ABSTRACT A MONOGRAPH OF THE GENUS MAACKIA By Carolyn K. Levings The genus Maackia (Papilionoideae, Fabaceae) comprises trees and shrubs native to China, Japan, Korea, Russia, and Taiwan. Morphologically, Maackia appears most closely related to Cladrastis. However, molecular data indicate that Maackia is distantly related to that genus, being more closely allied to some Sophora species, Salweenia, and Euchresta, whereas Cladrastis is basal to the subfamily. Individual floras have mentioned the genus, but none have included all the species in Maackia, and no previous monograph exists. Over 600 specimens were obtained on loan from herbaria around the world. Separate analyses of fruiting and flowering specimens were conducted, because flowers and fruit do not occur simultaneously on one plant. From 130 specimens, 77 vegetative characters, 16 pod and seed characters, and 56 floral characters were measured. Data were analyzed using NTSYS – pc and MINITAB. Eleven species are recognized, one of which is new, another newly named, and a new subspecies combination made. A MONOGRAPH OF THE GENUS MAACKIA A Thesis Submitted to the Faculty of Miami University in partial fulfillment of the requirements for the degree of Master of Science Department of Botany By Carolyn Kathleen Levings Miami University Oxford, Ohio 2006 Advisor _________________________________ Michael A. Vincent Reader__________________________________ R. James Hickey Reader__________________________________ Nancy Smith–Huerta TABLE OF CONTENTS Section Page Introduction............................................................................................1 General Habitats.....................................................................................2 Fossil History.........................................................................................4 Phytochemistry ......................................................................................7 Uses........................................................................................................7 Nodulation..............................................................................................8 Horticulture............................................................................................8 Herbivory and Diseases .......................................................................11 Morphology..........................................................................................11 Chromosome Counts............................................................................20 Related Genera.....................................................................................20 Rationale for this Study .......................................................................23 Methods................................................................................................25 Results..................................................................................................34 Discussion............................................................................................50 Taxonomic Treatment..........................................................................55 Literature Cited ....................................................................................95 Appendix 1. Floral and vegetative data used in analyses. .................103 Appendix 2. Fruit and vegetative data used in analyses. ...................105 Appendix 3. Complete list of specimens examined...........................115 ii LIST OF TABLES Table Page 1. Horticultural information for cultivated Maackia species.............10 2. Characters of genera reportedly related to Maackia......................22 3. Comprehensive list of names published in the genus Maackia, both extant and fossil taxa..............................................................24 4. Description of measurements taken for analytical and descriptive purposes .........................................................................................27 5. Diagnostic characters of Maackia species .....................................52 iii LIST OF FIGURES Figure Page 1. Map of East Asia....................................................................................3 2. Fossils from temperate East Asia...........................................................6 3. Habit of Maackia spp. grown in cultivation ........................................14 4. Bark images of cultivated Maackia species from Arnold Arboretum............................................................................................15 5. Hairs from leaves of pubescent members of Maackia.........................16 6. Opened calyces from putative Maackia species ..................................17 7. Pollen from Maackia species...............................................................18 8. Anatomical images from Maackia spp ................................................19 9. Diagram of floral characters used for data analysis.............................31 10. Diagram of fruit characters used for data analysis...............................32 11. Diagram of vegetative characters used for data analysis.....................33 12. Projection plot of floral characters according to their respective Eigen values from PCA ......................................................36 13. PCA projection plot of flowering specimens.......................................37 14. Projection plot of fruit characters according to their respective Eigen values from PCA ......................................................38 15. PCA projection plot of fruit specimens ...............................................39 16. Projection plot of floral characters according to their respective Eigen values from PCA ......................................................40 17. Phenogram resulting from PCA of data from flowering specimens of Maackia, excluding M. tenuifolia ..................................41 18. Projection plot of floral characters according to their respective Eigen values from PCA ......................................................42 19. Phenogram resulting from PCA of data from fruiting specimens of Maackia, excluding M. tenuifolia ..................................43 20. PCA projection plot of floral characters used in analysis of M. amurensis f. amurensis and M. amurensis f. pilosella ...................44 21. PCA tree plot of M. amurensis f. amurensis and M. amurensis f. pilosella based on flowering specimens..........................45 22. Photographs of opened calyces from Maackia amurensis showing greater fusion of teeth in M. amurensis f. pilosella and var. stenocarpa..............................................................................46 23. Projection plot of characters used in analysis of M. amurensis var. amurensis and M. amurensis var. stenocarpa according to their respective Eigen values from PCA......................................................................................................47 24. PCA tree plot of M. amurensis var. amurensis and M. amurensis var. stenocarpa based on fruiting specimens .....................48 25. SEM micrographs of pollen of Maackia species .................................49 26. Key characters of Maackia species......................................................58 27. Distribution map of M. amurensis Rupr. & Maxim ............................64 iv 28. Distribution of M. amurensis ssp. amurensis and M. amurensis ssp. pilosella on the Korean Peninsula...............................65 29. Distribution map of Maackia australis (Dunn) Takeda.......................68 30. Distribution map of M. chekiangensis .................................................70 31. Distribution map of Maackia fauriei ...................................................72 32. Distribution map of Maackia floribunda .............................................75 33. Distribution map of Maackia hupehensis ............................................78 34. Distribution map of Maackia hwashanensis........................................80 35. Diagnostic characters of Maackia japonica.........................................83 36. Distribution map of Maackia japonica................................................84 37. Diagnostic characters of Maackia nakaii.............................................86 38. Distribution map of Maackia nakaii....................................................87 39. Distribution map of Maackia taiwanensis ...........................................89 40. Distribution map of M. tashiroi ...........................................................92 v ACKNOWLEDGEMENTS There are so many people I would like to thank for getting me through these last 2 ½ years, that another 100 pages would have to be added to get them all in. My memory is now shot, and if I forgot a name, please don’t take offence. It was unintended and thank you for all your help. First I would like to thank my committee for all their work. Mike, I won’t be able to comprehend all you’ve done for me for years to come. I thank you for all your patience (especially during my mulish periods), letting me bring art in to the research, and everything you’ve taught me. Jim, I don’t know what hobbies I would have had to develop if I didn’t have you to tease. Even though I never mentioned it, I appreciate all the attention to detail that only a pteridologist specializing in a hideously deformed grass would possess. Nancy, thanks for bringing in a non-taxonomy perspective. My thesis would not have been as well rounded without