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Vegetation of the Great Smoky Mountains Author(S): RH Whittaker Source Vegetation of the Great Smoky Mountains Author(s): R. H. Whittaker Source: Ecological Monographs, Vol. 26, No. 1 (Jan., 1956), pp. 1-80 Published by: Ecological Society of America Stable URL: http://www.jstor.org/stable/1943577 Accessed: 03-09-2015 19:05 UTC Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at http://www.jstor.org/page/ info/about/policies/terms.jsp JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact [email protected]. Ecological Society of America is collaborating with JSTOR to digitize, preserve and extend access to Ecological Monographs. http://www.jstor.org This content downloaded from 198.21.188.189 on Thu, 03 Sep 2015 19:05:27 UTC All use subject to JSTOR Terms and Conditions VEGETATION OF THE GREAT SMOKY MOUNTAINS1 R. H. WHITTAKER Biology Department,Brooklyn College, Brooklyn10, N. Y. TABLE OF CONTENTS PAGE PAGE I. GRADIENT ANALYSIS .......................... 1 Zonation ................................. 34 36 Introduction ..... ................ 1 Ecotones ...................................... Nature of the Study...................... 1 Climax Patterns and Their Comparison............ 37 Literature on Area..................... 2 Considerationsof Logic and Method.............. 40 Acknowledgments ............................ 2 Geology and Climate................. 2 Conclusion .................................. 43 Methods .................4.............. ....... 4 III. VEGETATION TYPES AND THEIR DISTRIBUTIONAL Field Transects and Tree Classes.............. 4 RELATIONS ........... ...................... 43 Site-Samples and Composite Transects.......... 6 Bases of Recognizing and Describing Types ........ 43 Distributions of Species along the Moisture Vegetation Types of the Great Smoky Mountains ... 45 Gradient ................................... 8 1. Cove Hardwoods Forest..................... 45 in the and Trends in Relation to the Moisture Gradient...... 10 Mixed Mesophytic Smokies Growth-Forms .............. ................. 10 Cumberlands .............. ............... 46 2. Eastern Hemlock Forest..................... 48 Coverages ................................... 10 48 Diversities .......... 10 3. Gray Beech Forest......................... Forest.............. 49 Sizes and Numbers of Stems of Trees.......... 11 4. Red Oak-Pignut Hickory 5. Chestnut Forest 49 Self-Maintenanceof Stands.................... 11 Oak-Chestnut ................ 6. Chestnut Oak-Chestnut Heath ............... 50 High-Elevation Deciduous Forests................ 13 7. Red Oak-ChestnutForest .................... 51 Distributionsof Species in Relation to Elevation... 14 8. White Oak-ChestnutForest .................. 51 Mesic Sites ................................. 14 Pine Stands and Their Maintenance . 51 Submesic Sites ............................... 15 9. Virginia Pine Forest........................ 52 Subxeric Sites ........ ..................... 15 10. Pitch Pine Heath......................... 52 Xeric Sites .15 11. Table Mountain Pine Heath . 53 Trends in Relation to Elevation.16 12. Grassy Bald .............................. 53 Growth-Forms.16 The Southern Appalachian Subalpine Tree Statures and Stratal Coverages............. 17 Forest Center ............................ 53 Diversity and Environmental Favorableness... 18 13. Red Spruce Forest . 54 14. Fraser Fir Forest . 54 Forests .18 Spruce-Fir 15. Heath Bald . 55 Stratal Distributions.19 ................. Trends .20 The Balds as Topographic Climaxes? ... 56 Relations of Species to Unions and Associations 21 Distributional Relations ........................ 57 Continuityof Vegetation Types................ 21 The Mosaic Chart ......... ................... 57 Nature of Species Groupings................... 21 Distributions of Types....................... 59 Dominance in Relation to CommunityComposition 23 Distribution of Subalpine Forests.............. 60 Relation of the Vegetation Pattern to Those of Distributional ............ 24 Summary Groupings of Other Mountain Ranges................. 61 II. DISCUSSION: AN INTERPRETATION OF SUMMARY ..................................... 62 VEGETATION PATTERNING ..................... 25 LITERATURE CITED .......... 63 Distributionsof Species and the Study of Genecology 25 APPENDIXES .......... 69 The Association-UnitTheory and Individualistic Hypothesis .......... ...................... 30 A. Population Charts for Major Tree Species 69 The DistributionalBasis of Community-Types...... 32 Note on Supplementary Publication of Appendixes Gradation and the Grouping of Species......... 32 B and C . .... ... 69 I. GRADIENT ANALYSIS for a study of this vegetation. The work was origi- nally intendedto provide informationon the vege- INTRODUCTION tation for the sake of its own interestand as a basis NATUREOF THE STUDY for studies in animal ecology (Whittaker1952). A major purpose of both this and the precedingstudy, The Great Smoky Mountains of Tennessee and however,was use of the complex pattern of natural North Carolina support vegetation which is particu- communitiesin the Great Smoky Mountainsfor re- larly rich in species and varied in community types. search into the theoryof communityunits or asso- In the summer of 1947 field work was carried out ciations. For this purpose, the approach to vegeta- IBased on a thesis (Whittaker 1948); a contributionfrom tation was based on samplingwithout regard to ap- the Department of Zoology, University of Illinois, Urbana, and the Biology Department, Brooklyn College. parent associations and analysis of the samples in This content downloaded from 198.21.188.189 on Thu, 03 Sep 2015 19:05:27 UTC All use subject to JSTOR Terms and Conditions 2 R. H. WHITTAKER Ecological R. II. WHITTAKER ~~~~~~~~~~Vol.Monographs26, No. 1 relationto environmentalgradients. It was felt that ACKNOWLEDGMENTS relative validity of vegetationtypes should emerge While at the Universityof Illinois, the authorwas from data impartiallyobtained, and that the rela- aided by the suggestionsand criticismsof S. C. Ken- tions of types to one anothershould be revealed in deigh and A. G. Vestal. The park naturalistof the the studyof theirpopulations in relationto environ- Great Smokies,Arthur Stupka, gave the author the mental gradients. The work thus departs from the cooperationand benefitof broad knowledgeof the traditional approach of studyingintuitively recog- mountainswhich he extendsto studentsin the area. nized types or associations; it is an experimentin Help with the identificationof plant specimenswas population analysis of a whole vegetationpattern. given by Stupka and by A. J. Sharp and R. E. The first part of the monographdescribes results Shanks, who checked all determinations.Responsi- of the analysis in termsof relationsof species popu- bility for statementsof distribution,based on field lations to one anotherand environmentalgradients, determinations,remains with the author. A number and trendsin communitycomposition and structure of people have read part or all of the manuscript along environmentalgradients. The second part in- and offeredcomments on it: E. L. Braun, H. E. terpretsthe vegetationas a complexpattern, within Brewer,H. K. Buechner,W. H. Camp, A. Cronquist, which vegetationtypes may be understoodthrough R. Daubenmire,F. E. Egler, H. A. Gleason, A. R. the distributionalrelations of species populations. Kruckeberg,H. L. Mason, R. E. Shanks,A. F. Sharp, The third part presents a more conventionalde- and A. Stupka. The authoris especiallyindebted to scriptionof vegetationtypes and considersthe re- W. H. Camp for his suggestionsand for information lationsof theseto topography. The studyas a whole which permittedinterpretation of genetic and dis- thus seeks to analyze, interpret,and describe the tributionalphenomena. Cost of publication of the complex vegetational mantle of the Great Smoky tables and chartshas been met in part by a grant-in- Mountains. aid fromthe Society of the Sigma Xi. LITERATURE ON AREA GEOLOGY AND CLIMATE A series of studiesby Cain deal withvegetation of The Great SmokyMountains are part of the Blue the Smokies-the heath balds (1930b), subalpine Ridge Province,a systemof mountainsof great an- forests (1935), and cove hardwoods (1943), floristic tiquity. This part of the Southern Appalachians affinities(1930a), soil reaction (1931), Raunkiaer comprisestwo major ranges,the Blue Ridge proper life-forms(1945), and bryophyteunions (Cain & and the Unaka Mountains, along with their con- Sharp 1938). A number of vegetationtypes were necting cross-ranges (Fenneman 1938). The two describedby Cain et al. (1937). The grassy balds main ranges lie parallel,from northeast to southwest, were reportedon by Camp (1931, 1936) and Wells with the IJnakas,of which the Smokies are part, to (1936a, 1936b, 1937); the subalpine forests were the north. The drainage, north from the divide of recentlydescribed by Oosting& Billings (1951) and the Blue Ridge, is northwestinto the Great Valley of the beech gaps by Russell (1953). These papers and the Tennessee River; and rivers flowingfrom the the descriptionin Braun (1950) are the extentof the Blue Ridge to the Great Valley throughthe Unakas literaturedealing specificallywith the vegetationof cut the latter into a series of segmentsdivided by the Smokies. Other
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