STRUCTURAL PATTERNS OF TROPICAL BARKS

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1 STRUCTURAL PATTERNS OF TROPICAL BARKS by Professor Dr. INGRID ROTH Universidad Central de Venezuela, Caracas With 282 figures UNIVERSITATS- BIBLIOTHCK 1981 GEBRUDER BORNTRAEGER BERLIN STUTTGART

2 Introduction 1 Material and methods 2 Sample collection 2 Preparation of permanent slides 4 A word on plant identification in the Tropics 6 On the origin of vulgar and some,,latin" plant names 7 A. General observations The concept of "bark" Definition of cortex and bark Definition of soft bast and hard bast Definition of inner, middle and outer bark Definition of the rhytidome Criteria of bark characterization 16 B. Tissues composing the bark Sieve elements General considerations Patterns produced by sieve tube arrangement Especially large sieve elements Phloem parenchyma The common phloem parenchyma Specialized parenchyma types Parenchyma distribution Crystalliferous parenchyma Storage parenchyma Mechanical system Fibers Sclereids Hard bast sclereids Stone cells of the dilatation tissue Sclerenchyma and crystal formation General hard bast arrangement 50

3 XII 4. Secretory system General aspects Types of secretory structures Secretory cells and cavities Secretory canals Size of secretory canals Peculiarities of secretory canals Frequency of secretory canals Spatial expansion of secretory canals Intercommunicating systems of secretory cells Content of secretory cells Site of exudation Periodicity of exudation Phloem rays Ray width Homo-and heterogeneity of rays Frequency of rays Some peculiarities of rays Aggregate and fusion rays Physiology of rays Patterns produced by rays Secondary formation of stone cells in the rays Storied ray structure Special examples Special structures and inclusions in the rays Ray dilatation and dilatation meristems Storied structure Periderm Cork or phellem Cork width Composing cell types Arrangement of cells Cork prickles Phelloderm Phelloderm width Cell components Arrangement of cells Periodic increments Relations between cork and phelloderm Phylogenetic view points Function of the phelloderm Ill 7.3 Lenticels Outer appearance, frequency, distribution Inner structure 113

4 7.3-3 Stratification of lenticels Annual periodicity of lenticel layering Rhytidome Rhytidome formation Course of the periderms Number of developing periderms Rhytidome width Composing elements Periodic increments Permanence of cork and phelloderm on the bark Periodicity of periderm formation Relations between outer aspect and inner structure of the rhytidome. 126 C. Relations between outer bark aspect and inner structure Color Lenticels Outer bark pattern Cork prickles and other marks Consistence of the bark General considerations 133 D. Function of the bark in relation to structure 140 E. Changes in the non-conducting phloem Sieve tube collapse and parenchyma "inflation" Dilatation growth Secondary formation of stone cells Other changes 158 F. Special part Description of the studied families 159 Annonaceae 160 Capparidaceae 163 Flacourtiaceae 165 Vochysiaceae 171 Guttiferae 175 Quiinaceae 183 Caryocaraceae 183 Bombacaceae 185 Sterculiaceae!90 Tiliaceae *92 Elaeocarpaceae J 97 Humiriaceae 201 Rutaceae 203 Simaroubaceae 2 6 XIII

5 XIV Burseraceae 208 Meliaceae 213 Dichapetalaceae 218 Olacaceae 218 Opiliaceae 219 Celastraceae 221 Rhamnaceae 226 Sapindaceae 230 Sabiaceae 235 Anacardiaceae 235 Mimosaceae 241 Genus Inga Genus Pithecellobium Genus Strypknodendron Genus Parkia Genus Enterolobium 249 Caesalpiniaceae 251 Papilionaceae 261 Pterocarpus Lonchocarpus Clathrotropis Andira Ormosia Centrolobiumparaense Alexa imperatricis Dipteryx odorata 278 Swartzia schomburgkii Platymisciumpinnatum Diplotropispurpurea Hymenolobium Machaerium "Roble bianco" General conclusions concerning bark structure of Papilionaceae Comparison between the three families of Leguminosae 285 Rosaceae 286 Rhizophoraceae 295 Combretaceae 297 Myrtaceae 300 Lecythidaceae 308 Melastomaceae 312 Araliaceae 315 Rubiaceae 317 Sapotaceae 326 Genus Pouteria Other genera 329 Apocynaceae 333 Boraginaceae 338 Bignoniaceae 340 Verbenaceae 343 Nyctaginaceae 347 Polygonaceae 348 Myristicaceae 351 Lauraceae 355 Hemandiaceae 360 Euphorbiaceae 361 Species with a stratified secondary phloem 364, Species with alternating lenticular hard bast plates 365, Species with scattered solitary fibers 370

6 XV Moraceae 375 Lacistemaceae 382 Families with the same structural peculiarities 382 Families with a very regular bark structure 383 Families with secretory canals Tables of studied families 386 Annonaceae Capparidaceae Flacourtiaceae Vochysiaceae Guttiferae Quiinaceae Caryocaraceae Bombacaceae Sterculiaceae Tiliaceae Elaeocarpaceae Humiriaceae Rutaceae Simaroubaceae Burseraceae Meliaceae Dichapetalaceae Olacaceae Opiliaceae Celastraceae Rhamnaceae Sabiaceae Sapindaceae Anacardiaceae Mimosaceae Caesalpiniaceae Papilionaceae Rosaceae Rhizophoraceae Combretaceae Myrtaceae Lecythidaceae Melastomaceae 406 Araliaceae Rubiaceae Sapotaceae Apocynaceae Boraginaceae Bignoniaceae Verbenaceae 411- Nyctaginaceae Polygonaceae Myristicaceae Hernandiaceae Lauraceae Euphorbiaceae Moraceae Lacistemaceae Preparation of a key for the identification of families, genera and species. 417 G. Characters of taxonomic importance in bark structure Keys and classifications Key to sclerenchyma distribution Ray types Disposition of secretory cells, cavities and canals Disposition of canal sheath parenchyma Disposition of crystals Lenticel frequency and distribution Composition and arrangement of the hard bast Other criteria of taxonomic use Taxonomic importance of the rays Periderm properties of taxonomic value Cork Phelloderm Lenticels Number of periderms Rhytidome structure General observations 463 H. Ecological aspects Total batk width Rhytidome width Cork width 472

7 XVI 4. Phelloderm width Inner bark width General discussion of bark width and habitats Ecological formations of importance in these studies Relations between structural characteristics of the bark and special habitats General conclusions Comparison of the number of species and number of individuals in the humid forest of Venezuelan Guayana Growth ring formation in the bark Considerations on growth ring formation Special examples of growth ring formation in the bark General conclusions Barks from trees of other, mostly drier habitats Special examples General conclusions 538 I. Phylogenetic aspects 546 K. The main patterns in bark formation and their origin Pattern formation through growth activities in the cambium Pattern formation through dilatation growth Formation of horizontal stories Occurrence of net-like patterns The ontogenetic sequence The tendency to transform irregular patterns into regular ones by "synergetic" processes Repetition Pattern formation by combination of different elements 563 L. Economic bark utilization 565 Bibliography 572 Author index 589 Index of scientific plant names 593 Index of vernacular plant names 602 Subject index 606

of flow,rs and fruits

Roland Keller Identifi~cation of tropical woodyi plants in the absence of flow,rs and fruits I I A field guide Springer Basel AG Author: Dr. Roland Keller Institut de Botanique Systematique et de Geobotanique