Revisional study of neotropical Beilschmiedia species (Lauraceae ...

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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
Page 1 and 2: KURENAI : Kyoto University Researc
Page 3 and 4: Contents Abstract Chapter 1 Chapter
Page 5: Chapter 1 General Introduction Laur
Page 9 and 10: Akkermans", if it is not a type, Be
Page 11 and 12: Straightness of the indument son1et
Page 13 and 14: Staminodia in the fourth whorl are
Page 15 and 16: elongs not to B. ova/is but to B. a
Page 17 and 18: Many species occur fron1 premontane
Page 19 and 20: Although I did not make an intensiv
Page 21 and 22: 12a. Inflorescences racen1ose, more
Page 23 and 24: 1. Beilscluniedia alloiophylla (Rus
Page 25 and 26: slopeS of creek, NE slope of Altos
Page 27 and 28: Rica. This group appears to have re
Page 29 and 30: anay, and he stated that the Colotn
Page 31 and 32: Phenology: collected in flower in M
Page 33 and 34: angustifolia has pubescent pistils,
Page 35 and 36: The specimen with a label saying "B
Page 37 and 38: 23, 1990 (fr.), Estinoza45 (CR, INB
Page 39 and 40: Beilschmiedia rohliana Lasser, Bol.
Page 41 and 42: (fl.), Haber 11837 (MO); La Peii.a
Page 43 and 44: (F); Con. Ceylan, path La Esmeralda
Page 45 and 46: Colombia (Cuatrecasas 22220, 22438)
Page 47 and 48: locality, Nov. 29- Dec. 18, 1964 (s
Page 49 and 50: Among Brazilian Beifschmiedia speci
Page 51 and 52: van der Werff et al. 12951 (holotyp
Page 53 and 54: inside; starnens 9, filaments ca. 0
Page 55 and 56: ecause the flowers of the type coll
Page 57 and 58:
This species also shares a similar
Page 59 and 60:
synonym of B . sulcata, supposedly
Page 61 and 62:
Additional specimen examined: Brazi
Page 63 and 64:
Cristobal leaving main Hwy 100.8 mi
Page 65 and 66:
pubescent with erect, curly to wavy
Page 67 and 68:
Tree, to 20 m tall. Terminal buds p
Page 69 and 70:
16 x (5) 6-10 em; base obtuse to cu
Page 71 and 72:
Alajuela: Canton de Alfaro Ruiz, Pa
Page 73 and 74:
There are several collections from
Page 75 and 76:
Spiritus (Las Villas): same mountai
Page 77 and 78:
(fl.), Ekman Hl4565 (MO), Apr. 3, 1
Page 79 and 80:
316, 1906, Little Jr. 6293, Aguilar
Page 81 and 82:
Cerro Hueco, ca. 700 malt., Feb. 4,
Page 83 and 84:
ecause there is a distribution gap
Page 85 and 86:
Beilschmiedia immersinervis also ap
Page 87 and 88:
hairs. Fruits ellipsoid, ca. 5 x 3
Page 89 and 90:
Additional specimens examined: Cost
Page 91 and 92:
2, 1994 (young fl.), van der Werff
Page 93 and 94:
In the illustration of Ruiz and Pav
Page 95 and 96:
Table 1. List of the Beilschmiedia
Page 97 and 98:
Legends of figures Fig. 1. Five typ
Page 99:
4 A B c . w ; . -)1 . ..... J.f D f
Page 102 and 103:
o Beilschmiedia berteroana * B. bre
Page 104 and 105:
o Beilschmiedia hexanrhera * B. hon
Page 106 and 107:
0 o Beilschmiedia ovaLioides *B. ov
Page 108 and 109:
C::::J.o•' :: ~ ·a ' \) ' ,o 0 .
Page 110 and 111:
Chapter 3. Leaf Anatomy of neotropi
Page 112 and 113:
3. Leaf sections The technique used
Page 114 and 115:
16), B. ovalis (Fig. 18), B. peruiu
Page 116 and 117:
smooth. However, in B. angustielfip
Page 118 and 119:
uneven-sized subsidiary cells. This
Page 120 and 121:
Midrib. The most distinct character
Page 122 and 123:
This grouping is more or less suppo
Page 124 and 125:
In leaf sectional study, neotropica
Page 126 and 127:
Species of group 2 have leaves oppo
Page 128 and 129:
Table 1. List of samples for venati
Page 130 and 131:
0\ Table 2. Continued. B. emarginat
Page 132 and 133:
Legends of figures Fig. 1-25. Venat
Page 134 and 135:
Fig. 122-143. Midrib sections of ne
Page 136 and 137:
2 132
Page 138 and 139:
10 1 1 12 \ 14 134
Page 140 and 141:
136
Page 142 and 143:
138
Page 144 and 145:
140
Page 146 and 147:
142
Page 148 and 149:
144
Page 150 and 151:
146
Page 152 and 153:
148
Page 154 and 155:
150
Page 156 and 157:
152
Page 158 and 159:
154
Page 160 and 161:
References Allen, C. K. 1945. Studi
Page 162:
Meissner, C. F. 1864. Lauraceae. Pp
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