Revisional study of neotropical Beilschmiedia species (Lauraceae ...

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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
Page 1 and 2: KURENAI : Kyoto University Researc
Page 3: Contents Abstract Chapter 1 Chapter
Page 7 and 8: Chapter 2 Revisional study of neotr
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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Revisional study of neotropical Beilschmiedia species (Lauraceae ...

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