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ROBERT H. WHITTAKER December27, I920-October20, :1980 BY WALTER E. WESTMAN, ROBERT K. FEET, AND GENE E. LIKENS RosERT HARDING WHITTAKER was one of the preemi- nent community ecologists of the twentieth century. By studying the interactions of plant populations at the biogeo- chemical, species, ant! community levels, he macle contribu- tions to basic knowleclge in several subclisciplines of biology. He clevelopec} new approaches for the analysis of plant communities ant] provided exemplary insight into the pat- terns of composition, procluctivity, anc} diversity of land plants. He brought clarity to such disparate fields as the cias- sification ant! ordination of plant communities, plant succes- sion, allelochemistry, evolution ant! measurement of species diversity, niche theory, and the systematics of kingdoms of organisms. In several influential monographs he detailect the vegetational patterns of various montane regions of the United States, and cluring the last six years of his life- extencled his research to Mediterranean- and arid-climate re- gions of the United States, Israel, Australia, and South Africa. Whittaker's most cited work is his unclergracluate text- book, Communities and Ecosystems ( 1970,3; seconc! edition, 1975,3), which not only introduced thousands of students throughout the world to ecology but also proviclec! a succinct summary of a highly diverse literature and new insights use- ful to professional ecologists. 425

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426 BIOGRAPHICAL MEMOIRS EDUCATION AND EARLY LIFE Robert H. Whittaker, the youngest of three children, was born on December 27, 1920, in Wichita, Kansas, to Clive Charles and Adeline Harding Whittaker. His mother en- couraged Whittaker's abiding interest in languages, while his father stimulated an early interest in natural history. Whittaker entered Washburn Municipal College (now University) in Topeka, Kansas, in 1938. He received a Bach- elor of Arts degree in biology and languages in 1942 but postponed his plans to pursue graduate work in ecology to enlist in the Army. He was stationed in the United States and in England until 1946 as an Army-Air Force weather ob- server and forecaster. Upon his return to civilian life in 1946, he entered graduate school at the University of Illinois, where he completed his Ph.D. two-and-a-half years later. When Whittaker applied for graduate standing in the De- partment of Botany at Illinois, his application was denied because of insufficient background in botany, but he was ad- mitted to the Zoology Department and awarded a fellowship. In February 1946, he began his graduate studies under the direction of Victor Shelford, who retired from active teach- ing that summer. Charles Kendeigh replaced Shelford as Whittaker's adviser in September, and though Whittaker worked with him and acknowledged his debt, Whittaker was also heavily influenced by the University of Illinois botanist Arthur G. Vestal, whom Whittaker called his "second ad- viser.'' SCIENTIFIC WORK The Continuum of Plant Species Distribution Whittaker was particularly taken by classroom lectures in which Vestal questioned rigid Clementsian notions of plant association and discussed Gleason's opposing idea of individ-

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ROBERT H. WHITTAKER 427 ualistic species distributions. From later conversations it was apparent that Whittaker keenly appreciated Vestal's influ- ence in shaping his own theoretical approach and, in his later years at Cornell, was pleaser! to play a similar role for the graduate students of others. Whittaker's doctoral dissertation (194S,1) examined pat- terns of plant species change along an attitudinal gradient in the Great Smoky Mountains of Tennessee. In seeking to understand underlying patterns of species change, he plot- tect plant species' distributions along axes of environmental change. He then was able to show that the ecological impor- tance of plant species (as measurer! by density or cover) rose and fell in a Gaussian fashion along key environmental gra- rlients, with each species showing an in(livi(lualistic distribu- tion. Though Whittaker had hypothesized the occurrence of groups of coaciapted species with parallel distributions, what emerges! from his work was a validation of Gleason's hypoth- esis and rejection of his own: Most species were distributed independently along environmental gradients. The significance of this work was obvious. It supported the "continuum" concept of species distribution and ex- tencled the statistical basis for gradient analysis in general. W. H. Camp wrote Whittaker that his manuscript was "prob- ably the most important ecological paper of the present cen- tury" and that his method would revolutionize the field) Plant and Insect Population Patterns, and Element Cycling In 194S, Whittaker was appointed instructor in the De- partment of Zoology at Washington State College (now Uni- versity) in Pullman, Washington. While at Washington State ~ Despite this assessment, Whittaker's doctoral dissertation was not published until eight years later (1956,1). By that time along with J. T. Curtis and the Wisconsin school he had developed a series of detailed gradient analyses, but it took another ten to fifteen years before his thesis was widely accepted.

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428 BIOGRAPHICAL MEMOIRS he began field work on the vegetation of the Klamath region ant! Siskiyou Mountains of Oregon and California, including a comparative study of vegetation on serpentine and quartz- ctiorite soils. Returning to the original focus of his dissertation work, Whittaker completed a manuscript on foliage insects in the Great Smokies, building on his vegetation analysis there. At the same time he conducted a uniquely thorough study of copepod communities of small ponds in the Columbia basin. Whittaker left Washington State in 195 ~ to become a sen- ior scientist in the Hanford Laboratories Aquatic Biology Unit, Department of Radiological Sciences, in RichIand, Washington. Quick to see the value of radioactive tracers for unraveling complex ecological problems, he studied in petal! the movement of radioactive phosphorus in aquarium micro- cosms. His results were important to understanding the fate of raclionuclides in the environment and for evaluating the movement ant! storage of nutrients in ecosystems. At Han- ford (anc! later at Brookhaven National Laboratory with George Woodwell), he also contributes! to the first large-scale stucly of the effect of chronic gamma radiation on the struc- ture and function of forest ecosystems. While at the Hanford Laboratories, Bob met Clara Caroline BuehI, and the two were married on New Year's Day, 1953. Although Clara had an M.S. in biology, her role in the marriage soon became that of wife ant! mother rather than scientific collaborator. The Whittakers raiser! three sons: John Charles, Paul Louis, ant! Car] Robert. Dimension Analysis and the Classification of the Kingdoms In 1954, Whittaker was hired as an instructor in the De- partment of Biology of Brooklyn College, the City University of New York, where he wouIcl remain for ten years. During the summers he returned to the Great Smoky Mountains,

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ROBERT H. WHITTAKER 429 where he initiates! a major effort to obtain measurements of the biomass ant} productivity of the forest communities along an elevational gradient. Because he was interested in the entire production of plants above ground, he began to clevelop methods for mea- suring productivity of shrubs and herbs and other parts of trees in addition to trunks. He used a volumetric measure- ment based on growth rings and succeedecI, through labori- ous calculations, in obtaining productivity estimates for the major plant communities in the mountain range. His efforts provided a basis for the subsequent development of the di- mension analysis methodology still wiclely in use. Throughout his career in acIdition to conducting mode} studies of a variety of ecological systems he also maintainer! an interest in the problem of classification and speciation. In 1957 he proposed a new classification for the kingdoms of organisms baser! on the evolution of trophic structures and nutritional energy sources (1957,I). Later updated (1969,4), this system of classification eventually was accepted wiclely and used in biology textbooks. Desert and Forest: Structure and Function From 1963 to 1965, Whittaker and W. A. Niering pub- lished a series of stuclies of the Arizona Saguaro cactus de- sert among the first studies of a desert community to em- phasize functional rather than structural attributes. For this work the authors received the Ecological Society of America's 1966 Mercer Awarc! for the best paper published in the pre- cecling two years by a young ecologist. In ~ 964 another colleague and future collaborator, George M. Wooc~well, persuaclec! Whittaker to take a year's leave from Brooklyn College to work with him at Brookhaven National Laboratory in New York State. The two developed a profound! respect ant! fondness for each other, and

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430 BIOGRAPHICAL MEMOIRS throughout the 1960s the team of Whittaker and Woodwell was one of the most productive and influential in plant ecol- ogy. Together they produced eight papers on the surface area, biomass, production and nutrient flow, and effects of gamma radiation on structure and diversity of forested eco- systems in the Brookhaven oak-pine forest and surrounding vegetation. lust before leaving Brookhaven in 1966, Whittaker had initiated studies with Gene E. Likens and F. Herbert Bormann on the biomass, productivity, and nutrient con- tent of the Hubbard Brook Experimental Forest in New Hampshire. These subsequently led to two major mono- graphs about this northern hardwood forest ecosystem (1970,1; 1974,31. With Likens, Whittaker also compiled the widely cited summary tables of plant production, biomass, and associated characteristics for ecosystems of the world. Species Diversity, Ordination Methods In 1966 Whittaker decided to accept the offer of a pro- fessorship at the new Irvine campus of the University of Cali- fornia. He took up this new post with great enthusiasm and anticipation but was dismayed by the rapid pace of urbani- zation around Irvine. In September 1968 he accepted an in- vitation to move to Cornell University as professor of biology in the Section of Ecology and Systematics, where his last years were marked by a significant expansion ant! solidification of . his reputation. Once again pursuing his early interest in species diversity, Whittaker was stimulated in part by the attention G. E. Hutchinson, R. H. MacArthur, and their students had given to the topic. His concise paper in Science (1965,3) remains a classic review of the field. When general theories to explain patterns of plant species diversity did not emerge, Whittaker emphasized factors influencing local patterns, based on pe-

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ROBERT H. WHITTAKER 43 culiarities of site history and environment. In association with Hugh Gauch, Jr., and others he also explored techniques for ordinating species data techniques that helped computer- ize earlier gradient analyses he had developed along with I. T. Curtis and the Wisconsin school. TEACHER, DIPLOMAT, HONORED RESEARCHER At Irvine and Cornell, Whittaker had the opportunity to supervise graduate students for the Ph.D. for the first time. Of the twelve he trained, eight went on to complete their dissertations under his supervision. Through personal diplomacy, furthermore, he built bridges between American and European ecologists, calming the waters he himself hac! troubler! with his challenges to phytosociological theories and methods of classification. His reviews of classification ant! ordination studies and his global studies of diversity and productivity helped inspire North American ecologists to increase contacts and collaboration with ecologists beyond their borders. In his later years Robert Whittaker reaped the rewards of a prolific intellectual career. He enjoyed! a solid reputation among his peers, who elected! him vice president of the Eco- logical Society of America in 1971. He was elected to the National Academy of Sciences in 1974 and named Cornell's Charles A. Alexander Professor of Biological Sciences in 1976. Elected to the American Academy of Arts and Sciences in 1979, he also held honorary memberships in the British Ecological Society ant! the Swedish Phytogeographical Soci- ety. At the time of his death he was president of the American Society of Naturalists. HEALTH PROBLEMS In ~ 974 Whittaker's wife contracted cancer. CIara's struggle with the disease lasted three years, and at Christmas

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432 BIOGRAPHICAL MEMOIRS time in 1977, she finally succumbed. Though her prolonged illness upset Whittaker greatly, he remained stoically silent, and many of his students anc} colleagues were not aware of the events that were troubling him. Turning to his traditional values for support, he increaser} the intensity with which he pursued his work. Following CIara's death, Whittaker developed a close friendship with his doctoral student, Linda Olsvig. In Octo- ber 1979 the two were married, and Linda, taking an active interest in his research, accompanies! Whittaker into the field on visits to Israel and South Africa. There were no children from this marriage. Four months after his second! marriage Whittaker com- plainect of hip pain. X-rays revealecl cancer in hip and lungs, but he set himself to complete as much of his work as pos- sible. His health failer! in September ant! he flier! on October 20, 1980. Shortly before his death, the Ecological Society of America honored him with its highest award, that of Eminent Ecologist. IN CONCLUSION Difficult as it to assess which of Whittaker's many contri- butions to the science of ecology will prove most profound or long lasting, one hallmark stands out. Demonstrating the continuity of species' response to environmental gradients, he challenged the classificatory approach to vegetation struc- ture. Though Whittaker credited Ramensky, Gleason, Curtis, and McIntosh with much, it was his own theory, method, and empirical evidence that solidified gradient analysis into a scientifically accepted approach. IN THE PREPARATION OF THIS MEMOIR, the authors often re- ferred to a short biography by W. E. Westman and R. K. Feet pub- lished shortly after Whittaker's death, "Robert H. Whittaker

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ROBERT H. WHITTAKER 433 (1920-1980)The Man and His Work," Vegetatio 48(1982):97-122. A memorial volume has been published by Whittaker's students and colleagues: R. K. Feet, ea., Plant Community Ecology. Papers in Honor of Robert H. Whittaker (Dordrecht: Junk, 1985~. In 1975 Robert Whittaker supplied the National Academy with an autobio- graphical note, which remains on file with the Archives of the National Academy of Sciences in Washington, D.C.

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434 BIOGRAPHICAL MEMOIRS SELECTED BIBLIOGRAPHY 1944 With D. B. Stallings. Notes on seasonal variation in Lepidoptera. Entomol. News, 53~3~:67-71; (4~:87-92. 1948 A vegetation analysis of the Great Smoky Mountains (doctoral dis- sertation). University of Illinois, Department of Zoology. 1951 A criticism of the plant association and climatic climax concepts. Northwest Sci., 26: 17-31. 1952 A study of summer foliage insect communities in the Great Smoky Mountains. Ecol. Monogr., 22:1-44. 1953 A consideration of climax theory: the climax as a population and pattern. Ecol. Monogr., 23:41-78. 1954 Plant populations and the basis of plant indication. (German sum- mary.) Angew. Pflanzensoziol. (Wien), 1:183-206. The ecology of serpentine soils. I. Introduction. Ecology, 35:258- 59. The ecology of serpentine soils. IV. The vegetational response to serpentine soils. Ecology, 35:275 - 88. 1956 Vegetation of the Great Smoky Mountains. Ecol. Monogr., 26:1- 80. In honor of Edwin Aichinger. Review of Festscrhift fur Edwin Aich- inger zum 60 Geburtstag. 1954. Ecology, 37:296 -97. A new Indian Ecological Journal. Review of Bulletin of the Indian Council of Ecological Research, vol. 1. Ecology, 37:628.

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ROBERT H. WHITTAKER 1957 435 Recent evolution of ecological concepts in relation to the eastern forests of North America. Am. I. Bot., 44: 197-206. Also in: Fifty Years of Botany: Golden Jubilee Volume of the Botanical Society of America, ea., W. C. Steere, New York: McGraw-Hill, pp. 340- 58. The kingdoms of the living world. Ecology, 38:536-38. Review of H. Ellenberg. 1950 - 1954. Gradient analysis in agricul- turalecology. Landwirtsch. Pflansensoziol. Ecology, 38:363-64. Two ecological glossaries and a proposal on nomenclature. Ecology, 38:371. 1958 With C. W. Fairbanks. A study of plankton copepod communities in the Columbia Basin, southeastern Washington. Ecology, 39:46-65. Also in: Readings in Population and Community Ecology, ea., W. E. Hazen, Philadelphia: W. B. Saunders, pp. 369-88. A manual of phytosociology. Review of F. R. Bharucha and W. C. de Leeuw, A Practical Guide to Plant Sociology for Foresters and Agriculturalists ( 1 95 7~ . Ecology, 3 8: 1 8 2 . The Pergamon Institute and Russian journals of ecology. Ecology, 39: 182-83. 1959 On the broad classification of organisms. Q. Rev. Biol.,34:210-26. 1960 Ecosystem. In: McGraw-Hill Encyclopedia of Science and Technology, New York: McGraw-Hill, pp. 404-8. Vegetation of the Siskiyou Mountains, Oregon and California. Ecol. Monogr., 30:279-338. A vegetation bibliography for the northeastern states. Review of F. E. Egler, A Cartographic Guide to Selected Regional Vegetation Literature Where Plant Communities Have Been Described (1959~. Ecology, 41:245-46. 1961 Estimation of net primary production of forest and shrub com- munities. Ecology, 42: 177-80.

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436 BIOGRAPHICAL MEMOIRS Experiments with radio-phosphorus tracer in aquarium micro- cosm. Ecol. Monogr., 31:157-88. Vegetation history of the pacific coast states and the "central" sig- nificance of the Klamath Region. Madrono, 16:5-23. New serials. Ecology, 42:616. The chemostat as a model system for ecological studies. In: Modern Methods in the Study of Microbial Ecology, ed. T. Rosswell, Uppsala: Swedish National Sciences Research Council, pp. 347-56. 1962 Classification of natural communities. Bot. Rev., 28:1-239. Re- printed: New York: Arno Press (1977~. Net production relations of shrubs in the Great Smoky Mountains. Ecology, 43:357-77. With V. Garfine. Leaf characteristics and chlorophyll in relation to exposure and production in Rhododendron maximum. Ecology, 43: 190-25. The pine-oak woodland community. Review of I. T. Marshall, Birds of Pine-Oak Woodland in Southern Arizona and Adjacent Mexico 1957~. Ecology, 43: 180-81. 1963 With W. A. Niering and C. H. Lowe. The saguaro: A population in relation to environment. Science, 142: 15-23. Essays on enchanted islands. Review of G. E. Hutchinson, The En- chanted Voyage and Other Studies (19621. Ecology, 44:425. Net production of heath bards and forest heaths in the Great Smoky Mountains. Ecology, 44:176-82. With N. Cohen and J. S. Olson. Net production relations of three tree species at Oak Ridge, Tennessee. Ecology, 44:806-10. With W. A. Niering. Vegetation of the Santa Catalina Mountains. Prog. Agric. Ariz., 15 :4-6. 1964 With W. A. Niering. Vegetation of the Santa Catalina Mountains, Arizona. I. Ecological classification and distribution of species. I. Ariz. Acad. Sci., 3:9-34.

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ROBERT H. WHITTAKER 1965 437 With W. A. Niering. The saguaro problem and grazing in south- western national monuments. Natl. Parks Mag., 39:4-9. Branch dimensions and estimation of branch production. Ecology, 46:365-70. Dominance and diversity in land plant communities. Science, 147:250-60. With W. A. Niering. Vegetation of the Santa Catalina Mountains, Arizona. II. A gradient analysis of the south slope. Ecology, 46:429-52. With W. A. Niering. The saguaro problem and grazing in south- western national monuments. Nat. Parks Mag., 39:4-9. 1966 Forest dimensions and production in the Great Smoky Mountains. Ecology, 47:103-21. With G. M. Woodwell and W. M. Malcolm. A-bombs, bugbombs, and us. NAS-NRC Symposium on "The Scientific Aspects of Pest Control," the Brookhaven National Laboratory, U.S. Atomic Energy Commission. Washington, D.C.: Atomic Energy Commission. 1967 Ecological implications of weather modification. In: Ground Level Climatology, ed. R. H. Shaw, Washington, D.C.: AAAS, pp. 367- 84. Gradient analysis of vegetation. Biol. Rev., 42: 207- 64. With G. M. Wooclwell. Surface area relations of woody plants and forest communities. Am. }. Bot., 54:931-39. With G. M. Woodwell. Primary production and the cation budget of the Brookhaven forest. In: Symposium on Primary Productivity and Mineral Cycling in Natural Ecosystems, ed. H. E. Young, Orono: University of Maine Press, pp. 151-66. 1968 With I. Frydman. Forest associations of southeast Lublin Province, Poland. (German summary.) Ecology, 49:896-908. With S. W. Buol, W. A. Niering, and Y. H. Havens. A soil and

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438 BIOGRAPHICAL MEMOIRS vegetation pattern in the Santa Catalina Mountains, Arizona. Soil Sci., 105:440-50. With W. A. Niering. Vegetation of the Santa Catalina Mountains, Arizona. III. Species distribution and floristic relations on the north slope. }. Ariz. Acad. Sci., 5:3-21. With W. A. Niering. Vegetation of the Santa Catalina Mountains, Arizona. IV. Limestone and acid soils. I. Ecol., 56:523-44. With G. M. Woodwell. Effects of chronic gamma irradiation on plant communities. Q. Rev. Biol., 43:42-55. With G. M. Woodwell. Primary production in terrestrial ecosys- tems. Am. Zool., 8:19-30. 1969 A view toward a National Institute of Ecology. Ecology, 50: 169-70. Een nieuwe indeling van de organismen. Nat. Tech., 37:124-32. Evolution of diversity in plant communities. In: Diversity and Sta- bility in Ecological Systems, Brookhaven Symposia in Biology, Upton, Near York: Brookhaven Natl. Lab. Publ. 50175 (C-56), No. 22, pp. 178-95. New concepts of kingdoms of organisms. Science, 163: 150-60. With G. M. Woodwell. Structure, production, and diversity of the oak-pine forest at Brookhaven, New York. I Ecol., 57: 155-74. 1970 With F. H. Bormann, T. G. Siccama, and G. E. Likens. The Hub- bard Brook ecosystem study: Composition and dynamics of the tree stratum. Ecol. Monogr., 40:373-88. With W. L. Brown and T. Eisner. Allomones and kairomones: Transspecific chemical messengers. BioScience, 20:21-22. Communities and Ecosystems. New York: Macmillan. ~ Japanese edi- tion, Tokyo, 1974.) Neue Einteilung der Organismenreiche. Umschau, 16:514-15. Taxonomy. In: McGraw-Hill Yearbook of Science and Technology 1970, New York: McGraw-Hill, pp. 365-69. The biochemical ecology of higher plants. In: Chemical Ecology, eds. E. Sondheimer and J. B. Simeone, New York: Academic Press, pp. 43-70. The population structure of vegetation. In: Gesellschaftsmorpholog~e (Strakturforschung) (German summary), ed. R. Tuxen, Ber.

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ROBERT H. WHITTAKER 439 Symp. Int. Ver. Vegetationskunde, Rinteln, 1966. The Hague: Junk., pp. 39-62. With G. M. Woodwell. Ionizing radiation and the structure and functions of forests. In: Gesellschaftsmorphologie (Straklurfor- schungJ (German summary), ed. R. Tuxen, Ber. Symp. Int. Ver. Vegetationskunde, Rinteln, 1966. The Hague: Junk, pp. 334- 39. 1971 With P. P. Feeny. Allelochemics: Chemical interactions between spe- cies. Science, 171 :757-70. With G. M. Woodwell. Evolution of natural communities. In: Ecosystem Structure and Function, Proc. 31st Ann. Biol. Colloq., ed. I. A. Wiens, Corvallis: Oregon State University Press. pp. 137-56. With G. M. Woodwell. Measurement of net primary production of forests. In: Productivity of Forest Ecosystems (French summary), ed. P. Duvigneaud, Proc. Brussels Symp., 1969 Paris: UNESCO, pp. 159-75. With P. F. Brussard, A. Levin, and L. N. Miller. Redwoods: A pop- ulation model debunked. Science, 175 :435-36. Dry weight, surface area, and other data for individuals of three tree species at Oak Ridge, Tennessee. In: Dryweight and Other Data for Trees and Woody Shrubs of the Southeastern United States, (Publ. ORNL-IBP-71-6), eds. P. Sollins and R. M. Anderson, Oak Ridge, Tennessee: Oak Ridge National Laboratory, pp.37- 38. The chemistry of communities. In: Biochemical Interactions Among Plants. Washington, D.C.: National Academy of Sciences, pp. 10-18. 1972 With H. G. Gauch,Jr. Coenocline simulation. Ecology, 53:446-51. With H. G. Gauch, {r. Comparison of ordination techniques. Ecol- ogy, 53:868-75. Convergences of ordination and classification. In: Basic Problems and Methods in Phytosociology (German summary), ed. R. Tuxen, Ber. Symp. Int. Ver. Vegetationskunde, Rinteln, 1970. The Hague: Junk, pp. 39-55.

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440 BIOGRAPHICAL MEMOIRS Evolution and measurement of species diversity. Taxon, 21:213- 51. A hypothesis rejected: The natural distribution of vegetation. In: Botany: An Ecological Approach, eds. W. A. Jensen and F. B. Salis- bury, Belmont, Calif.: Wadsworth, Inc., pp. 689-91. Also in: Biology, ed. W. A. Jensen et al., Belmont, Calif.: Wadsworth, Inc., 1979, pp. 474-76. 1973 With G. Cottam and F. G. Golf. Wisconsin comparative ordination. In: Ordination and Classification of Communities, ed. R. H. Whit- taker, The Hague: Junk, pp. 193-221. Approaches to classifying vegetation. In: Handbook of Vegetation Sci- ence, V. Ordination and Classification of Vegetation, ed. R. H. Whit- taker, The Hague: Junk, pp. 325-54. Community, biological. In: Encyclopaedia Britannica, 15th ea., pp. 1027-35 Direct gradient analysis: Results. In: Handbook of Vegetation Science, V. Ordination and Classification of Vegetation, ed. R. H. Whittaker, The Hague: Junk, pp. 9-31. Dominance-types. In: Handbook of Vegetation Science, V. Ordination and Classification of Vegetation, ed. R. H. Whittaker, The Hague: Junk, pp. 389-92. (Editor). Handbook of Vegetation Science, V. Ordination and Classifica- tion of Vegetation. The Hague: Junk. Introduction. In: Handbook of Vegetation Science, V. Ordination and Classification of Vegetation, ed. R. H. Whittaker, The Hague: Junk, pp. 1-6. With H. G. Gauch, fir. Evaluation of ordination techniques. In: Handbook of Vegetation Science, V. Ordination and Classification of Vegetation, ed. R. H. Whittaker, The Hague: Junk, pp.289-321. With S. A. Levin and R. B. Root. Niche, habitat, and ecotope. Am. Nat., 107:321-38. With G. E. Likens. Carbon in the biota. In: Carbon and the Biosphere, Symp. 2d. Cong. Am. Inst. of Bio. Sci., Miami, Florida, 1971. Human Ecol., 1 :301-2. With G. E. Likens. Primary production: The biosphere and man. In: Handbook of Vegetation Science, ed. R. H. Whittaker, The Hague: Junk, pp. 55-73.

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ROBERT H. WHITTAKER 1974 441 With H. G. Gauch, fir., and G. B. Chase. Ordination of vegetation samples by Gaussian species distributions. Ecology, 55: 1382-90. Climax concepts and recognition. In: Handbook of Vegetation Science, VIII. Vegetation Dynamics, ed. R. Knapp, pp. 139-54. With F. H. Bormann, G. E. Likens, and T. G. Siccama. The Hub- bard Brook ecosystem study: Forest biomass and production. Ecol. Monogr., 44:233-54. 1975 With H. Leith (eds.~. The Primary Productivity of the Biosphere. New York: Springer-Verlag. With W. E. Westman. The pygmy forest region of northern Cali- fornia: Studies on biomass and primary productivity. }. Ecol., 62:493-520. Communities and Ecosystems, 2d ed. New York: Macmillan. ~ Japanese ea., Tokyo, 1978.) Functional aspects of succession in deciduous forests. In: SuLzes- sionsforschung (German summary), ed. W. Schmidt., Ber. Symp. Int. Ver. Vegetationskunde, Rinteln, 1973, pp. 377-405. The design and stability of plant communities. In: Unifying Concepts in Ecology (Rep. Plenary Sessions, 1st Int. Cong. Ecology, The Hague, 1974), eds. W. H. van Dobben and R. H. Lowe- McConnell, The Hague: Junk, and Wageningen: Pudoc, pp. 169-81. vegetation and parent material in the western United States. In: Vegetation and Substrate (German summary), ed. H. Dierschke, Ber. Symp. Int. Ver. Vegetationskunde, Rinteln, 1969, pp. 443- 65. With S. A. Levin (eds.~. Niche: Theory and Application, Benchmark Papers in Ecology. Stroudsburg: Dowden, Hutchinson, and Ross. With S. A. Levin and R. B. Root. On the reasons for distinguishing "niche, habitat, and ecotope." Am. Nat., 109:479-82. With G. E. Likens. The biosphere and man. In: Primary Productivity of the Biosphere, eds. H. Leith and R. H. Whittaker, New York: Springer-Verlag, pp. 305-23. With P. L. Marks. Methods of assessing terrestrial productivity. In:

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442 BIOGRAPHICAL MEMOIRS Primary Productivity of the Biosphere, eds. H. Leith and R. H. Whittaker, New York: Springer-Verlag, pp. 55-118. With W. A. Niering. Vegetation of the Santa Catalina Mountains, Arizona. V. Biomass, production, and diversity along the ele- vation gradient. Ecology, 56:771-90. With G. M. Woodwell and R. A. Houghton. Nutrient concentra- tions in plants in the Brookhaven oak-pine forest. Ecology, 56:318-32. 1976 With H. G. Gauch, Jr. Simulation of community patterns. Vegeta- tio, 33:13-16. With R. B. Hanawalt. Altitudinally coordinated patterns of soils and vegetation in the San tacinto Mountains, California. Soil Sci.,121:114-24. With S. R. Kessell. Comparisons of three ordination techniques. Vegetatio, 32:21-29. 1977 With H. G. Gauch, fir., and T. R. Wentworth. A comparative study of reciprocal averaging and other ordination techniques. i. Ecol., 65:157-74. With R. B. Hanawalt. Attitudinal patterns of Na, K, Ca and Mg in soils and plants in the San iacinto Mountains, California. Soil Sci., 123:25-36. With R. B. Hanawalt. Attitudinal gradients of nutrient supply to plant roots in mountain soils. Soil Sci., 123:85-96. With I. Noy-Meir. Continuous multivariate methods in community analysis: Some problems and developments. Vegetatio, 33:79- 98. Animal effects on plant species diversity. In: Vegetation and Fauna, ed. R. Tuxen, Ber. Symp. Int. Ver. Vegetationskunde, Rinteln, 1976. Vaduz, The Netherlands: Cramer, pp. 409-25. Broad classification: The kingdoms and the protozoans. In: Para- sitic Protozoa, ed. I. Krier, New York: Academic Press, vol. 1, pp. 1-34. Evolution of species diversity in land communities. In: Evolutionary Biology, eds. M. K. Hecht, W. C. Steere, and B. Wallace, New York: Plenum Publishing Corp, vol. 10, pp. 1-67.

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ROBERT H. WHITTAKER 443 With S. A. Levin. The role of mosaic phenomena in natural com- munities. Theor. Pop. Biol., 12: 117-39. 1978 With I. Noy-Meir. Recent developments in continuous multivariate techniques. In: Ordination of Plant Communities, ed. R. H. Whit- taker, The Hague: Junk, pp. 337-78. (Editor). Classification of Plant Communities. The Hague: Junk. With H. G. Gauch, fir. Evaluation of ordination techniques. In: Ordination of Plant Communities, ed. R. H. Whittaker, The Hague: Junk, pp. 277-336. With L. Margulis. Protist classification and the kingdoms of organ- isms. BioSystems, 10:3-18. With G. M. Woodwell, W. A. Reiners, G. E. Likens, C. C. Delwiche, and D. B. Botkin. The biota and the world carbon budget. Science, 199: 141-46. Review of Terrestrial Vegetation of California, eds. M. G. Barbour and J. Major, Vegetatio, 38:124-25. 1979 With Z. Naveh. Measurements and relationships of plant species diversity in Mediterranean shrublands and woodlands. In: Eco- logical Diversity in Theory and Practice, eds. F. Grassle, G. P. Patil, W. Smith, and C. Taillie, Fairland, Md.: Int. Co-op. Pub. House, pp.219-39. With L. S. Olsvig and J. F. Cryan. Vegetational gradients of the pine plains and barrens of Long Island. In: Pine Barrens: Eco- system and Landscape, ed. R. T. T. Forman, New York: Academic Press, pp. 265-82. With S. R. Sabo. Bird niches in a subalpine forest: An indirect ordination. PNAS, 76:1338-42. With A. Shmida. Convergent evolution of deserts in the old and new world. In: Werden and Vergehen non Ppanzengesellschaften, eds., O. Wilmanns and R. Tuxen, Ber. Symp. Int. Ver. Vegeta- tionskunde, Rinteln, 1978, Vaduz, The Netherlands: Cramer, pp. 437-50. Vegetational relationships of the pine barrens. In: Pine Barrens: Ecosystem and Landscape, ea., R. T. T. Forman, New York: Academic Press, pp. 315-31.

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444 BIOGRAPHICAL MEMOIRS With L. E. Gilbert and I. H. Connell. Analysis of two-phase pattern in a mesquite grassland, Texas. I. Ecol., 67:935-52. With D. Goodman. Classifying species according to their demo- graphic strategy. I. Population fluctuations and environmental heterogeneity. Am. Nat., 113: 185-200. With G. E. Likens, F. H. Bormann, l. S. Eaton, and T. G. Siccama. The Hubbard Brook ecosystem study: Forest nutrient cycling and element behavior. Ecology, 60:203-20. With Z. Naveh. Analysis of two-phase patterns. In: Contemporary Quantitative Ecology and Related Econometrics, eds. G. P. Patil and M. Rosenzweig, Fairland, Md.: Int. Co-op. Pub. House, pp. 157-65. With W. A. Niering and M. D. Crisp. Structure, pattern, and di- versity of a mallee community in New South Wales. Vegetatio, 39:65-76. 1980 With Z. Naveh. Structural and floristic diversity of shrublands and woodlands in northern Israel and other Mediterranean areas. Vegetatio, 41: 171-90. 1981 With H. G. Gauch, fir., and S. B. Singer. A comparative study of nonmetric ordinations. J. Ecol., 69: 135 -52. With A. Shmida. Pattern and biological microsite effects in two shrub communities, southern California. Ecology, 62:234-51. With K. D. Woods. Canopy-understory interaction and the internal dynamics of mature hardwood and hemlock-hardwood forests. In: Forest Success~on: Concepts and Application, eds. D. West, H. H. Shugart and D. B. Botkin, New York: Springer-Verlag, pp. 305-23. With H. G. Gauch, fir. Hierarchical classification of community data. I. Ecol., 69:537-57. 1984 With }. Morris and D. Goodman. Pattern analysis in savanna wood- lands at Nylsvley, South Africa. Mem. Bot. Surv. S. Africa, 49.

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