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Title Revisional Study of Neotropical Beilschmiedia Species (Lauraceae Revisional study of neotropical Beilschmiedia species Title (Lauraceae) with special reference to leaf anatomy( Dissertation_全文 ) Author(s) Nishida, Sachiko Citation 京都大学 Issue Date 1998-03-23 URL https://doi.org/10.11501/3135589 Right Type Thesis or Dissertation Textversion author Kyoto University 7 Revisional study of neotropical Beilschmiedia species (Lauraceae) with special reference to leaf anatomy by Sachiko Nishida March,1998 Contents Abstract u Chapter 1 General Introduction Chapter 2 Revisional study of neotropical Beilsch1niedia species Introduction - Taxonomic history 3 Materials and Methods 3 Morphology 5 Anatomy 10 Palynology 11 Karyology 12 Distribution 12 Reproductive biology and seed dispersal 13 Systematic position of the species in the Lauraceae 14 Relationships within neotropical Beilschmiedia species 15 Systematic treatment 15 Tables 91 Legends of figures 93 Figures 94 Chapter 3 Leaf anaton1y of neotropical Beilschntiedia species Introduction 106 Materials and Methods 107 Results 108 1. Venation patterns 108 2. Cuticular characters 111 3. Leaf sections 114 Discussion 116 Tables 124 Legends of figures 128 Figures 131 Acknowledgments 155 References 156 Abstract Lauraceae contain n1any species which are important constituents in tropical forests both ecologically and econon1ically, however, taxonomy of the family is still not sufficiently clarified at the genus or species level. Beilschmiedia is an exan1ple of such a poorly understood genus. Taking advantage of recent intensive collecting of neotropical Lauraceae species, this study revised neotropical Beilschmiedia species, with general descriptions of morphology, anatomy, palynology, karyology, distribution, reproductive biology, seed dispersal, systematic position of the species in the family, and the systematic treatment. As a result, 27 species were recognized with four species newly described and two species newly con1bined in the genus. Additionally, leaf anatomy of neotropical Beilschmiedia species was employed for better understanding of the species relationships. In leaf anatomy, groupings of cuticular characters was accepted as representing the species relationships, which resulted to recognize five groups among the species. 11 Chapter 1 General Introduction Lauraceae are the largest family in the order Laurales of the Magnoliidae (sensu Cronquist 1981), with about 50 genera. It is a pantropical family with a few temperate members, and main centers of the diversity are the Indomalayan region and Central to South America (Rohwer 1993 b). All species are trees or shrubs except for the species of Cassytha which are herbaceous parasites. Although the family contains tnany species which are very important constituents in tropical forests ecologically and econon1ically, taxonomy of the family is still not sufficiently clarified at genus or species level. TI1e number of the species in the family is still difficult to estimate, probably between 2500 and 3500 (Rohwer 1993 b). In the neotropics, Lauraceae have a reputation of being one of the most difficult families in angiosperms (Burger 1988). It is due to the following facts; that the generic concepts are still not clear in some cases, that the species are difficult to recognize, and that the representation in herbarium collections is poor (Burger 1988). Systematic revisions of the family with more intensive collecting is sorely needed. Beilschmiedia is an exan1ple of such a poorly understood genus. This genus is pan tropical, distributed fron1 Mexico and southern islands in Japan at the northern limit and to central Chile and New Zealand at the southern limit. It comprises about 250 species and is usually distinguished frotn the other laurel genera by a con1bination of the following characters; bisexual and trin1erous flowers, six equal to sub-equal tepals, six to nine fertile stamens, 2-celled anthers, stan1inal glands only in the third whorl, shallow receptacle, and fruit without cupule. Since Meissner (1864) revised the entire family, no revision for the whole genus Beilschmiedia has been made. Regional treatments were made by Liou (1934) for the species in China and Indochina, by Robyns and Wilczek (1951) for the African species, by Fouilloy (1974) for the species in Cameroon, by Li et al. ( 1982) for the Chinese species, by Wright (1984) for the species in New Zealand, and by Hyland (1989) for the Australian species. For the neotropical species, Kostermans 's work (1938) was the most recent comprehensive revision, and Allen ( 1945) treated the Mexican and Central American species. These revisional works for neotropical Beilschmiedia species were not based on abundant n1aterial, 7 of 15 species in Kostermans 's revision and 4 of 8 species in Allen's revision were known only from the type collections. And since Kostermans' s revision, 15 names have been already described under Beilschmiedia in the neotropics. A revision for neotropical Beilschmiedia species, being up to date and with n1ore material, is needed for an understanding of the whole genus and also the neotropical Lauraceae. Fortunately, as Burger (1988) mentions, Latin American botanists are actively collecting neotropical plants including Lauraceae these days. Especially, van der Werff in the Missouri Botanical Garden has been working on a number of projects on a systematic study of Lauraceae coupled with intensive collecting of the family in the neotropics. I had an opportunity to study at Missouri Botanical Garden from fall in 1994 to spring in 1997, and took the advantage to work on neotropical Beilschmiedia species under van der Werff's supervision. In this study I revised neotropical Beilschmiedia species in which I recognize 27 species, with general descriptions of morphology, anatomy, palynology, karyology, distribution, reproductive biology, seed dispersal, and systematic position of the species in the fmnily in the second chapter. And for better understanding of relationships of the species, I employed leaf anatomy in the third chapter. Neotropical Beilschmiedia species have relatively similar flowers and fruits except for a few species, and it is more reasonable to study vegetative n1orphology rather than reproductive morphology for systen1atics of the species. An1ong neotropical Beilscluniedia species, there appear to be two groups, one group of the species alternate leaves and fine ramification (ramification is here defined as ultimate branching patterns in leaf veins), and another group of the species with opposite leaves and coarse ratnification. Preliminary observation (Christophel, pers. comm.) indicated that the two groups also differ in characters of the cuticle. Therefore, I investigated leaf anaton1ical characters (venation patterns, cuticular and leaf section characters) for all the neotropical Beilschrniedia species available in order to discuss the following; 1) whether the cuticular and leaf section characters support the grouping based on phyllotaxis and venation patterns, and 2) how we can incorporate leaf-anatomical characters in the classification of neotropical Beilschmiedia species. These studies would not only help to know neotropical Beilschmiedia species further, but would also give an important step to understand whole Beilschmiedia, one of the largest genera in Lauraceae. 2 Chapter 2 Revisional study of neotropical Beilschnliedia species Introduction - Taxonon1ic history The genus Beilschmiedia was described by Nees (1831), with two Asian species B . raxburghiana and B. fagifoiia. Nees (1833) described f!ujefandia with two neotropical species, H. pendufa and H. thamaea. Since then 12 species had been described under Hujefandia. But Hemsley (1882) transferred the type species H . pendula (as well as H. thomaea) under Beifschmiedia. Mez ( 1889) restored Hufelandia to generic rank, but Kostermans (1938) included Hufelandia in Beifschmiedia again. In this study, I follow Kostem1ans 's sense about the generic treatment, because Beilschmiedia in Asia where the type species of the genus occurs is poorly known and it is impossible to clarify the relationship between the neotropical and Asian species before a better understanding of the Asian species. In addition to Hujefandia Nees, Beifschmiedia has another synonym, Beffota Gay, for the Chilean species. Kostern1ans (1938) listed other three synonyms, Bafdu Nees (non Feuilee), Baldus 0. Kuntze (non Adanson) and Wimmeria Nees ex Meissner, but they are not valid or legitimately published. First, Boldu Nees is a superfluous name. And Baldus is the natne which Kuntze did not desctibe but only reestablished from Baldus Molina, and Baldus Molina actually belongs to the Monimiaceae. Wimm.eria is the name written on a specimen label of B . pendula and not validly published. Gay (1849) described a new genus Bellata with one species, Bellata miersii. The genus Gay described, Bellota, was combined in Beilsch1niedia by Kostern1ans ( 1938). For the palaeotropical species, Beilschmiedia has the following synonyms; Nesadaphne Hooker f., Bernieria Baillon, Tyloste1non Engler, Afrodaphne Stapf, Thouvenotia Danguy, Purkayasthaea Purkayastha, and Lauramerrillia C. K. Allen. Kostem1ans (1952) con1bined a neotropical genus, Anaueria, in Beilschmiedia. But Richter ( 1981) found that Anaueria differs strongly fron1 Beilschmiedia in wood anaton1y; this, combined with differences in flower and fruit morphology, resulted in the current recognition of Anaueria as a distinct genus. Materials and Methods 3 Most of the revisional study was done at Missouri Botanical Garden (MO), St. Louis,
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