617-638; F. A. Bazzaz and S. T. A. Pickett, Annu. Rev. Ecol. Syst. 11, 287 (1980); J. S. Denslow, Biotropica (Suppl.) 12, 47 (1980); G. H. Orians, Trop. Ecol. 23, 255 (1982); _, in The Ecology of Natural Disturbance and Patch Dynamics, S. T. A. Pickett and P. S. White, Eds. (Academic Press, New York, 1985), pp. 307-323; Annu. Rev. Ecol. Syst. 18, 431 (1987); T. C. Whitmore, Ecology 70, 536 (1989); M. D. Swaine and T. C. Whitmore, Vegetatio 75, 81 (1988); A. Brandani et al., J. Trop Ecol. 4, 99 (1988); P. M. Vitousek and J. S. Denslow, J. Ecol. 74, 1167 (1986); J. S. Denslow et al., Oecologia 74, 370 (1987); Ecology 71, 165 (1990); Ecology 66, 682 (1985); E. W. Schupp et al., Ecology 70, 562 (1989);
9. D. R. Strong, J. Biogeogr. 4, 215 (1977); R. E. Ricklefs, Am. Nat. 1111, 376 (1977); G. S. Hartshorn, in Tropical Trees as Living Systems, P. B. Tomlinson and M. H. Zimmerman, Eds. (Cambridge Univ. Press, New York, 1978), pp. 617-638; F. A. Bazzaz and S. T. A. Pickett, Annu. Rev. Ecol. Syst. 11, 287 (1980); J. S. Denslow, Biotropica (Suppl.) 12, 47 (1980); G. H. Orians, Trop. Ecol. 23, 255 (1982); _, in The Ecology of Natural Disturbance and Patch Dynamics, S. T. A. Pickett and P. S. White, Eds. (Academic Press, New York, 1985), pp. 307-323; Annu. Rev. Ecol. Syst. 18, 431 (1987); T. C. Whitmore, Ecology 70, 536 (1989); M. D. Swaine and T. C. Whitmore, Vegetatio 75, 81 (1988); A. Brandani et al., J. Trop Ecol. 4, 99 (1988); P. M. Vitousek and J. S. Denslow, J. Ecol. 74, 1167 (1986); J. S. Denslow et al., Oecologia 74, 370 (1987); Ecology 71, 165 (1990); Ecology 66, 682 (1985); E. W. Schupp et al., Ecology 70, 562 (1989);
10. D. R. Strong, J. Biogeogr. 4, 215 (1977); R. E. Ricklefs, Am. Nat. 1111, 376 (1977); G. S. Hartshorn, in Tropical Trees as Living Systems, P. B. Tomlinson and M. H. Zimmerman, Eds. (Cambridge Univ. Press, New York, 1978), pp. 617-638; F. A. Bazzaz and S. T. A. Pickett, Annu. Rev. Ecol. Syst. 11, 287 (1980); J. S. Denslow, Biotropica (Suppl.) 12, 47 (1980); G. H. Orians, Trop. Ecol. 23, 255 (1982); _, in The Ecology of Natural Disturbance and Patch Dynamics, S. T. A. Pickett and P. S. White, Eds. (Academic Press, New York, 1985), pp. 307-323; Annu. Rev. Ecol. Syst. 18, 431 (1987); T. C. Whitmore, Ecology 70, 536 (1989); M. D. Swaine and T. C. Whitmore, Vegetatio 75, 81 (1988); A. Brandani et al., J. Trop Ecol. 4, 99 (1988); P. M. Vitousek and J. S. Denslow, J. Ecol. 74, 1167 (1986); J. S. Denslow et al., Oecologia 74, 370 (1987); Ecology 71, 165 (1990); Ecology 66, 682 (1985); E. W. Schupp et al., Ecology 70, 562 (1989);
11. D. R. Strong, J. Biogeogr. 4, 215 (1977); R. E. Ricklefs, Am. Nat. 1111, 376 (1977); G. S. Hartshorn, in Tropical Trees as Living Systems, P. B. Tomlinson and M. H. Zimmerman, Eds. (Cambridge Univ. Press, New York, 1978), pp. 617-638; F. A. Bazzaz and S. T. A. Pickett, Annu. Rev. Ecol. Syst. 11, 287 (1980); J. S. Denslow, Biotropica (Suppl.) 12, 47 (1980); G. H. Orians, Trop. Ecol. 23, 255 (1982); _, in The Ecology of Natural Disturbance and Patch Dynamics, S. T. A. Pickett and P. S. White, Eds. (Academic Press, New York, 1985), pp. 307-323; Annu. Rev. Ecol. Syst. 18, 431 (1987); T. C. Whitmore, Ecology 70, 536 (1989); M. D. Swaine and T. C. Whitmore, Vegetatio 75, 81 (1988); A. Brandani et al., J. Trop Ecol. 4, 99 (1988); P. M. Vitousek and J. S. Denslow, J. Ecol. 74, 1167 (1986); J. S. Denslow et al., Oecologia 74, 370 (1987); Ecology 71, 165 (1990); Ecology 66, 682 (1985); E. W. Schupp et al., Ecology 70, 562 (1989);
12. D. R. Strong, J. Biogeogr. 4, 215 (1977); R. E. Ricklefs, Am. Nat. 1111, 376 (1977); G. S. Hartshorn, in Tropical Trees as Living Systems, P. B. Tomlinson and M. H. Zimmerman, Eds. (Cambridge Univ. Press, New York, 1978), pp. 617-638; F. A. Bazzaz and S. T. A. Pickett, Annu. Rev. Ecol. Syst. 11, 287 (1980); J. S. Denslow, Biotropica (Suppl.) 12, 47 (1980); G. H. Orians, Trop. Ecol. 23, 255 (1982); _, in The Ecology of Natural Disturbance and Patch Dynamics, S. T. A. Pickett and P. S. White, Eds. (Academic Press, New York, 1985), pp. 307-323; Annu. Rev. Ecol. Syst. 18, 431 (1987); T. C. Whitmore, Ecology 70, 536 (1989); M. D. Swaine and T. C. Whitmore, Vegetatio 75, 81 (1988); A. Brandani et al., J. Trop Ecol. 4, 99 (1988); P. M. Vitousek and J. S. Denslow, J. Ecol. 74, 1167 (1986); J. S. Denslow et al., Oecologia 74, 370 (1987); Ecology 71, 165 (1990); Ecology 66, 682 (1985); E. W. Schupp et al., Ecology 70, 562 (1989);
13. D. R. Strong, J. Biogeogr. 4, 215 (1977); R. E. Ricklefs, Am. Nat. 1111, 376 (1977); G. S. Hartshorn, in Tropical Trees as Living Systems, P. B. Tomlinson and M. H. Zimmerman, Eds. (Cambridge Univ. Press, New York, 1978), pp. 617-638; F. A. Bazzaz and S. T. A. Pickett, Annu. Rev. Ecol. Syst. 11, 287 (1980); J. S. Denslow, Biotropica (Suppl.) 12, 47 (1980); G. H. Orians, Trop. Ecol. 23, 255 (1982); _, in The Ecology of Natural Disturbance and Patch Dynamics, S. T. A. Pickett and P. S. White, Eds. (Academic Press, New York, 1985), pp. 307-323; Annu. Rev. Ecol. Syst. 18, 431 (1987); T. C. Whitmore, Ecology 70, 536 (1989); M. D. Swaine and T. C. Whitmore, Vegetatio 75, 81 (1988); A. Brandani et al., J. Trop Ecol. 4, 99 (1988); P. M. Vitousek and J. S. Denslow, J. Ecol. 74, 1167 (1986); J. S. Denslow et al., Oecologia 74, 370 (1987); Ecology 71, 165 (1990); Ecology 66, 682 (1985); E. W. Schupp et al., Ecology 70, 562 (1989);
14. D. R. Strong, J. Biogeogr. 4, 215 (1977); R. E. Ricklefs, Am. Nat. 1111, 376 (1977); G. S. Hartshorn, in Tropical Trees as Living Systems, P. B. Tomlinson and M. H. Zimmerman, Eds. (Cambridge Univ. Press, New York, 1978), pp. 617-638; F. A. Bazzaz and S. T. A. Pickett, Annu. Rev. Ecol. Syst. 11, 287 (1980); J. S. Denslow, Biotropica (Suppl.) 12, 47 (1980); G. H. Orians, Trop. Ecol. 23, 255 (1982); _, in The Ecology of Natural Disturbance and Patch Dynamics, S. T. A. Pickett and P. S. White, Eds. (Academic Press, New York, 1985), pp. 307-323; Annu. Rev. Ecol. Syst. 18, 431 (1987); T. C. Whitmore, Ecology 70, 536 (1989); M. D. Swaine and T. C. Whitmore, Vegetatio 75, 81 (1988); A. Brandani et al., J. Trop Ecol. 4, 99 (1988); P. M. Vitousek and J. S. Denslow, J. Ecol. 74, 1167 (1986); J. S. Denslow et al., Oecologia 74, 370 (1987); Ecology 71, 165 (1990); Ecology 66, 682 (1985); E. W. Schupp et al., Ecology 70, 562 (1989);
15. D. R. Strong, J. Biogeogr. 4, 215 (1977); R. E. Ricklefs, Am. Nat. 1111, 376 (1977); G. S. Hartshorn, in Tropical Trees as Living Systems, P. B. Tomlinson and M. H. Zimmerman, Eds. (Cambridge Univ. Press, New York, 1978), pp. 617-638; F. A. Bazzaz and S. T. A. Pickett, Annu. Rev. Ecol. Syst. 11, 287 (1980); J. S. Denslow, Biotropica (Suppl.) 12, 47 (1980); G. H. Orians, Trop. Ecol. 23, 255 (1982); _, in The Ecology of Natural Disturbance and Patch Dynamics, S. T. A. Pickett and P. S. White, Eds. (Academic Press, New York, 1985), pp. 307-323; Annu. Rev. Ecol. Syst. 18, 431 (1987); T. C. Whitmore, Ecology 70, 536 (1989); M. D. Swaine and T. C. Whitmore, Vegetatio 75, 81 (1988); A. Brandani et al., J. Trop Ecol. 4, 99 (1988); P. M. Vitousek and J. S. Denslow, J. Ecol. 74, 1167 (1986); J. S. Denslow et al., Oecologia 74, 370 (1987); Ecology 71, 165 (1990); Ecology 66, 682 (1985); E. W. Schupp et al., Ecology 70, 562 (1989);
16. D. R. Strong, J. Biogeogr. 4, 215 (1977); R. E. Ricklefs, Am. Nat. 1111, 376 (1977); G. S. Hartshorn, in Tropical Trees as Living Systems, P. B. Tomlinson and M. H. Zimmerman, Eds. (Cambridge Univ. Press, New York, 1978), pp. 617-638; F. A. Bazzaz and S. T. A. Pickett, Annu. Rev. Ecol. Syst. 11, 287 (1980); J. S. Denslow, Biotropica (Suppl.) 12, 47 (1980); G. H. Orians, Trop. Ecol. 23, 255 (1982); _, in The Ecology of Natural Disturbance and Patch Dynamics, S. T. A. Pickett and P. S. White, Eds. (Academic Press, New York, 1985), pp. 307-323; Annu. Rev. Ecol. Syst. 18, 431 (1987); T. C. Whitmore, Ecology 70, 536 (1989); M. D. Swaine and T. C. Whitmore, Vegetatio 75, 81 (1988); A. Brandani et al., J. Trop Ecol. 4, 99 (1988); P. M. Vitousek and J. S. Denslow, J. Ecol. 74, 1167 (1986); J. S. Denslow et al., Oecologia 74, 370 (1987); Ecology 71, 165 (1990); Ecology 66, 682 (1985); E. W. Schupp et al., Ecology 70, 562 (1989);
17. D. R. Strong, J. Biogeogr. 4, 215 (1977); R. E. Ricklefs, Am. Nat. 1111, 376 (1977); G. S. Hartshorn, in Tropical Trees as Living Systems, P. B. Tomlinson and M. H. Zimmerman, Eds. (Cambridge Univ. Press, New York, 1978), pp. 617-638; F. A. Bazzaz and S. T. A. Pickett, Annu. Rev. Ecol. Syst. 11, 287 (1980); J. S. Denslow, Biotropica (Suppl.) 12, 47 (1980); G. H. Orians, Trop. Ecol. 23, 255 (1982); _, in The Ecology of Natural Disturbance and Patch Dynamics, S. T. A. Pickett and P. S. White, Eds. (Academic Press, New York, 1985), pp. 307-323; Annu. Rev. Ecol. Syst. 18, 431 (1987); T. C. Whitmore, Ecology 70, 536 (1989); M. D. Swaine and T. C. Whitmore, Vegetatio 75, 81 (1988); A. Brandani et al., J. Trop Ecol. 4, 99 (1988); P. M. Vitousek and J. S. Denslow, J. Ecol. 74, 1167 (1986); J. S. Denslow et al., Oecologia 74, 370 (1987); Ecology 71, 165 (1990); Ecology 66, 682 (1985); E. W. Schupp et al., Ecology 70, 562 (1989);
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20. F. A. Bazzaz, Plants in Changing Environments (Cambridge Univ. Press, Cambridge, 1996);
21. D. Tilman, Plant Strategies and the Dynamics and Structure of Plant Communities (Princeton Univ. Press, Princeton, NJ, 1988);
22. D. A. Clark and D. B. Clark, Ecol. Monogr. 62, 315 (1992).
23. The climate, soils, and an overview of the ecology of the BCI forest can be found in E. G. Leigh Jr. et al., Eds. The Ecology of a Tropical Forest: Seasonal Rhythms and Long-Term Changes (Smithsonian Institution Press, Washington, DC, 1982) and in T. Croat, The Flora of Barro Colorado Island (Stanford Univ. Press, Stanford, CA, 1978).
24. The climate, soils, and an overview of the ecology of the BCI forest can be found in E. G. Leigh Jr. et al., Eds. The Ecology of a Tropical Forest: Seasonal Rhythms and Long-Term Changes (Smithsonian Institution Press, Washington, DC, 1982) and in T. Croat, The Flora of Barro Colorado Island (Stanford Univ. Press, Stanford, CA, 1978).
25. R. Condit et al., Bioscience 42, 822 (1992).
26. Canopy heights were classified into five height classes: 0 m (canopy ≤ m), 2 m (canopy > 2 and ≤ 5 m), 5m (canopy > 5and ≤ 10 m), 10 m (canopy > 10 and ≤ 20 m), and > 20 m. Gaps in our analysis all had canopy heights in the 0-m or 2-m height classes; that is, foliage may have been present between 0 and 5 m above the ground, but not above. This height was appropriate for the 1 cm dbh sapling, which had a mean height of 2.3 m.
27. S. P. Hubbell and R. B. Foster, Rev. Biol. Trop. (Suppl. 1) 35, 22 (1987); C. W. Weiden, S. W. Hewett, S. P. Hubbell, R. B. Foster, Ecology 72, 35 (1991); R. C. Condit, S. P. Hubbell, R. B. Foster, Ecol. Monogr. 65, 419 (1995); J. Trop. Ecol. 12, 231 (1996).
28. S. P. Hubbell and R. B. Foster, Rev. Biol. Trop. (Suppl. 1) 35, 22 (1987); C. W. Weiden, S. W. Hewett, S. P. Hubbell, R. B. Foster, Ecology 72, 35 (1991); R. C. Condit, S. P. Hubbell, R. B. Foster, Ecol. Monogr. 65, 419 (1995); J. Trop. Ecol. 12, 231 (1996).
29. S. P. Hubbell and R. B. Foster, Rev. Biol. Trop. (Suppl. 1) 35, 22 (1987); C. W. Weiden, S. W. Hewett, S. P. Hubbell, R. B. Foster, Ecology 72, 35 (1991); R. C. Condit, S. P. Hubbell, R. B. Foster, Ecol. Monogr. 65, 419 (1995); J. Trop. Ecol. 12, 231 (1996).
30. S. P. Hubbell and R. B. Foster, Rev. Biol. Trop. (Suppl. 1) 35, 22 (1987); C. W. Weiden, S. W. Hewett, S. P. Hubbell, R. B. Foster, Ecology 72, 35 (1991); R. C. Condit, S. P. Hubbell, R. B. Foster, Ecol. Monogr. 65, 419 (1995); J. Trop. Ecol. 12, 231 (1996).
31. The number of transitions between canopy height classes was counted over the 13-year period from 1982-95 at each 5-m grid point. The mean of the number of transitions was computed for the 16 sample points within each 20 m by 20 m quadrat.
32. Gap sizes were measured by the number of contiguous 5 m by 5 m quadrats (sharing a side) with canopies < 5 m in height.
33. Our hypothesis for higher species richness on the slopes is that slopes are more favorable dry-season habitats for seedlings. Soil moisture levels on the slopes remain high longer into the dry season than on the plateau, due to many seepage areas and springs [P. Becker et al., J. Trop. Ecol. 4, 173 (1988)].
34. K. E. Harms, thesis, Princeton University (1997); _ and S. J. Wright, unpublished data.
35. K. E. Harms, thesis, Princeton University (1997); _ and S. J. Wright, unpublished data.
36. B. Wechsler, senior thesis, Princeton University (1995); W. E. Pugh, senior thesis, Princeton University (1996); E. Carlisle, senior thesis, Princeton University (1997); S. Finklestein, senior thesis, Princeton University (1998).
37. B. Wechsler, senior thesis, Princeton University (1995); W. E. Pugh, senior thesis, Princeton University (1996); E. Carlisle, senior thesis, Princeton University (1997); S. Finklestein, senior thesis, Princeton University (1998).
38. B. Wechsler, senior thesis, Princeton University (1995); W. E. Pugh, senior thesis, Princeton University (1996); E. Carlisle, senior thesis, Princeton University (1997); S. Finklestein, senior thesis, Princeton University (1998).
39. B. Wechsler, senior thesis, Princeton University (1995); W. E. Pugh, senior thesis, Princeton University (1996); E. Carlisle, senior thesis, Princeton University (1997); S. Finklestein, senior thesis, Princeton University (1998).
40. D. Tilman, Ecology 75, 25 (1994);
41. G. C. Hurtt and S. W. Pacala, J. Theor. Biol. 176, 1 (1995).
42. S. P. Hubbell, R. B. Foster, R. Condit, unpublished data.
43. J. Weiner, Ecology 66, 743 (1985); Trends Ecol., Evol. 5, 360 (1990).
44. J. Weiner, Ecology 66, 743 (1985); Trends Ecol., Evol. 5, 360 (1990).
45. 6 seeds in the seed traps over the years. We thank D. Hamill, S. Hewett, B. Minor, and U. Smith for database management during the first three censuses, P. Phillips for Fig. 1, and J. Dalling, A. Herre, and P. Phillips for comments on the manuscript. We thank NSF, the Smithsonian Tropical Research Institute, the Geraldine R. Dodge Foundation, EarthWatch, the Guggenheim Foundation, the Pew Charitable Trusts, the John D. and Katherine T. MacArthur Foundation, and the Andrew W. Mellon Foundation, as well as numerous other private donors, for their generous support of this research.