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Transcript of Vegetation of the Great Smoky Mountains R. H. Whittaker ... Mixed Mesophytic in the Smokies and...

  • Vegetation of the Great Smoky Mountains

    R. H. Whittaker

    Ecological Monographs, Vol. 26, No. 1. (Jan., 1956), pp. 1-80.

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  • VEGETATION OF THE GREAT SJlOKY lllOUSTAINS1

    R. H. WHITTAKER Biology Department. Brooklyn College. Brooklyn 10. X . Y

    T A B L E OF C O N T E S T S

    PAGE

    Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1

    Nature o f the Study . . . . . . . . . . . . . . . . . . . . . . . . . . 1

    Literature on Area . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

    Acknowledgments . . . . . . . . . . . . . . . . . . . . . . 2

    Geology and Climate . . . . . . . . . . . . . . . . . . . . . . . . . . 2

    Methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

    Field Transects and Tree Classes . . . . . . . . . . . . . . 4

    Site-Samples and Composite Transects . . . . . . . . . . 6

    Distributions o f Species along the Moisture Gradient . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8

    Trends i n Relation t o the Moisture Gradient . . . . . . 10

    Growth-Forms . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10

    Coverages . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10

    Diversities . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10

    Sizes and Numbers of Stems of Trees . . . . . . . . . . 11

    Self-Maintenance of Stands . . . . . . . . . . . . . . . . . . . . 11

    High-Elevation Deciduous Forests . . . . . . . . . . . . . . . . 13

    Distributions o f Species i n Relation t o Elevation . . .

    Xesic Sites . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Submesic Sites . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

    Subxeric Sites . . . . . . . . . . . . . . . . . . . . . . . . .

    Xeric Sites . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

    Trends i n Relation to Elevation . . . . . . . . . . . . . . . . Growth-Forms . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tree Statures and Stratal Coverages . . . . . . . . . . . . . Diversity and Environmental Favorableness . . .

    Spruce-Fir Forests . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

    Stratal Distributions . . . . . . . . . . . . . . . . . . . . . . . . . .

    Trends . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

    Relations o f Species to Unions and Associations . . . . Continuity of Vegetation Types . . . . . . . . . . . . . . . . Nature of Species Groupings . . . . . . . . . . . . . . . Dominance i n Relation t o Community Composition

    Summary o f Distributional Groupings . . . . . . . . . . . .

    I1. D I S C U S S I O N : OFA N INTERPREPATION

    \ T F X 3 ~ ~ ~ ~ ~ . . . . . . . . . . . . . . . . . . . . .~ PATTERNING~ Distributions o f Species and the Study o f Genecology

    T h e Association-Unit Theory and Individualistic Hypothesis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

    T h e Distributional Basis o f Community.Types . . . . . . Gradation and the Grouping o f Species . . . . . . . . .

    I . G R A D I E N T A N A L Y S I S

    INTRODUCTION

    NATURE O F T H E STUDY

    The Great Snloky Mountains o f Tennessee and Nor th Carolina s u p p o r t vegetat ion w h i c h i s particu- ' " 1 ~ '''' i n s ~ e c i e s and varied in c o m m u n i t y t y p e s . I n t h e summer o f 1947 field w o r k \&*as carried o u t

    l Based on a thesis (Whittaker 1 9 4 8 ) ; a contr~bution from the Department of Zoology. University of Illinois. LTrbana. and the Biology Department. Brooklyn College .

    Zonation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34

    Ecotones . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36

    Climax Patterns and Their Comparison . . . . . . . . . . . . 37

    Considerations o f Logic and Zlethod . . . . . . . . . . . . . . 40

    Conclusion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43

    111. \ T E G ~ ~ . 4 T 1 0 ~ T H E I R DISTRIBUTIONAL T Y P E SA X D

    RELATIONS. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43 Bases o f Recognizing and Describing Types . . . . . . . . 43 Vegetat ion Types o f the Great Smoky Mountains . . . 45

    1. Cove Hardwoods Forest . . . . . . . . . . . . . . . . . . . . . 45 Mixed Mesophytic i n the Smokies and

    Cumberlands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46 2. Eastern Hemlock Forest . . . . . . . . . . . . . . . . . . . . . 48

    3. Gray Beech Forest . . . . . . . . . . . . . . . . . . . . . . . . . 48

    4. Red Oak-Pignut Hickory Forest . . . . . . . . . . . . . . 49

    5. Chestnut Oak-Chestnut Forest . . . . . . . . . . . . . . . . 49

    6. Chestnut Oak-Chestnut Heath . . . . . . . . . . . . . . . 50

    7. Red Oak-Chestnut Forest . . . . . . . . . . . . . . . . . . . . .51

    8. Whi te Oak-Chestnut Forest . . . . . . . . . . . . . . . . . .

    Pine Stands and Their Naintenance . . . . . . . . . . . . .

    9. Virginia Pine Forest . . . . . . . . . . . . . . . . . . . . . . . .

    10. Pitch Pine Heath . . . . . . . . . . . . . . . . . . . . .

    11. Table Vounta in Pine Heath . . . . . . . . . . . . . . . .

    12. Grassy Bald . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . The Southern Appalachian Subalpine

    Forest Center . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13. Red Spruce Forest . . . . . . . . . . . . . . . . . . . . . . . . .

    14. Fraser Fir Forest . . . . . . . . . . . . . . . . . . . . . . . . . .

    15. Heath Bald . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

    The Balds as Topographic Climaxes? . . . . . . . . . . . .

    Distributional Relations . . . . . . . . . . . . . . . . . . . . . . .

    The Xosaic Chart . . . . . . . . . . . . . . . . . . . . . . . . . . . .

    Distributions of Types . . . . . . . . . . . . . . . . . . . . .

    Distribution of flubalpine Forests . . . . . . . . . . . . . .

    Relation of the Vegetation Pattern to Those

    of Other V o u n t a i n Ranges . . . . . . . . . . . . . . . . .

    Suncnca~. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Y LITERATURECITED. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . APPENDIXES. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. Population Charts for Major Tree Species . . . . . . Kote on Supplementary Publication o f Appendixes

    B and C . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

    f o r a s t u d y o f th i s vegetat ion . T h e w o r k w a s origi- na l ly intended t o provide i n f o r m a t i o n o n t h e vege- ta t ion f o r t h e sake o f i t s o w n interest and as a basis f o r s tudies i n animal ecology ( W h i t t a k e r 1952). A m a j o r purpose o f bo th th i s and t h e preceding s tudy . hou-ever. u-as use o f t h e complex pa t tern o f natural communities in the Great Smoky Mountains for re- search into t h e theory comlnunity units o r asso-

    . F~~ th i s purpose. the approachto vegeta- tat ion w a s based o n sampling wi thout regard t o a p - parent associations and analysis o f t h e samples i n

  • relation to environmental gradients. I t was felt that relative validity of vegetation types should emerge from data impartially obtained, and that the rela- tions of types to one another should be revealed in the study of their populations in relation to environ- mental gradients. The work thus departs from the traditional approach of studying intuitively recog- nized types or associations; i t