Revisional study of neotropical Beilschmiedia species (Lauraceae ...

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Systematic position of the species in Lauraceae As mentioned in the introduction, I treated the species studied here as Beilschmiedia Nees (1833). Mez (1889) recognized Hufelandia distinct from Beilschmiedia mainly by absence of an involucre (consisting of scales and subtending young inflorescences), but Kostern1ans (1938) regarded the involucre character unimportant and combined Hufelandia in Beilschmiedia. I thus far do not know if the involucre character is taxonomically important and if the species with involucres are distinct enough to be separated from the neotropical species. These questions should be answered after a careful study of palaeotropical Beilschmiedia species. In a preliminary observation of the palaeotropical species, the involucre character did not appear consistently and there were several species identified as Beilschmiedia apparently congeneric to the neotropical species. Beilschmiedia had long been placed in the tribe Perseae Nees based on thyrsoid inflorescences without involucres. Kostermans ( 1957) regarded development or lack of cupules as an in1portant character for the Lauraceae systen1atics, and placed Beilschmiedia closed to the genera such as Endiandra, Mezilaurus, Potmneia, Persea and Phoebe. Hutchinson (1964) regarded nUinber of anther cells as a more important character, and placed Beilschtniedia in the tribe Apollonieae Hutchinson with such genera as Endiandra, Aniba, Mezilaurus, Licaria and Endlicheria. Richter (1981) published wood and bark anaton1y of Lauraceae, in which he found three large groups of genera. He placed Beilschmiedia in a group with Endiandra, Potameia, Triadodaphne, Cryptocarya and Ravensara (see the section wood anatomy). Cryptocarya has the fruits completely enclosed by cupules and had been regarded a distantly related genus to Beilschmiedia by Kostermans ( 1957). VanderWerff & Richter (1996) reviewed these classifications and concluded androecial characters such as number of anther cells were not suitable to use in a classification of Lauraceae. Instead they proposed a classification based on the inflorescence structure and the wood and bark anatomy. In their study Beilschmiedia was placed in the tribe Cryptocaryeae Nees, distinguished by several characters such as paniculate inflorescences with the ultimate divisions not strictly cymose, marginal parenchyma, nonseptate fibers with conspicuously bordered pits, and exclusively simple vessel perforations in secondary xylem. 14
Although I did not make an intensive study for the systematic position of neotropical Bei!schmiedia species in Lauraceae, my preliminary con1parison of the Lauraceae genera indicated that the closest genus of Beilschmiedia in the neotropics might be Cryptocarya. Relationships within neotropical Beilscluniedia species Relationships within neotropical Beilschmiedia species is discussed with their leaf anaton1y in the third chapter of this study. As the result of the study, they are divided into five groups tnainly in their cuticular characters. See the third chapter for detail. Systematic treatn1ent Beilscluniedia Nees in Wallich, Pl. Asiat. rar. 2: 61, 69. 1831. TYPE: Beilschtniedia roxburghiana Nees. Hufelandia Nees, Plantarun1 Laurinarun1 Secundum Affinitates Naturales Expositio: 11. 1833. TYPE: Hufelandia pendula Nees. Bellota Gay, Historia Fisica y Politica de Chile, Botanica 5: 297. 1849. TYPE: Bello! a miersii Gay. Trees or rarely shrubs. Leaves alternate or opposite, rarely clustered, pinnately veined. Inflorescences axillary, paniculate or racen1ose, terminal branches of the panicles not strictly cymose. Flowers bisexual; tepals 6, equal or subequal, usually deciduous; stan1ens 9 or 6 (the third whorl stan1inodial), filaments usually shorter than the anthers; anthers usually 2- locular, the first and second whorls introrse, the third whorl extrose to almost introrse, staminodia 3 in the fourth whorl or absent, or rarely 6 in the third and fourth whorls, the staminodia representing the third whorl columnar, the staminodia representing fourth whorl cordate to triangular in outline; ovary superior; receptacle flat to shallowly cup-shaped. Fruit elJipsoid, pyrifom1, or spherical, usually purple-black, free on a scarcely thickened pedicel. 15
Page 1 and 2: KURENAI : Kyoto University Researc
Page 3 and 4: Contents Abstract Chapter 1 Chapter
Page 5 and 6: Chapter 1 General Introduction Laur
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: Many species occur fron1 premontane
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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