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, ~" " •. """~" > ACKN OWLEDGE1VIENTS This thesis has been an interna.tional effort, produced with trle cooperation of a grea.t many people in Japan~ Korea, England, and the United States. With no .i.l1tentionof diminishing the cor:tributi.qn of anyone not. mentioned below, I would like to single out a. few people for special thanks. I wish to thank the IJcmgwood Program at the University of Delavv"are for th8 award of the fell.owE}hip which helped to support this project. I am particularly indebted to my thesis committee--Dr. Richard W. Lighty, Mr. William H. Frederick, ,Jr., and Dr. Donald Huttleston-- for their very patient assistance. In Japan I wish to thank Dr. Sumihiko Hatsushima, Dr. FU"TIioMaekawa., Mr. Tadanori 'I'animura, Mr. Eiji Yamaha.ta, and Mr. NIatoshi Yoshida for patiently respondi.ng to my endless questions and for providing much of the research material cited herein. In addition, Mr. Yoshimichi Hirose, Mr. Mikinori Ogisu, Dr. Yotaro rrsuka.moto, and iii Dr. Ma,sato Yokol have earned my gratitude for directing me to useful research material. Among the many people who helped me in Japan, I must single out the efforts of Dr. Toshio Ando who so often arranged my itinerary in Japan and helped me with the translation of Japanese source material. For the privilege of examining herbarium specimens I wish to thank Dr. TchaT1;gBok 1,ee and h.is staff at Seoul. National University, Korea, and Mr. Ian Beier and his staff at the Royal Botanic Gardens, Kew, England. My thanks too to Mr. Yong-,Jun Chang (Korea), Mr. Donald Emminger (United States), and Mr. Eiji Yamahata (Japan) for the preparation of the illustrations which have added greatly to the value of this paper. iv < i I TABLE OF CONTENTS ACKNOVIT,EDGEMENTS •• •••••••••••• • •• iii TABLE OF CONTENTS ................ v LIST OF TABLES •• . • • • · ..... viii LIST OF FIGURES ....... · ..... ix INTRODUCTION • . ••• a •••• 1 PART ONE. 'J:iHE GENUS ASARUM AND ITS JAPANESE SPECIES Cha.pter I. CHARACTERISTICS OF THE GENUS ASARUM 5 II. THE DISTRIBUTION AND PALEO- PHyrrOGEOGRAPHY OF ASARUM •• · .... 18 III. THE INTEGRITY OF THE GENUS ASARUM • • 26 IV • THE KARYOLOGY OF ASARUM ••••••• .37 V. MIGRATION AND SPECIATION OF JAPAJlfESE ASARUM •••••••••••••••• 43 VI. THE JAPANESE SPECIES OF ASARUM • • •• .50 Introduction • • • • •••••• · .. .50 The Sieboldii-Caulescens group · · • • 52 The Sakawanum-Asnerum group · • • • • 61 rrhe Curvastigma group • • • • • • • · 78 The Ki us ian Uill group • • • • • • · • • 87 I I v / ( The Nipponicum group ••••••••• 98 The Ryukyu Islands group ••••••• 117 FOOTNOTES, PART I •••••••••••••••• 128 BIBLIOGRAPHY, PART I • • •• • • • • • • • • • •• 138 APPENDIX 1. J.VIAPOF JAPAN .0 •••••••••• 143 APPENDIX 2. A KEY TO THE ASARUM SPEC IES OF THE JAPANESE ISLANDS FROM HOKKAIDO TO THE TOKARA STRAIT •••••••••• 145 APPENDIX 3. PRECIS OF DESCRIPTIONS OF ASARUM SPECIES REPORTED FROM ,JAPAN •• • •• 153 Bi.bliography ••••••••• • •• 172 PART TWO. JAPANESE ASARUM CULTIVAJZS AND THE HISTORY OF THEIR CULTIVATION Chapter VII. INTRODUCTION • • • • • • •• • •••• 17.5 VIII. THE ASARUM MON ••••••• • •••• 178 IX. THE DEVELOPMffilTOF KOTEN ENGEl • • •• 195 X. ASARUM CULTIVATION AS A PART OF KOTEN ENGEl •••••••••• • •• 215 XI • CONCLUSION • • • • • • • •• • • • •• 243 FOOTNOTES, PART II •••••••••••••• 245 BIBLIOGRAPHY, PART II • ••••••••• t:, •• 250 APPENDIX ii-, A SELECTED GLOSSARY OF THE CHARACTER- ISTICS (GEl) OF ASARUM CULTIVARS •• 25.3 Bi.bliography •••••••••••• 255 APPENDIX 5. DESCRIPTIONS OF KOTEN ENGEl ASARUM CULTIVARS •••••••••••••• 256 Bibliogra.phy •••••••••••• 274 APPENDIX 6. ASARUl\1MElKAN RANKINGS (1976) • • •• 275 vi APPENDIX 7. TECHNIQUES OF ASARUM CUIJTIVATION •••••••• • •••• 278 Bibliography ••••••• • ••••• 284 APPENDIX 8. ASARUM SPECIES IN JAPANESE GARDENS 285 APPENDIX 9. JAPANESE ASARUM IN AMERICAN GARDEN S •••••••••• • •••• 287 SUPPLEMENTAlJ BIBLl OGRAPHY •••••••••• 't • 289 vii LIST OF TABLES Table 1. Morphological Variation among Asarum Species •• 12 2. Chart of Geological Eras, Periods, and Epochs •••••••••••••••••••• 20 J . Maekawa's Separation of Asarum, Sensu Lato • • 28 viii LIST OF FIGURES Figure 1. A Diagramatic Representation of the Aristolochiaceae •••••••••• • • • • • 6 2. Saruma henryi ••••••••••••••••• 9 3. Some Parts of the Asarum Flower • • • • • • • • 11 l}. Possible Routes of Pri.mary Migration of Asarum 2..5 5. Possi.ble Subsequent Route of Migration of Asarum from Asi.a to North America ••••••• 25 6. Examples of Flower Structure Whi.ch Illustrate Maekawa's Divisions of Asarum, ~ensu Lato ••• 30 7. Asarum albomaculatum from Taiwan ""'" 3J 8. Asarum geophilum from Southern China ••••• 33 Comparison of Hexas,tylis shuttleworthii and Heterotropa takaoi •••••••••••• • • 34 10. Comparison of Two Species within the Genus Heterotropa: H. takaoi and !i. sakawana •••• 35 11. Asarum Arranged by Chromosome Number on a Map of the Modern World •••••••••••••• 41 12. The Probable Routes of Asarum Migration into tTapan •••••••••••••••••• 0 •• l~6 13. Stamen Movement in the Sieboldii-Caulescens Group ••••• •••••••••••••• • • 54 ix ( 11j.. A Comparison of the Flowers of Asarum sieboldii, A. heterotropoides, and !::.. caulescens .•••••••••••••• 54 15. Asarum dimidianum ••••••••••••• 55 16. Asarum Species in the Sieboldii-Caulescens Group and the Sakawanum-Asperum Group ••• 57 Distribution of Asarum heterotropoides, A. sieboldii, A. dimidianum, and I. caulescens - ••••••••••••• • • .59 18. Distri.buti.onof Asarum costatum, A. sakawa- --num, -A. minamitanianum, and -A. stellatum •• 64 19. Distribution of Asarum as'Perum, !::.. hexalobum, A. hexalobum yare perfectum, and A. crassum 66 20. Comparison of Calyx Development in Asarum mina!nitanianum , A. stellatQm, A. sakawanum, and A. costaturn- ••••••• - ••••••• 67 21. Flower Structure of Asarum costatum • ••• 68 22. Flower Structure of Asarum sakawanum •• • • 69 23. Asarum minamitanianum ••••••••••• 70 24. Photographs of Asarum Species in the Sakaw-anum-Asperum Group and the Kiusianum Group •••••••••• • • • • 72 25. Flower Structure of Asarum hexalobum • • • • 74 26. Flower Structure of Asarum crassum • • t • 7.5 27 • Asarum crassum • ••••••••• • • • • • 76 28. Distribution of Asarum tamaense, A. muramatsui, A. curvastigma, A. asaroides, A. unzen, and !::.. satsumense •••••••• 80 29. A Comnarison of the Flowers of Asarum curvastigma, A. tamaense, A. satsumense, and A. asaroides •••••••••••••••• 81 30. Photographs of Asarum Species in the Curvastigma Group ••••••••• • ••• 83 x 31. Flower Structure of Asarum curvastigma •• , 86 32. Distribution of Asarum kiusianum var. kiusianum, ~. kiusianum var. melanosiphon, ( A. kiusianum yare tubulosum, A. kumageanurn, A. subglobusum, ~. trigy~um, and i A. yakusimense ••••••••••••••• 88 33. Photographs of Asarum Species in the KiusianumGroup ••••••••••• • • • • 92 34. Flower Structure of Asarum subglobosum •• , 94 35. Asarum kinoshitai • . , ..... • ••• 36. Flower Structure of Asarwn kumageanum •••• 96 37. A Comparison of the Flowers of Asaru.'IJ1 kumageanum, ~. yakusimense, and A. lutchuense •••••••••••••••• 99 38. Distribution of Asarum nipponicum yare nipponicum, ~. nipponicum yare kooyanum, A. nipponicum yare brachypodion, ~. nipponicum var. nankaiense, ~. savatieri, and A. bl ume i ••••••••••••••• 102 .39. Distribution of Asarum nipponicum var. rigescens and ~. megacalyx ••••••••• 1.03 40. Distribution of Asarum takaoi and ~. fauriei 104 41. Flower Structure of Asarum nipponicum var. nipponicum ••••••••••••••••• 107 42. Photographs of Asaru.rnSpecies i.n the Nipponicum_ Group •••••••••• • • • • 108 43. Style Shapes and Reticulation Patterns of Three Nipponicum Group Asarum Species •• · . 110 44. Asarum nitl'Ponicum yare brachypodion . • • 111 45. Asarum nipponicum yare nankaiense • ••• • • 113 46. Flower Structure of Asarum takaoi yare hi sauc hi i ••••••••••••••• ••• 115 xi Asarum Distribu·tion in the Ryukyu Islands ••••••••••••• • » ••• 118 48. Styles and Stamens of Six Ryukyu Islands Asarum Species •••••••••••••• 121 49. Photographs of Asarum Species in the Ryukyu Islands Group ••••••••••• 123 50. Asarum leptophyllum ••••••••• • •• 12.5 51. Flower Structure of Asarum dissitum • • •• 126 52. Typical Japanese Kamon Used as Decoration 179 53. Two Designs of Asarum caulescens Used by the Matsuo Shrine in Kyoto •••••••• 185 54. Two Designs of Asarum caulescens Used by the Kamigamo Shrine i.nKyoto ••••••• 187 55. A Drawing of Asarum europaeum from the German Latin Herbarius •••••••••• 188 .56. Derivative Asarum Crests (Kamon) • •••• 193 57. A Structure for the Displa.y of Koten Engei Plants •••••••••••••••••• 204 58. A Structure for the Storage and Display of Koten Enge~ Plants ••••••••••• 205 59. Three Examples of Typical Koten Engei Plan·ts . • •• 212 60. Three Examples of Typical Koten Engei Plan ts ••••••••••••••• • •• 214 61. Leaf Characteristi.cs (Gei) of Asarum Cultivars •••••• :--:-•••••• • •• 221 62. A Tradi.tiona.lAsarum Pot of the Na.minishiki Design • • • • • • • • • • • • • • • • • • 224 63. Six Cultivars of Asarum • • • • • • • • • • 228 64. Six Cultivars of Asarum • • • • • • • • • • 230 6.5 • Six Cultivars of Asarum • • • • • • • • • • 232 xii 66. The Japanese Asarum Society Meikan: the Order of Rankings of One Hundred Cultivars i.n1976 •••••••••• •••• 236 r I A Cartoonist's Representati.on of the I Maturation of Plant Interest in Japan •••• 240 xii.i INTRODUCTION The genus Asarum sensu lato* has historically received more attention from the Japanese than from Westerners. In recent times Japanese enthusiasm for Asarum has been most obvious in two