Systematics and Biogeography of Aralia L. (Araliaceae)
Systematics and Biogeography of Aralia L. (Araliaceae)
Systematics and Biogeography of Aralia L. (Araliaceae)
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SMITHSONIAN INSTITUTION<br />
Contributions from the United States National Herbarium<br />
Volume 57: 1-172<br />
<strong>Systematics</strong> <strong>and</strong> <strong>Biogeography</strong> <strong>of</strong> <strong>Aralia</strong> L. (<strong>Aralia</strong>ceae):<br />
Revision <strong>of</strong> <strong>Aralia</strong> Sects. <strong>Aralia</strong>, Humiles, Nanae, <strong>and</strong><br />
Sciadodendron<br />
by<br />
Jun Wen<br />
Department <strong>of</strong> Botany<br />
National Museum <strong>of</strong> Natural History<br />
Washington, DC<br />
2011
ABSTRACT<br />
Wen, Jun. <strong>Systematics</strong> <strong>and</strong> <strong>Biogeography</strong> <strong>of</strong> <strong>Aralia</strong> L. (<strong>Aralia</strong>ceae): Revision <strong>of</strong> <strong>Aralia</strong> Sects. <strong>Aralia</strong>,<br />
Humiles, Nanae, <strong>and</strong> Sciadodendron. Contributions from the United States National Herbarium, volume<br />
57, 172 pages. This treatment provides a revision <strong>of</strong> four sections <strong>of</strong> <strong>Aralia</strong> L. (<strong>Aralia</strong>ceae). It is the third <strong>and</strong> last <strong>of</strong> the <strong>Aralia</strong><br />
monographic series by the author. The first was on <strong>Aralia</strong> sect. Pentapanax (Seem.) Wen (19 species, Wen 2002); <strong>and</strong> the second<br />
was on <strong>Aralia</strong> sect. Dimorphanthus (Miq.) Miq. (29 spp., Wen 2004). This revision treats <strong>Aralia</strong> sects. <strong>Aralia</strong> (14 spp.), Humiles (3<br />
spp.), Nanae (1 sp.), <strong>and</strong> Sciadodendron (5 spp.). A taxonomic key to all sections <strong>of</strong> <strong>Aralia</strong> is provided. Species keys are provided<br />
for each <strong>of</strong> the three sections with multiple species. As typified by <strong>Aralia</strong> fargesii Franch., <strong>Aralia</strong> sect. Anomalae Harms is now<br />
placed in synonymy <strong>of</strong> <strong>Aralia</strong> sect. <strong>Aralia</strong>. <strong>Aralia</strong> sect. Nanae is the only monotypic section <strong>of</strong> the genus, consisting <strong>of</strong> <strong>Aralia</strong><br />
nudicaulis L. Marchal’s genus Coudenbergia is merged with <strong>Aralia</strong> sect. Sciadodendron in this study. Detailed descriptions on the<br />
morphology <strong>and</strong> ecology, illustrations, <strong>and</strong> distribution maps are provided for each taxon <strong>of</strong> the four sections (23 species). The<br />
phylogeny <strong>of</strong> <strong>Aralia</strong> based on sequences <strong>of</strong> the internal transcribed spacer (ITS) regions <strong>of</strong> nuclear ribosomal DNA <strong>and</strong> three<br />
chloroplast markers, including the ndhF gene, the trnL-F region, <strong>and</strong> the atpB-rbcL spacer, is presented. An early biogeographic<br />
radiation <strong>of</strong> <strong>Aralia</strong> is hypothesized. Taxa <strong>of</strong> <strong>Aralia</strong> sect. Humiles do not form a clade with those <strong>of</strong> the Asian <strong>Aralia</strong> sect. Pentapanax.<br />
These two sections were once treated as constituting the genus “Pentapanax,” <strong>and</strong> they are shown here to be non-monophyletic <strong>and</strong><br />
best treated as belonging to two different sections. <strong>Aralia</strong> bahiana J. Wen is herein described as a new species from Bahia, Brazil.<br />
<strong>Aralia</strong> sect. Sciadodendron is a new nomenclatural combination with the genus Sciadodendron as its basionym. The newly<br />
lectotypified names include (accepted names in boldface): <strong>Aralia</strong> cachemirica Decne., A. californica S. Watson, A. californica<br />
var. acuminata S. Watson ex Howell, A. chilapensis Sessé & Moc., A. continentalis Kitagawa, A. cordata Thunb., A. edulis Sieb.<br />
& Zucc., A. fluminensis Glaz., A. henryi Harms, A. humilis Cav., A. pilosa Franch., A. pubescens DC., A. racemosa L. var.<br />
sachalinensis Regel, Megalopanax rex Ekman ex Harms, Pentapanax ulei Harms, <strong>and</strong> Sciadodendron excelsum Griseb.<br />
Key words: <strong>Aralia</strong>, <strong>Aralia</strong> sect. <strong>Aralia</strong>, <strong>Aralia</strong> sect. Dimorphanthus, <strong>Aralia</strong> sect. Humiles, <strong>Aralia</strong> sect. Nanae, <strong>Aralia</strong> sect.<br />
Pentapanax, <strong>Aralia</strong> sect. Sciadodendron, <strong>Aralia</strong>ceae, biogeography, systematics.<br />
DATE OF PUBLICATION: March 2011<br />
Cover Design by Alice R. Tangerini: front <strong>Aralia</strong> tibetana G. Hoo illustrated by ART; back <strong>Aralia</strong> bahiana<br />
J. Wen illustrated by ART.<br />
Contributions from the United States National Herbarium (ISSN 0097-1618) Department <strong>of</strong> Botany, National<br />
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CONTENTS<br />
INTRODUCTION<br />
TAXONOMIC HISTORY<br />
Generic concept <strong>and</strong> historical infrageneric classifications<br />
<strong>Aralia</strong> from the New World<br />
<strong>Aralia</strong> from the Old World<br />
PHYLOGENETIC RELATIONSHIPS<br />
MORPHOLOGICAL CHARACTERS<br />
Habit<br />
Underground systems<br />
Prickles<br />
Leaf architecture <strong>and</strong> morphology<br />
Trichomes <strong>and</strong> vesture<br />
Cuticles<br />
Pedicel articulation<br />
Inflorescence<br />
Bracts <strong>and</strong> bracteoles<br />
Flowers<br />
Fruits<br />
POLLEN MORPHOLOGY<br />
CHROMOSOME NUMBERS<br />
INFRAGENERIC CLASSIFICATION<br />
I. <strong>Aralia</strong> sect. <strong>Aralia</strong><br />
II. <strong>Aralia</strong> sect. Humiles<br />
III. <strong>Aralia</strong> sect. Nanae<br />
IV. <strong>Aralia</strong> sect. Pentapanax<br />
V. <strong>Aralia</strong> sect. Sciadodendron<br />
VI. <strong>Aralia</strong> sect. Dimorphanthus<br />
KEY TO SECTIONS OF ARALIA<br />
REPRODUCTIVE BIOLOGY<br />
SPECIES CONCEPTS<br />
DISTRIBUTION AND BIOGEOGRAPHY<br />
Early evolutionary radiation in <strong>Aralia</strong><br />
<strong>Aralia</strong> sect. <strong>Aralia</strong><br />
<strong>Aralia</strong> sect. Humiles<br />
<strong>Aralia</strong> sect. Sciadodendron<br />
TAXONOMY<br />
ARALIA L. SECT. ARALIA<br />
Key to species <strong>of</strong> <strong>Aralia</strong> sect. <strong>Aralia</strong><br />
1. <strong>Aralia</strong> californica<br />
2. <strong>Aralia</strong> racemosa<br />
5<br />
7<br />
7<br />
8<br />
9<br />
10<br />
13<br />
13<br />
13<br />
13<br />
13<br />
13<br />
13<br />
14<br />
15<br />
22<br />
22<br />
22<br />
22<br />
26<br />
28<br />
28<br />
28<br />
28<br />
29<br />
29<br />
30<br />
31<br />
31<br />
32<br />
32<br />
32<br />
33<br />
33<br />
33<br />
34<br />
34<br />
35<br />
36<br />
42
3. <strong>Aralia</strong> bicrenata<br />
4. <strong>Aralia</strong> cordata<br />
5. <strong>Aralia</strong> schmidtii<br />
6. <strong>Aralia</strong> taiwaniana<br />
7. <strong>Aralia</strong> continentalis<br />
8. <strong>Aralia</strong> cachemirica<br />
9. <strong>Aralia</strong> tibetana<br />
10. <strong>Aralia</strong> fargesii<br />
11. <strong>Aralia</strong> atropurpurea<br />
12. <strong>Aralia</strong> apioides<br />
13. <strong>Aralia</strong> glabra<br />
14. <strong>Aralia</strong> henryi<br />
ARALIA SECT. NANAE<br />
15. <strong>Aralia</strong> nudicaulis<br />
ARALIA SECT. HUMILES<br />
Key to species <strong>of</strong> <strong>Aralia</strong> sect. Humiles<br />
16. <strong>Aralia</strong> humilis<br />
17. <strong>Aralia</strong> regeliana<br />
18. <strong>Aralia</strong> scopulorum<br />
ARALIA SECT. SCIADODENDRON<br />
Key to species <strong>of</strong> <strong>Aralia</strong> sect. Sciadodendron<br />
19. <strong>Aralia</strong> soratensis<br />
20. <strong>Aralia</strong> warmingiana<br />
21. <strong>Aralia</strong> rex<br />
22. <strong>Aralia</strong> excelsa<br />
23. <strong>Aralia</strong> bahiana<br />
EXCLUDED NAMES<br />
ACKNOWLEDGEMENTS<br />
LITERATURE CITED<br />
ALPHABETICAL INDEX TO COLLECTORS<br />
INDEX TO SCIENTIFIC NAMES<br />
COLOR PLATES 1-16<br />
68<br />
73<br />
77<br />
80<br />
83<br />
87<br />
90<br />
94<br />
96<br />
100<br />
103<br />
106<br />
109<br />
109<br />
118<br />
118<br />
118<br />
127<br />
131<br />
133<br />
134<br />
134<br />
138<br />
142<br />
142<br />
149<br />
151<br />
151<br />
152<br />
158<br />
170<br />
49
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
<strong>Systematics</strong> <strong>and</strong> <strong>Biogeography</strong> <strong>of</strong> <strong>Aralia</strong> L. (<strong>Aralia</strong>ceae):<br />
Revision <strong>of</strong> <strong>Aralia</strong> Sects. <strong>Aralia</strong>, Humiles, Nanae, <strong>and</strong> Sciadodendron<br />
<strong>Aralia</strong> L. is one <strong>of</strong> the approximately 50 genera<br />
<strong>of</strong> the predominantly tropical <strong>Aralia</strong>ceae (or the<br />
ginseng plant family). The genus consists <strong>of</strong> 71<br />
species <strong>and</strong> is the fifth largest in the <strong>Aralia</strong>ceae<br />
(following Schefflera J. R. Forst. & G. Forst.,<br />
Polyscias J. R. Forst. & G. Forst., Oreopanax<br />
Decne. & Planch., <strong>and</strong> Dendropanax Decne. &<br />
Planch.). <strong>Aralia</strong> plays an important role in<br />
underst<strong>and</strong>ing the diversification <strong>of</strong> <strong>Aralia</strong>ceae<br />
because it represents most north temperate species<br />
<strong>of</strong> the family. Harms (1898) considered <strong>Aralia</strong> as<br />
one <strong>of</strong> the centers <strong>of</strong> diversity within <strong>Aralia</strong>ceae<br />
<strong>and</strong> suggested that it played a critical role in leading<br />
to other lineages in <strong>Aralia</strong>ceae <strong>and</strong> Apiaceae.<br />
Recent phylogenetic analyses <strong>of</strong> the core<br />
<strong>Aralia</strong>ceae suggest that <strong>Aralia</strong> is closely related to<br />
Panax L. The <strong>Aralia</strong> – Panax clade constitutes<br />
one <strong>of</strong> the three major clades in the core <strong>Aralia</strong>ceae<br />
(Wen et al. 2001; Plunkett et al. 2004). Within<br />
<strong>Aralia</strong>ceae, <strong>Aralia</strong> <strong>and</strong> Panax are perhaps most<br />
closely related to Polyscias, Pseudopanax K. Koch,<br />
<strong>and</strong> their close relatives (Wen et al. 2001; Plunkett<br />
et al. 2004).<br />
The phylogenetic position <strong>of</strong> <strong>Aralia</strong> in the<br />
<strong>Aralia</strong>ceae <strong>and</strong> its relationship to Panax were<br />
variously hypothesized prior to modern<br />
phylogenetic analyses. Seemann (1868) treated<br />
<strong>Aralia</strong> <strong>and</strong> Panax in different tribes. Harms (1898)<br />
suggested a close relationship between <strong>Aralia</strong> <strong>and</strong><br />
Panax. Emphasizing a single-character taxonomy,<br />
Hutchinson (1967) placed <strong>Aralia</strong> <strong>and</strong> Panax once<br />
again in different tribes. But this treatment has<br />
been criticized as “unnatural” (Hoo <strong>and</strong> Tseng<br />
by<br />
J. Wen 1<br />
INTRODUCTION<br />
1 Department <strong>of</strong> Botany, National Museum <strong>of</strong> Natural History, MRC-166, Smithsonian Institution, P.O. Box 37012, Washington,<br />
D.C. 20013-7012, U.S.A., wenj@si.edu<br />
1978, p.178). On the other h<strong>and</strong>, some workers<br />
(e.g., Decaisne <strong>and</strong> Planchon 1854; Clarke 1879;<br />
Burkill 1902) treated Panax as a synonym <strong>of</strong><br />
<strong>Aralia</strong>. Based on the 5-merous flowers <strong>and</strong><br />
articulated pedicels, Harms (1898) <strong>and</strong> Eyde <strong>and</strong><br />
Tseng (1971) suggested that Stilbocarpa (Hook.<br />
f.) Decne. & Planch. from New Zeal<strong>and</strong> <strong>and</strong><br />
adjacent isl<strong>and</strong>s may be closely related to <strong>Aralia</strong><br />
<strong>and</strong> Panax. Vegetatively Stilbocarpa shares many<br />
similarities with some early-diverging lineages <strong>of</strong><br />
Apiaceae. Mitchell et al. (1999) have shown that<br />
Stilbocarpa is nested in a clade <strong>of</strong> Apiaceae with<br />
Azorella <strong>and</strong> Schizeilema from the Southern<br />
Hemisphere, very distinct from any araliaceous<br />
genera.<br />
As slightly modified from Wen (1993), <strong>Aralia</strong><br />
is here defined by the following characters: 1-4pinnately<br />
compound leaves, articulated pedicels,<br />
5-12-merous flowers, imbricate aestivation,<br />
smooth seed surface, <strong>and</strong> smooth endosperm.<br />
Recent phylogenetic analyses have tested the<br />
<strong>Aralia</strong> generic concept, <strong>and</strong> major lineages or<br />
clades within <strong>Aralia</strong> have been delimited (Wen<br />
2001a, also see Wen 2002, 2004). Species <strong>of</strong> <strong>Aralia</strong><br />
are now classified into six sections (see Wen 2002,<br />
2004) primarily based on characters including<br />
habit, presence/absence <strong>of</strong> prickles, leaf<br />
architecture, trichome types, <strong>and</strong> presence/absence<br />
<strong>of</strong> bracts at the base <strong>of</strong> the inflorescence. These<br />
sections are: <strong>Aralia</strong> sect. <strong>Aralia</strong> (14 spp.), sect.<br />
Dimorphanthus (29 spp.), sect. Humiles Harms (3<br />
spp.), sect. Nanae Harms (1 sp.), sect. Pentapanax<br />
(Seem.) J. Wen (19 spp.), <strong>and</strong> sect. Sciadodendron<br />
5
6<br />
(Griseb.) J. Wen (5 spp.). Because <strong>of</strong> the large<br />
number <strong>of</strong> species involved, revisions <strong>of</strong> these<br />
sections have been published in three treatments.<br />
The present monograph <strong>of</strong> <strong>Aralia</strong> is the third <strong>and</strong><br />
the last <strong>of</strong> the series (see Wen 2002 for the revision<br />
<strong>of</strong> <strong>Aralia</strong> sect. Pentapanax; <strong>and</strong> Wen 2004 for the<br />
treatment <strong>of</strong> <strong>Aralia</strong> sect. Dimorphanthus).<br />
<strong>Aralia</strong> is widely distributed in different areas<br />
<strong>of</strong> Asia <strong>and</strong> the New World. Asia has relatively<br />
high species diversity for <strong>Aralia</strong> (57 <strong>of</strong> the 71<br />
species), whereas only 14 species occur in the New<br />
World. The 57 Asian species belong to three<br />
sections (sect. <strong>Aralia</strong>, sect. Dimorphanthus, <strong>and</strong><br />
sect. Pentapanax) with only sect. Pentapanax<br />
endemic to Asia; the other two sections are disjunct<br />
between Asia <strong>and</strong> North America. Despite the<br />
relatively low species diversity, the New World<br />
species <strong>of</strong> <strong>Aralia</strong> belong to five sections with three<br />
endemic to the New World (sect. Humiles, sect.<br />
Nanae, <strong>and</strong> sect. Sciadodendron) <strong>and</strong> two sections<br />
disjunct with Asia (sect. <strong>Aralia</strong> <strong>and</strong> sect.<br />
Dimorphanthus). The high species richness <strong>of</strong><br />
<strong>Aralia</strong> in Asia is largely due to the greater number<br />
<strong>of</strong> species in two sections: sects. <strong>Aralia</strong> (11 out <strong>of</strong><br />
14 in Asia) <strong>and</strong> Dimorphanthus (27 out <strong>of</strong> 29 in<br />
Asia).<br />
<strong>Aralia</strong> has a long taxonomic history (Wen <strong>and</strong><br />
Reveal 1992). It was first reported from the New<br />
World by Cornut in 1635. Several species were<br />
introduced into cultivation in Europe in the 1600’s<br />
<strong>and</strong> 1700’s. Tournefort (1700) coined the name<br />
for the genus from the old French-Canadian name<br />
“aralie” (Fernald 1950). Several species <strong>of</strong> <strong>Aralia</strong><br />
have been cultivated in various countries in Europe<br />
<strong>and</strong> Asia. Commonly cultivated <strong>Aralia</strong> species<br />
include: A. cordata Thunb., A. elata (Miq.) Seem.,<br />
A. cachemirica Decne., A. stipulata Franch., <strong>and</strong><br />
sometimes A. spinosa L., A. racemosa L., <strong>and</strong> A.<br />
californica S. Watson (also see Harms 1897;<br />
Rehder 1900; Wen 2002, 2004).<br />
Systematic studies <strong>of</strong> <strong>Aralia</strong> provide a muchneeded<br />
framework for further biogeographic<br />
studies, especially on the origin <strong>and</strong> development<br />
<strong>of</strong> the eastern Asian <strong>and</strong> eastern North American<br />
disjunct distribution, <strong>and</strong> the diversification <strong>of</strong> taxa<br />
within each <strong>of</strong> the two continents. Two sections<br />
<strong>of</strong> <strong>Aralia</strong> (sects. <strong>Aralia</strong> <strong>and</strong> Dimorphanthus) <strong>and</strong><br />
their close relative Panax (Wen et al. 2001) are<br />
disjunctly distributed in eastern Asia <strong>and</strong> eastern<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
North America (Wen 1991; Wen <strong>and</strong> Zimmer 1996;<br />
Wen et al. 1998; Wen 1999, 2000a, 2001a). The<br />
disjunct distributions <strong>of</strong> congeneric plants between<br />
eastern Asia <strong>and</strong> eastern North America have<br />
fascinated botanists since the Linnaean period (e.g.,<br />
Halenius 1750; Gray 1840, 1846, 1859, 1878; Hu<br />
1935, 1936; Hara 1952, 1956, 1972; Li 1952, 1972;<br />
Graham 1972; Boufford <strong>and</strong> Spongberg 1983; Wu<br />
1983; Tiffney 1985a, 1985b; Parks <strong>and</strong> Wendel<br />
1990; Wen 1998, 1999, 2001b). Many studies have<br />
documented the disjunct distributional patterns in<br />
various taxa. Phylogenetic analyses have now been<br />
conducted for many plant genera or sections (see<br />
review in Wen 1999, 2001b; Manos <strong>and</strong> Donoghue<br />
2001). Both molecular <strong>and</strong> fossil data suggest<br />
multiple origins <strong>of</strong> the disjunct pattern in the<br />
Tertiary (Tiffney 1985a, 1985b; Wen et al. 1996;<br />
Wen 1999; Xiang et al. 2000). Few studies,<br />
however, provide taxonomic treatments or<br />
monographs <strong>of</strong> genera that show this disjunct<br />
pattern.<br />
With the completion <strong>of</strong> the revision <strong>of</strong> <strong>Aralia</strong><br />
sect. Pentapanax (Wen 2002) <strong>and</strong> sect.<br />
Dimorphanthus (Wen 2004), the largest <strong>and</strong> most<br />
challenging yet-to-be monographed section is<br />
<strong>Aralia</strong> sect. <strong>Aralia</strong>. It consists <strong>of</strong> 14 species with<br />
three species in North America <strong>and</strong> eleven species<br />
in Asia, primarily in the Sino-Himalayan floristic<br />
region extending from eastern Russia southward<br />
to Taiwan, <strong>and</strong> from Japan westward to Kashmir,<br />
India <strong>and</strong> western Pakistan. Mainl<strong>and</strong> China has<br />
relatively high species diversity, harboring six <strong>of</strong><br />
the eleven Asian species (with A. cachemirica, A.<br />
taiwaniana, A. cordata, A. glabra <strong>and</strong> A. schmidtii<br />
excepted). <strong>Aralia</strong> cordata occurs throughout<br />
Japan, <strong>and</strong> A. taiwaniana is restricted to the<br />
mountains <strong>of</strong> central Taiwan. Species <strong>of</strong> the section<br />
form a monophyletic group with synapomorphies<br />
including ternately compound leaves.<br />
This treatment includes a comprehensive<br />
revision <strong>of</strong> four sections <strong>of</strong> <strong>Aralia</strong> (sects. <strong>Aralia</strong>,<br />
Humiles, Nanae <strong>and</strong> Sciadodendron), with detailed<br />
documentation <strong>of</strong> the morphology, ecology,<br />
distribution, <strong>and</strong> uses <strong>of</strong> the 23 species belonging<br />
to these sections, presentation <strong>of</strong> a multigene<br />
phylogeny <strong>of</strong> <strong>Aralia</strong>, a synopsis <strong>of</strong> the classification<br />
<strong>of</strong> the genus, <strong>and</strong> discussions <strong>of</strong> the group’s<br />
biogeographic diversification.
Generic concept <strong>and</strong> historical infrageneric<br />
classifications<br />
<strong>Aralia</strong> (<strong>Aralia</strong>ceae) consists <strong>of</strong> 71 species<br />
distributed in eastern <strong>and</strong> southeastern Asia <strong>and</strong><br />
the Americas. <strong>Aralia</strong> is defined as those species<br />
<strong>of</strong> <strong>Aralia</strong>ceae with pinnately to quadri-pinnately<br />
compound leaves, articulated petioles, 5-12merous<br />
flowers, imbricate petals, <strong>and</strong> articulated<br />
pedicels (as modified from Wen 1991, 1993).<br />
Previous workers provided various infrageneric<br />
classifications <strong>of</strong> <strong>Aralia</strong> (e.g., Persoon 1805;<br />
Sprengel 1825; Blume 1826; de C<strong>and</strong>olle 1830;<br />
Decaisne <strong>and</strong> Planchon 1854; Miquel 1863; Harms<br />
1896; Nakai 1927; Hoo <strong>and</strong> Tseng 1965).<br />
However, their systems are <strong>of</strong> limited value<br />
because <strong>of</strong> the problematic generic limits <strong>of</strong> <strong>Aralia</strong>,<br />
<strong>and</strong> the utilization <strong>of</strong> only one or two characters<br />
for infrageneric classification (see Wen 1993 for<br />
more detailed discussions). This current definition<br />
<strong>of</strong> <strong>Aralia</strong> recognizes the following<br />
synapomorphies: pinnately compound leaf<br />
architecture, presence <strong>of</strong> stipules, 5-12-locular<br />
ovaries, smooth seed surface, <strong>and</strong> flattened seeds.<br />
Its closest relative, the genus Panax possesses<br />
palmately compound leaves, absence <strong>of</strong> stipules,<br />
2-4-locular ovary, rough seed surfaces, <strong>and</strong> nonflattened<br />
seeds.<br />
Most early workers (e.g., Persoon 1805;<br />
Sprengel 1825; Blume 1826; de C<strong>and</strong>olle 1830)<br />
followed the broad Linnaean concept <strong>of</strong> <strong>Aralia</strong>,<br />
which stressed reproductive characters in defining<br />
the genus. It is now realized that floral characters<br />
are relatively conserved in <strong>Aralia</strong>ceae (Philipson<br />
1979; Wen et al. 2001) <strong>and</strong> the sole use <strong>of</strong> these<br />
characters is not effective in delimiting genera in<br />
the family. The Linnaean concept <strong>of</strong> <strong>Aralia</strong> had<br />
thus been ab<strong>and</strong>oned by later workers (e.g., Miquel<br />
1863; Seemann 1868; Li 1942; Hoo <strong>and</strong> Tseng<br />
1978; Philipson 1979; Shang 1985b; Wen 1993).<br />
Most previous systems <strong>of</strong> <strong>Aralia</strong> emphasized<br />
only one or two characters (e.g., Persoon 1805;<br />
Sprengel 1825; Blume 1826; de C<strong>and</strong>olle 1830;<br />
Decaisne <strong>and</strong> Planchon 1854; Nakai 1927; Hoo <strong>and</strong><br />
Tseng 1965). Because different workers stressed<br />
different characters, their systems are very<br />
different. These systems resulting from the<br />
overemphasis on one or two characters are<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
TAXONOMIC HISTORY<br />
apparently artificial <strong>and</strong> <strong>of</strong> limited predictive value.<br />
Harms (1896) constructed his classification <strong>of</strong><br />
<strong>Aralia</strong> based on several characters, but with an<br />
emphasis on inflorescence structure. The sections<br />
in Harms’ system appear more natural than those<br />
in previous systems. Harms followed Miquel’s<br />
(1863) generic concept <strong>of</strong> <strong>Aralia</strong>. Furthermore,<br />
Harms’ sect. Capituligerae was established based<br />
only on its capitate inflorescences. This character<br />
appears to have evolved at least twice in <strong>Aralia</strong>:<br />
once in the A. dasyphylla <strong>and</strong> A. urticifolia<br />
complex, <strong>and</strong> a second time in A. dasyphylloides.<br />
<strong>Aralia</strong> sect. Capituligerae is thus not<br />
monophyletic, <strong>and</strong> Harms’ system <strong>of</strong> the genus<br />
st<strong>and</strong>s in need <strong>of</strong> revision.<br />
Li <strong>and</strong> Xiang (1992) proposed a new<br />
classification <strong>of</strong> <strong>Aralia</strong>, in which two subgenera<br />
<strong>and</strong> eight sections were recognized. These workers<br />
circumscribed all herbaceous taxa in <strong>Aralia</strong> as<br />
constituting their new subgenus Paralia Shang &<br />
X. P. Li, <strong>and</strong> placed all woody, prickly members in<br />
subgenus <strong>Aralia</strong>. However, the treatment <strong>of</strong> the<br />
herbaceous members in a new subgenus is<br />
illegitimate because the type species <strong>of</strong> <strong>Aralia</strong>, A.<br />
racemosa, belongs to this group (cf. Wen et al.<br />
1998). Subgenus “<strong>Aralia</strong>” sensu Li <strong>and</strong> Xiang<br />
(1992) is equivalent to Miquel’s sect.<br />
Dimorphanthus. Li <strong>and</strong> Xiang (1992) proposed<br />
four new sections for <strong>Aralia</strong>: sects. Undulatae C.<br />
B. Shang & X. P. Li, Glaucae C. B. Shang & X. P.<br />
Li, Echinatae C. B. Shang & X. P. Li, <strong>and</strong><br />
Tomentosae C. B. Shang & X. P. Li. These new<br />
sections were largely based on presence or absence<br />
<strong>of</strong> leaflet pubescence, leaflet margin, presence or<br />
absence <strong>of</strong> prickles on petioles <strong>and</strong> inflorescence,<br />
<strong>and</strong> presence or absence <strong>of</strong> tomentose or strigose<br />
pubescence. Our phylogenetic analysis suggests<br />
that none <strong>of</strong> the sections recognized by Li <strong>and</strong><br />
Xiang is monophyletic (Wen 2004).<br />
Wang <strong>and</strong> Hu (2001) proposed another<br />
classification <strong>of</strong> <strong>Aralia</strong>, in which two subgenera<br />
<strong>and</strong> six sections were recognized. Four <strong>of</strong> the<br />
sections are herein treated as synonyms <strong>of</strong> sect.<br />
Dimorphanthus. Two new sections described in<br />
Wang <strong>and</strong> Hu (2001) are illegitimate because they<br />
were based on type species <strong>of</strong> previously described<br />
sections. The type <strong>of</strong> sect. Laxipaniculae Z. Z.<br />
Wang was designated to be A. spinifolia, which is<br />
7
8<br />
also the type <strong>of</strong> sect. Echinatae C. B. Shang & X.<br />
P. Li (Li <strong>and</strong> Xiang 1992). <strong>Aralia</strong> undulata is the<br />
type for sect. Undulatae C. B. Shang & X. P. Li as<br />
well as for sect. Glabrae Z. Z. Wang.<br />
Harms’ (1896) classification <strong>of</strong> <strong>Aralia</strong> has been<br />
most influential. Hoo <strong>and</strong> Tseng (1965, 1978)<br />
basically followed Harms (1896), but split Harms’<br />
sect. Arborescentes into two sections based on<br />
slight differences in inflorescence structure.<br />
Species <strong>of</strong> Harms’ sects. Anomalae <strong>and</strong><br />
Genuinae are similar: herbaceous, unarmed,<br />
branched, leaves tripinnate (on lower portion <strong>of</strong><br />
stem) to pinnate (on upper portion <strong>of</strong> stem), flowers<br />
small, <strong>and</strong> inflorescence terminal <strong>and</strong> axillary.<br />
Harms established these two sections based on<br />
slight differences in inflorescence pattern, i.e., sect.<br />
Anomalae with umbellate overall inflorescence<br />
structure <strong>and</strong> sect. Genuinae with paniculate<br />
overall structure. It is now realized that<br />
intermediates exist between Harms’ two “sections”.<br />
<strong>Aralia</strong> cordata (sect. Genuinae) sometimes has an<br />
umbellate structure, <strong>and</strong> A. fargesii (sect.<br />
Anomalae) sometimes demonstrates a paniculate<br />
pattern <strong>of</strong> inflorescence. Our phylogenetic analysis<br />
suggests that species <strong>of</strong> Anomalae <strong>and</strong> Genuinae<br />
constitute a monophyletic group. Harms’ sects.<br />
Anomalae <strong>and</strong> Genuinae are, therefore, combined<br />
here into one section, <strong>Aralia</strong>.<br />
Marchal (1879a) regarded his Coudenbergia<br />
(monotypic then) as a close relative <strong>of</strong> <strong>Aralia</strong>, but<br />
differing from the latter by the former’s 8-merous<br />
flowers (vs. 5-merous), the recurved oblong to<br />
linear anthers (vs. straight anthers), <strong>and</strong> concave<br />
floral disc with adnated edges (vs. conical to flat<br />
floral disc with free edges). These differences<br />
indeed hold true if Coudenbergia warmingiana is<br />
the sole member <strong>of</strong> the genus. However, there is<br />
little doubt that Coudenbergia angelicifolium <strong>and</strong><br />
C. ulei (now as the synonym <strong>of</strong> <strong>Aralia</strong><br />
warmingiana, which is the accepted name for<br />
Coudenbergia warmingiana) are very closely<br />
related <strong>and</strong> they should be treated as congeners.<br />
With the exp<strong>and</strong>ed concept <strong>of</strong> the Coudenbergia<br />
group, Coudenbergia <strong>and</strong> <strong>Aralia</strong> become hardly<br />
distinguishable. In terms <strong>of</strong> the number <strong>of</strong> floral<br />
parts, Coudenbergia angelicifolium, C. ulei <strong>and</strong> C.<br />
warmingiana are 5, 6, <strong>and</strong> 8-merous, respectively.<br />
Flowers <strong>of</strong> <strong>Aralia</strong> s. str. are 5-(6)-merous. Thus,<br />
based on floral parts, <strong>Aralia</strong> <strong>and</strong> Coudenbergia can<br />
not be well separated. The discovery <strong>of</strong> <strong>Aralia</strong><br />
weberbaueri Harms made the limits between<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
<strong>Aralia</strong> <strong>and</strong> Coudenbergia even more indistinct.<br />
The floral structure <strong>of</strong> Harms’ <strong>Aralia</strong> weberbaueri<br />
is very similar to that <strong>of</strong> the other <strong>Aralia</strong> species,<br />
with more or less distinct styles <strong>and</strong> 5merous floral<br />
parts. But it has a conical to flat floral disc <strong>and</strong> is<br />
vegetatively very similar to the South American<br />
Coudenbergia species. Even Harms himself (1917)<br />
was not certain about his placement <strong>of</strong> this species<br />
in <strong>Aralia</strong> or in Pentapanax (Harms merged<br />
Coudenbergia with Pentapanax in his worldwide<br />
monograph <strong>of</strong> <strong>Aralia</strong>ceae in 1898). He, thus,<br />
described it as <strong>Aralia</strong>? weberbaueri. In his original<br />
description, he presented the hypothesis that <strong>Aralia</strong><br />
weberbaueri was an intermediate between <strong>Aralia</strong><br />
<strong>and</strong> Pentapanax sensu Harms (as he treated<br />
Coudenbergia under the synonymy with<br />
Pentapanax). Because the difference between<br />
Coudenbergia <strong>and</strong> <strong>Aralia</strong> is minor <strong>and</strong> the<br />
intermediate forms are distinguishable, I treated<br />
Coudenbergia under synonymy with the Linnaean<br />
<strong>Aralia</strong> (Wen 1993). This treatment is now<br />
supported by phylogenetic data (Wen 2001a, 2002,<br />
2004, this study).<br />
Sciadodendron, a monotypic genus consisting<br />
<strong>of</strong> S. excelsum from Central America, the<br />
Caribbean to northern South America, was<br />
described as closely related to <strong>Aralia</strong>, but differing<br />
in its more numerous floral parts, more enlarged<br />
anthers, straight (vs. recurved) filaments, <strong>and</strong> nonarculated<br />
pedicels (Grisebach 1858; Gentry 1993;<br />
Wen 1993). Recent phylogenetic analysis using<br />
the ITS sequences <strong>of</strong> the nuclear ribosomal DNA<br />
(Wen 2001a) suggested that Sciadodendron is<br />
nested within <strong>Aralia</strong>. Wen (2002) formally merged<br />
it with <strong>Aralia</strong>.<br />
<strong>Aralia</strong> from the New World<br />
Linnaeus (1753) accounted for three <strong>Aralia</strong><br />
species from eastern North America: A. nudicaulis,<br />
A. racemosa <strong>and</strong> A. spinosa. The early history <strong>of</strong><br />
these species was discussed in detail by Wen <strong>and</strong><br />
Reveal (1992). Subsequently it was shown that<br />
they belong to three different sections. <strong>Aralia</strong><br />
nudicaulis <strong>and</strong> A. racemosa are treated in this<br />
monograph, <strong>and</strong> A. spinosa was treated in Wen<br />
(2004). <strong>Aralia</strong> californica <strong>and</strong> A. bicrenata were<br />
later reported from the western part <strong>of</strong> North<br />
Amerca <strong>and</strong> both species are members <strong>of</strong> sect.<br />
<strong>Aralia</strong> commonly compared with the eastern North<br />
American A. racemosa (Smith 1944).
The Sessé <strong>and</strong> Mociño expedition in 1787-1803<br />
to Mexico <strong>and</strong> nearby regions resulted in the<br />
publication <strong>of</strong> eight species <strong>of</strong> <strong>Aralia</strong>: A.<br />
chilapensis <strong>and</strong> A. pinnata published in Sessé <strong>and</strong><br />
Mociño (1888), <strong>and</strong> A. fruticosa, A. lobata, A.<br />
longifolia, A. ovata, A. tuxtlensis <strong>and</strong> A. sp. in their<br />
Flora Mexicana (Sessé <strong>and</strong> Mociño 1894). Only<br />
their A. chilapensis <strong>and</strong> A. pinnata truly belong to<br />
the genus <strong>Aralia</strong> with both herein treated as the<br />
synonyms <strong>of</strong> A. humilis. The other six species<br />
belong to Oreopanax <strong>and</strong> Dendropanax (see<br />
McVaugh 2000).<br />
Cavanilles (1797) described his <strong>Aralia</strong> humilis<br />
based on cultivated material from the Madrid<br />
Botanical Garden. In the original description, it<br />
was noted that it flowered every year in October<br />
<strong>and</strong> had fruits in 1792. McVaugh (2000) found no<br />
type material in the Sessé <strong>and</strong> Mociño Herbarium<br />
nor in the general Madrid herbarium. Most likely,<br />
the plant was grown from seeds collected in<br />
Mexico during the Sessé <strong>and</strong> Mociño Expedition.<br />
The other major expedition to Mexico at the time<br />
was the Malaspina Expedition, during which<br />
botanists Née <strong>and</strong> Haenke collected in western <strong>and</strong><br />
central Mexico in 1791. Apparently Cavanilles’<br />
<strong>Aralia</strong> humilis was planted in the Madrid Botanical<br />
Garden earlier than 1791.<br />
De C<strong>and</strong>olle’s <strong>Aralia</strong> pubescens was described<br />
based on material growing in the Botanical Garden<br />
in Montpellier, France (De C<strong>and</strong>olle 1813). Later<br />
in his Prodromus, De C<strong>and</strong>olle (1830) also<br />
included <strong>Aralia</strong> scabra Presl ex DC. as a synonym<br />
<strong>of</strong> A. pubescens. St<strong>and</strong>ley (1924) expressed<br />
uncertainty whether A. pubescens should be treated<br />
as a synonym <strong>of</strong> A. humilis. Smith (1944) defined<br />
A. humilis broadly <strong>and</strong> treated A. chilapensis, A.<br />
pinnata, A. brevifolia, A. pubescens <strong>and</strong> A. scabra<br />
as its synonyms.<br />
Grisebach (1858) described Sciadodendron as<br />
a new monotypic genus from Panama, consisting<br />
<strong>of</strong> S. excelsum. It was regarded as a close relative<br />
<strong>of</strong> <strong>Aralia</strong>, but was differentiated from the latter by<br />
Sciadodendron’s 10-12-locular ovaries. Donnell<br />
Smith (1910) reported Reynoldsia americana from<br />
Peninsula Nicoya <strong>of</strong> Costa Rica. Sciadodendron<br />
excelsum <strong>and</strong> Reynoldsia americana were shown<br />
to be identical along with Rusby’s (1920)<br />
Pentapanax granatensis (Harms 1928; Smith<br />
1936). Wen (2002) merged Sciadodendron with<br />
<strong>Aralia</strong> <strong>and</strong> made the nomenclatural combination.<br />
Marchal (1879a) described three species <strong>of</strong><br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
<strong>Aralia</strong>: A. regeliana <strong>and</strong> A. brevifolia from Mexico<br />
<strong>and</strong> A. soratensis from Bolivia. He also reported<br />
Coemansia warmingiana (= <strong>Aralia</strong> warmingiana)<br />
as a new species from Minas Gerais <strong>of</strong> Brazil.<br />
Harms (1908) described Pentapanax ulei from<br />
Bahia, Brazil. This name was placed as a synonym<br />
<strong>of</strong> <strong>Aralia</strong> warmingiana by Wen (1993). Harms<br />
(1918) described <strong>Aralia</strong>? weberbaueri from Peru<br />
as a species very similar to <strong>Aralia</strong> soratensis. He<br />
also compared his new species with Pentapanax<br />
angelicifolius Griseb. from Argentina. He<br />
suggested that A. soratensis <strong>and</strong> A. weberbaueri<br />
were intermediates between <strong>Aralia</strong> <strong>and</strong><br />
Pentapanax. Wen (1993) placed A. weberbaueri<br />
<strong>and</strong> Pentapanax angelicifolius as synonyms <strong>of</strong> A.<br />
soratensis, which has nomenclatural priority.<br />
Harms (1924) also reported Ekman’s Megalopanax<br />
rex Ekman ex Harms, which was recently merged<br />
with <strong>Aralia</strong> (Wen 1993).<br />
St<strong>and</strong>ley (1924) recognized five species <strong>of</strong><br />
<strong>Aralia</strong> from Mexico: A. scopulorum, A. regeliana,<br />
A. humilis, A. pubescens, <strong>and</strong> A. racemosa. His A.<br />
racemosa clearly represented A. bicrenata. He<br />
differentiated A. pubescens from A. humilis by the<br />
pubescence on the pedicels. He defined A. humilis<br />
as possessing glabrous pedicels.<br />
Smith (1944) enumerated the North American<br />
species <strong>of</strong> <strong>Aralia</strong>ceae <strong>and</strong> recognized the following<br />
eight species <strong>of</strong> <strong>Aralia</strong> from North America: A.<br />
racemosa, A. californica, A. nudicaulis, A. spinosa,<br />
A. hispida, A. humilis, A. regeliana <strong>and</strong> A.<br />
scopulorum. He also treated Sciadodendron <strong>and</strong><br />
Megalopanax as distinct genera from <strong>Aralia</strong><br />
because they have more numerous floral parts (7-<br />
12 vs 4-6). He distinguished Sciadodendron from<br />
Megalopanax by the former’s nonarticulated<br />
pedicels <strong>and</strong> slightly divided styles (vs. articulated<br />
pedicels <strong>and</strong> firmly connate styles. <strong>Aralia</strong><br />
bicrenata was placed as a synonym <strong>of</strong> A. racemosa.<br />
<strong>Aralia</strong> from the Old World<br />
Species <strong>of</strong> <strong>Aralia</strong> from the Old World belong<br />
to three sections: <strong>Aralia</strong>, Dimorphanthus <strong>and</strong><br />
Pentapanax. The latter two sections have been<br />
recently monographed <strong>and</strong> the taxonomic history<br />
<strong>of</strong> the species has been reviewed by Wen (2002,<br />
2004). For regional interests, the readers are<br />
referred to Miquel (1840, 1856a, 1856b, 1857,<br />
1863), Ridley (1922), Blume (1826), Blanco<br />
(1877), van Steenis (1948), Backer <strong>and</strong> Bakhuizen<br />
9
10<br />
van den Brink (1965), <strong>and</strong> Philipson (1951, 1977,<br />
1979) for southeast Asia; Li (1942), Ho (1952),<br />
Hoo <strong>and</strong> Tseng (1965, 1978), Ling (1977, 1987),<br />
Shang (1985a, 1985b, Li <strong>and</strong> Xiang (1992), Huang<br />
(1993), Wen (1994), <strong>and</strong> Wang <strong>and</strong> Hu (2001) for<br />
China; Nakai (1927) <strong>and</strong> Lee (1993) for Korea;<br />
Ohwi (1984) for Japan; Wallich (1831-32), Don<br />
(1834), Clarke (1879), Mizushima (1966), <strong>and</strong> Wen<br />
et al. (2002) for the Himalayan region; <strong>and</strong> Bui<br />
(1964) <strong>and</strong> Ha (1974) for Indochina. The first<br />
Asian <strong>Aralia</strong> species was described by Linnaeus<br />
(1753) based on collections made by P. Osbeck<br />
near Canton, China (Bretschneider 1898; Hansen<br />
<strong>and</strong> Moule 1973), which was <strong>Aralia</strong> chinensis <strong>of</strong><br />
sect. Dimorphanthus. Below I provide only a brief<br />
history <strong>of</strong> the taxonomic work <strong>of</strong> <strong>Aralia</strong> sect. <strong>Aralia</strong><br />
in the Old World.<br />
Thunberg (1784) described <strong>Aralia</strong> cordata <strong>of</strong><br />
Japan, the first Asian species <strong>of</strong> sect. <strong>Aralia</strong>. The<br />
same species was later described as A. edulis <strong>and</strong><br />
A. nutans by Siebold <strong>and</strong> Zuccarini (1837) <strong>and</strong><br />
Franchet <strong>and</strong> Savatier (1878), respectively.<br />
Although <strong>Aralia</strong> cordata was mostly regarded as<br />
an endemic species to Japan earlier in its history<br />
(Thunberg 1784; Persoon 1805; De C<strong>and</strong>olle 1830;<br />
Don 1834; Miquel 1863; Franchet <strong>and</strong> Savatier<br />
1875), a few workers thought that it also occurred<br />
in continental eastern Asia (e.g., Harms 1896;<br />
Nakai 1909; Li 1942). Kitagawa (1935) published<br />
<strong>Aralia</strong> continentalis based on specimens from<br />
Manchuria <strong>of</strong> northeastern China as well as Korea.<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
He noted differences between A. cordata <strong>and</strong> A.<br />
continentalis in overall inflorescence architecture,<br />
pediclel length, thickness <strong>and</strong> pubescence, <strong>and</strong> the<br />
connation <strong>of</strong> styles. The separation <strong>of</strong> the two taxa<br />
was first proposed in Russian literature according<br />
to Pojarkova (1973) <strong>and</strong> was accepted by a few<br />
later workers (e.g., Pojarkova 1950, 1973;<br />
Kitagawa 1979; Lee 1993). Hoo <strong>and</strong> Tseng (1978)<br />
somehow recognized both A. continentalis <strong>and</strong> A.<br />
cordata from China.<br />
Matsumura (1899) reported <strong>Aralia</strong> glabra, the<br />
second recognized species <strong>of</strong> sect. <strong>Aralia</strong> from<br />
Japan based on collections made near Nikko.<br />
Harms (1896) described A. henryi from Central<br />
China with the specimen A. Henry 6655 as the type.<br />
Franchet (1896) reported his A. pilosa from Central<br />
China, also citing A. Henry 6655 as one <strong>of</strong> the two<br />
syntypes. Franchet’s name appeared one day later<br />
than Harms’ (September 16 vs. 15, 1896) <strong>and</strong> the<br />
species is thus recognized as A. henryi. In the same<br />
paper, Franchet (1896) described two additional<br />
species <strong>of</strong> sect. <strong>Aralia</strong>: A. atropurpurea <strong>and</strong> A.<br />
fargesii from China.<br />
Hoo <strong>and</strong> Tseng (1965) described A. tibetana<br />
from Tibet, China. The species was later recorded<br />
to occur also in India <strong>and</strong> Nepal (Wen et al. 2002).<br />
Liu <strong>and</strong> Lu (1976) described A. taiwaniana from<br />
Central Taiwan. Other Asian taxa <strong>of</strong> the section<br />
include A. schmidtii (Pojarkova 1950) <strong>and</strong> A.<br />
cachemirica from the Himalaya (Decaisne 1844).<br />
PHYLOGENETIC RELATIONSHIPS<br />
A phylogenetic analysis is herein presented to<br />
help identify <strong>and</strong> test main evolutionary lineages<br />
within <strong>Aralia</strong>. The sampling includes 48 species<br />
<strong>of</strong> <strong>Aralia</strong>ceae, representing all previously<br />
recognized sections <strong>of</strong> <strong>Aralia</strong>, its close relative<br />
Panax, <strong>and</strong> representatives <strong>of</strong> the Polyscias clade.<br />
The Polyscias clade was hypothesized to be closely<br />
related to the <strong>Aralia</strong>-Panax clade. Five species <strong>of</strong><br />
the Asian core <strong>Aralia</strong>ceae clade (see Wen et al.<br />
2001) were included as outgroups. A detailed<br />
phylogenetic analysis <strong>of</strong> the <strong>Aralia</strong>-Panax clade<br />
will be published in a separate paper (J. Wen, in<br />
prep.). Herein I outline only the main findings.<br />
Phylogenetic analyses were based on four<br />
molecular markers: the internal transcribed spacer<br />
(ITS) regions <strong>of</strong> nuclear ribosomal DNA <strong>and</strong> three<br />
chloroplast markers including the ndhF gene, the<br />
trnL-F region, <strong>and</strong> the atpB-rbcL spacer. The<br />
analyses were performed with PAUP* (version 4.0,<br />
Sw<strong>of</strong>ford 2003) using heuristic searches with<br />
MULPARS <strong>and</strong> furthest addition sequence options.<br />
Clade support for monophyletic groups revealed<br />
in the maximally parsimonious tree(s) (MPTs) was<br />
examined with 500 bootstrap replicates<br />
(Felsenstein 1985) with the r<strong>and</strong>om addition <strong>and</strong><br />
the heuristic search options using parsimony.
The parsimony analysis generated 816 MPTs<br />
with a total length <strong>of</strong> 1389 steps, a consistency<br />
index <strong>of</strong> 0.69 (0.54 excluding uninformative<br />
characters), <strong>and</strong> a retention index <strong>of</strong> 0.73. The strict<br />
consensus tree with bootstrap support <strong>and</strong> the 50%<br />
majority-rule consensus tree are presented in Figs.<br />
1 <strong>and</strong> 2, respectively.<br />
The phylogenetic analysis supports a close<br />
relationship between <strong>Aralia</strong> <strong>and</strong> Panax with 100%<br />
bootstrap support. Within <strong>Aralia</strong>, six major<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
11<br />
lineages are recognizable, <strong>and</strong> these six groups<br />
corresponding to the currently recognized sections<br />
<strong>of</strong> the genus (Figs. 1 <strong>and</strong> 2). The monotypic sect.<br />
Nanae (A. nudicaulis <strong>of</strong> North America) is herein<br />
suggested to be most closely related to the New<br />
World endemic sect. Humiles. Xiang <strong>and</strong> Li (1990)<br />
merged <strong>Aralia</strong> scopulorum <strong>of</strong> New World sect.<br />
Humiles with the Asian sect. Pentapanax, <strong>and</strong> this<br />
analysis does not support the monophyly <strong>of</strong> the<br />
sect. Pentapanax – sect. Humiles group (Figs. 1-<br />
Fig. 1. Strict consensus tree <strong>of</strong> maximum parsimony analysis <strong>of</strong> <strong>Aralia</strong> <strong>and</strong> its close relatives based on sequences <strong>of</strong> the internal<br />
transcribed spacer (ITS) regions <strong>of</strong> nuclear ribosomal DNA <strong>and</strong> three chloroplast markers including the ndhF gene, the trnL-<br />
F region, <strong>and</strong> the atpB-rbcL spacer (816 maximally parsimonious trees, a total length <strong>of</strong> 1389 steps, a consistency index <strong>of</strong><br />
0.69, <strong>and</strong> a retention index <strong>of</strong> 0.73). The bootstrap support <strong>of</strong> 500 replicates is indicated above the clades.
12<br />
2). Nevertheless the relationships among the six<br />
sections <strong>of</strong> the genus are still poorly resolved.<br />
Additional molecular markers will be employed<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
to resolve the relationships among the major<br />
lineages.<br />
Fig. 2. The 50% majority-rule consensus tree <strong>of</strong> the parsimony analysis <strong>of</strong> <strong>Aralia</strong> <strong>and</strong> its close relatives based on sequences <strong>of</strong><br />
the internal transcribed spacer (ITS) regions <strong>of</strong> nuclear ribosomal DNA <strong>and</strong> three chloroplast markers including the ndhF<br />
gene, the trnL-F region, <strong>and</strong> the atpB-rbcL spacer (816 maximally parsimonious trees, a total length <strong>of</strong> 1389 steps, a consistency<br />
index <strong>of</strong> 0.69, <strong>and</strong> a retention index <strong>of</strong> 0.73).
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
MORPHOLOGICAL CHARACTERS<br />
This section is intended to discuss the<br />
variation <strong>of</strong> the characters used to delimit taxa; <strong>and</strong><br />
to define the characters used in the descriptions<br />
<strong>and</strong> keys.<br />
Habit<br />
<strong>Aralia</strong> species are perennial herbs, shrubs or<br />
trees. Species <strong>of</strong> <strong>Aralia</strong> sect. <strong>Aralia</strong> are perennial<br />
herbs. <strong>Aralia</strong> nudicaulis <strong>of</strong> sect. Nanae has been<br />
described as being herbaceous, yet it has an<br />
extensive underground rhizome system <strong>and</strong> is thus<br />
essentially woody. Members <strong>of</strong> <strong>Aralia</strong> sect.<br />
Dimorphanthus consist <strong>of</strong> shrubs, treelets or trees,<br />
occasionally woody climbers, or semishrubs (i.e.,<br />
herbaceous with a woody base). <strong>Aralia</strong> sect.<br />
Pentapanax <strong>of</strong>ten has epiphytic members. Species<br />
<strong>of</strong> sect. Sciadodendron are <strong>of</strong>ten trees or sometimes<br />
large shrubs, whereas members <strong>of</strong> sect. Humiles<br />
are <strong>of</strong>ten shrubs to sometimes small trees.<br />
Underground systems<br />
<strong>Aralia</strong> nudicaulis (sect. Nanae) is highly clonal<br />
with branched horizontal rhizomes. Most species<br />
<strong>of</strong> sect. <strong>Aralia</strong> have rhizomes <strong>and</strong> well-developed<br />
roots. Some species <strong>of</strong> <strong>Aralia</strong> sect. Dimorphanthus<br />
have been observed to form large clones (e.g.,<br />
<strong>Aralia</strong> elata, A. hispida, A. spinosa, <strong>and</strong> A.<br />
stipulata), <strong>and</strong> it is presumed that they possess<br />
rhizomes. This character has not been used<br />
taxonomically, because most species have not been<br />
well documented for their underground features.<br />
Prickles<br />
All members <strong>of</strong> <strong>Aralia</strong> sect. Dimorphanthus<br />
bear prickles on their stems. The presence <strong>of</strong><br />
prickles is herein regarded as a synapomorphy <strong>of</strong><br />
the section (Wen 2004). Taxa <strong>of</strong> other sections are<br />
unarmed.<br />
Leaf architecture <strong>and</strong> morphology<br />
Leaf architecture is considered to be<br />
taxonomically useful in <strong>Aralia</strong>. <strong>Aralia</strong> sect. <strong>Aralia</strong><br />
<strong>and</strong> sect. Nanae have ternately compound leaves.<br />
Leaves <strong>of</strong> sect. Dimorphanthus have mostly<br />
13<br />
bipinnate, but sometimes 3- or rarely 4-pinnate<br />
leaves. <strong>Aralia</strong> sect. Sciadodendron usually has 3or<br />
4-pinnate leaves. <strong>Aralia</strong> sect. Humiles <strong>and</strong> sect.<br />
Pentapanax usually have pinnate to bipinnate<br />
(rarely 3-pinnate) leaves. Taxa with 2-4-pinnate<br />
leaves usually have a pair <strong>of</strong> accessory leaflets or<br />
pinnae at each division <strong>of</strong> the rachis (see Fig. 1 in<br />
Wen 2004).<br />
Trichomes <strong>and</strong> vesture<br />
Vesture is taxonomically useful in <strong>Aralia</strong>. The<br />
terminology used in this revision follows Lawrence<br />
(1951), Harrington <strong>and</strong> Durrell (1957), <strong>and</strong> Stearn<br />
(1983), <strong>and</strong> is defined as below (examples refer to<br />
the abaxial leaflet surface if not specified<br />
otherwise): ciliate, margins with s<strong>of</strong>t hairs; pilose,<br />
with sparse slender s<strong>of</strong>t hairs; pubescent, with short<br />
s<strong>of</strong>t straight hairs (e.g., in A. thomsonii); scabrous,<br />
with short, coarse, stiff white hairs, usually with a<br />
slightly swollen base, rough to touch (e.g., on the<br />
adaxial leaflet surface <strong>of</strong> A. atropurpurea);<br />
tomentose, with dense, more or less interwoven,<br />
short s<strong>of</strong>t hairs (e.g., A. dasyphylloides). Vesture<br />
is usually regarded as useful in differentiating<br />
species, but less so phylogenetically because <strong>of</strong><br />
uncertainty in establishing transformation series.<br />
The trichome types in <strong>Aralia</strong> may be roughly<br />
grouped into four types: (1) branched (Fig. 3A,<br />
e.g., on lower leaf surface <strong>of</strong> A. humilis); (2) long<br />
<strong>and</strong> s<strong>of</strong>t (Fig. 3B, e.g., in A. nudicaulis); (3)<br />
biseriate (Fig. 3C, e.g., in A. racemosa, <strong>and</strong> A.<br />
cordata); <strong>and</strong> (4) multiseriate (Fig. 3D, e.g., in A.<br />
elata). Also see Figs. 4-8 for detailed illustrations<br />
<strong>of</strong> trichome morphology <strong>of</strong> <strong>Aralia</strong> apioides, A.<br />
atropurpurea, A. cordata, A. racemosa, <strong>and</strong> A.<br />
humilis.<br />
Cuticles<br />
Two main cuticular patterns were reported on<br />
the abaxial leaflet surface in members <strong>of</strong> sect.<br />
Dimorphanthus: (1) striate, <strong>and</strong> (2) coronulate (see<br />
Wen 2004). This terminology follows Hardin <strong>and</strong><br />
Beckmann (1982). The coronulate pattern is found<br />
only in <strong>Aralia</strong> sect. Dimorphanthus, including in<br />
the following species: A. bipinnata, A.<br />
dasyphylloides, A. debilis, A. decaisneana, A. elata,
14<br />
A. gintungensis, A. <strong>of</strong>ficinalis, A. spinosa, A.<br />
stipulata, <strong>and</strong> A. undulata. The abaxial surface <strong>of</strong><br />
all other <strong>Aralia</strong> species is striate, suggesting the<br />
coronulate pattern as the derived character state in<br />
the genus. The adaxial leaflet surface has the striate<br />
cuticle pattern throughout the genus <strong>Aralia</strong> (Figs.<br />
9-10).<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
Pedicel articulation<br />
Pedicel articulation has been regarded as an<br />
important taxonomic character in <strong>Aralia</strong>ceae. In<br />
<strong>Aralia</strong>, there is a secondary loss <strong>of</strong> pedicel<br />
articulation in A. excelsa <strong>and</strong> its close relative, A.<br />
bahiana, a new taxon reported in this study.<br />
Fig. 3. Trichome types on lower leaflet surface in <strong>Aralia</strong>: A. Branched (A. humilis). B. Long <strong>and</strong> s<strong>of</strong>t (A. nudicaulis). C.<br />
Biseriate (A. racemosa). D. Multiseriate (A. elata).
Inflorescence<br />
The presence or absence <strong>of</strong> a main<br />
inflorescence axis was in the past considered to be<br />
important taxonomically in <strong>Aralia</strong> (Harms 1896;<br />
Hoo <strong>and</strong> Tseng 1965). Harms (1896) used this<br />
character to distinguish two sections: sect.<br />
Genuinae (= sect. <strong>Aralia</strong>) with a well-developed<br />
main axis, <strong>and</strong> sect. Anomalae without a main axis.<br />
Hoo <strong>and</strong> Tseng (1965) employed this character to<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
Fig. 4. Epidermal characters <strong>of</strong> <strong>Aralia</strong> apioides: A-C. Trichomes on pedicels. D. Trichome on lower leaflet surface. E.<br />
Trichome on upper leaflet surface. F. Stomates on lower leaflet surface (Wen et al. 1241, US).<br />
15<br />
differentiate sect. Arborescentes Harms<br />
(synonymous with sect. Dimorphanthus,<br />
characterized by having a main axis) from sect.<br />
Digitatipanicula Hoo (without a main axis). In<br />
contrast to Harms (1896) <strong>and</strong> Hoo <strong>and</strong> Tseng<br />
(1965), I regard the presence or absence <strong>of</strong> a main<br />
inflorescence axis as significant only at the species<br />
level, <strong>and</strong> so it is not used here to delimit sections<br />
(also see Wen 2004). Several species (e.g., <strong>Aralia</strong><br />
fargesii) show an intermediate inflorescence
16<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
Fig. 5. Epidermal characters <strong>of</strong> <strong>Aralia</strong> atropurpurea: A&B. Trichome on pedicels. C&D. Trichomes on upper leaflet surface.<br />
E&F. Trichome on lower leaflet surface (Wen et al. 926, US).
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
Fig. 6. Epidermal characters <strong>of</strong> <strong>Aralia</strong> taiwaniana: A&B. Trichomes on pedicels. C. Trichomes on lower leaflet vein. D.<br />
Trichome <strong>and</strong> cuticle on upper leaflet surface. E. Trichome on lower leaflet surface. F. Trichomes <strong>and</strong> stomates on lower<br />
leaflet surface (Wen 9424, US).<br />
17
18<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
Fig. 7. Epidermal characters <strong>of</strong> <strong>Aralia</strong> racemosa: A&B. Trichomes on leaf petiole. C&D. Trichome on upper leaflet surface.<br />
E&F. Trichomes, cuticle <strong>and</strong> stomates on lower leaflet surface (Wen 9697, US).
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
Fig. 8. Epidermal characters <strong>of</strong> <strong>Aralia</strong> humilis: A. Trichomes on lower leaflet surface. B-D. Trichomes on upper leaflet surface.<br />
E. Cuticle on upper leaflet surface, noting the presence <strong>of</strong> stomates. F. Cuticle <strong>and</strong> stomates on lower leaflet surface.<br />
19
20<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
Fig. 9. Variation <strong>of</strong> striated type <strong>of</strong> leaf cuticles: A. <strong>Aralia</strong> soratensis, upper surface. B. A. soratensis, lower surface. C. A.<br />
racemosa, upper surface. D. A. racemosa, lower surface midvein with trichomes.
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
Fig. 10. Variation <strong>of</strong> striated type <strong>of</strong> leaf cuticles: A. <strong>Aralia</strong> humilis, upper surface. B. A. humilis, lower surface. C. A. nudicaulis,<br />
lower surface. D. A. nudicaulis, upper surface.<br />
21
22<br />
architecture between the two extreme conditions.<br />
Bracts <strong>and</strong> bracteoles<br />
The size, shape, pubescence <strong>and</strong> persistence <strong>of</strong><br />
bracts <strong>and</strong> bracteoles in the inflorescence are useful<br />
for delimiting taxa in <strong>Aralia</strong>.<br />
Flowers<br />
Floral morphology is relatively constant in<br />
<strong>Aralia</strong> as well as it is throughout <strong>Aralia</strong>ceae (Eyde<br />
<strong>and</strong> Tseng 1971; Wen 2004). Most species <strong>of</strong><br />
<strong>Aralia</strong> possess 5 minute triangular to rounded<br />
sepals, 5 petals that are ovate or nearly so, 5<br />
stamens, 5-locular ovaries, <strong>and</strong> styles that are<br />
distinct or connate at the base. <strong>Aralia</strong> sect.<br />
Sciadodendron shows more variation in floral<br />
parts, ranging from having 5-merous to 10- or even<br />
12-merous flowers. <strong>Aralia</strong> nudicaulis <strong>and</strong> A.<br />
scopulorum occasionally have 6-merous flowers<br />
with the predominant condition <strong>of</strong> being 5-merous.<br />
The condition with more than five petals appears<br />
to represent secondary increase. In <strong>Aralia</strong> excelsa<br />
<strong>of</strong> sect. Sciadodendron, the author observed the<br />
later developmental split <strong>of</strong> 4-5 petals into 8-10<br />
separate petal units.<br />
<strong>Aralia</strong> L. — Pollen mostly subprolate in<br />
equatorial view, triangular to rounded triangular<br />
in polar view; 3-colporate, the colpal margins<br />
incurved, sometimes with a colpal ridge, costa<br />
ectocolpi variable, from poorly developed (A.<br />
californica, A. humilis) to more prominent (A.<br />
leschenaultii, A. nudicaulis); the endoaperture<br />
mostly lalongate, frequently with diffuse lateral<br />
margins (A. hispida excepted), costa endocolpi<br />
variable, small but sharply delimited (A.<br />
californica) to more prominent (A. leschenaultii);<br />
the tectum variable, complete, <strong>and</strong> punctate,<br />
incomplete, <strong>and</strong> finely reticulate with perforations<br />
larger at the poles (A. spinosa <strong>and</strong> other taxa in<br />
sect. Dimorphanthus), incomplete <strong>and</strong> perforate to<br />
irregularly perforate, or weakly striato-reticulate<br />
(most species in sects. <strong>Aralia</strong>, Humiles <strong>and</strong><br />
Pentapanax). In TEM, the nonapertural endexine<br />
thin; the apertural endexine thickened at the colpi,<br />
sometimes (partially) filling the arch <strong>of</strong> the<br />
ectexine, sometimes with a gap or marked thinning<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
POLLEN MORPHOLOGY<br />
Filament length shows interspecific variation<br />
in most cases. In the dioecious <strong>Aralia</strong> nudicaulis,<br />
the male flowers have significantly longer<br />
filaments than the female flowers.<br />
The division <strong>of</strong> styles is useful to differentiate<br />
taxa. Species in <strong>Aralia</strong> have completely free or<br />
distinct styles (e.g., in A. nudicaulis) to nearly<br />
completely united styles (e.g., in A. excelsa).<br />
Shape <strong>of</strong> the floral disc may prove to be<br />
taxonomically <strong>and</strong>/or phylogenetically significant.<br />
In sect. <strong>Aralia</strong>, A. atropurpurea has a projected<br />
floral disk, whereas A. californica has a flat floral<br />
disk. In sect. Dimorphanthus, some species such<br />
as A. armata, A. bipinnata, A. chinensis, A.<br />
dasyphylloides, A. malabarica, A. searelliana <strong>and</strong><br />
A. spinifolia have projected floral discs, whereas<br />
in other species (such as A. foliolosa), they are flat.<br />
This character may be important for differentiating<br />
closely related species, but it clearly varies within<br />
sections.<br />
Fruits<br />
Fruit size <strong>and</strong> shape may be used to differentiate<br />
species. Most <strong>Aralia</strong> species have dark purple to<br />
blackish fruits. <strong>Aralia</strong> henryi is the only taxon <strong>of</strong><br />
the genus possessing bright red fruits.<br />
at the boundary <strong>of</strong> nonapertural <strong>and</strong> apertural<br />
endexines (A. californica; A. hispida, A.<br />
leschenaultii); the foot layer thin to thick; the<br />
columellae very short, sometimes elongated at the<br />
poles (conspicuously so in sect. Dimorphanthus);<br />
the tectum variable, continuous, or nearly so to<br />
more discontinuous. Many <strong>Aralia</strong> species<br />
examined have a weakly striato-reticulate tectum<br />
(sects. <strong>Aralia</strong>, Humiles, <strong>and</strong> Pentapanax). <strong>Aralia</strong><br />
nudicaulis is a morphologically very distinct<br />
species within <strong>Aralia</strong> <strong>and</strong> is the sole member <strong>of</strong><br />
sect. Nanae (Harms 1896; this study). Its very thin<br />
exine may represent an extreme reduction. On the<br />
other h<strong>and</strong>, pollen <strong>of</strong> <strong>Aralia</strong> sect. Dimorphanthus<br />
is thicker-walled <strong>and</strong> has elongated columellae at<br />
the poles (cf. Wen <strong>and</strong> Nowicke 1999). Taxa <strong>of</strong><br />
sect. Dimorphanthus also have larger tectal<br />
perforations at the poles (see Wen <strong>and</strong> Nowicke<br />
1999; Wen 2004). See Figs. 11-15 for pollen<br />
characteristics <strong>of</strong> <strong>Aralia</strong>.
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
Fig. 11. Pollen morphology <strong>of</strong> <strong>Aralia</strong>. A-D. A. nudicaulis (Moyer s.n., Pennsylvania, US). E-H. A. californica (Robbins 2066,<br />
US)<br />
23
24<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
Fig. 12. Pollen morphology <strong>of</strong> <strong>Aralia</strong>. A&B. A. soratensis (Venturi 1081, US). C-E. A. bicrenata (Nelson 6141, US).
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
Fig. 13. Pollen morphology <strong>of</strong> <strong>Aralia</strong>. A-C. A. warmingiana (Smith & Klein 14120, US). D-G. A. humilis (McGill & Lehto<br />
L20373, US).<br />
25
26<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
Fig. 14. Pollen morphology <strong>of</strong> <strong>Aralia</strong>. A-D. A. regeliana (McVaugh 10348, US).<br />
I herein provide an update on the chromosomal<br />
evolution in <strong>Aralia</strong> sect. <strong>Aralia</strong>, <strong>and</strong> sect. Nanae.<br />
The base chromosome number <strong>of</strong> <strong>Aralia</strong> has been<br />
suggested to be x = 12 (Wen 2002). The readers<br />
are also referred to Yi et al. (2004) for more<br />
information on chromosomal evolution in<br />
CHROMOSOME NUMBERS<br />
<strong>Aralia</strong>ceae, <strong>and</strong> to Wen (2004) for an overview <strong>of</strong><br />
chromosome numbers in <strong>Aralia</strong>. Both diploids <strong>and</strong><br />
tetraploids have been reported from sect. <strong>Aralia</strong><br />
<strong>and</strong> sect. Nanae (see Table 1). To date all counts<br />
for members <strong>of</strong> sect. Dimorphanthus (<strong>Aralia</strong> elata<br />
var. elata, A. elata var. m<strong>and</strong>shurica, A. hispida,
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
Fig. 15. Pollen morphology <strong>of</strong> <strong>Aralia</strong>. A-F. A. scopulorum (Purpus s.n., Jan-Mar 1898, US).<br />
<strong>and</strong> A. spinosa) <strong>and</strong> sect. Pentapanax (A. franchetii,<br />
A. leschenaultii, <strong>and</strong> A. parasitica) have been 2n<br />
= 24. Chromosomal numbers <strong>of</strong> sect. Humiles <strong>and</strong><br />
27<br />
sect. Sciadodendron have not been reported at<br />
present.
28<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
INFRAGENERIC CLASSIFICATION<br />
Based primarily on the results from the<br />
phylogenetic analyses <strong>and</strong> our current<br />
underst<strong>and</strong>ing <strong>of</strong> the morphological variation in<br />
<strong>Aralia</strong>, a new classification <strong>of</strong> the genus is herein<br />
formally described with six sections recognized.<br />
I. <strong>Aralia</strong> sect. <strong>Aralia</strong><br />
<strong>Aralia</strong> sect. Anomalae Harms, Bot. Jahrb. 23: 12,<br />
1896.<br />
<strong>Aralia</strong> sect. Genuinae Harms, Bot. Jahrb. 23: 15,<br />
1896.<br />
Branched herbs with rhizomes. Leaves<br />
stipulate, bi or tripinnate on lower portion <strong>of</strong> stem,<br />
pinnate to bipinnate on upper portion <strong>of</strong> stem,<br />
trichomes simple. Umbels aggregated into<br />
umbellate or paniculate structure, <strong>and</strong><br />
inflorescence usually robust, terminal <strong>and</strong> axillary.<br />
Flowers small (1-2 mm in diameter), 5-merous.<br />
Fruits dark purple or occasionally red.<br />
Number <strong>of</strong> species: 14.<br />
Type species: A. racemosa L.<br />
Distribution: eastern Asia, the Himalayas,<br />
eastern North America, <strong>and</strong> western North<br />
America.<br />
List <strong>and</strong> distribution <strong>of</strong> species in <strong>Aralia</strong> sect.<br />
<strong>Aralia</strong><br />
1. <strong>Aralia</strong> californica S. Watson<br />
California & Oregon<br />
2. <strong>Aralia</strong> bicrenata Wooton & St<strong>and</strong>l.<br />
SW US & N Mexico<br />
3. <strong>Aralia</strong> racemosa L.<br />
E N America<br />
4. <strong>Aralia</strong> cordata Thunb.<br />
Japan & Korea<br />
5. <strong>Aralia</strong> schmidtii Pojark.<br />
E Russia<br />
6. <strong>Aralia</strong> taiwaniana Y. C. Liu & F. Y. Lu<br />
Taiwan<br />
7. <strong>Aralia</strong> continentalis Kitag.<br />
China, Korea & E Russia<br />
8. <strong>Aralia</strong> cachemirica Decne.<br />
Himalaya<br />
9. <strong>Aralia</strong> tibetana G. Hoo<br />
E Himalaya<br />
10. <strong>Aralia</strong> fargesii Franch.<br />
China<br />
11. <strong>Aralia</strong> atropurpurea Franch.<br />
China<br />
12. <strong>Aralia</strong> apioides H<strong>and</strong>.-Mazz.<br />
China<br />
13. <strong>Aralia</strong> glabra Matsum.<br />
Japan<br />
14. <strong>Aralia</strong> henryi Harms<br />
China.<br />
II. <strong>Aralia</strong> sect. Humiles Harms<br />
<strong>Aralia</strong> sect. Humiles Harms, Bot. Jahrb. 23: 13.<br />
1896.<br />
Unarmed branched shrubs. Leaves pinnate or<br />
bipinnate (A. regeliana), stipulate; trichomes on<br />
lower leaf surface branched, stomates on both<br />
abaxial <strong>and</strong> adaxial leaf surfaces. Umbels 5-15,<br />
loosely clustered into panicles, inflorescence<br />
terminal. Flowers 5 (-6)-merous. Fruits dark<br />
purple.<br />
Number <strong>of</strong> species: 3.<br />
Type species: A. humilis Cav.<br />
Distribution: Central to North America.<br />
List <strong>and</strong> distribution <strong>of</strong> species in <strong>Aralia</strong> sect.<br />
Humiles<br />
1. <strong>Aralia</strong> humilis Cav.<br />
C & N America<br />
2. <strong>Aralia</strong> regeliana Marchal<br />
Mexico<br />
3. <strong>Aralia</strong> scopulorum Br<strong>and</strong>egee<br />
Baja California<br />
III. <strong>Aralia</strong> sect. Nanae Harms<br />
<strong>Aralia</strong> sect. Nanae Harms, Bot. Jahrb. 23: 12, 1896.<br />
Herbs with long horizontal rhizomes,<br />
acaulescent, dioecious. Leaves ternately<br />
compound, solitary, arising from the tip <strong>of</strong> the short<br />
rhizome, trichomes simple <strong>and</strong> slender,<br />
occasionally branching at the tip. Inflorescence<br />
solitary, arising from the base, umbels (2) 3 (7) in<br />
a cluster with a long scape. Flowers 5 or<br />
occasionally 6-merous, styles 5 (-6), free. Fruits
lackish purple.<br />
Number <strong>of</strong> species: 1.<br />
Type species: <strong>Aralia</strong> nudicaulis L.<br />
Distribution: North America.<br />
IV. <strong>Aralia</strong> sect. Pentapanax (Seem.) J. Wen<br />
<strong>Aralia</strong> sect. Pentapanax (Seem.) J. Wen, Cathaya<br />
11-12: 31. 2002.<br />
Pentapanax Seem., J. Bot. 2: 294. 1864. Type<br />
species: Pentapanax leschenaultii (DC.) Seem.<br />
Parapentapanax Hutch., Gen. Fl. Pl. 2: 56. 1967.<br />
Type species: Parapentapanax racemosus<br />
(Seem.) Hutch.<br />
Unarmed shrubs or trees, branched, glabrous<br />
or with simple trichomes. Leaves pinnate, rarely<br />
bipinnate or tripinnate (A. wilsonii <strong>and</strong> A. plumosa),<br />
stipulate to exstipulate. Umbels or occasionally<br />
racemes (A. gigantean, A. lihengiana, <strong>and</strong> A.<br />
subcordata), then aggregated into panicles or<br />
umbellate panicles, inflorescence terminal,<br />
subtended by persistent bracts at the base. Flowers<br />
5-merous, styles connate, or connate at the base,<br />
rarely free. Fruits dark purple.<br />
Number <strong>of</strong> species: 19.<br />
Type species: <strong>Aralia</strong> leschenaultii (Seem.) J.<br />
Wen<br />
Distribution: widely distributed in Asia,<br />
especially in the Sino-Himalayan region such as<br />
northern India, Nepal, <strong>and</strong> western China, also<br />
extending to Java in Indonesia, northern Thail<strong>and</strong>,<br />
<strong>and</strong> northern Vietnam.<br />
List <strong>and</strong> distribution <strong>of</strong> species in <strong>Aralia</strong> sect.<br />
Pentapanax<br />
1. <strong>Aralia</strong> wilsonii Harms<br />
China<br />
2. <strong>Aralia</strong> plumosa H. L. Li<br />
China<br />
3. <strong>Aralia</strong> caesia H<strong>and</strong>.-Mazz.<br />
China<br />
4. <strong>Aralia</strong> delavayi J. Wen<br />
China<br />
5. <strong>Aralia</strong> shangiana J. Wen<br />
China<br />
6. <strong>Aralia</strong> franchetii J. Wen<br />
China<br />
7. <strong>Aralia</strong> tomentella Franch.<br />
China<br />
8. <strong>Aralia</strong> stellata (King) J. Wen<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
29<br />
China & N Thail<strong>and</strong><br />
9. <strong>Aralia</strong> castanopsisicola (Hayata) J. Wen<br />
Taiwan, China<br />
10. <strong>Aralia</strong> parasitica (D. Don) J. Wen<br />
China & Himalaya<br />
11. <strong>Aralia</strong> laevis J. Wen<br />
SE Asia<br />
12. <strong>Aralia</strong> verticillata (Dunn) J. Wen<br />
China & N Vietnam<br />
13. <strong>Aralia</strong> hypoglauca (C. J. Qi & T. R. Cao) J.<br />
Wen & Y.-F. Deng<br />
China<br />
14. <strong>Aralia</strong> glabrifoliolata (C. B. Shang) J. Wen<br />
China<br />
15. <strong>Aralia</strong> leschenaultii (DC.) J. Wen<br />
E Asia<br />
16. <strong>Aralia</strong> kingdon-wardii J. Wen, Lowry & Esser<br />
Himalaya<br />
17. <strong>Aralia</strong> gigantea J. Wen<br />
Himalaya<br />
18. <strong>Aralia</strong> lihengiana J. Wen, L. Deng & X. Shi<br />
W China<br />
19. <strong>Aralia</strong> subcordata (Don) J. Wen<br />
Himalaya<br />
V. <strong>Aralia</strong> sect. Sciadodendron (Griseb.) J. Wen<br />
<strong>Aralia</strong> sect. Sciadodendron (Griseb.) J. Wen, comb.<br />
et stat. nov.<br />
Sciadodendron Griseb., Bonpl<strong>and</strong>ia 6: 7. 1858.<br />
Type species: Sciadodendron excelsum Griseb.<br />
Coudenbergia Marchal, Bull. Acad. Roy. Sci.<br />
Belgique 47: 514, 1879. Type species:<br />
Coudenbergia warmingiana (Marchal)<br />
Marchal.<br />
Megalopanax Ekman ex Harms, Notizbl. Bot. Gart.<br />
Berlin 9: 122, 1924. Type species:<br />
Megalopanax rex Ekman ex Harms.<br />
Unarmed glabrous trees or large shrubs,<br />
branched. Leaves 3- to 4- or 2-pinnate. Umbels<br />
clustered into panicles, several inflorescences<br />
usually aggregated at tip <strong>of</strong> stem, with persistent<br />
bracts at the base. Flowers 5-, 6-10-(12)-merous,<br />
styles connate or at least at the base. Fruits dark<br />
purple.<br />
Number <strong>of</strong> species: 5.<br />
Type species: <strong>Aralia</strong> excelsa (Griseb.) J. Wen<br />
Distribution: Central <strong>and</strong> South America.<br />
List <strong>and</strong> distribution <strong>of</strong> species in <strong>Aralia</strong> sect.
30<br />
Sciadodendron<br />
1. <strong>Aralia</strong> soratensis Marchal<br />
S America<br />
2. <strong>Aralia</strong> warmingiana (Marchal) J. Wen<br />
S America<br />
3. <strong>Aralia</strong> rex (Ekman ex Harms) J. Wen<br />
Cuba<br />
4. <strong>Aralia</strong> excelsa (Griseb.) J. Wen<br />
C & S America<br />
5. <strong>Aralia</strong> bahiana J. Wen<br />
E Brazil<br />
VI. <strong>Aralia</strong> sect. Dimorphanthus (Miq.) Miq.<br />
<strong>Aralia</strong> sect. Dimorphanthus (Miq.) Miq., Ann.<br />
Mus. Lugd.bat. 1: 6. 1863.<br />
<strong>Aralia</strong> sect. Arborescentes Harms, Bot. Jahrb. 23:<br />
16. 1896.<br />
<strong>Aralia</strong> sect. Capituligerae Harms, Bot. Jahrb. 23:<br />
19. 1896.<br />
<strong>Aralia</strong> sect. Digitatipanicula Hoo in Hoo & Tseng,<br />
Acta Phytotax. Sin., suppl. 1: 170. 1965.<br />
Armed shrubs or trees, <strong>and</strong>romonoecious or<br />
hermaphrodite. Leaves bipinnate, sometimes 3pinnate<br />
to 4-pinnate (e.g., A. armata, A.<br />
finlaysoniana <strong>and</strong> A. ferox), stipulate, trichomes<br />
simple. Umbels numerous, aggregated into a<br />
paniculate to umbellate structure, inflorescence<br />
large (20150 cm long) <strong>and</strong> terminal. Flowers 5merous,<br />
styles free or connate at the base. Fruits<br />
dark purple to black.<br />
Number <strong>of</strong> species: 29.<br />
Type species: <strong>Aralia</strong> elata (Miq.) Miq.<br />
Distribution: disjunct between temperate <strong>and</strong><br />
tropical Asia <strong>and</strong> eastern North America.<br />
List <strong>and</strong> distribution <strong>of</strong> species in <strong>Aralia</strong> sect.<br />
Dimorphanthus<br />
1. <strong>Aralia</strong> spinosa L.<br />
E North America<br />
2. <strong>Aralia</strong> hispida Vent.<br />
E North America<br />
3. <strong>Aralia</strong> stipulata Franch.<br />
China<br />
4. <strong>Aralia</strong> gintungensis C. Y. Wu<br />
China<br />
5. <strong>Aralia</strong> undulata H<strong>and</strong>.-Mazz.<br />
China<br />
6. <strong>Aralia</strong> <strong>of</strong>ficinalis Z. Z. Wang<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
China<br />
7. <strong>Aralia</strong> debilis J. Wen<br />
China<br />
8. <strong>Aralia</strong> echinocaulis H<strong>and</strong>.-Mazz.<br />
China<br />
9a. <strong>Aralia</strong> elata (Miq.) Seem. var. elata<br />
China & Japan<br />
9b. <strong>Aralia</strong> elata (Miq.) Seem. var. m<strong>and</strong>shurica<br />
(Rupr. & Maxim.) J. Wen<br />
China & E Russia<br />
9c. <strong>Aralia</strong> elata (Miq.) Seem. var. ryukyuensis J.<br />
Wen<br />
S Japan<br />
9d. <strong>Aralia</strong> elata (Miq.) Seem. var. inermis<br />
(Yanagita) J. Wen<br />
S Japan<br />
10. <strong>Aralia</strong> dasyphylloides (H<strong>and</strong>.-Mazz.) J. Wen<br />
China<br />
11. <strong>Aralia</strong> scaberula G. Hoo<br />
China<br />
12a. <strong>Aralia</strong> bipinnata Blanco var. bipinnata<br />
The Philippines & Taiwan<br />
12b. <strong>Aralia</strong> bipinnata Blanco var. apoensis (Elmer)<br />
J. Wen<br />
The Philippines<br />
13. <strong>Aralia</strong> decaisneana Hance<br />
Taiwan, China<br />
14. <strong>Aralia</strong> finlaysoniana (Wall. ex Don) Seem.<br />
China & Indochina<br />
15. <strong>Aralia</strong> spinifolia Merr.<br />
China<br />
16. <strong>Aralia</strong> merrillii C. B. Shang<br />
SE Asia<br />
17. <strong>Aralia</strong> ferox Miq.<br />
SE Asia<br />
18. <strong>Aralia</strong> foliolosa Seem. ex C. B. Clarke<br />
China & Himalaya<br />
19. <strong>Aralia</strong> montana Blume<br />
SE Asia<br />
20. <strong>Aralia</strong> frodiniana J. Wen<br />
Sulawesi<br />
21. <strong>Aralia</strong> malabarica Bedd.<br />
S India<br />
22. <strong>Aralia</strong> armata (Wall. ex Don) Seem.<br />
China & Himalaya<br />
23. <strong>Aralia</strong> chinensis L.<br />
China<br />
24. <strong>Aralia</strong> hiepiana J. Wen & Lowry<br />
Vietnam<br />
25. <strong>Aralia</strong> thomsonii Seem. ex C. B. Clarke<br />
China & Indochina
26. <strong>Aralia</strong> vietnamensis T.-D. Ha<br />
China & Indochina<br />
27. <strong>Aralia</strong> searelliana Dunn<br />
China & N Vietnam<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
28. <strong>Aralia</strong> dasyphylla Miq.<br />
SE Asia<br />
29. <strong>Aralia</strong> urticifolia Miq.<br />
Java<br />
KEY TO SECTIONS OF ARALIA<br />
1. Herbs or sometimes with woody rhizomes ........................................................................................ 2<br />
1. Shrubs or trees ....................................................................................................................………… 3<br />
2. Acaulescent dioecious herbs with long horizontal woody rhizomes ...................................... Sect. Nanae<br />
2. Caulescent <strong>and</strong>romonoeceous herbs with short <strong>and</strong> thick rhizomes ..…............................... Sect. <strong>Aralia</strong><br />
3. Shrubs or trees with prickles on stems ...................................................................... Sect. Dimorphanthus<br />
3. Shrubs or trees with unarmed stems .................................................................................................. 4<br />
4. Trichomes on lower leaflet surface branched ....................................................................... Sect. Humiles<br />
4. Trichomes on lower leaflet surface simple if they are present ............................................................ 5<br />
5. Leaves 3-4-pinnately compound, ovaries 5-10 (-12)-locular, from Central & S America ...........................<br />
.................................................................................................................................. Sect. Sciadodendron<br />
5. Leaves 1-2-pinnately or occasionally 3-pinnately compound, ovaries 5-locular, from eastern to<br />
southeastern Asia ......................................................................................................... Sect. Pentapanax<br />
The reproductive biology <strong>of</strong> a few <strong>Aralia</strong><br />
species has been documented. Possible<br />
evolutionary pathways leading to various<br />
reproductive systems have been proposed for the<br />
New Caledonian <strong>Aralia</strong>ceae (Schlessman et al.<br />
1990, 2001). A range <strong>of</strong> reproductive systems are<br />
present in <strong>Aralia</strong>, e.g., dioecious,<br />
<strong>and</strong>romonoecious, <strong>and</strong> hermaphroditic (Wen<br />
2004).<br />
Lovell (1898) noted that prot<strong>and</strong>rous<br />
dichogamy is strongly developed in <strong>Aralia</strong><br />
racemosa, a member <strong>of</strong> the herbaceous <strong>Aralia</strong> sect.<br />
<strong>Aralia</strong>. At the beginning <strong>of</strong> anthesis, the styles are<br />
initially united <strong>and</strong> only about 1 mm long <strong>and</strong> the<br />
stamens are about 3 mm long with the anthers<br />
horizontal or inclining outward. Small dipterans<br />
visit the flowers, thrusting their heads between the<br />
stamens or the bodies <strong>of</strong> the larger ones passing<br />
through them. With the stamens <strong>and</strong> petals falling<br />
away, the styles elongate to about 3 mm long <strong>and</strong><br />
become reflexed finally. Abundant nectar is<br />
REPRODUCTIVE BIOLOGY<br />
31<br />
produced <strong>and</strong> can be seen as minute drops on the<br />
floral disk. The honey-bee was observed to make<br />
40 or so visits in one minute, <strong>and</strong> Lucilia cornicina<br />
was seen to visit 26 flowers in the same time.<br />
Lovell (1898) reported that a large number <strong>of</strong> less<br />
specialized hymenopterans, such as ichneumonflies<br />
<strong>and</strong> wasps visited the flowers.<br />
The reproductive biology <strong>of</strong> <strong>Aralia</strong> nudicaulis<br />
(sect. Nanae) has been well studied (e.g., Barrett<br />
<strong>and</strong> Helenurm 1981; Barrett <strong>and</strong> Thomson 1982;<br />
Bawa et al. 1982; Edwards 1984; Flanagan <strong>and</strong><br />
Moser 1985; Thomson <strong>and</strong> Brunet 1990). <strong>Aralia</strong><br />
nudicaulis is dioecious in the boreal <strong>and</strong> temperate<br />
forests <strong>of</strong> North America. The species is pollinated<br />
mostly by bumble bees. The female ramets have<br />
fewer flowers per umbel than the male ramets <strong>and</strong><br />
reach peak flowering before the male ramets. The<br />
sex-related differences in the frequency <strong>of</strong><br />
flowering have been examined in detail in the<br />
species by Bawa et al. (1982). Bawa et al. (1982)<br />
also noted that a large proportion <strong>of</strong> sexually
32<br />
mature ramets may not flower in a given year.<br />
<strong>Aralia</strong> hispida (sect. Dimorphanthus) is<br />
<strong>and</strong>romonoecious <strong>and</strong> maintains outcrossing<br />
through synchronous dichogamy (Thomson <strong>and</strong><br />
Barrett 1981; Thomson et al. 1982; Thomson 1988;<br />
Thomson et al. 1990). This species blooms in three<br />
ranks (primary, secondary, <strong>and</strong> tertiary) <strong>of</strong> umbel<br />
orders. All flowers <strong>of</strong> one order complete opening<br />
before flowers <strong>of</strong> the next order start. The flowers<br />
In this treatment, species have been delimited<br />
using a phylogenetic concept sensu Nixon <strong>and</strong><br />
Wheeler (1990). All species <strong>of</strong> <strong>Aralia</strong> are<br />
recognized by a unique combination <strong>of</strong> character<br />
states in comparable individuals. Once again, I<br />
view species as hypotheses based on available<br />
characters (also see Wen 2002, 2004). The<br />
circumscription may be revised as needed when<br />
additional data become available.<br />
Species-delimiting characters must show<br />
fixation throughout the distributional range. Some<br />
“species” were based on extreme conditions <strong>of</strong><br />
certain characters. For example, <strong>Aralia</strong> pubescens<br />
was described by de C<strong>and</strong>olle (1813) based on the<br />
highly pubescent specimens from Chiapas. When<br />
collections <strong>of</strong> <strong>Aralia</strong> humilis were examined<br />
throughout its distributional range, I realized that<br />
pubescence on leaves, inflorescence <strong>and</strong> pedicels<br />
is highly variable in this species. It is difficult to<br />
discern any discontinuity in this character nor any<br />
geographic integrity <strong>of</strong> the pubescent “form.”<br />
Therefore I have treated <strong>Aralia</strong> humilis broadly <strong>and</strong><br />
placed A. pubescens as a synonym <strong>of</strong> A. humilis.<br />
<strong>Aralia</strong> bicrenata was previously recognized as<br />
a variety or a subspecies <strong>of</strong> A. racemosa or simply<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
SPECIES CONCEPTS<br />
are prot<strong>and</strong>rous, with the flowers open first as<br />
males. After 5-7 days, the styles elongate <strong>and</strong><br />
diverge, signaling the functioning female-phase <strong>of</strong><br />
the flower. The female-phase flowers are crosspollinated<br />
before the openning <strong>of</strong> flowers <strong>of</strong> the<br />
next order. Pollinators <strong>of</strong> A. hispida are primarily<br />
Bombus spp. (Thomson et al. 1982; Thomson<br />
1988).<br />
treated as a synonym <strong>of</strong> A. racemosa. Our field<br />
studies in Colorado, New Mexico, Arizana, <strong>and</strong><br />
Texas suggest that A. bicrenata differs from A.<br />
racemosa in its inflorescence structure, leaflet<br />
margin, stipule morphology, <strong>and</strong> stem color. Our<br />
herbarium work also shows that these<br />
morphological differences are consistent<br />
throughout the distributional range <strong>of</strong> both taxa.<br />
Geographically there is no overlap between the two<br />
taxa. Varieties are considered by this author to have<br />
one or a few morphological distinctions that have<br />
geographic integrity (Lewis 1955; Stuessy 1990).<br />
Unlike species, some geographic overlap may<br />
occur among other conspecific varieties, but such<br />
overlap is common at this level in the hierarchy<br />
(Lawrence 1951; Stuessy 1990). In my previous<br />
revisions <strong>of</strong> <strong>Aralia</strong> such as in sect. Dimorphanthus,<br />
morphological differences among conspecific<br />
varieties are mostly related to pubescence, shape,<br />
texture, number <strong>of</strong> lateral veins <strong>of</strong> leaflets, <strong>and</strong><br />
pedicel length. Considering the differences in<br />
several morphological characters <strong>and</strong> the lack <strong>of</strong><br />
geographic overlap, I recognize <strong>Aralia</strong> bicrenata<br />
as a species distinct from A. racemosa.<br />
DISTRIBUTION AND BIOGEOGRAPHY<br />
Early evolutionary radiation in <strong>Aralia</strong><br />
Our phylogenetic analyses using multiple<br />
markers have not generated a well resolved<br />
phylogeny <strong>of</strong> <strong>Aralia</strong> at “deep” level (see Figs. 1 &<br />
2). The sections I presently recognize are all<br />
supported as monophyletic groups, yet the<br />
relationships among them remain poorly resolved.<br />
I propose that the major lineages <strong>of</strong> the genus<br />
diversified as an evolutionary radiation in a very<br />
short period <strong>of</strong> time.<br />
It is significant that the New World harbors a<br />
relatively high diversity <strong>of</strong> major lineages even<br />
though it has only 14 <strong>of</strong> the 71 species <strong>of</strong> <strong>Aralia</strong>.
The New World species <strong>of</strong> the genus belong to five<br />
sections, <strong>of</strong> which three are endemic to the New<br />
World (sect. Humiles, sect. Nanae, <strong>and</strong> sect.<br />
Sciadodendron) <strong>and</strong> two sections disjunct with<br />
Asia (sect. <strong>Aralia</strong> <strong>and</strong> sect. Dimorphanthus). The<br />
57 Asian species belong to three sections (sect.<br />
<strong>Aralia</strong>, sect. Dimorphanthus, <strong>and</strong> sect.<br />
Pentapanax) with only sect. Pentapanax endemic<br />
to Asia, <strong>and</strong> the other two sections disjunct between<br />
Asia <strong>and</strong> North America. Most species in Asia are<br />
from two sections: sects. <strong>Aralia</strong> (eleven out <strong>of</strong> 14<br />
in Asia) <strong>and</strong> Dimorphanthus (27 out <strong>of</strong> 29 in Asia).<br />
<strong>Aralia</strong> sect. <strong>Aralia</strong><br />
This section has a wide intercontinental disjunct<br />
distribution in the northern hemisphere, with<br />
eleven species in eastern Asia <strong>and</strong> three species<br />
allopatrically distributed in North America. Within<br />
eastern Asia, species are primarily allopatric in<br />
distribution. Only distantly related taxa, such as<br />
A. continentalis, A. fargesii <strong>and</strong> A. henryi in Central<br />
China may occur in the same mountain. In North<br />
America, A. racemosa is in eastern North America,<br />
A. bicrenata is in southwestern North America <strong>and</strong><br />
northern Mexico, <strong>and</strong> A. californica is in California<br />
<strong>and</strong> southern Oregon. Wen et al. (1998) suggested<br />
the origin <strong>of</strong> the section in eastern Asia <strong>and</strong> that<br />
the ancester <strong>of</strong> the North American taxa migrated<br />
across the Bering l<strong>and</strong> bridge in the late Tertiary<br />
<strong>and</strong> diversified in North America. Within North<br />
America, an eastern <strong>and</strong> western North American<br />
biogeographic track is supported by the distribution<br />
<strong>of</strong> A. racemosa – A. bicrenata – A. californica. The<br />
eastern species A. racemosa occupies an extensive<br />
distributional area, whereas the western A.<br />
californica is more restricted to California <strong>and</strong><br />
southern Oregon. This may be due to the fact that<br />
A. californica has co-existed <strong>and</strong> co-adapted with<br />
the redwood <strong>and</strong> the Douglas fir forests, which are<br />
ecologically restricted in western North America.<br />
Within Asia, the <strong>Aralia</strong> cordata – A. taiwaniana<br />
- A. continentalis – A. schmidtii – A. tibetana – A.<br />
cachemirica group shows an eastern Asian<br />
biogeographic track spanning from Japan, across<br />
China <strong>and</strong> to the western Himalaya in the East –<br />
West direction. The <strong>Aralia</strong> cordata –A.<br />
continentalis group shows a close biogeographic<br />
relationship <strong>of</strong> the Sino-Japanese region as a<br />
biogeographic track. This track is also supported<br />
in <strong>Aralia</strong> by the distribution <strong>of</strong> A. glabra in Japan<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
33<br />
<strong>and</strong> A. fargesii in central China.<br />
The phylogeographic relationship <strong>of</strong> the <strong>Aralia</strong><br />
cordata complex (A. cordata, A. taiwaniana, A.<br />
continentalis, A. schmidtii, A. cachemirica <strong>and</strong> A.<br />
tibetana) needs to be carefully examined. The six<br />
species occur allopatrically <strong>and</strong> are<br />
morphologically similar. <strong>Aralia</strong> taiwaniana may<br />
have been derived from A. cordata in Japan <strong>and</strong><br />
been preserved in the central mountains in Taiwan.<br />
<strong>Aralia</strong> schmidtii from the Sakhalin <strong>and</strong> Kuril<br />
isl<strong>and</strong>s in eastern Russia, north <strong>of</strong> the Japanese<br />
archipelago in the North Pacific may have its origin<br />
from A. cordata in J apan. The split between A.<br />
continentalis <strong>and</strong> A. cordata may be more ancient.<br />
Because <strong>of</strong> the presumably central role <strong>of</strong> A.<br />
cordata in giving rise to other species in the<br />
complex, it may be difficult to discern the<br />
evolutionary relationships among these species.<br />
<strong>Aralia</strong> sect. Humiles<br />
This section is distributed in relatively dry<br />
habitats from southern Arizona across Mexico to<br />
Guatemala, Honduras, <strong>and</strong> Nicaragua.<br />
Phylogenetic analysis has not been conducted for<br />
the section. Based on morphological comparisons,<br />
<strong>Aralia</strong> scopulorum from Baja California may be a<br />
derivative species <strong>of</strong> the more widespread <strong>Aralia</strong><br />
humilis, especially from populations <strong>of</strong> northern<br />
Mexico. Furthermore A. regeliana may also be a<br />
derivative <strong>of</strong> A. humilis, likely from a bipinnate<br />
form in central or southern Mexico. The two<br />
species in each presumed progenitor-derivative<br />
pair do not overlap in geographic distribution, <strong>and</strong><br />
allopatric speciation via geographic <strong>and</strong> habitat<br />
isolation may explain such a pattern.<br />
<strong>Aralia</strong> sect. Sciadodendron<br />
The five species <strong>of</strong> <strong>Aralia</strong> in this section are<br />
primarily allopatric in distribution in the<br />
neotropics. <strong>Aralia</strong> rex occurs in the Carribean<br />
isl<strong>and</strong> <strong>of</strong> Cuba; A. excelsa is distributed in Central<br />
America, extending south to Colombia <strong>and</strong><br />
Venezuela; A. soratensis is found in the Andes <strong>of</strong><br />
Peru, Bolivia, <strong>and</strong> Argentina; <strong>Aralia</strong> bahiana is a<br />
new species recorded only from Bahia, Brazil; <strong>and</strong><br />
A. warmingiana is primarily in the lowl<strong>and</strong> areas<br />
in Brazil <strong>and</strong> Paraguay. These five species are<br />
morphologically highly distinct <strong>and</strong> may have<br />
radiated early in the evolutionary history <strong>of</strong> the
34<br />
section, <strong>and</strong> developed in separate trajectories in<br />
Central <strong>and</strong> South America. <strong>Aralia</strong> bahiana <strong>and</strong><br />
A. excelsa both have non-articulated pedicels. This<br />
character state may be apomorphic in <strong>Aralia</strong> <strong>and</strong><br />
may be the synapomorphy for the two species. The<br />
geographically highly restricted A. bahiana may<br />
be a derivative <strong>of</strong> the more widespread A. excelsa.<br />
Nevertheless, it is also possible that A. bahiana<br />
was derived from A. warmingiana <strong>of</strong> Brazil.<br />
<strong>Aralia</strong> soratensis from South America is<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
TAXONOMY<br />
morphologically similar to the North American A.<br />
spinosa, which belongs to <strong>Aralia</strong> sect.<br />
Dimorphanthus (see Wen 2004). They differ<br />
primarily in the presence <strong>of</strong> prickles in A. spinosa,<br />
<strong>and</strong> in the more congested <strong>and</strong> more compact<br />
inflorescence <strong>of</strong> A. soratensis. The morphological<br />
similarities <strong>of</strong> the two species suggest the close<br />
relationship between sect. Sciadodendron <strong>of</strong> the<br />
neotropics <strong>and</strong> sect. Dimorphanthus primarily <strong>of</strong><br />
the northern hemisphere.<br />
<strong>Aralia</strong> L., Sp. Pl. 273. 1753.; Gen. Pl., ed. 5: 134. 1754. TYPE SPECIES: <strong>Aralia</strong> racemosa L.<br />
Dimorphanthus Miq., Comm. Phytogr. 3: 95. 1840. TYPE SPECIES: Dimorphanthus elatus Miq.<br />
Sciadodendron Griseb., Bonpl<strong>and</strong>ia 6: 7. 1858. TYPE SPECIES: Sciadodendron excelsum Griseb.<br />
Pentapanax Seem., J. Bot. 2: 294. 1864. TYPE SPECIES: Pentapanax leschenaultii (DC.) Seem.<br />
Coemansia Marchal, Bull. Acad. Roy. Sci. Belgique 47: 94. 1879. nom. illeg., non Tiegh. &<br />
Monnier in Ann. Sci. Nat. Bot., Ser. 5, 17: 392. 1873.<br />
Coudenbergia Marchal, Bull. Acad. Roy. Sci. Belgique 47: 514. 1879. TYPE SPECIES:<br />
Coudenbergia warmingiana (Marchal) Marchal.<br />
Acanthophora Merr., Philipp. J. Sci. 13: 316. 1918. nom. illeg., non J. V. Lamour. 1813.<br />
Neoacanthophora Bennet, Indian J. Forest. 2: 283. 1979. TYPE SPECIES: Neoacanthophora<br />
sc<strong>and</strong>ens (Merr.) Bennet.<br />
Cwangayana Rauschert, Taxon 31: 556. 1982. TYPE SPECIES: Cwangayana sc<strong>and</strong>ens (Merrill)<br />
Rauschert.<br />
Megalopanax Ekman ex Harms, Notizbl. Bot. Gart. Berlin 9: 122. 1924. TYPE SPECIES:<br />
Megalopanax rex Ekman ex Harms.<br />
Parapentapanax Hutch., Gen. Fl. Pl. 2:56. 1967. TYPE SPECIES: Parapentapanax racemosus<br />
(Seemann) Hutch.<br />
Trees, shrubs, lianas or herbs with rhizomes,<br />
armed or unarmed; leaves alternate, 14pinnate,<br />
rachis articulated, mostly stipulate; inflorescence<br />
terminal, paniculate or corymbose or umbellate,<br />
usually consisting <strong>of</strong> umbels, capitula or racemes,<br />
occasionally umbels solitary; flowers 512merous,<br />
with pedicels articulated below, only occasionally<br />
non-articulated; sepals 512; petals 5-12, imbricate;<br />
stamens 5-12, recurved in buds; ovary inferior,<br />
512locular, occasionally aborted into 3, with a<br />
ARALIA L. SECT. ARALIA<br />
<strong>Aralia</strong> sect. Genuinae Harms, Bot. Jahrb. 23: 15. 1896.<br />
<strong>Aralia</strong> sect. Anomalae Harms, Bot. Jahrb. 23: 12. 1896, synonym nov.<br />
floral disk, styles distinct or connate at the base or<br />
completely connate; fruit a berry, 512-locular, more<br />
or less globose.<br />
<strong>Aralia</strong> consists <strong>of</strong> 71 species disjunctly<br />
distributed from eastern Asia to southeastern Asia<br />
<strong>and</strong> the Americas from Canada to Argentina,<br />
Bolivia, Peru <strong>and</strong> Brazil. Eastern <strong>and</strong> southeastern<br />
Asia is especially rich in <strong>Aralia</strong> species (57 spp),<br />
whereas South America, North America <strong>and</strong><br />
Central America together have 14 species.
Branched unarmed herbs with rhizomes.<br />
Leaves alternate, stipulate, ternately bi- or<br />
tripinnate on lower portion <strong>of</strong> stem, pinnate to<br />
bipinnate on upper portion <strong>of</strong> stem, petiole<br />
articulated. Trichomes simple, biseriate.<br />
Inflorescence terminal <strong>and</strong> axillary, paniculately<br />
or umbellately arranged, with many umbels,<br />
pedicels articulated below the flowers. Sepals 5,<br />
minute, triangular or rounded. Petals 5, imbricate.<br />
Stamens 5. Ovary inferior, 5-locular, styles 5,<br />
connate at the base, floral disc small <strong>and</strong> flat. Fruit<br />
a berry, 5-seeded, dark purple when mature.<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
35<br />
Endosperm smooth.<br />
The section consists <strong>of</strong> 14 species disjunctly<br />
distributed between eastern Asia (11 spp.)<br />
including the Himalayas <strong>and</strong> North America (3<br />
spp.). Species <strong>of</strong> this section are perennial herbs<br />
primarily in mesic temperate forests.<br />
Species in the section are largely differentiated<br />
based on stipule morphology, leaf architecture,<br />
pubescence, leaflet size, shape <strong>and</strong> margin,<br />
inflorescence architecture, number <strong>of</strong> flowers per<br />
umbel, division <strong>of</strong> styles, <strong>and</strong> fruit size, shape <strong>and</strong><br />
color.<br />
Key to species <strong>of</strong> <strong>Aralia</strong> sect. <strong>Aralia</strong><br />
1. Fruits red; inflorescence with fewer than 10 umbels, each terminal umbel with fewer than 10 flowers<br />
..................................................................................................................................................... A. henryi<br />
1. Fruits dark purple; inflorescence with more than 20 umbels, each terminal umbel with more than 10<br />
flowers ............................................................................................................................................... 2<br />
2. Inflorescence a dense panicle with more than 25 primary branches ................................................... 3<br />
2. Inflorescence a loose panicle or an umbellate structure with fewer than 15 primary branches ......... 11<br />
3. Fruits 5.5-7 mm in diameter .................................................................................................. A. californica<br />
3. Fruits 3-5 mm in diameter .................................................................................................................. 4<br />
4. Inflorescence corymbose in architecture ............................................................................................ 5<br />
4. Inflorescence paniculate to racemose in architecture, primary bracts usually not leafy<br />
............................................................................................................................................................ 7<br />
5. Inflorescence with leaves <strong>and</strong> leafy bracts subtending the primary inflorescence branches; terminal<br />
umbels with 12-20 (25) flowers; in southwestern US <strong>and</strong> in northern Mexico ..................... A. bicrenata<br />
5. Inflorescence with leaves subtending the primary inflorescence branches, with the leaves decreasing<br />
in size drastically from lower part <strong>of</strong> the inflorescence to the upper part; terminal umbels with<br />
(25-) 35-60 flowers ........................................................................................................................... 6<br />
6. Leaflet margin finely serrulate, base <strong>of</strong> leaflets cordate, stipules lanceolate; endemic to the Sakhalin<br />
& Kuril isl<strong>and</strong>s <strong>of</strong> far eastern Russia ...................................................................................... A. schmidtii<br />
6. Leaflet margin doubly serrate, base <strong>of</strong> leaflets obliquely truncate to subcordate, rarely cordate, stipules<br />
narrowly triangular; endemic to Taiwan ............................................................................. A. taiwaniana<br />
7. Terminal umbels usually with fewer than 35 flowers ........................................................................ 8<br />
7. Terminal umbels usually with 35 or more flowers .............................................................................. 9<br />
8. Stem dark purple to greenish purple, without stiff hairs at the base <strong>of</strong> stem; stipules narrowly triangular,<br />
not leafy; flowers smaller with petals 1.4-1.6 mm long, 0.9-1.1 mm wide; eastern U.S. <strong>and</strong> Canada<br />
................................................................................................................................................ A. racemosa<br />
8. Stem green to greenish purple, with stiff hairs at the base <strong>of</strong> stem; stipules leafy; flowers longer with<br />
petals 1.8-2 mm long, 0.9-1.1 mm wide; mainl<strong>and</strong> China, Korea & eastern Russia ........ A. continentalis
36<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
9. The pair <strong>of</strong> leaflets below the terminal leaflet narrowly ovate, base rounded to subcordate<br />
........................................................................................................................................... A. cachemirica<br />
9. The pair <strong>of</strong> leaflets below the terminal leaflet ovate to broadly so, base cordate ............................... 10<br />
10. Leaves primarily tripinnately ternate; leaflets membranaceous; stipules not leafy ................ A. tibetana<br />
10. Leaves primarily bipinnately ternate; leaflets papery or chartaceous; stipules usually leafy on the<br />
lower part <strong>of</strong> the stem ................................................................................................................ A. cordata<br />
11. Leaflets glabrous throughout; pedicels 12-45 mm long ............................................................ A. glabra<br />
11. Leaflets pilose to pubescent; pedicels less than 15 mm long in general .......................................... 12<br />
12. Leaflet margin irregularly doubly serrate with deep teeth, sometimes lobed ........................ A. apioides<br />
12. Leaflet margin serrate to doubly so without deep teeth, never lobed ............................................... 13<br />
13. Pedicels 8-12 mm long in terminal umbels; styles completely distinct to the base ......... A. atropurpurea<br />
13. Pedicels 5-6 mm long in terminal umbels; styles connate to the middle or more ................. A. fargesii<br />
1. <strong>Aralia</strong> californica S. Watson — Fig. 16; color<br />
plate 1: A-F.<br />
<strong>Aralia</strong> californica S. Watson, Proc. Amer. Acad.<br />
Arts 11: 144. 1876. TYPE: U.S.A.<br />
CALIFORNIA: northern California, 1860-62,<br />
W. H. Brewer 747 (lectotype: US!, here<br />
designated; isolectotype: GH!, UC!).<br />
<strong>Aralia</strong> californica S. Watson var. acuminata S.<br />
Watson ex Howell, Fl. N. W. Amer. 1: 271.<br />
1898. TYPE: U.S.A. OREGON: southern<br />
Oregon, Siskiyou Mountains, Jul 1888, T.<br />
Howell 786 (lectotype: GH!, here designated).<br />
Robust spreading perennial herb, shrub-like,<br />
1.5-3.5 m tall, 1-3 m in spread, <strong>and</strong>romonoecious.<br />
Stem pilose, green to light purplish green, with 3-<br />
5 leaves; rhizomes thick <strong>and</strong> horizontal, bearing<br />
purple buds. Leaves 40-150 cm long, 35-160 cm<br />
wide, ternately compound, spreading, lower leaves<br />
tripinnate, uppermost ones pinnate; stipules 1.2-<br />
7.0 cm long, 1-7.0 cm wide, leafy on lower leaves,<br />
triangular to irregularly ovate on upper leaves;<br />
petioles (8) 15-35 cm long, sparsely pilose, green<br />
to purplish green; leaflets 8-45 cm long, 4.5-22<br />
cm wide, papery, ovate, narrowly ovate to broadly<br />
so, acuminate to acute at apex, subcordate, cordate<br />
to rounded at base, commonly oblique on lateral<br />
leaflets, serrate to doubly serrate at margin; lateral<br />
veins 9-12 on each side, conspicuous on both<br />
surfaces, green above <strong>and</strong> light green below,<br />
slightly pilose along the veins on both surfaces,<br />
petiolules 0.2-4, mostly 0.5-1.5 cm long, slightly<br />
pilose <strong>and</strong> thick. Inflorescence terminal <strong>and</strong><br />
axillary, pilose, green, sometimes purple, axillary<br />
inflorescences 20-30 cm long, terminal<br />
inflorescence 30-75 cm long, 10-25 cm wide,<br />
consisting <strong>of</strong> numerous umbels, twice or thrice<br />
branched, primary branches many, racemosely<br />
arranged on a main axis, <strong>of</strong>ten 3-8 forming a circle<br />
on the main axis, each primary branch 5-10 cm<br />
long, consisting <strong>of</strong> 3-10 umbels, with a few lateral<br />
umbels bearing no fruits (functionally male<br />
umbels); terminal umbels 30-75-flowered, pedicels<br />
12-20 mm long, pubescent; lateral umbels 20-30flowered,<br />
pedicels 6-12 mm long; bracts <strong>of</strong> primary<br />
branches linear to lanceolate or <strong>of</strong>ten leaf-like,<br />
pilose, the upper ones 10-20 (30) mm long, 1-2<br />
mm wide, the lower ones transitioning into a<br />
bipinnately ternate leaf or a pinnate leaf subtending<br />
the inflorescence branch; bracteoles linear, nearly<br />
glabrous <strong>and</strong> ciliate at margin, 2-2.5 mm long.<br />
Flowers cream white; calyx <strong>of</strong> 5 teeth, sepals 0.5-<br />
0.6 mm long <strong>and</strong> wide, triangular, persistent on<br />
fruits, with a conspicuous vascular bundle in the<br />
middle; petals 2-2.3 mm long, 1.1-1.3 mm wide,<br />
narrowly triangular to narrowly ovate, greenish<br />
white to white, with a conspicuous vascular bundle<br />
in the middle; stamens 5, recurved in buds,<br />
filaments 2.3-2.6 mm long; ovaries 5-locular,<br />
hypanthium with 10 conspicuous vascular bundles.<br />
Fruits 6-7 mm long, 5-6 mm wide, globose, dark<br />
purple to blackish, persistent styles 2.2-2.6 mm<br />
long, divided to the middle or only connate at the<br />
base.
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
Fig. 16. <strong>Aralia</strong> californica S. Watson. A. Habit with a leaf <strong>and</strong> an inflorescence. B. Leafy stipule. C. Close-up <strong>of</strong> leaflet margin<br />
<strong>and</strong> veinlets. D. Flower. E. Young fruit.<br />
37
38<br />
Common names: spikenard, California<br />
spikenard, Californian spikenard, California<br />
ginseng, elk clover, western aralia, <strong>and</strong> western<br />
spikenard.<br />
Uses: roots used as a tonic or for curing<br />
menstrual disorders.<br />
Phenology: flowering in June to July; fruiting<br />
in August to September.<br />
Distribution: California <strong>and</strong> southern Oregon<br />
<strong>of</strong> U.S.A. (Fig. 17).<br />
Ecology: in shady <strong>and</strong> moist habitats, e.g., in<br />
moist redwood, Douglas fir, <strong>and</strong>/or pine forests,<br />
mountain slopes <strong>and</strong> valleys, at the edge <strong>of</strong> forests,<br />
along river banks, shaded streams, creeks, <strong>and</strong><br />
ravines; 0-1830 m.<br />
Representative specimens examined: U.S.A.<br />
CALIFORNIA: Alameda Co., Strawberry Canyon,<br />
near creek, 4 Sep 1932, J. E. Adams 21 (UNCC);<br />
Oakl<strong>and</strong> Hills, ravines, Dr. Bol<strong>and</strong>er s.n. (NY);<br />
Berkeley, Strawberry Canyon, University <strong>of</strong><br />
California campus, 22 Aug 1939, H. P. Bracelin<br />
1092 (L, 3 sheets); lower Strawberry Canyon, 15<br />
Mar 1936, J. A. Ewan 9433 (UC); Oakl<strong>and</strong>, 9 Jul<br />
1881, young fr, M. E. Jones 2375 (NY, US);<br />
Oakl<strong>and</strong>, 9 Jul 1881, M. E. Jones 13414 (BM, NY,<br />
OS, PH, UC, US); Strawberry Canyon, Aug 1930,<br />
H. E. Parks 710 (UC); in creek bed west <strong>of</strong><br />
Botanical Garden, Strawberry Canyon, University<br />
<strong>of</strong> California Campus, Berkeley, 11 Aug 1943, B.<br />
Rodin 311 (UC); vicinity <strong>of</strong> Berkeley, Jul-Sep<br />
1906, H. A. Walker 401 (L, PH, UC); Strawberry<br />
Canyon, at the bottom <strong>of</strong> the canyon, moist area,<br />
growing with Rubus <strong>and</strong> ferns, large perennial 2-3<br />
m tall, leaves ca. 150 cm long, 160 cm wide, fruits<br />
dark purple, 12 Sep 1990, fr, J. Wen 747 (OS).<br />
Amador Co., Panther Creek, 4000 ft, Aug 1892, G.<br />
Hansen 325 (LE). Butte Co., at the Forbestown<br />
Reservoir, ca. 5 mi NE <strong>of</strong> Forbestown, 450 m, 24<br />
Jul 1983, L. & P. Ahart 4214 (MO, TEX); ca. 1 mi<br />
S <strong>of</strong> Ponderosa Dam across the south fork <strong>of</strong> the<br />
Feather River, ca. 3 mi NW <strong>of</strong> Forbestown, 1800<br />
ft, 1 Jul 1985, L. Ahart 5106 (MO). Colusa Co.,<br />
Paradise Creek at crossing <strong>of</strong> trail between Fouts<br />
Camp <strong>and</strong> Box Springs, Snow Mountain, 4500 ft,<br />
10 Jul 1972, A. Q. Howard 13 (JEPS). Contra<br />
Costa Co., Donner Canyon, 800-1000 ft, 21 Jun<br />
1933, M. L. Bowerman 233 (UC). Del Norte Co.,<br />
Smith River, 20 Jul 1935, G. N. Jones 7842 (GH, 2<br />
sheets); Patrick Creek, H. E. & S. E. Parks 24030<br />
(UC); along SR 199, ca. 9 miles east <strong>of</strong> jct with Rt<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
101, dry area, plant ca. 2 m tall, 15 Sep 1990, fr,<br />
fruit very dark purple, styles completely divided,<br />
J. Wen 762 (OS). El Dorado Co., north fork Webber<br />
Creek, ca. 3 mi E <strong>of</strong> Camino, 3000 ft, 16 Jul 1945,<br />
G. T. Robbins 2066 (GH, UC, US). Humboldt Co.,<br />
near Hoopa Reservation, Trinity River, 3 Sep 1948,<br />
A. M. Alex<strong>and</strong>er et al. 5587 (UC); Hoopa Valley,<br />
22 Sep 1920, D. Duncan s.n. (UC); Eel River, H.<br />
M. Hall s.n. (UC); east fork <strong>of</strong> Willow Creek, 1500<br />
ft, 1 Sep 1941, S. K. & C. C. Harris et al. 6976<br />
(GH, 2 sheets); Van Duzen River Valley, east <strong>of</strong><br />
Bridgeville, 2400 ft, 22 Aug 1975, fl, A. L. & H.<br />
N. Moldenke 30238 (TEX, US); Bull Creek, 7 Jul<br />
1911, fl, H. H. Smith 3942 (F, GH, MO, MU, NY,<br />
US); Mt. Chamisai, below summit, ca. 1900 ft, 5<br />
Aug 1965, R. F. Thorne et al. 35137 (BM, ENCB,<br />
UC); along Willow Creek, 1000 ft, 5 Jul 1911, J.<br />
P. Tracy 3320 (UC); Hoopa Valley, near Hoopa<br />
Reservation, abundant along creek, 14 Sep 1990,<br />
fr, fruits globose, green, milky white, purple to dark<br />
purple, J. Wen 755 (OS); same locality as Wen 755,<br />
plant smaller, fr & fl, hypanthium greenish white,<br />
sepals pale greenish, petals white, filaments long,<br />
anthers pale yellow, fruits dark purple, J. Wen 756<br />
(OS); Humboldt Redwoods State Park, ca. 4 miles<br />
N <strong>of</strong> Myers Flat, in redwood forest, ca. 2 m tall,<br />
1.4 m in spread, 15 Sep 1990, fr, J. Wen 757 (OS);<br />
Humboldt Redwoods State Park, along Pesula<br />
Road, abundant, inflorescence purple, 15 Sep 1990,<br />
J. Wen 758 (OS); Humboldt Redwoods State Park,<br />
Lower Bull Creek, along Bull Creek Road, inside<br />
Rockfeller Redwood Forest, 15 Sep 1990, J. Wen<br />
759 (OS); North Coast Range, Bull Creek –<br />
Petrolia Road, 12 mi W <strong>of</strong> Redwood Highway,<br />
1500 ft, 8 Sep 1937, C. B. Wolf 3060 (UC). Lake<br />
Co., Blue Lakes, 7 Sep 1929, J. W. Blankinship<br />
s.n. (MO, 2 sheets); Bear Creek, E <strong>of</strong> Timber Lake,<br />
Snow Mt., 5150 ft., 28 Jul 1981, L. R. Keckard et<br />
al. 5683 (JEPS). Los Angeles Co., Mt. Wilson,<br />
Little Santa Anita Canyon, 1350 ft, 31 Aug 1910,<br />
fr, S. F. Blake 837 (GH, US); moist bank in canon<br />
in Mt. Wilson, 25 Jul 1915, J. F. Macbride et al.<br />
882 (GH); Pasadena, 16 Sep 1892, fr, A. J.<br />
McClatchie s.n. (NY); Fallen Leaf Spring, W <strong>of</strong><br />
Sunset Peak, draining northward to Cow Canyon,<br />
4400 ft, seepage on north-facing shaded granitic<br />
slope along drainage with Alnus rhombifolia, Acer<br />
macrophyllum, Boykinia rotundifolia,<br />
Umbellularia californica, Ribes sp., 22 Nov 1991,<br />
past fr, T. S. Ross & A. H. Ross 6025 (F); mountain,<br />
woods, wet, rocky, Jul 1901, fl, G. B. Grant 4450
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
Fig. 17. Map <strong>of</strong> California <strong>and</strong> Oregon showing the distribution <strong>of</strong> <strong>Aralia</strong> californica S. Watson.<br />
39
40<br />
(NY, US). Marin Co., Hwy 1, 20 km W <strong>of</strong> the<br />
Tamalpias junction, E <strong>of</strong> Stinson Beach, coastal<br />
scrub/creekside vegetation, 75 m, herbaceous<br />
perennial to 2.5 m tall, under trees on moist sites,<br />
flowers white, 12 Jun 1986, B. Anderson 3041 (NY,<br />
2 sheets); Mt. Tamalpais, Jun 1915, floral buds, F.<br />
Beckwith 128 (NY); Mill Valley, 28 Aug 1892, F.<br />
T. Bioletti s.n. (UC); Mt. Tamalpais, Jun 1905, K.<br />
Br<strong>and</strong>egee s.n. (UC); Mt. Tamalpais, Br<strong>and</strong>egee<br />
s.n. (DS); Paper Mill Creek, Jul + Oct 1880, fr, J.<br />
W. Congdon s.n. (NY); Mt. Tamalpais, 2 Oct 1897,<br />
B. Davy s.n. (UC, 8 sheets); Mt. Tamalpais, 11 Jul<br />
1926, A. Eastwood 13922 (CAS); Lagunitas, Aug<br />
1876, fr, H. Edwards s.n. (NY); Phoenix Lake road<br />
1.5 mi below Lake Lagunitas, 8 Mar 1936, J. A.<br />
Ewan 9426 (UC); Mt. Tamalpais, on the railroad<br />
between Mill Valley trail <strong>and</strong> the water tank, 18<br />
Jun 1902, A. A. Heller 5717 (DS, E, F, GH, LE,<br />
MO, NY, PH, US); n slope <strong>of</strong> Bolinas Ridge, above<br />
Alpine Lake, shady canyons, 13 Aug 1949, fr, L.<br />
S. Lewis 49158 (NY); Mill Valley, 28 Aug 1892,<br />
fr, Michener & Bioletti s.n. (NY, 2 sheets); road<br />
west <strong>of</strong> Geronim, following creek at conspicuous<br />
water trough, 8 ft tall growing in moist area, 23<br />
Sep 1939, fr, D. G. Nelson 633 (NY); Bolinas<br />
Ridge, 14-16 Jun 1892, floral buds, E. Palmer 2408<br />
(NY, US); near Lagunitas Lake, Jun 1935, H. M.<br />
Pollard s.n. (UC); foot <strong>of</strong> Mt. Tamalpais, Jun 1881,<br />
V. Rattan s. n. (DS); Mill Valley, G. P. Rixford s.n.<br />
(CAS); north slope <strong>of</strong> Bolinas Ridge, above Alpine<br />
Lake, 13 Aug 1949, L. S. Rose 49158 (NY, UC);<br />
Lone Pine Beach, Tomales Bay, 25 Jul 1938, B. O.<br />
Schreiber 685 (UC); by a brook in the forest north<br />
<strong>of</strong> Mill Valley, 12 Jul 1913, W. N. Suksdorf 484<br />
(UC); Audubon Canyon Ranch, Pike County Gulch<br />
(Galloway Canyon), on western slope <strong>of</strong> Bolinas<br />
ridge, ca. 3 mi N <strong>of</strong> Stinson Beach, alt. 10-25 ft,<br />
17 Aug 1977, G. H. True 8455 (CAS); Mt.<br />
Tamalpais, Jul 1880, fl, G. R. Vasey 233 (NY); Mill<br />
Valley, 4 Jul 1907, H. A. Walker s.n. (US); Mt.<br />
Tamalpais State Forest, at the bottom <strong>of</strong> a creek,<br />
under the canopy <strong>of</strong> Alnus trees, spreading plant<br />
ca. 1.5 m tall, rhizomes big <strong>and</strong> thick, bearing<br />
purple buds, 13 Sep 1990, fr, J. Wen 748 (OS); Mt.<br />
Tamalpais State Park, near a stream, plant ca. 3 m<br />
tall, 12 Sep 1990, fr, J. Wen 751 (OS). Mendocino<br />
Co., Mendocino, Aug 1898, fr, H. E. Brown 903<br />
(F, NY, US); Round Valley, 440 m, V. K. Chestnut<br />
220 (US, 2 sheets); Glen Blair, Jul 1894, A.<br />
McCallum s.n. (CAS); Big River, Jul 1903, fl, J.<br />
McMurphy 4 (NY, US); under redwoods along river<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
road just W <strong>of</strong> confluence <strong>of</strong> Gualala & North Fork<br />
Gualala rivers, ca. 6.5 km E <strong>of</strong> town <strong>of</strong> Gualala, 3<br />
Aug 1981, G. L. Smith & C. R. Wheeler 7288 (CAS,<br />
2 sheets); Bear Creek <strong>of</strong>f Usal Creek, 1500 ft, in<br />
shade, in ravine on SW slope in redwood tan oak,<br />
madrono DF, logged forest, 3 Aug 1959, fl, L. C.<br />
Wheeler 7626 (F); 1875, G. R. Vasey, s.n. (US, 2<br />
sheets); Redwood Highway – Rockport Road, 2<br />
mi W <strong>of</strong> S fork <strong>of</strong> Eel River, 300 ft, 30 Jun 1934,<br />
C. B. Wolf 5831 (GH, TEX). Monterey Co.,<br />
Jamesburg, W <strong>of</strong> Church Creek divide, 3700 ft, 25<br />
Oct 1932, R. St. John 312 (UC); headwaters <strong>of</strong><br />
Arroyo Seco, Santa Lucia Mountains, 25 Oct 1930,<br />
H. L. Mason 5759 (UC). Napa Co., near Haven<br />
Place, above St. Helena, 10 Sep 1967, F. R. Fosberg<br />
48647 (US); Howell Mountain, beds <strong>of</strong> Moore’s<br />
Creek, 1500 ft, 6 Jul 1899, J. P. Tracy 416 (UC);<br />
Bothe Napa Valley State Park, Richey Creek, at<br />
the bottom <strong>of</strong> stream right near water, moist <strong>and</strong><br />
rocky area, occasional, plant 1-1.5 m tall, common<br />
name as alder-clover by the rangers, 13 Sep 1990,<br />
fr, J. Wen 752 (OS); Bothe Napa Valley State Park,<br />
Upper Valley Creek, at bottom <strong>of</strong> stream, abundant,<br />
13 Sep 1990, fr, J. Wen 753 (OS). Nevada Co.,<br />
Alpha Road 1.5 miles south <strong>of</strong> Washington, 3300<br />
ft, shady north slope, yellow pine forest, 23 Aug<br />
1965, fr, G. H. True & J. T. Howell 2453 (CAS).<br />
Orange Co., Silverado Canyon, Santa Ana Mts.,<br />
Clevel<strong>and</strong> National Forest, Trabuco District, 4200<br />
ft, E. W. Lathrop 7656 (UC); Trabuco, 3800 ft, 9<br />
Oct 1912, fr, H. H. Smith 5357 (F). Placer Co.,<br />
Canyon Creek, Dutch Flat, 14 Jul 1934, fl, F. G.<br />
MacFadden s.n. (NY). Plumas Co., Mill Creek,<br />
Jun 1877, fl, R. M. Austin s.n. (NY); Mill Creek,<br />
Sep 1896, R. M. Austin 512 (US); canõn just W <strong>of</strong><br />
Quiney, 3700 ft, 4 Aug 1912, H. M. Hall 9383 (UC,<br />
2 sheets); Downieville, 1 mi S <strong>and</strong> 1/3 mi W <strong>of</strong><br />
Nelson Pt., 4300 ft, E. Nourse 34 (UC); Forest<br />
Lodge, 3400 ft, 2 Aug 1941, H. S. Reed s.n. (UC);<br />
1 st Water Trough Creek, N Greenville, 4000 ft, with<br />
Acer, Alnus <strong>and</strong> Cornus, occasional, 24 Jul 1931,<br />
fl, L. W. Swift 40 (CAS). San Bernardino Co., Lyrte<br />
Creek – Falls, 25 Jul 1901, fl, L. R. Abrams 1954<br />
(NY); S. B. Parish, s.n. (UC); San Bernardino Mts.,<br />
Jul 1881, fl & young fr, S. B. & W. F. Parish 437<br />
(F, NY, US); San Gabriel Mts., Angeles National<br />
Forest, ca. 1 mi up Icehouse Canõn, ca. 5700 ft, R.<br />
F. Thorne 35425 (UC); on La Porte Rd., 10 km S<br />
<strong>of</strong> jct with Hwy 70, 39 o 53’N, 120 o 52’W, 1400 m,<br />
associated with Acer, Alnus, Pseudotsuga, <strong>and</strong><br />
Pinus, growing at edge <strong>of</strong> creek in pine forest,
hizomatous herb ca. 1.6 m tall, 3 Aug 1994, young<br />
fr, J. B. Walker et al. 872 (NY). San Diego Co., in<br />
ravine on S side <strong>of</strong> Pauma Creek, Palomar<br />
Mountain, 33 o 21’N, 116 o 56’W, 4200 ft, 30 Jul<br />
1968, C. V. Meyer s.n. (UC, US, 2 sheets). San<br />
Mateo Co., woodside, 4 Jul 1919, fl, E. Walther<br />
s.n. (CAS). Santa Clara Co., Loma Prieta Peak,<br />
Aug 1903, young fr, A. D. E. Elmer 4987 (CAS,<br />
MO, NY, UC, US); foothills <strong>of</strong> the eastern side <strong>of</strong><br />
the Santa Cruz Mountains, 5 mi S <strong>of</strong> center <strong>of</strong> Palo<br />
Alto, along Los Trancos Creek, moist canyon in<br />
foothill woodl<strong>and</strong>, 600-1800 ft, 4 Jul 1974, R.<br />
Martineau 359 (DS); Loma Prieta, Santa Cruz<br />
Mountains, 3200 ft, 15 Aug 1939, E. H. Nelson<br />
s.n. (UC, 2 sheets); the Santa Cruz Peninsula,<br />
Steven’s Creek, 10 Jul 1907, J. D. R<strong>and</strong>all 6 (DS,<br />
2 sheets, MU); 2 mi S <strong>of</strong> Saratoga Springs, ca. 1000<br />
ft, 3 Jul 1966, L. S. Rose 66045 (AUT, ENCB, GH);<br />
eastern slope <strong>of</strong> Mount Hamilton, 3300 ft, 22 May<br />
1936, H. K. Sharsmith 3721 (UC, 2 sheets), 3000<br />
ft., 3883 (UC). Santa Cruz Co., Redwood forest,<br />
J. Ball s.n. (US); Waddell Creek, 23 Sep 1901, W.<br />
R. Dudley s.n. (DS); near Glenwood, Santa Cruz<br />
Mtns, bottom <strong>of</strong> shady ravine, 4 Jun 1931, fl, F. R.<br />
Fosberg S5286 (NY); Boulder Creek, 16 Oct 1938,<br />
L. S. Rose 38304 (B, 2 sheets, MO, 3 sheets, UC, 2<br />
sheets); Big Basin Redwoods State Park, along<br />
Opal Creek, 30 Sep 1950, J. H. Thomas 2439<br />
(MU); 1.3 mi below Empire Grade on Alba Rd.,<br />
1800 ft, J. H. Thomas 3467 (DS, 2 sheets, UC);<br />
Big Basin Redwoods State Park, near a stream<br />
close to the Ranger Station, shady area, plant ca. 2<br />
m tall, 17 Sep 1990, J. Wen 774 (OS); along Empire<br />
Grade Rd., ca. 1 mi NMW <strong>of</strong> Santa Cruz (jct. <strong>of</strong><br />
Empire Grade Rd. <strong>and</strong> Hellier Ave.), roadside <strong>and</strong><br />
near a ravine, plant ca. 1.5 m tall, 17 Sep 1990, J.<br />
Wen 775 (OS); along SR 9, 3.5 mi from Santa Cruz<br />
toward Felton, on hillside <strong>of</strong> Redwood forest,<br />
shady, plant ca. 1.5 m tall, 17 Sep 1990, J. Wen<br />
776 (OS). Shasta Co., Shasta National Forest,<br />
along Castle Creek ca. 1 mi W <strong>of</strong> Castle Crags<br />
State Park, 21 Sep 1972, R. F. Thorne et al. 42376<br />
(BM, ENCB, KLU, MO, UNCC). Sierra Co.,<br />
1875, J. G. Lemmon 564 (GH); at Indian Hill loop,<br />
ca. 4 mi NE <strong>of</strong> Camptonville, 3100 ft, 20 Aug 1982,<br />
fl, M. S. Taylor & R. Wessel 5028 (CAS); northern<br />
Sierra Nevada, north fork <strong>of</strong> Yuba River, 5 miles<br />
above Downieville, 3500 ft, arid transition –<br />
Pseudotsuga, wet slopes, rocky loam, shade,<br />
perennial 6 ft, spread 6 ft, 6 Aug 1937, young fr,<br />
C. B. Wolf 9047 (NY, 2 sheets). Siskiyou Co., 6<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
41<br />
mi W from Red Bank F. S. Camp, on road to<br />
Somesbar from Sawyers Bar, 2000 ft, 24 Oct 1959,<br />
P. C. Everett et al. 23942 (UC); near Cantarra, 10<br />
Jul 1905, L. Krautter s.n. (BM, 2 sheets); along<br />
Cade Creek near its mouth, 3 miles east <strong>of</strong> Happy<br />
Camp, 8 Jul 1940, fl, M. Ownbey & F. G. Meyer<br />
2221 (GH, MO, 2 sheets, NY); Klamath National<br />
Forest, along Indian Creek Rd., 8.2 miles NW <strong>of</strong><br />
Happy Camp, dry habitat, granite bedrock, plant<br />
1-1.5 m tall, ca. 1 m in spread, 16 Sep 1990, fr, J.<br />
Wen 771 (OS); along SR 96, 4.4 miles E <strong>of</strong> Happy<br />
Camp, near a small creek, locally abundant, plant<br />
1.5-2 m tall, 16 Sep 1990, J. Wen 772 (OS); Seiad<br />
Valley, along SR 96, ca. 2.5 miles E <strong>of</strong> Seiad Creek<br />
between milestones 63 <strong>and</strong> 64, in a small creek,<br />
poor soil, rocky area, 16 Sep 1990, fr, J. Wen 773<br />
(OS); gravel bar in S fork <strong>of</strong> Salmon R, 9.5 mi<br />
above Cecilville, 3500 ft, 22 Jul 1955, fl, I. L.<br />
Wiggins 13455 (NY). Sonoma Co., Stewart Pt.,<br />
Jul 1923, A. Griffin s.n. (CAS). Trinity Co., Trinity<br />
Mountains, Dedrick, 2500 ft., 13 Aug 1948, A. M.<br />
Alex<strong>and</strong>er et al. 5480 (UC); creek bank 30 miles<br />
west <strong>of</strong> Weaverville, 1500 ft, 18 Jul 1930, fl, L.<br />
Benson 2196 (NY); North Fork Trinity River, Hobo<br />
Gulch Camp <strong>and</strong> vicinity, 18 miles NW <strong>of</strong><br />
Weaverville, near mouth <strong>of</strong> Keystone Meadows<br />
Gulch, moist shaded creek bank, 2950 ft, 13 Aug<br />
1971, fl, E. Carter 331 (CAS). Tehama Co., <strong>of</strong>f<br />
Hwy 32, Potato Patch Campground in Lassen<br />
National Forest, beside Deer Creek in shady mixed<br />
deciduous & coniferous forest, 24 Jun 2003, J. Wen<br />
7104 (US). Tulare Co., Sequoia National Park, 24<br />
Aug 1966, fr, J. Redden s.n. (ASU). Yuba Co.,<br />
near Slate Creek, yellow pine forest, 1200 m, 25<br />
Jul 1980, fl, L. Ahart 2483 (CAS). Bolmer Bay, 9<br />
Apr 1854, J. M. Bigelow s.n. (NY). Upper<br />
Sacramento, Wilkes Expedition s.n. (US, 2 sheets).<br />
OREGON: Curry Co., mouth <strong>of</strong> Waters Creek,<br />
near Agness, 22 Aug 1938, G. M. Hansen s.n. (UC);<br />
along Cheteu River 7 mi above Harbor, 19 Jul<br />
1919, fl, M. E. Peck 8914 (GH, MO, NY). Douglas<br />
Co., east fork <strong>of</strong> north fork <strong>of</strong> Umpqua River, 6-10<br />
miles east <strong>of</strong> Peel, 1500 ft, 14 Jul 1898, fl, E. I.<br />
Applegate 2703 (NY, US, 2 sheets); Umpqua<br />
National Forest, North Umpqua River road at<br />
Fairview Creek, 1115 ft, 20 Jul 1950, E. C. Earle<br />
4596 (PH); Coast Range, along Umqua River, 10<br />
Aug 1880, G. Engelmann s.n. (MO); West Fork,<br />
Cow Creek Canyon, 9 Aug 1898, F. A. Walpole 57<br />
(US). Jackson Co., Sykes Creek, near Wimer, 12<br />
Jul 1892, E. W. Hammond 168 (MO, NY, US).
42<br />
Josephine Co., Steve Peak, Siskiyou Mountains,<br />
Applegate River watershed, E. I. Applegate 6528<br />
(DS); south slope <strong>of</strong> Sexton Mt., 21 Aug 1928, N.<br />
P. Gales 325 (MO, 2 sheets); near McMullin Creek<br />
on McMullin Creek Rd., 6.5 mi E <strong>of</strong> US 199 <strong>and</strong><br />
Selma, 7 mi N <strong>of</strong> Cave Creek jct., 2 Aug 1997, fl<br />
& young fr, W. Hess et al. 7817 (F, NY); Grayback<br />
Forest Camp, 8 mi west <strong>of</strong> Oregon Caves National<br />
Monument, 6 Jul 1939, fl, C. L. Hitchcock & J. S.<br />
Martin 5191 (NY); Siskiyou Mountains, Jun 1884,<br />
T. Howell s.n. (LE); along streams near Oregon<br />
Caves, Siskiyou Mts., 6000 ft, 31 Jul 1935, fl, J.<br />
W. Thompson 12422 (GH, NY, UC, US); along SR<br />
46, 0.2 mi E <strong>of</strong> Grayback Creek (or 10.2 miles E<br />
<strong>of</strong> Cave Jct.), on the bed <strong>of</strong> a small creek, limestone,<br />
15 Sep 1990, fr, J. Wen 763 (OS); along SR 46, on<br />
hillside between milestones 12 <strong>and</strong> 13, near a small<br />
ravine, locally abundant, about 100 individuals in<br />
the ravine, 1.5-2.5 m tall, fruits dark purple to<br />
black, styles connate at base, 16 Sep. 1990, fr, J.<br />
Wen 764 (OS); along SR 46, on hillside between<br />
milestones 16 <strong>and</strong> 17, locally abundant, 16 Sep<br />
1990, fr, J. Wen 765 (OS); Oregon Caves, near a<br />
small creek, plant 1.5-2 m tall, 16 Sep 1990, fr, J.<br />
Wen 766 (OS). Lane Co., Salt Creek, forest above<br />
Oakridge, Jul 1922, L. Constance s.n. (UC); Gold<br />
Hill, deep canyon, moderately moist, dense shade,<br />
SE slope, coniferous forest, Pseudotsuga-Acer<br />
circinatum-Corylus, 8 Sep 1962, fl, L. E. Detling<br />
9040 (RSA). Linn Co., along US hwy 20, 32 mi<br />
W <strong>of</strong> junction with state hwy 22, 1000 ft, perennial<br />
to 3 m tall, 24 Jul 1962, fl, A. S. Barclay et al.<br />
1456 (LL); western foothills <strong>of</strong> the Cascade Range<br />
on the N side <strong>of</strong> Green Peter Reservoir, along<br />
Quartzville Road 4.4 miles NE <strong>of</strong> the Thistle Creek<br />
Boat Ramp, flowers white, mature fruits black<br />
purple, perennial 2.2 m tall, aromatic, on roadside<br />
with Alnus, Hypericum, Pteridium, Pseudotsuga,<br />
Polystichum, <strong>and</strong> Agrostis, 317 m, 17 Aug 1999, fl<br />
& fr, R. R. Halse 5634 (NY, 2 sheets, RSA); near<br />
Cascadia, 22 Sep 1952, J. Seibert s.n. (GH).<br />
Multnomah Co., Portl<strong>and</strong>, Foley Springs Road,<br />
moist creek banks, 16 Jul 1903, M. W. Gorman<br />
1668 (US).<br />
Cultivated material: U.S.A. CALIFORNIA:<br />
Alameda Co., Berkeley, the garden <strong>of</strong> Anson <strong>and</strong><br />
Anita Blake, 29 Jun 1941, N. F. Bracelin 1439 (L);<br />
13 Sep 1942, N. F. Bracelin 2290 (L, NY);<br />
2. <strong>Aralia</strong> racemosa L. — Fig. 18; color plate 2:<br />
A-F.<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
<strong>Aralia</strong> racemosa L., Sp. Pl.: 273. 1753. TYPE:<br />
U.S.A. VIRGINIA: Herb. Linn. 394.6<br />
(lectotype: LINN!, designated by Wen <strong>and</strong><br />
Reveal 1992).<br />
<strong>Aralia</strong> racemosa var. foliosa Vict. & J. Rousseau,<br />
Contr. Inst. Bot. Univ. Montréal 36: 37. 1940.<br />
<strong>Aralia</strong> racemosa L. f. foliosa (Vict. & J.<br />
Rousseau) Scoggan, Fl. Canada 1: 52. 1978.<br />
TYPE: Canada. QUÉBEC: Lac-Saint-Jean<br />
Co., Pointe-Bleue, à découvert sur les berges<br />
calcaires du lac Saint-Jean, élément du taillis<br />
de Sambucus, Viburnum, etc, 25 Jul 1935,<br />
Marie-Victorin, Roll<strong>and</strong>-Germain & Meilleur<br />
44562 (MT!).<br />
Perennial herb, 0.6-2.5 m tall with large<br />
aromatic roots. Stem pilose, dark purple to<br />
greenish purple, with 4-5 leaves; rhizomes thick<br />
<strong>and</strong> horizontal. Leaves 50-83 cm long, 54-87 cm<br />
wide, ternately compound, spreading, purple at<br />
nodes; stipules 1.2-1.5 cm long, 0.4-0.9 cm wide,<br />
narrowly triangular to lanceolate, not leafy,<br />
glabrous, ciliate at margin, thick papery; petioles<br />
15-30 cm long, purple; leaflets 7-20.5 cm long,<br />
4.5-14 cm wide, thin papery, the pair <strong>of</strong> leaflets<br />
below the terminal leaflet ovate, other leaflets ovate<br />
to broadly so, acute at apex, subcordate to cordate<br />
at base, commonly oblique on lateral leaflets,<br />
doubly serrate at margin; lateral veins 8-9 on each<br />
side, conspicuous on both surfaces, leaflets green,<br />
nearly glabrescent on upper surface, pilose on<br />
lower surface, petiolules 0-3 cm long, pilose.<br />
Inflorescence terminal <strong>and</strong> axillary, pubescent;<br />
axillary inflorescences 6-21 cm long; terminal<br />
inflorescence 15-40 cm long, 3-10 cm wide,<br />
consisting <strong>of</strong> numerous umbels, primary<br />
inflorescence branches 15-25, racemosely arranged<br />
on a main axis, <strong>of</strong>ten 3-5 forming a circle on the<br />
upper part <strong>of</strong> the main inflorescence axis, each<br />
primary branch 3-14.5 cm long, consisting <strong>of</strong> 1-<br />
10 umbels, with a few lateral umbels bearing no<br />
fruits (functionally male umbels); terminal umbels<br />
20-25-flowered, pedicels 6-8 mm long, pubescent,<br />
tip enlarged conspicuously at anthesis, somewhat<br />
slightly enclosing the base <strong>of</strong> the hypanthium;<br />
lateral umbels 10-20-flowered, pedicels 3-5 mm<br />
long; bracts <strong>of</strong> primary branches 3-5 mm long, 0.5-<br />
1 mm wide, linear, more or less pilose, ciliate at<br />
margin; bracteoles 1.0-1.5 mm long, 0.2-0.3 mm<br />
wide, linear, ciliate at margin. Sepals 0.2-0.3 mm<br />
long <strong>and</strong> wide, triangular, glabrous, persistent on
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
Fig. 18. <strong>Aralia</strong> racemosa L. A. Habit with a leaf <strong>and</strong> a terminal <strong>and</strong> an axillary inflorescence. B. Root <strong>and</strong> rhizome. C. Leaf<br />
base showing triangular stipules. D. Flower. E. Fruit.<br />
fruits; petals 1.4-1.6 mm long, 0.9-1.1 mm wide,<br />
ovate, greenish white to white, with a conspicuous<br />
vascular bundle in the middle, spreading to<br />
recurved at anthesis; stamens 5, erect at anthesis,<br />
filaments 1.3-1.5 mm long, anthers 0.4-0.5 mm<br />
long, 0.2-0.3 mm wide, oblong, whitish yellow;<br />
ovaries 5-locular, styles 0.6-0.7 mm long at<br />
anthesis, base <strong>of</strong> styles more or less enlarged,<br />
43<br />
appearing like a stylopodium. Fruits 4-4.5 mm<br />
long, 3.5-4.5 mm wide, globose, dark purple to<br />
purple, turning dark maroon-red when mature,<br />
persistent styles divided to the middle, 1.3-1.5 mm<br />
long. Seeds 2-2.2 mm long, 1-1.2 mm wide, 0.2-<br />
0.25 mm thick, kidney-shaped, whitish gray,<br />
smooth.
44<br />
Common names: spikenard, American<br />
spikenard, Indian root, spignet, spiceberry, pettymorrel,<br />
life-<strong>of</strong>-man, <strong>and</strong> old man’s root.<br />
Uses: aromatic roots used in syrup <strong>of</strong> spikenard,<br />
a tonic, or for cough <strong>and</strong> irritation <strong>of</strong> the bronchopulmonary<br />
tract.<br />
Phenology: flowering in June to August;<br />
fruiting in August to October.<br />
Distribution: widely distributed in eastern<br />
U.S.A. <strong>and</strong> Canada, westward to South Dakota <strong>and</strong><br />
North Dakota, south to Georgia <strong>and</strong> Alabama (Fig.<br />
19).<br />
Ecology: rich deciduous forests, wooded<br />
slopes, ravines, moist bluffs, riverside forests,<br />
creeksides, <strong>and</strong> streamsides; 100-1500 m.<br />
Representative specimens examined: Canada.<br />
Georgeville, 18 Aug 1886, J. R. Churchill s.n.<br />
(MO). NEW BRUNSWICK: ca. 20 mi NE <strong>of</strong><br />
Fredericton, 10 Aug 1955, H. J. Scoggan 12795<br />
(W); Hillsborough, ca. 15 miles SSE <strong>of</strong> Moncton,<br />
18 Aug 1955, H. J. Scoggan & D. Erskine 12953<br />
(W). Kent Co., Bass River, 1868, J. Fowler s.n.<br />
(F). NOVA SCOTIA: Hants Co., Gypsum Cliff,<br />
Five Mile River, 1 Sep 1927, fr, A. R. Prince & C.<br />
E. Atwood 764 (DS). ONTARIO: Bruce Peninsula,<br />
Stokes Bay, mixed rocky woods, 9 Jul 1934, fl, P.<br />
V. Krotkov 9263 (NY, US). Carleton Co., Ottawa<br />
Dist., Taylor Hill, 25 Jul 1953, L. Jenkins 3227<br />
(W); Ile Bizard, 4 Aug 1929, P. Louis-Marie s.n.<br />
(CAS, W). Hastings Co., abundant in scrub<br />
hardwood, N side <strong>of</strong> highway, 1 mi W <strong>of</strong> Coe Hill,<br />
25 Aug 1959, fr, C. E. Heidenreich 437 (MO).<br />
Kingston, Muskoka, Aug 1893, fl, J. Fowler s.n.<br />
(MO); Kingston, 25 Jul 1893, fl, J. Fowler s.n.<br />
(US). Leeds Co., Rideau Park, Rideau Ferry, Aug<br />
1898, young fr, T. W. Edmondson 1333 (NY).<br />
Swamp, Golden Lake, 27 Jul 1899, fl, L. M.<br />
Umbach s.n. (F). Near Sucker Lake, 4 Aug 1904,<br />
W. F. Wright 163 (US), 166 (2 sheets, US). Thunder<br />
Bay Dist., N48 o 12’, W90 o 05’, opposite gravel pit,<br />
3.5 km S on East Arrow Lake Road, Hardwick<br />
Twp., 19 Jul 1988, young fr, C. E. Garton 24121<br />
(NY). QUÉBEC: Argenteuil Co., Carillon, along<br />
rt 344, just E <strong>of</strong> Carillon, N45 o 33.832’,<br />
W74 o 22.094’, 51 m, 11 Aug 2008, fl, herb 1-1.5 m<br />
tall, petals white, J. Wen 10489 (US). Bonaventure<br />
Co., ravine, bluffs, west <strong>of</strong> Carleton, 26 Jul 1904,<br />
fl, J. F. Collins & A. S. Pease s.n. (CAS); talus<br />
decouverts sur les bords de la riviere Restigouche,<br />
1 Aug 1936, fl, F. Marie-Victorin et al. 48990 (F,<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
NY, US). Brome Co., Glen Sutton, talus dominant<br />
un ruisseau, avec Rubus odoratus, 9 Aug 1942, fl<br />
& young fr, F. F. Marie-Victorin et al. 56252 (US).<br />
Cabano, edge <strong>of</strong> woods, 6 Aug 1956, floral buds,<br />
H. J. Scoggan 13255 (NY). Gatineau Co., Ottawa<br />
Dist., Hull Twp., Gatineau Park, E <strong>of</strong> Ridge Rd.,<br />
23 Jul 1941, fl, H. A. Senn et al. 1073 (MO).<br />
Labelle Co., Belleville, 10 Jul 1942, F. F. Lucien<br />
& Louis-Marie 1202 (CAS); Frog Lake, Blue Sea<br />
Lake, 17 Aug 1922, M. O. Malte 700/22 (W);<br />
Nomingue, forêt des Laurentides, 2 Aug 1947, F.<br />
Roll<strong>and</strong>-Germain 3200 (DS, F, NY, U, US, W);<br />
Nomingue, 24 Jul 1929, fl, E. Roy 1203 (US).<br />
Montreal Isl<strong>and</strong> Co., Montreal, 12 Aug 1882, C.<br />
Mohr s.n. (US). Papineau Co., Montebello, along<br />
an unpaved road, just N <strong>of</strong> rt 148, between rt 323<br />
<strong>and</strong> rt 321, ca 0.5 km west <strong>of</strong> the jct with rt 323,<br />
near Papineauville secteur Sainte-Angélique,<br />
N45 o 39.988’, W74 o 57.842’, 168 m, 11 Aug 2008,<br />
herb ca 1 m tall <strong>and</strong> 1 m in diameter, stem <strong>and</strong><br />
petiole purple, J. Wen 10482 (US); near the county<br />
line <strong>of</strong> Les Laurentides Co., along local rt 7, ca 5<br />
km NE <strong>of</strong> Duhamel, N46 o 05.750’, W75 o 08.727’,<br />
229 m, 11 Aug 2008, fl, herb ca 1.5 m tall, petals<br />
white, anthers white, J. Wen 10483 (US). Rouville<br />
Co., NE corner <strong>of</strong> Yamaska Mountain, 960-1180<br />
ft, dry-mesic deciduous forest <strong>of</strong> Acer saccharum<br />
– Ostrya virginiana – Tilia americana, 3 Aug 1962,<br />
fl, A. Auclair & A. Walther 57 (NY). Pickanock,<br />
Quebec, 16 Aug 1894, fr, J. Macon s.n. (US).<br />
U.S.A. ALABAMA: Marshall Co., shaded cool<br />
moist s<strong>and</strong>y ravine by trail to boat l<strong>and</strong>ing on<br />
Reservoir, Bucks Pocket State Park, 10 Aug 1973,<br />
fl, R. Kral 51097 (MO). ARKANSAS: Newton<br />
Co., narrow ravine with extensive s<strong>and</strong>stone<br />
exposures along upper reaches <strong>of</strong> Terrapin Branch,<br />
700-800 ft, scattered along lower slope, 24 Jul<br />
1971, fl, P. L. Redfearn Jr & W. Weber 27516<br />
(NCU). Pope Co., Nogo, 9 Jul 1941, G. M. Merrill<br />
583 (TEX). CONNECTICUT: Fairfield Co.,<br />
Bridgeport, 5 Aug 1886, fl, A. L. Winton s.n. (YU).<br />
Litchfield Co., Washington, 18 Aug 1919, young<br />
fr, A. W. Eames s.n. (YU); Kent, Macedonia Brook<br />
State Park, in woods, 23 Aug 1958, young fr, J.<br />
Ebinger 857 (YU); Woodbury, 30 Sep 1905, fr, dry<br />
rocky woods, Eames & Godfrey 269 (YU);<br />
Salisbury, Tom’s Mountain, woody roadside, 27<br />
Aug 1915, fl & young fr, C. C. Godfrey s.n. (YU).<br />
Middlesex Co., Chester, thicket, 1 Aug 1932, fl,<br />
R. H. Burton s.n. (YU); Killingworth, 5 Sep 1932,<br />
fl & young fr, E. B. Harger s.n. (YU); East
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
Fig. 19. Map <strong>of</strong> North America showing the distribution <strong>of</strong> <strong>Aralia</strong> racemosa L.<br />
Haddam, 16 Aug 1923, fl, J. F. Smith s.n. (YU);<br />
town <strong>of</strong> Killingworth, rich shaded soil by small<br />
pond NW <strong>of</strong> junction <strong>of</strong> routes 80 <strong>and</strong> 81, 28 Jun<br />
1975, G. C. Tucker 36 (YU). New Haven Co.,<br />
Waterbury, rich woods, 8 Aug 1908, young fr, A.<br />
E. Blewitt 2587 (YU); Milford, Meadows End, 27<br />
Aug 1909, H. S. Clark s.n. (YU); Meriden, Aug<br />
1856, young fr, D. C. Eaton s.n. (YU); New Haven,<br />
D. C. Eaton s.n. (LE); New Haven, 5 Aug 1873, fl,<br />
G. W. Harris s.n. (YU); New Haven, 1874, G. R.<br />
Kleeberger s.n. (CAS). New London Co., Franklin,<br />
Dragon’s Den, 27 Aug 1885, fr, G. Waldo s.n. (YU).<br />
Toll<strong>and</strong> Co., near the town <strong>of</strong> Willington, along rt.<br />
45<br />
320, ca. 1 mi S <strong>of</strong> jct with rt 74, at the edge <strong>of</strong><br />
deciduous forest, 14 May 1991, J. Wen 796 (A).<br />
DELAWARE: ex Herb. W. M. Canby s.n. (P).<br />
DISTRICT OF COLUMBIA: 26 May 1880, E. S.<br />
Burgess s.n. (NY). GEORGIA: Murray Co.,<br />
Cohutta Wildlife Management Area, ca. 8 mi E <strong>of</strong><br />
Eton, along West Cowpens Rd. to FS 17, NE <strong>of</strong><br />
Grassy Mountain, stem red, infrequent, 10 Sep<br />
1982, in late fl., N. C. Coile 3376 (NCSC); moist<br />
s<strong>and</strong>y loam under white pine – hemlock, ravine<br />
drainage north to damsite, Carters Dam area, 16<br />
Jul 1973, fl, R. Kral 50610 (MO). Rabun Co.,<br />
northern flank <strong>of</strong> Big Mt. South <strong>of</strong> Rock Gorge,
46<br />
along jeep trail, 1850 ft., 21 Jul 1968, in fl., D. M.<br />
DuMond 1307 (NCSC); Patterson Gap Road above<br />
Moon Valley, moist edge <strong>of</strong> woods <strong>and</strong> road<br />
embankment at edge <strong>of</strong> Happins-so property, 13<br />
Jul 1991, fl, Stiles 409 (MO). Union Co., Sosebee<br />
Cove, along rocky stream, on slope <strong>of</strong> Sosebee<br />
Cove, scattered, 11 Jun 1991, J. Wen et al. 982<br />
(A). White Co., 4 miles N <strong>of</strong> Helen along rt 75, at<br />
the base <strong>of</strong> a big s<strong>and</strong>stone rock, moist area, plant<br />
1.9 m tall, 5 Sep 1986, fr, J. Wen 148 (OS); 7 miles<br />
N <strong>of</strong> the town Helen along Rt 75, 5 Sep 1986, fr,<br />
fruits purple, J. Wen 149 (OS). ILLINOIS: Beverly<br />
Hills, 24 Sep 1903, fr, R. Bebb 1572 (F). Highl<strong>and</strong><br />
Park, W. C. Ohlendorf 4154 (F). Ravinia, Silva<br />
Ravinia, N Chicago area, 4 Sep 1905, fr, F. C. Gates<br />
864 (F); Skokie Marsh, Ravinia, 24 Aug 1911, fr,<br />
E. E. Sherff s.n. (F). Bureau Co., vic. De Peu, along<br />
rte. 29, about 5 mi N <strong>of</strong> jct. <strong>of</strong> Rte 29 <strong>and</strong> rte 26,<br />
abundant on wooded slope with rich black soil, 4<br />
Sep 1981, fr, D. D. Soejarto 5446 (F). Carroll Co.,<br />
along Sunset Trail in woods, Mississippi Palisades<br />
State Park, 30 Aug 1955, J. W. Thieret 1578 (F).<br />
Cook Co., Palos Twp., Swallow Cliffs Forest<br />
Preserve, SW <strong>of</strong> jct US Hwy 45 <strong>and</strong> State Hwy 83,<br />
scarce in transitional oak-maple forest at slope<br />
crest, S <strong>of</strong> toboggan run, 22 Aug 1997, young fr, T.<br />
G. Lammers 10297 (F). Gallatin Co., the Pounds,<br />
SSE <strong>of</strong> Gibsonia, base <strong>of</strong> cliff in woods, 27 Aug<br />
1956, J. W. Thieret 2427 (F). Grundy Co., Moris,<br />
Mason Creek, between Gooselake Praire preserve<br />
<strong>and</strong> Moris, a clump <strong>of</strong> 30-50 plants, 23 Jul 1980,<br />
fl, D. D. Soejarto 5133 (F). Jackson Co., on moist<br />
s<strong>and</strong>stone ledge, 30 Jul 1967, young fr, R. T.<br />
Johnson 419 (NY). Jo Daviess Co., in woods N <strong>of</strong><br />
dirt road, NE corner <strong>of</strong> sec 28, Derinda Twp., 4.5<br />
miles N <strong>of</strong> north section <strong>of</strong> Mississippi Palisades<br />
State Park, 30 Aug 1955, J. W. Thieret 1590 (F).<br />
Kane Co., Elgin, Trout Park Woods, wet to swampy<br />
forest floor, rich woods, 23 Aug 1979, fl, D. D.<br />
Soejarto 5034 (F, 2 sheets); Elgin, 20 Aug 1918,<br />
young fr, H. C. Benke 362 (F). Lasalle Co., the<br />
Illinois State Park at Starved Rock, May-Sep 1921,<br />
F. Thone 97 (F). Peoria Co., NW <strong>of</strong> Mossville,<br />
wooded hillside, 4 Sep 1949, V. H. Chase 10781<br />
(F, 3 sheets); Peoria Heights, 17 Aug 1953, young<br />
fr, V. H. Chase 13596 (LL). Tazewell Co., 2 mi E<br />
<strong>of</strong> Upper Bridge (Peoria), 5 Sep 1920, V. H. Chase<br />
3455 (DS, W). Union Co., 27 Jul 1878, fl, G. H.<br />
French s.n. (US). INDIANA: Bluffton, 25 Aug<br />
1897, fl, C. C. Deam s.n. (F). Jennings Co., rich<br />
woods, 6 Aug 1932, fl, S. McCoy 585 (NY).<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
Laporte Co., Michigan City, thickets, 15 Jul 1903,<br />
C. D. Mell 130 (US). Martin Co., bluffs <strong>of</strong> White<br />
River near point locally known as Old Man’s Nose,<br />
6 miles NE <strong>of</strong> Shoals, dry, sunny, 22 Aug 1935, fr,<br />
R. M. Tryon Jr. 1935 (F). Monroe Co., near a small<br />
stream, 14 miles southeast <strong>of</strong> Bloomington, 18 Jul<br />
1946, fl, V. Foley s.n. (TEX). Porter Co., in woods<br />
near Long Lake near the town <strong>of</strong> Black Hawk<br />
Beach, 14 Jul 1952, floral buds, F. A. Swink 1465<br />
(F); in shaded ground <strong>of</strong> a decadent red mapleyellow<br />
birch bog between Route 20 <strong>and</strong> the<br />
Baltimore & Ohio Railroad, just E <strong>of</strong> County Line<br />
Road SW <strong>of</strong> Ogden Dunes, 27 Jul 1952, fl, F. A.<br />
Swink 1587 (F). Wells Co., Harrison Twp., in<br />
woods, 29 Aug 1905, fl, C. C. Deam 495 (US).<br />
IOWA: Black Hawk Co., red oak forest, 9 Jul 1929,<br />
fl, M. Burk 505 (MO, 2 sheets). Clayton Co., deep<br />
upl<strong>and</strong> woods at Pike’s Peak McGregor, 18 Jul<br />
1927, fl, B. Shimek s.n. (MO); upl<strong>and</strong> woods, Pike’s<br />
Peak McGregor, 8 Aug 1921, fr, B. Shimek s.n.<br />
(US). Dickinson Co., wooded bluff, west side <strong>of</strong><br />
Little Sioux River, 3.5 mi north <strong>of</strong> west from<br />
Milford, 1 Jul 1918, fl, B. Shimek s.n. (MO);<br />
ungrazed woods along Little Sioux River, 2 miles<br />
W <strong>of</strong> Milford, Okoboji Twp., 7 Aug 1953, young<br />
fr, R. F. Thorne 13220 (US). Fayette Co., spreading<br />
herb 3-4.5 ft tall, 15 Jul 1894, fl, B. Fink 250 (US,<br />
on the same sheet with nearly mature fruits<br />
collected on 31 Jul 1894). Hardin Co., Aug 1876,<br />
young fr, M. E. Jones s.n. (NY). Van Buren Co.,<br />
Bentonsport, rich woods, Aug 1926, fr, E. W.<br />
Graves s.n. (MO). KANSAS: Doniphan Co., 5<br />
miles north <strong>and</strong> 2 miles east <strong>of</strong> Troy, thicket, rich<br />
moist loam, scattered in local colonies, 3 Sep 1952,<br />
fr, W. H. Horr 4243 (LL, NY). KENTUCKY:<br />
Barren Co., deciduous woods, Bonayre Forest, ca.<br />
5 mi W <strong>of</strong> Glasgow, 29 Sep 1970, K. A. Nicely<br />
3126 (NCSC). Carter Co., Carter Caves State Park,<br />
E end <strong>of</strong> bridge over Tygarts Creek, NW exposure,<br />
Ky 182, Cumberl<strong>and</strong> Plateau Province, 1 Jul 1961,<br />
floral buds, E. M. Browne & E. T. Browne Jr 4417<br />
(NCU). Edmondson Co., Chameleon Springs, 13<br />
Aug 1899, fr, S. F. Price s.n. (MO). Estill Co., 5<br />
mi NW <strong>of</strong> Irvine, 14 Aug 1939, fr, M. E. Wharton<br />
5177a (MO). Harlan Co., on Big Black Mountain,<br />
Aug 1893, T. H. Kearney Jr. 148 (F, 2 sheets, MO,<br />
NY, US). Hart Co., Big Woods, mesophytic woods,<br />
8 Sep 1940, fr, E. L. Braun 3579 (US). Lewis Co.,<br />
rich wooded north slope 1 mile east <strong>of</strong> Vanceburg,<br />
1 Aug 1939, young fr, M. E. Wharton 4992 (NCU).<br />
Menifee Co., Red River Gorge, Daniel Boone
National Forest, in maple-hickory-Magnolia<br />
deciduous forest, associated with Carpinus<br />
caroliniana, Acer, Magnolia tripetala,<br />
Rhododendron maximum, Circaea, Hydrangea,<br />
Impatiens, <strong>and</strong> Parthenocissus quinquefolia, 29<br />
Jun 1991, J. Wen 1071 (A). Pike Co., Pine<br />
Mountain, near Bald Knob, 2600 ft, 30 Aug 1981,<br />
fl, F. Levy s.n. (NCU). Powell Co., 1.8 miles east<br />
<strong>of</strong> Kentucky routes 11-15 on Kentucky route 77,<br />
Nada Tunnel in Daniel Boone National Forest, 1<br />
Sep 1974, young fr, D. E. Boufford et al. 15452<br />
(NCU). Wayne Co., Beaver Creek, SW <strong>of</strong><br />
Monticello, 12-14 Jul 1937, floral buds, L. B. Smith<br />
& A. R. Hodgdon 3935 (F, NY, US). Wolfe Co., in<br />
rich moist woods along Swift Camp Creek, 3 Sep<br />
1969, fr, P. D. Higgins 1827 (NCU). MAINE:<br />
Aroostook Co., 0.4 mi W <strong>of</strong> Center Brook, T11R15,<br />
region <strong>of</strong> Clayton Lake, 8 Aug 1967, F. C. Seymour<br />
& H. K. Svenson 25868 (CAS, MO); Mars Hill, 23<br />
Aug 1967, fr, F. C. Seymour 26295 (MO). Hancock<br />
Co., Mount Desert Isl<strong>and</strong>, Southwest Valley, 29<br />
Aug 1889, fr, E. L. R<strong>and</strong> s.n. (CAS); Mt. Desert<br />
Isl<strong>and</strong>, 29 Aug 1891, fr, J. H. Redfield s.n. (UC).<br />
Knox Co., in rich woods, 4 miles northwest <strong>of</strong><br />
Washington, 19 Aug 1929, young fr, J. A.<br />
Steyermark 1017 (MO); Camp Medomak, near<br />
Washington, 22 Jul 1930, fl, J. A. Steyermark 4115<br />
(F). Lincoln Co., Monhegan Isl<strong>and</strong>, woods,<br />
between ledges, 3 Sep 1921, J. R. Churchill s.n.<br />
(MO). Somerset Co., Athens, moist thickets, 14<br />
Aug 1888, fl, H. H. Tucker s.n. (F). MARYLAND:<br />
Baltimore Co., near Baltimore, 1873, E. Foreman<br />
s.n. (NY); Gunpowder Falls State Park, area<br />
between end <strong>of</strong> Schroeder Ave. <strong>and</strong> Perry Hall Rd.,<br />
Perry Hall, north-facing slope, shaded rocky<br />
habitat, near a river, locally rare, 22 May 1991, J.<br />
Wen 817 (A). Garrett Co., Swanton, at RR crossing<br />
near Crabtree Creek, foot <strong>of</strong> Backbone Mtn., 2300-<br />
2500 ft, fruits purple, occasional herb 1.5 m tall,<br />
wet woods, roadside, 20 Sep 1980, fr, S. R. Hill<br />
9576 (NY). Harford Co., rocky banks along<br />
Susquehanna River, at old Conowingo Bridge, SE<br />
<strong>of</strong> Flintville, 5-6 Aug 1927, fr, J. W. Adams & G.<br />
Hopkins 897 (MO). MASSACHUSETTS:<br />
Berkshire Co., Cheshire, 26 Jul 1915, fl, J. R.<br />
Churchill s.n. (MO); Aug 1898, J. M. Milligan s.n.<br />
(US). Hampshire Co., Florence, woodl<strong>and</strong>, 3 Sep<br />
1977, H. E. Ahles 84804 (ASU, BKF); Quincy,<br />
Blue Hill Reservation, 3 Sep 1917, S. F. Blake 6725<br />
(LL). Hampton Co., wooded bank <strong>of</strong> Chicopee<br />
River, Chicopee Falls, 16 Jul 1950, fl, T.<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
47<br />
Delevoryas s.n. (TEX); Granville, 20 Sep 1913,<br />
sterile, F. C. Seymour 52 (MO, 2 sheets).<br />
Middlesex Co., Hopkinton, moist open woods, 20<br />
Aug 1911, fr, C. H. Knowlton s.n. (ASU). Norfolk<br />
Co., Canton, 31 Jul 1901, Clark s.n. (LL); Milton,<br />
15 Jul 1903, floral buds, J. Murdoch 1367 (F).<br />
Worcester Co., Petersham, Turnpike Road in shady<br />
site, 29 Jul 1947, fr, J. E. Canright 327a (ASU),<br />
327 (ASU). Unknown locality, Chapman s.n.<br />
(CAS). MICHIGAN: Alger Co., Miner’s Castle,<br />
Munising, 30 Jul 1940, fl, A. Ch<strong>and</strong>ler 816 (MO);<br />
3.2 mi N <strong>of</strong> jct with SR 28 on Koski Road, mixed<br />
hardwood forest, 46 o 27’39"N, 86 o 47’47"W, 270 m,<br />
shrub 1 m tall, 28 Aug 1993, fr, H. H. Schmidt &<br />
M. Merello 1082 (MO). Baraga Co., near Otter<br />
Lake area, at the Access site <strong>of</strong> DNR <strong>of</strong> the Otter<br />
River Dam, in maple-Tilia-Fraxinus mixed<br />
deciduous forest, associated with Tilia americana,<br />
Acer sp., <strong>Aralia</strong> nudicaulis, Corylus cornuta,<br />
Prunus serotina, Prunus sp., Fraxinus sp.,<br />
Osmorhiza claytonii, O. longistylis, Parthenocissus<br />
quinquefolia, Trillium sp., <strong>and</strong> Cornus sp., 27 Jun<br />
1999, perennial 0.6-0.9 m tall, stem purplish, 3-4<br />
ternately compound leaves per individual plant,<br />
abundant, J. Wen 4714 (F, US). Berrien Co., New<br />
Buffalo Twp., red <strong>and</strong> white oak woods, 19 Aug<br />
1950, in fl., G. W. Parmelee 1769 (MSC).<br />
Chebuygan Co., Douglas Lake, 7 Jul 1911, in fl.,<br />
F. C. Gates 130 (MSC). Chippewa Co., deciduous<br />
woodl<strong>and</strong>, roadside ca. 5 miles east <strong>of</strong> ferry, N end<br />
<strong>of</strong> Sugar Isl<strong>and</strong>, 1 Sep 1947, R. McVaugh 8794<br />
(DS). Ingham Co., Michigan State Univ., Water<br />
Quality Area, S<strong>and</strong>hill Woodlot, 20 Aug 1971, in<br />
fl., D. M. Valasek 889 (MSC); Ironwood, 28 Jul<br />
1909, J. Clemens s.n. (CAS). Jackson Co.,<br />
Waterloo Twp., in slight depression under closed<br />
canopy <strong>of</strong> black <strong>and</strong> white oaks, 6 Aug 1948, in<br />
fl., G. W. Parmelee 687 (MSC). Keweenaw Co.,<br />
W side <strong>of</strong> Bare Mountain, E <strong>of</strong> Bete Grise, in old<br />
logging road/trail in deep forest, 6 Jul 1999, in fl.,<br />
M. Chamberl<strong>and</strong> 950 (MSC). Macomb Co., Bruce<br />
Twp., Thuja swamp, 29 Aug. 1950, in young fr., G.<br />
W. Parmelee 1911 (MSC). Mason Co., Hamlin<br />
Lake, Ludington, 20 Aug 1910, young fr., R. W.<br />
Chaney 187 (F, NY). St. Clair Co., Port Huron, 5<br />
Jul 1895, fl, C. K. Dodge s.n. (TEX).<br />
MINNESOTA: Becker Co., along SR 113 S <strong>of</strong><br />
Itasca State Park, 12 Jul 1963, C. R. Janssen 498<br />
(U). Clearwater Co., Itasca State Park, along road<br />
from main drive to the Campgrounds, 14 Aug 1962,<br />
C. R. Janssen 257 (U); mixed forest, NW part Bear
48<br />
Paw Point, Itasca State Park, 20 Jul 1962, fl, R. F.<br />
Thorne 31188 (DS). Cass Co., Gull Lake, Jul 1893,<br />
C. A. Ballard s.n. (LE). Itasca Co., woods at the n<br />
end <strong>of</strong> Bowstring Lake, 27 Aug 1930, fr, N.<br />
Hotchkiss & P. Jones 4068 (US). Wabasha Co.,<br />
on the way from Lake City to Wabasha, 19 miles S<br />
<strong>of</strong> Red Wing along SR 61 between milestones 70<br />
<strong>and</strong> 71, or 3 miles S <strong>of</strong> Lake City, N44 o 25.294’,<br />
W92 o 13.089’, 230 m, 5 Sep 2007, fr, perennial herb<br />
with horizontal rhizome, fruits light purple turning<br />
to purple, J. Wen 9949 (US). MISSISSIPPI:<br />
Grenada Co., Camp McCain, at 0.2 mi S Grant<br />
Road <strong>and</strong> 0.03 mi E <strong>of</strong> western boundary <strong>of</strong> Camp<br />
McCain, T22N R5E Sect. 35 NW/4 <strong>of</strong> SE/4 <strong>of</strong> NE/<br />
4, upper creek terrace, near base <strong>of</strong> mesic slope,<br />
24 Jul 1996, sterile, C. T. Bryson 15469 (MO).<br />
Tallahatchie Co., 7.2 mi N jct MS Hwys 32 & 35,<br />
N <strong>of</strong> Charleston, E <strong>of</strong> MS Hwy 32, ravine behind<br />
Mt. Zion M. B. Church, 14 Sep 1990, fr, C. T.<br />
Bryson 10410 (MO). MISSOURI: Callaway Co.,<br />
crevices <strong>of</strong> north-facing limestone bluffs along<br />
Stinson Creek, T47N, R9W, sect. 34, 3 mi SE <strong>of</strong><br />
Pulton, 10 Sep 1937, fr, J. A. Steyermark 26090<br />
(F). De Kalb Co., at base <strong>of</strong> wooded n-facing slope<br />
bordering Lost Creek, 2 mi N <strong>of</strong> Maysville, 21 Jun<br />
1938, floral buds, J. A. Steyermark 5946 (F).<br />
Douglas Co., N-facing limestone bluffs with<br />
Roubidoux s<strong>and</strong>stone above, along Rippee Creek,<br />
2.5-3 mi SE <strong>of</strong> Sweden, 1 Sep 1951, fr, J. A.<br />
Steyermark 72525 (F). Pulaski Co., clefts <strong>of</strong><br />
limestone bluffs by Peterson Cave along<br />
Hasconade River, sect. 8, 5 mi NW <strong>of</strong> Waynesville,<br />
24 Aug 1937, fr, J. A. Steyermark 25258 (F). Ralls<br />
Co., Ilasco, 13 Aug 1915, fl, J. Davis 4853 (F).<br />
Texas Co., N-facing wooded limestone exposures<br />
along North Prong <strong>of</strong> Jacks Fork, 4 mi NE <strong>of</strong> Clear<br />
Springs, 11 Aug 1951, fr, J. A. Steyermark 72377<br />
(F). NEBRASKA: Nebraska City, Aug 1889, T.<br />
A. Williams s.n. (US). NEW HAMPSHIRE:<br />
Carroll Co., Chocorua, 5 Aug 1889, G. J. Peirce<br />
s.n. (DS). Coos Co., White Mountains National<br />
Forest, by Townline Brook, ca. 4 miles NW <strong>of</strong> jct<br />
with St. 16, 071 14.75W, 044 21.67N, mixed<br />
hardwood-coniferous forest with deep duff, with<br />
Betula lenta, Tsuga, Picea, Acer rubrum,<br />
Viburnum, Clintonia, 12 Jul 1999, fl, W. Hess et<br />
al. 8770 (F). Nelson, 18 Jul 1932, C. F. Batchelder<br />
s.n. (MO). NEW JERSEY: Mercer Co., Princeton,<br />
5 Aug 1851, young fr, A. Schott s.n. (F). Morris<br />
Co., Morris Plains, rocky woods, 13 Aug 1905, fr,<br />
K. K. Mackenzie 1607 (MO). Passaic Co.,<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
Ringwood jc, rocky woods, Sep 1908, K. K.<br />
Mackenzie 3895 (MO). Somerset Co., Watchung,<br />
sprawling herb in woods, 29 Jul 1949, floral buds,<br />
H. N. Moldenke 20270 (NY). NEW YORK:<br />
Broome Co., Binghamton, Glenmoor Ravine, Lat.<br />
42.6N, Long. 75.54N, 1882, C. F. Millspaugh s.n.<br />
(F); cliff along stream east-southeast <strong>of</strong> Union<br />
Center, Susquehanna River Drainage, 18 Sep 1949,<br />
fr, R. T. Clausen 7904 (NY). Chemung Co., the<br />
upper waters <strong>of</strong> the Susquehanna, Elmira, 5 Aug<br />
1896, fl, T. F. Lucy 8779 (F). Essex Co., open<br />
woods E <strong>of</strong> Newcomb, 15 Aug 1940, fr, H. D.<br />
House 27588 (MO). Oneida Co., Trenton Falls,<br />
woods, Sep 1871, fr, J. H. Redfield 2036 (MO).<br />
Rensselaer Co., Troy, 1828-1834, J. Hall s.n. (F).<br />
Sullivan Co., Cochecton, Aug 1887, J. Scheruk s.n.<br />
(MO). Tompkins Co., Ithaca, Beebe Lake, 8 Aug<br />
1920, W. C. Muenscher & A. R. Bechtel 477 (CAS).<br />
Warren Co., moist woods, n <strong>of</strong> Bolton L<strong>and</strong>ing,<br />
Lake George, 27 Sep 1945, fr, H. D. House 29957<br />
(TEX). Washington Co., damp woods, Lake<br />
George, 3 Sep 1941, fr, H. D. House 28182 (TEX).<br />
NORTH CAROLINA: Alleghany Co., Blue Ridge<br />
Parkway, Doughton Park, L. A. Mullen II 590<br />
(NCSC). Ashe Co., wooded edge <strong>of</strong> Long Hope<br />
Waterfalls, 10 Aug 1969, young fr, J. K. Moore<br />
2488 (NCU). Avery Co., Lost Cove, along road<br />
464A near the gate, 3000 ft., 2 Aug 1986, in young<br />
fr., T. L. Bradshaw s.n. (NCSC); along Blue Ridge<br />
Parkway, ca 0.7 mi N <strong>of</strong> Graybeard Mtn View<br />
Lookout, 28 Apr 2002, sterile, J. Wen 6250 (US).<br />
Buncombe Co., vicinity <strong>of</strong> Montreat, thick woods,<br />
23 Aug 1913, young fr, P. C. St<strong>and</strong>ley & H. C.<br />
Bollman 9982 (US). Burke Co., near jct <strong>of</strong> Shinny<br />
Creek <strong>and</strong> Jacob Fork River, 21 Sep 1975, in fr.,<br />
L. Smith 45 (NCSC). Cherokee Co., Joe Brown<br />
Highway (SR 1326), gap at north end <strong>of</strong> Buck<br />
Knob, between Unaka <strong>and</strong> Violet, steep forested<br />
slopes, 16 Jul 1998, sterile, B. A. Sorrie 9845<br />
(NCU, 2 sheets). Forsyth Co., wooded slope, 2.4<br />
miles north <strong>of</strong> Stanleyville on US 52, 17 May 1958,<br />
sterile, H. E. Ahles 40851 (NCU). Graham Co.,<br />
common along trail to Stratton Bald in Joyce<br />
Kilmer Memorial Forest ca. 15 mi W <strong>of</strong><br />
Robbinsville, 26 Aug 1955, in young fr., R. L.<br />
Wilbur 4592 (NCSC). Haywood Co., Mt. Sterling,<br />
near Waterville, 0.2 mi NW <strong>of</strong> the town Mt. Sterling<br />
on the road to Davenport, 2 Sep 1986, J. Wen et al.<br />
139 (OS). Henderson Co., growing in hardwoods<br />
on mountain side, 2 miles NW <strong>of</strong> Fruitl<strong>and</strong> in<br />
Baldwin Gap, 2 Sep 1956, fr, D. Pittillo 346 (NY).
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
49
50<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong>
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
51
52<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong>
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
53
54<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong>
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
55
56<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong>
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
57
58<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong>
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
59
60<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong>
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
61
62<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong>
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
63
64<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong>
Jackson Co., rich cove, head <strong>of</strong> Short<strong>of</strong>f Creek, E<br />
slope <strong>of</strong> Short<strong>of</strong>f Mt., 26 Aug 1951, W. B. Fox &<br />
R. K. Godfrey 5378 (NCSC); Wolf Creek Preserve<br />
on Cullowhee Mt., 3600 ft, 26 Sep 1967, J. D.<br />
Pittillo 2877 (LL, NCU, U); Whiterock Road (Co.<br />
1164), plants scattered in moist coves along edge<br />
<strong>of</strong> road, plant 0.5-1.5 m tall, 10 Aug 1970, fl, D.<br />
Pittillo 3161 (MO, NCU); N <strong>of</strong> Cashiers, along<br />
NC 1120, near Cedar Creek, 30 May 1991, J. Wen<br />
895 (A). Macon Co., in bramble thickets, cut-over<br />
hemlock forest, between Cole Mt. Gap <strong>and</strong> Yellow<br />
Mt. Gap, R. K. Godfrey & J. E. Oconnell 51881<br />
(NCSC); near the Coweeta Hydrological<br />
Laboratory, ca. 15 miles W <strong>of</strong> Highl<strong>and</strong>s, on the<br />
W side <strong>of</strong> US 23, rich woods, 29 May 1991, J.<br />
Wen 883 (A); Dry Falls 3 miles W <strong>of</strong> Highl<strong>and</strong>s on<br />
SR 64, abundant along the trail down to the falls,<br />
31 May 1991, J. Wen 906 (A); Highl<strong>and</strong>s area,<br />
along the road to Glen Falls, ca 0.2 mi from jct<br />
with Rt 106, 29 Apr 2002, sterile, J. Wen 6270 (US).<br />
Madison Co., 8.4 mi W <strong>of</strong> SR 209 on the Forest<br />
Service road to Max Patch, 35 o 46’50"N,<br />
82 o 57’42"W, 1140 m, shrub 1.5 m tall, corollas<br />
cream colored, 11 Aug 1994, fl, J. S. Miller & N.<br />
Snow 8345 (MO, 2 sheets). McDowell Co., near<br />
Marion, in the property <strong>of</strong> Richard Weaver, ca. 6<br />
miles S <strong>of</strong> Marion, ca. 2 miles along Polly Spout<br />
Rd. from its jct with NC 221, 28 May 1991, J. Wen<br />
880 (A). Mitchell Co., wooded slope, Iron<br />
Mountain Gap on NC 26 (northwest <strong>of</strong> Buladean),<br />
25 Sep 1958, fr, H. E. Ahles 49872 (NCU). Orange<br />
Co., dry rocky woods on Enu River, 3.5 miles E <strong>of</strong><br />
Hillsboro, 21 Jul 1952, fl, A. E. Radford 6258<br />
(NCU). Polk Co., along old logging road from<br />
Green River to Long Ridge, margins <strong>of</strong> logging<br />
trail in cut over woods, 1400 ft., 7 Sep 1973, in fr.,<br />
D. M. DuMond 1809 (NCSC); common in cool<br />
mountain forests, in rich loam, Vaughan’s Gap,<br />
Rocky Spur, 14 Aug 1921, fl, D. C. Peattie 1212A<br />
(F). R<strong>and</strong>olph Co., foot <strong>of</strong> Carroway Mt., 20 May<br />
1959, sterile, L. Melvin 3393 (NCU). Swain Co.,<br />
mixed deciduous forest, near NC 28, 1.5 miles<br />
south <strong>of</strong> Lauada, 24 Aug 1956, fr, A. E. Radford<br />
17228 (NCU). Transylvania Co., moist cove, in<br />
gap near N.C. Rt. 178, 4.5 mi S <strong>of</strong> Rosman, 27<br />
Aug 1951, in young fr., W. B. Fox & R. K. Godfrey<br />
5415 (NCSC). Watauga Co., mixed woods, 2.5<br />
miles north <strong>of</strong> S<strong>and</strong>s on NC 194, 23 Sep 1958, fr,<br />
H. E. Ahles 49417 (ASU); low rocky bank <strong>of</strong><br />
Watauga River, near Shull’s Mill, 8 Sep 1949, in<br />
fr., W. B. Fox & R. K. Godfrey 3388 (NCSC); Rich<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
65<br />
Mountain N <strong>of</strong> Boone, in mixed forest, 27 May<br />
1991, plant 1.2 m tall, stem purple, stipule small,<br />
not leafy, J. Wen 855 (A); on the N side <strong>of</strong> Rich<br />
Mountain Rd, ca 1 mile E <strong>of</strong> jct with US Rt 421, 1<br />
Jun 1994, J. Wen 1514 (US). Wilkes Co., on shady<br />
moist road bank, west slope, dirt road from Gilreath<br />
P. O. to Hidnite, 7.6 miles south <strong>of</strong> Sunl<strong>and</strong><br />
Orchards, 23 Jul 1940, Radford & Stewart 1769<br />
(TEX). Yancey Co., wooded cove, 3.8 miles north<br />
<strong>of</strong> Swiss, 16 Jul 1958, fl, H. E. Ahles 46779 (NCU,<br />
3 sheets). OHIO: Butler Co., Oxford, along<br />
Tallaw<strong>and</strong>a, 3 Aug 1910, fl, L. Braun s.n. (MO).<br />
Champaign Co., Cedar Swamp, 12 Aug 1917,<br />
young fr, E. C. Leonard 1699 (US, 2 sheets), 6<br />
Sep 1920, fr, E. C. Leonard 2101 (US, 2 sheets).<br />
Coshocton Co., in open woodl<strong>and</strong>, North<br />
Appalachian Experimental Watershed, near<br />
Coshocton, 2 Aug 1942, H. N. Moldenke 13532<br />
(MEXU). Fairfield Co., Ohio State University<br />
(OSU) Property, Clear Creek, 10 May 1986, J. Wen<br />
84 (OS); Clear Creek, OSU property, 16 Jul 1986,<br />
fl, J. Wen 95 (OS); Clear Creek, 25 Sep 1988, fr, J.<br />
Wen 625 (OS, US); Clear Creek, in mixed<br />
deciduous forest, plant 1.5 m tall, stem highly zigzag,<br />
25 Sep 1990, J. Wen 783 (OS). Hamilton Co.,<br />
mesophytic woods, 7 Jul 1903, fl, E. L. Braun s.n.<br />
(US). Hocking Co., Conkel’s Horrow, in s<strong>and</strong>stone<br />
area, associated with Tsuga canadensis, Ulmus<br />
americana, <strong>and</strong> ferns, 4 Jul 1986, J. Wen 87 (OS).<br />
Licking Co., Blackh<strong>and</strong> Gorge, on s<strong>and</strong>stone cliff,<br />
30 Jul 1986, J. Wen 109 (OS). Lorain Co., Oberlin,<br />
4 Aug 1894, fl, A. E. Ricksecker s.n. (US). Portage<br />
Co., infrequent, N-facing hemlock slope on S side<br />
<strong>of</strong> South Fork <strong>of</strong> Eagle Creek, 300 yards W <strong>of</strong><br />
Wadsworth Rd., Windham Twp., 30 Jun 1978,<br />
floral buds, B. A. Andreas & T. S. Cooperrider 2186<br />
(US). Vinton Co., Jackson Twp., along Co. Rd.<br />
17, 0.5 mi north <strong>of</strong> jct with Co. Rd. 44, mesic forest,<br />
6 Jul 1986, fl, J. Wen & T. Lammers 88 (OS).<br />
PENNSYLVANIA: Bedford Co., mountains <strong>of</strong> S.<br />
Pennsylvania, in the vicinity <strong>of</strong> Hyndman, 19-23<br />
Aug 1890, fr, J. K. Small s.n. (F). Lancaster Co.,<br />
on the Welsh Mountains, between Churchtown <strong>and</strong><br />
Beartown, 7 Sep 1892, young fr, 500 ft, J. K. Small<br />
s.n. (F). Lehigh Co., woods 1 mile NW <strong>of</strong> Lowhill,<br />
17 Aug 1950, young fr, R. L. Schaeffer Jr. 34359<br />
(US); 1.25 miles NE <strong>of</strong> Lynnport, 25 Aug 1951,<br />
young fr, R. L. Schaeffer Jr. 37866 (US). Lycoming<br />
Co., Deer Hole Mountain, South Williamsport to<br />
summit, 200-400 m, 21 Sep 1928, fr, W. W.<br />
Eggleston & J. W. Kelly 22748 (US). Mifflin Co.,
66<br />
10 Aug 1867, fl, J. T. Rothrock s.n. (F). Monroe<br />
Co., 13-15 Jul 1889, N. L. Britton et al. s.n. (F).<br />
Perry Co., Marysville, 9 Aug 1888, fl, J. K. Small<br />
s.n. (F). Philadelphia, rich woods, 15 Aug 1888,<br />
fl, J. B. Brinton s.n. (F). Pocono Plateau, Jul-Aug<br />
1904, J. W. Harshberger s.n. (F, MO). RHODE<br />
ISLAND: Providence Co., near Limestone Rock,<br />
2 Oct 1910, fr, J. M. Greenman 2554 (MO).<br />
SOUTH CAROLINA: Greenville Co., rich woods,<br />
Hogback Mt., 2500 ft, D. C. Peattie 1244 (F),<br />
1244A (F). Oconee Co., open wooded bank <strong>of</strong><br />
Thompson River, above Jocassee, 21 Aug 1951,<br />
in young fr., R. K. Godfrey & J. E. O’Connell 52001<br />
(NCSC). SOUTH DAKOTA: Grant Co., dark<br />
woods west side <strong>of</strong> Big Stone Lake, 27 Jul 1916,<br />
fl, W. H. Over 1951 (US); Big Stone, 23 Aug 1892,<br />
fr, T. A. Williams s.n. (US). Marshall Co., Sieche<br />
Hollow, 29 Jun 1959, floral buds, B. Ailts 234<br />
(ASU). Minnehata Co., 3 mi S <strong>of</strong> Garretson,<br />
densely wooded Sioux quartzite bluffs, relatively<br />
rare in moist ravines, 10 Jul 1965, floral buds, L.<br />
J. Harms 2673 (NY). Roberts Co., common in<br />
damp rich woods, Jul 1922, fl, W. H. Over 14415<br />
(US). Rare in Acer saccharum woods, Sieche<br />
Hollow, Sisseton, 31 May 1957, sterile, O. A.<br />
Stevens & D. R. Moir 1690 (US). TENNESSEE:<br />
Blount Co., along edge <strong>of</strong> woods at base <strong>of</strong><br />
Chilhowee Mountain beside Reed Creek Road east<br />
<strong>of</strong> Wall<strong>and</strong> near Sevier County line west <strong>of</strong><br />
Waldens Creek, 9 Aug 1994, young fr, R. D.<br />
Thomas et al. 141164 (NY). Cocke Co., 5.6 mi SE<br />
<strong>of</strong> Del Rio on SR 107 in Cherokee National Forest,<br />
35 o 51’41"N, 082 o 58’58"W, 600 m, 19 Aug 1977,<br />
fr, J. S. Miller et al. 8889 (MO); ca. 10 miles E <strong>of</strong><br />
NC State line, just west <strong>of</strong> Pigeon River from I40<br />
to Waterville, E <strong>of</strong> Carolina Power <strong>and</strong> Light Co.<br />
Water Plant, roadside wet area, ca. 1 m tall, 1.5 m<br />
in spread, 1 Sep 1986, fr, J. Wen et al. 138 (OS).<br />
Green Co., Yellow Spring Rd., 29 Jul 1965, L. A.<br />
Mullen II s.n. (NCSC). Grundy Co., creekside<br />
forest in Fiery Gizzard Cove, near Grundy State<br />
Forest, 20 Aug 1964, fl, R. C. Clark 1165 (NCU).<br />
Monroe Co., Cherokee National Forest, along<br />
Miller Cemetery Road south <strong>of</strong> Tellico River <strong>and</strong><br />
Tenn 165 east <strong>of</strong> Tellico Plains, 20 Jul 1996, fl, R.<br />
D. Thomas & B. R. Thomas 149978 (NY). Putnam<br />
Co., E <strong>of</strong> Monterey on escarpment, s<strong>and</strong>stone<br />
bluffs by I-40, 2 Aug 1973, fl, R. Kral 51004 (MO,<br />
2 sheets). Sevier Co., near Gatlinburg, Great<br />
Smoky National Park, Ramsay Creek, 19 Oct 1944,<br />
fr, G. S. Miller 2101 (US). Tipton Co., heavily<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
wooded hillsides at Bear Creek, turn S on farm<br />
road about 2.6 miles W <strong>of</strong> road jct at Drummonds,<br />
creek at bottom <strong>of</strong> hill on west side <strong>of</strong> cultivated<br />
fields, 1 Sep 1968, young fr, S. J. Warrington 373<br />
(NCU). Unicoi Co., Limestone Cove, along rt 173,<br />
ca 5 miles E <strong>of</strong> Unicoi, roadside woods,<br />
N36 o 10.553’, W82 o 17.427’, 683 m, 31 Aug 2008,<br />
fr, perennial ca 1.5 m tall, fruits purple, flowers<br />
whitish, J. Wen 10507 (US). Wayne Co., Natchez<br />
Trace Parkway, bottoml<strong>and</strong> forest, 23 Jul 1947, fl,<br />
W. B. McDougall 1413 (US). VERMONT:<br />
Bennington Co., Manchester, 28 Jul 1898, fl, M.<br />
A. Day 77 (US); Pownal Twp., along paved road,<br />
1.7 miles north <strong>of</strong> North Pownal Post Office, mesic<br />
forest, associated with Impatiens, Sambucus<br />
canadensis, <strong>and</strong> Quercus, 9 Aug 1986, J. Wen et<br />
al. 130 (OS); along SR 9, opposite to Woodford<br />
State Park, moist woods, 9 Aug 1986, J. Wen et al.<br />
132 (OS); Woodford State Park, at the edge <strong>of</strong><br />
Quercus-Larix forest, 9 Aug 1986, J. Wen et al.<br />
134 (OS). Caledonia Co., Peacham, 10 Sep 1883,<br />
F. Blanchard s.n. (DS); Peacham, 29 Jul 1892, F.<br />
Blanchard s.n. (DS, F, MO). Chittenden Co.,<br />
wooded bank, Williston, 240 ft, 29 Jul 1911, fl, S.<br />
F. Blake 2329 (LL). Franklin Co., rich woods at<br />
northern end <strong>of</strong> Metcalf Pond, 25 Sep 1964, fr, F.<br />
C. Seymour & W. D. Countryman 22569 (MO).<br />
Lamoille Co., in deep wet woods, Smiggler’s<br />
Notch, Mount Mansfield, 24 Aug 1976, fl, A. L. &<br />
H. N. Moldenke 31095 (LL). Rutl<strong>and</strong> Co.,<br />
Middletown Springs, D. S. Carpenter s.n. (CAS);<br />
Br<strong>and</strong>on, rich woods, 500 ft, 11 Aug 1923, D. L.<br />
Dutton s.n. (DS). Windham Co., Jamaica, in dense<br />
woods, 26 Aug 1935, H. N. Moldenke 8741 (F,<br />
MO). VIRGINIA: Augusta Co., summit <strong>of</strong> Elliott<br />
Knob, 1360 m, 4 Jul 1937, floral buds, H. A. Allard<br />
3158 (US). Bedford Co., Peaks <strong>of</strong> Otter, 1871, A.<br />
H. Curtiss s.n. (LE, MO, NY). Fauquier Co., the<br />
Plains, Kinloch Farm, 15 Sep 1984, young fr., T.<br />
Plowman 13862 (F). Giles Co., Mountain Lake,<br />
12 Aug 1950, in fl., B. W. Wells s.n. (NCSC); along<br />
rt 700 near Mountain Lake, 9 Sep 1986, J. Wen et<br />
al. 182 (OS). Loudoun Co., wooded bluff, Goose<br />
Creek on Co. Rt. 621 (vic. Of Evergreen Mills),<br />
11 Sep 1965, sterile, H. E. Ahles 61512 (NCU, 2<br />
sheets). Madison Co., Stony Man Mountain, rocky<br />
woods, 2000 ft, 25 Aug 1901, young fr, W. Palmer<br />
& W. H. King 142 (US). Nelson Co., along road to<br />
Spy Rock above Fish Hatchery, rich soil, ravine,<br />
R. S. Freer 2442 (US). Page Co., Stony Man<br />
Mountain, rocky woods, 2500 ft, rare, 24 Aug 1901,
fl, W. Palmer & W. H. King 126 (US); Matthew’s<br />
Arm Rd., roadside, 14 Aug 1938, young fr, E. H.<br />
Walker 2677 (US); Stony Man Mountain & vicinity<br />
in the Blue Ridge, near Luray, 20 Aug 1901, fr,<br />
3500 ft, E. S. Steele 25 (MO, NY). Rappahannock<br />
Co., Shen<strong>and</strong>oah National Park, on Appalachian<br />
trail north <strong>of</strong> Pinnacle Mountain, on rocky slope<br />
near crest <strong>of</strong> Blue Ridge, 21 Sep 1941, fr, E. H.<br />
Walker 3212 (US). Shen<strong>and</strong>oah Co., at the top <strong>of</strong><br />
Short Mountain, along Rt. 374, roadside, near Mt.<br />
Jackson, large perennial about 2.5 m tall, terminal<br />
infl 40 cm long, 26 cm wide, berries purple, 10<br />
Sep 1994, fr, J. Wen 1800 (US); Short Mountain,<br />
near the top, 22 Sep 1994, J. Wen 1824 (US).<br />
Smyth Co., on Nick’s Creek, at base <strong>of</strong> Pine Glade<br />
Mountain, 5 Aug 1892, fl, 2500 ft, J. K. Small s.n.<br />
(F, MO). Surry Co., rich calcareous wooded gullies<br />
along James River, Claremont Wharf, 23 Aug 1938,<br />
young fr, M. L. Fernald & B. Long 9112 (US).<br />
Warren Co., Linden, G. Thompson Wildlife<br />
Management Area, along main trail near Parking<br />
Area 6, in woods, N38 o 57’13.3", W78 o 01’37.4",<br />
648 m, 8 May 2007, sterile, herb ca 1 m tall, stem<br />
purple, J. Wen 9697 (US); near Rangeview<br />
Overlook on Skyline Dr., Shen<strong>and</strong>oah National<br />
Park, ca 12 miles S <strong>of</strong> Dickey Ridge Visitor Center,<br />
N 38 o 45’58.6", W78 o 13’42.8", 850 m, 4 Jul 2007,<br />
herb ca 1.5 m tall, stem dark purple, J. Wen 9841<br />
(US). Washington Co., ravine just north <strong>of</strong><br />
Mendota, 6 Sep 1967, young fr, A. M. Harvill<br />
17768 (NCU). Wythe Co., on Reed Creek, at base<br />
<strong>of</strong> Lower Rocks, 23 Jul 1892, fl, 2000 ft, J. K. Small<br />
s.n. (F). WEST VIRGINIA: Greenbrier Co., rocky<br />
woods, frequent, 2 Sep 1903, fr, K. K. Mackenzie<br />
429 (NY). Hampshire Co., hill about 2 miles<br />
northeast <strong>of</strong> Levels, 1 Aug 1967, young fr, R. M.<br />
Downs 1580 (NCU). Mineral Co., mixed<br />
deciduous woods on north slope facing north<br />
branch <strong>of</strong> Potomac River, scattered outcrops <strong>of</strong><br />
Greenbrier limestone in varicolored shales <strong>and</strong><br />
s<strong>and</strong>stones, ca. 1 mile southeast <strong>of</strong> Piedmont, 9<br />
Aug 1969, sterile, R. M. Downs 8124 (NCU).<br />
Pendleton Co., North Fork Mountain, west-facing<br />
slope, 28 Aug 1945, fl, J. W. Roller 348 (US).<br />
Putnam Co., wooded margin <strong>of</strong> Lake Washington,<br />
near Hurricane, 20 Jul 1940, F. A. Gilbert et al.<br />
876 (DS, MO, 2 sheets). Summers Co., from WV<br />
Hwy 20 in Hinton, follow Hwy 3 west about 1.25<br />
miles, south <strong>of</strong> Beech Run Creek, 3 Aug 2001,<br />
young fr, P. Diamond 1297 (NCU). Tucker Co.,<br />
Canaan Valley, 900-1200 m, 3 Aug 1945, fl, H. A.<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
67<br />
Allard 11526 (US, 2 sheets). Wetzel Co., woods at<br />
mouth <strong>of</strong> Brown’s Run, 2 mi E <strong>of</strong> Littleton, 8 Aug<br />
1931, O. Haught 737 (DS, 2 sheets). Locality<br />
unknown in West Virginia, Brookside, 17 Aug<br />
1898, fl, H. W. Olds s.n. (US). WISCONSIN:<br />
woods along lake, S <strong>of</strong> Milwaukee, 11 Aug 1902,<br />
young fr, A. Chase 1949 (F). Woods, Williams<br />
Bay, 21 Aug 1897, young fr, L. M. Umbach s.n. (F,<br />
2 sheets, MO). Brown Co., Oneida Reservation, 3<br />
Aug 1886, young fr, J. H. Schuette s.n. (F).<br />
Chippewa Co., along rte M on the way from<br />
Holcombe to New Auburn, ca 1 mi E <strong>of</strong> the junction<br />
with county Rte E, or 11.7 miles east <strong>of</strong> the junction<br />
with rte 40, N45 o 14.073’, W91 o 13.940’, 395 m, in<br />
maple-birch-Populus forest, 3 Sep 2007, fr,<br />
perennial ca 1.5 m tall, stem dark purple, fruits<br />
light purple, J. Wen 9931 (US). Door Co., Ellison<br />
Bay, 31 Jul 1887, young fr, J. H. Schuette s.n. (F);<br />
Sister Bay, 2 Aug 1887, J. H. Schuette s.n. (US).<br />
Shawano Co., 4 mi SSE <strong>of</strong> Bonduel, on farm <strong>of</strong><br />
Lorenz Liesner, 30 Sep 1984, past fr, R. Liesner<br />
17208 (MO). Walworth Co., in woods <strong>of</strong> Covenant<br />
Harbor near Lake Geneva, 4 Jul 1953, floral buds,<br />
F. A. Swink 2268 (F).<br />
Cultivated specimens: U.S.A. NEW YORK:<br />
New York Botanical Garden, Herbaceous Gounds,<br />
29 Aug 1907, fl, N. Taylor & R. C. Schneider 0619<br />
(NY). PENNSYLVANIA: Berks Co., L<strong>and</strong>is Store,<br />
District Township, a perennial herb 8 dm tall,<br />
leaves compound, flowers white, 30 Jul 1973, fl,<br />
W. C. Brumbach 8414 (NY). Lancaster Co.,<br />
Rawlinsville, 1883, J. Galen 1047 (US).<br />
Marie-Victorin <strong>and</strong> Rousseau (1940) described<br />
<strong>Aralia</strong> racemosa var. foliosa Vict. & J. Rousseau<br />
based on the presence <strong>of</strong> leafy bracts or leaf-like<br />
structures on the inflorescence in some collections<br />
from Québec, Canada. This character is<br />
occasionally observed in specimens from other<br />
areas (e.g., Raud s.n. collected from Hancock Co.<br />
<strong>of</strong> Maine on 29 Aug 1889; <strong>and</strong> Clemens s.n.<br />
collected from Ironwood, Michigan on 28 Jul<br />
1909). Scoggan (1978) proposed to recognize it<br />
as a form: <strong>Aralia</strong> racemosa L. f. foliosa (Vict. & J.<br />
Rousseau) Scoggan. I have found that the species<br />
is quite variable <strong>and</strong> I prefer not to recognize forms<br />
in this treatment.<br />
<strong>Aralia</strong> racemosa is easily distinguished from<br />
A. californica by the former’s non-leafy stipules,<br />
much narrower inflorescence (3-10 cm vs. 10-25
68<br />
cm wide), fewer flowers (20-25 vs. 30-75) per<br />
terminal umbel, shorter pedicels (6-8 mm vs. 12-<br />
20 mm long) on terminal umbels, <strong>and</strong> smaller fruits<br />
(4-4.5 vs. 6-7 mm long). The distinction between<br />
A. racemosa <strong>and</strong> A. bicrenata is discussed under<br />
A. bicrenata.<br />
3. <strong>Aralia</strong> bicrenata Wooton & St<strong>and</strong>l. — Fig. 20;<br />
color plate 3: A-G.<br />
<strong>Aralia</strong> bicrenata Wooton & St<strong>and</strong>l., Contr. U. S.<br />
Natl. Herb 16: 157. 1913. <strong>Aralia</strong> racemosa<br />
subsp. bicrenata (Wooton & St<strong>and</strong>l.) S. L.<br />
Welsh & J. T. Atwood, Great Basin Naturalist<br />
35: 333. 1975. TYPE: U.S.A. NEW<br />
MEXICO: Socorro Co., near Holts Ranch in<br />
the Mogollon Mountains, 20 Jul 1900, fl., E.<br />
O. Wooton s.n. (holotype: US!, US Herb. #<br />
563963; isotype: US!, US Herb. # 736551).<br />
<strong>Aralia</strong> arizonica Eastw., Proc. Calif. Acad. Sci.,<br />
IV, 20: 148. 1931. TYPE: U.S.A. ARIZONA:<br />
Cochise Co., Chiricahua Mountains, Cave<br />
Creek Canyon, 6-8000 ft., 26-29 Jun 1927, J.<br />
A. Kusche s.n. (holotype: CAS!, photo at US!).<br />
Robust perennial herb, 1-2.2 m tall,<br />
<strong>and</strong>romonoecious. Stem sparsely pubescent on the<br />
older parts, pubescent on the younger parts, green<br />
to greenish purple, with 4-6 leaves; rhizomes thick<br />
<strong>and</strong> horizontal. Leaves 73-120 cm long, 60-118<br />
cm wide, ternate, spreading; stipule 3.2-8.9 cm<br />
long, 1.1-5.8 cm wide, leafy; petioles 25-45 cm<br />
long, light green to purplish green; leaflets 6-14<br />
cm long, 3-8 cm wide, thin papery to papery, the<br />
pair <strong>of</strong> leaflets below the terminal leaflet narrowly<br />
ovate, other leaflets ovate to broadly so, acuminate<br />
to acute at apex, truncate, broadly acute to<br />
subcordate at base, commonly oblique on lateral<br />
leaflets, doubly serrate at margin; lateral veins 8-<br />
9 on each side, conspicuous on both surfaces, green<br />
<strong>and</strong> sparsely pilose on veins <strong>and</strong> veinlets on the<br />
adaxial surface, light green <strong>and</strong> pilose on veins<br />
<strong>and</strong> veinlets on the abaxial surface, petiolules 0.2-<br />
4 cm long, pilose. Inflorescence 35-50 cm long,<br />
30-50 cm wide, terminal <strong>and</strong> axillary, the transition<br />
between the axillary inflorescence below <strong>and</strong> the<br />
upper terminal inflorescence gradual, essentially<br />
the inflorescence terminal with the lower primary<br />
branches subtended by bipinnately ternate leaves<br />
<strong>and</strong> the upper primary branches subtended by leafy<br />
bracts, the overall architecture corymbose,<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
consisting <strong>of</strong> 10-20 primary branches, primary<br />
branches <strong>of</strong>ten 3-5 forming one or two circles on<br />
the upper part <strong>of</strong> the main inflorescence axis, each<br />
primary branch 8-30 cm long, consisting <strong>of</strong> 3-30<br />
umbels, with some lateral umbels functionally<br />
male; terminal umbels 12-20 (-25)-flowered,<br />
pedicels 7-12 mm long, pubescent, tip enlarged;<br />
lateral umbels 8-15-flowered, pedicels 3-6 mm<br />
long; bracts <strong>of</strong> the upper primary branches leafy<br />
to lanceolate or linear, the leafy ones varied in size,<br />
1-5 cm long, 0.4-3 cm wide, the lanceolate to linear<br />
true bracts 5-12 mm long, 1-3 mm wide, pilose,<br />
ciliate at margin; bracteoles 1.1-2.0 mm long, 0.4-<br />
0.7 mm wide, lanceolate, pilose, ciliate at margin.<br />
Sepals 0.4-0.6 mm long <strong>and</strong> wide, triangular,<br />
persistent on fruits; petals 1.9-2.0 mm long, 1.1-<br />
1.3 mm wide, ovate, greenish white to white, with<br />
a conspicuous vascular bundle in the middle;<br />
stamens 5, filaments 2-2.4 mm long, anthers<br />
oblong, 0.9-1 mm long, 0.6-0.7 mm wide; ovaries<br />
5-locular, styles 0.6-0.8 mm long at anthesis, base<br />
<strong>of</strong> styles slightly enlarged. Fruits 3.5-4.5 mm in<br />
diameter, globose, occasionally ovoid globose,<br />
dark purple, persistent styles divided to 1/3 to the<br />
base, 1.5-1.7 mm long. Seeds 2.5-2.9 mm long,<br />
1.5-2.0 mm wide, 0.4-0.5 mm thick, kidneyshaped,<br />
whitish gray, smooth.<br />
Common name: spikenard.<br />
Phenology: flowering in July to August; fruiting<br />
in August to October.<br />
Distribution: in Arizona <strong>and</strong> New Mexico, rare<br />
in Colorado (La Plata Co. <strong>and</strong> Mineral Co.), Utah,<br />
<strong>and</strong> Texas (a single population on Mt. Livermore,<br />
Jeff Davis Co.) <strong>of</strong> U.S.A. as well as in northern<br />
Mexico (Chihuahua, Coahuila, <strong>and</strong> Sonora) (Fig.<br />
21).<br />
Ecology: deciduous or mixed broadleaf <strong>and</strong><br />
coniferous forests, edges <strong>of</strong> forests, along streams,<br />
on rocky cliffs in moist habitats, <strong>and</strong> roadsides near<br />
forests; 1500-3000 m.<br />
Representative specimens examined. Mexico.<br />
CHIHUAHUA: Mpio. Madera, arroyo de las<br />
Garrochas, ejido “El Largo-Madera”, 18 Oct 1990,<br />
fl, 2600 m, A. Benítez P. 2930 (MEXU); Mpio. De<br />
Madera, arroyo de las Garrochas, ejido El Largo,<br />
bosque mixto de coníferas y latifoliadas, 2600 m,<br />
28 Aug 1990, fl, O. Bravo Bolaños 1314 (MEXU);<br />
Mpio. De Bocoyna, Rio Oteros, ca. 10 mi W <strong>of</strong><br />
Creel, near bend in river at st<strong>and</strong> <strong>of</strong> Picea
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
Fig. 20. <strong>Aralia</strong> bicrenata Wooton & St<strong>and</strong>l. A. Leaf with an accessory pinna. B. Inflorescence with transitional leaves <strong>and</strong> leafy<br />
bracts subtending the primary branches. C. Root <strong>and</strong> rhizome. D. Leafy stipules. E. Upper non-leafy stipules. F. Umbel.<br />
G. Floral bud. H. Flower. I. Flower after anthesis. J. Young fruit. K. Fruit.<br />
69
70<br />
chihuahuana, 7600 ft, 30 Jul 1974, R. A. Bye 6628<br />
(MEXU); Chuichupa, Sierra Madre Mts., 21 Sep<br />
1903, fr, 7000 feet, M. E. Jones s.n. (RSA); Largo<br />
Canyon, Sierra Madre Mts., 7000 ft, 26 Sep 1903,<br />
fr, M. E. Jones s.n. (BM, RSA, 2 sheets, US);<br />
Chuchuichupa, Aug-Sep 1936, fr, H. LeSueur 940<br />
(F, MO); Salto de Bapuora, 20 Jul 1937, fl, H.<br />
LeSueur 1400 (F, MO); Sierra Madre, Jun-Jul 1899,<br />
E. W. Nelson 6141 (K, US); 5 mi SE <strong>of</strong> Colonia<br />
Garcia in the Sierra Madres, 7500 ft, 27 Jul 1899,<br />
fl, C. H. T. Townsend & C. M. Barber 183 (BM, 2<br />
sheets, F, G, MEXU, MO, MSC, P, 2 sheets, US,<br />
WU). COAHUILA: ca. 20 (air) miles ESE <strong>of</strong><br />
Boquillas in Sierra del Carmen, ca. 2.5 miles E <strong>of</strong><br />
Rancho El Jardin in Canyon Hundido on the<br />
northernmost north-facing igneous canyon <strong>of</strong> the<br />
Sierra del Carmen, infrequent in shaded temperate<br />
forest with mid-canyon, with Quercus, Pinus,<br />
Prunus, Viburnum, <strong>and</strong> Cornus, near spring, 6600<br />
ft, 27 Jul 1973, young fr, 29 o 06’N, 102 o 37’W, J.<br />
Henrickson 11442 (RSA); Canyon Hundido on N<br />
side <strong>of</strong> Pico de Centinela, Sierra del Jardin, 8 km<br />
E <strong>of</strong> Rancho El Jardin by winding road, 29 o 06’-<br />
29 o 08’N, 102 o 37’-102 o 38’W, 1500-2250 m, 27 Jul<br />
1973, fl, M. C. Johnston et al. 11819 (F, MEXU,<br />
MO); Del Carmen Mts., 12 Sep 1936, fl, E. G.<br />
Marsh Jr. 828 (F); Sierra Maderas del Carmen,<br />
Cañon El Dos, in middle, deep, very mesic part <strong>of</strong><br />
canyon, from ca. 1 mi above jct with Cañon El<br />
Oso up to washed-out bridge ca. 1/2 mi down from<br />
Campo El Dos, mesic woods <strong>of</strong> Quercus<br />
hypoleucoides, Pinus ponderosa, P. stromiformis,<br />
Tilia, Acer, Pseudotsuga, Cupressus, Salix<br />
lasiolepis, <strong>and</strong> Ostrya, rhyolitic area, 28 o 59’N,<br />
102 o 35-36’W, 2000-2200 m, 3 Aug 1974, fl, T.<br />
Wendt & A. Adamcewicz 462 (MEXU). SONORA:<br />
upper Arroyo Frijolito, north slope <strong>of</strong> Cerro de las<br />
Flores, 30 o 56’N, 109 o 57’30"W, 2300 m, growing<br />
on north-facing slope, above arroyo bottom, across<br />
from base <strong>of</strong> steep cliffs with Abies concolor,<br />
Symphoricarpos oreophilus, Geranium<br />
richarsonii, <strong>and</strong> Acer gr<strong>and</strong>identatum, common,<br />
9 Oct 1992, fr, M. Fishbein et al. 704 (MEXU);<br />
Arroyo Frijolito, 30 o 56’30"N, 109 o 57’W, 2200 m,<br />
growing at base <strong>of</strong> limestone walls in narrow<br />
canyon bottom in pine-oak woods with Acer<br />
gr<strong>and</strong>identatum, Bromus, Robinia neomexicana,<br />
Habenaria limosa <strong>and</strong> Smilacina stellata, common,<br />
9 Oct 1992, fr, M. Fishbein et al. 725 (ASU,<br />
MEXU, MO). U.S.A. ARIZONA: Apache Co.,<br />
Ryan Ranch, Apache Res., 2 Oct 1927, fr, G. J.<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
Harrison 4849 (US); Badger Lake, near McNary,<br />
7200 ft, Jun 1956, floral buds, J. A. McCleary s.n.<br />
(ASU). Cochise Co., Chiricahua Mountains,<br />
Chaperon Canyon, shady gulch along Living<br />
Brook, at road Cold Living Brook, 7300 ft, 25 Aug<br />
1907, young fr, J. C. Blumer 1625 (F, NY, US),<br />
1625a (US); Miller Cañon, Huachuca Mts., 6 May<br />
1919, sterile, R. D. Camp 38-3 (F); Miller’s Cañon,<br />
Huachuca Mts., 5 Aug 1909, L. N. Goodding 324<br />
(CAS, G, NY); Miller Cañon, 2 Aug 1909, fl, along<br />
moist canyons, L. N. Goodding s.n. (NY); Ramsey<br />
Cañon, Huachuca Mountains, 29 Sep 1929, fr, M.<br />
E. Jones 24945 (NY, RSA); Huachuca Mts., Miller<br />
Cañon, 7500 ft, 4-5 ft tall, in moist situations, 28<br />
Sep 1916, fr, F. Shrieve 5087 (US); Coronado<br />
National Forest in Chiricahua Mountains, at the<br />
west end <strong>of</strong> Foresr Road 713, along a creek,<br />
N31 o 52.758’, W109 o 14.920’, 1975 m, moist rocky<br />
habitat in Pseudotsuga-Pinus-Juglans mixed<br />
forest, associated with Pseudotruga, Pinus,<br />
Juglans, Thalictrum, Quercus, Rubus, Aquilegia,<br />
<strong>and</strong> Rhamnus, 14 Jul 2000, fl, J. Wen 4981 (F, US).<br />
Coconino Co., western foothill region <strong>of</strong> San<br />
Francisco Peak, 2300 m, 21 Aug 1901, fr, J. B.<br />
Leiberg 5883 (US); Oak Creek near Flagstaff,<br />
shaded soil in cañon, 3-7 ft high, 29 Jul 1891, fl,<br />
D. T. MacDougal 465 (US); Oak Creek Canyon,<br />
0.5 miles north <strong>of</strong> Cave Spring Forest Camp, along<br />
stream, 28 Jun 1969, floral buds, Pinkava et al.<br />
16143 (ASU, RSA); Sycamore Canyon Wilderness<br />
Area, Kaibab National Forest, Sycamore Canyon,<br />
ca. 0.5 mi E <strong>of</strong> Sycamore Falls, at the bottom <strong>of</strong><br />
the canyon, on rocky cliffs <strong>and</strong> s<strong>and</strong>y soil,<br />
N35 o 08.193’, W112 o 01.112’, 1930 m, associated<br />
with Pseudotsuga, Acer negundo, Quercus sp.,<br />
Parthenocissus vitacea, Vitis sp., Cornus, Humilus,<br />
Acer glabrum, Robinia pseudoacasia, Ribes, <strong>and</strong><br />
Thalictrum, 15 Jul 2000, fl, J. Wen 4982 (F, US);<br />
SW <strong>of</strong> US route 89A in Sterling Canyon ca. 3.10<br />
km NW <strong>of</strong> the summit <strong>of</strong> Ritter Butte, in rocky<br />
alluvial soil (basalt, limestone, s<strong>and</strong>stone) near<br />
permanent spring in canyon bottom, shaded<br />
riparian/mixed conifer community with Cornus,<br />
Equisetum, Acer <strong>and</strong> Pinus, 5775 ft, 19 Sep 1995,<br />
fr, M. D. Windham 95-268 (NY); Sycamore Canyon<br />
Wilderness Area, Big Spring Canyon, 1.9 km NNW<br />
<strong>of</strong> White Horse Lake, 6280 ft, canyon bottom,<br />
rooted on north-facing limestone rock face, fruits<br />
blue-black, sweet but with bitter aftertaste, 20 Sep<br />
1992, fr, T. Wright 92-332 (ASU, RSA). Gila Co.,<br />
Tonto forest area, along creek – Workman Falls,
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
Fig. 21. Map <strong>of</strong> U.S.A. <strong>and</strong> Mexico showing the distribution <strong>of</strong> <strong>Aralia</strong> bicrenata Wooton & St<strong>and</strong>l.<br />
very scarce with Abies, Acer, Cornus, Cimicifuga,<br />
25 Aug 1976, fl, R. K. Gierisch 3789 (ASU, UNM,<br />
2 sheets); Workman Creek area near falls, Sierra<br />
Anchas Mountains, mixed conifer-oak hillside near<br />
swiftly flowing stream, 7000 ft, 29 Jul 1968, fl, C.<br />
Pase & D. Keil 3461 (RSA); near waterfalls <strong>of</strong><br />
Workman Canyon, rich moist area beside the<br />
Workman Creek, in mixed Douglas fir, Acer, <strong>and</strong><br />
Alnus forest, plant 2-5.5 ft tall, basal leaves ca. 1<br />
m long <strong>and</strong> 1 m wide, young fruits purplish green,<br />
18 Sep 1999, J. Wen 4972 (CS, F, US). Graham<br />
Co., Mt. Graham, 9500 ft, 12 Aug 1934, fl, T. H.<br />
Kearney & R. H. Peebles 9901 (US); Mt. Graham,<br />
22 Jul 1927, fl, R. H. Peebles et al. 4486 (US);<br />
Pinaleno Mts., Swift Trail (Hwy 366), Twilight<br />
Canyon, 13.5 mi W <strong>of</strong> jct <strong>of</strong> Hwy 666, 1.4 mi W <strong>of</strong><br />
Arcadia Campground, permanent stream with<br />
rocky, mossy banks, maple, box elder, Ribes, 2195<br />
m, 28 Jul 1975, fl, T. Reeves 3889 (ASU, 2 sheets).<br />
71<br />
Pima Co., along the trail to Potato Patch, Santa<br />
Catalina Mountains, 7700 ft, 12 Oct 1968, fr, V. L.<br />
Bohrer 1263 (NCU); Pinal Mountains, southcentral<br />
Arizona, south <strong>of</strong> the town <strong>of</strong> Globe,<br />
Sixshooter Canyon Trail, on north side <strong>of</strong> Pinal<br />
Peak, mixed conifer forest, over 2 m tall, 7200 ft,<br />
10 Jul 1982, floral buds, S. Forbes 1702 (ASU);<br />
Santa Catalina Mts, 5-10 Sep 1904, fr, D. Griffiths<br />
7122 (MO); Sierra Ancha Mountains <strong>and</strong><br />
Wilderness Area, 2.25 mi E <strong>of</strong> Young hwy, <strong>of</strong>f FSR<br />
487, 2133 m, near Workman Creek, with Douglasfir,<br />
Acer gr<strong>and</strong>identatum, infrequent, 22 Aug 1984,<br />
young fr, W. Hodgson 3230 (ASU); Sierra Ancha<br />
Wilderness Area in Tonto National Forest,<br />
Reynold’s Creek below Knoll’s Hole Spring, along<br />
trail 150, growing with Pinus ponderosa, Abies<br />
concolor, Pseudotsuga menziesii, Alnus<br />
oblongifolia <strong>and</strong> Acer gr<strong>and</strong>identatum, along<br />
creek, ca. 3 m tall, 7000 ft, 29 Aug 1992, fr, G. J.
72<br />
Imdorf 988 (ASU); Sierra Ancha, Workman Creek<br />
below falls, 5500 ft, 9 Sep 1964, fr, C. P. Pase<br />
1413 (ASU); in rocky gorge in Soldier Camp above<br />
dam, Santa Catalina Mts., 7400 ft, 27 Jul 1944, fl,<br />
W. S. Phillips 2467 (NY). Navaho Reservation,<br />
only in wet caves, total shade, Jul 1916, young fr,<br />
Collector unknown s.n. (NY). COLORADO: La<br />
Plata Co., Elbert Creek ca. 1.2 mi SW <strong>of</strong> conf with<br />
Sawmill Creek <strong>and</strong> Animas River, 7500 ft, 11 Jul<br />
2000, fl, J. Wen 4977 (F, US). Mineral Co., San<br />
Juan Mtns., 5 miles below summit Wolf Creek<br />
Pass-west side, 8000-8500 ft, 28 Jul 1928, fl, C.<br />
B. Wolf 3071 (CAS, RSA, 2 sheets). NEW<br />
MEXICO: Catron Co., Gila National Forest, along<br />
drainage beginning NNE <strong>of</strong> Bearwallow Lookout<br />
Mt., 8160-9800 ft, rich coniferous forest with<br />
Picea, Abies, Pinus, Pseudotsuga, Populus,<br />
Quercus, Acer, 33 o 27’10"-33 o 28’15", 108 o 40’00"-<br />
108 o 40’40", corolla white, scarce, 20 Jul 1984, fl,<br />
T. F. Daniel & S. Nelson 3572 (ASU); the Mogollon<br />
Mts <strong>of</strong> the Gila Wilderness, Gila National Forest,<br />
in the vicinity <strong>of</strong> Mogollon & Whitewater Baldy,<br />
Black, & Sacaton Mts., Silver Drip Cabin trail, by<br />
Big Dry Creek from Apache Springs, perennial<br />
near creek, 4 ft tall, 7500 ft, 1 Sep 1968, fl, W.<br />
Hess 2367 (NCU); ca. 1.5 mi E <strong>of</strong> Mogollon along<br />
route 159, shady moist area under pine-Juglans-<br />
Alnus mixed forest, associated with Parthenocissus<br />
vitacea, Rubus sp., Potentilla sp., Monarda sp.,<br />
Toxicodendron radicans, <strong>and</strong> Rosa sp.,<br />
N33 o 23.037’, W108 o 46.392’, 2090 m, along a<br />
creek on roadside, 13 Jul 2000, fl, J. Wen 4979 (F,<br />
US), 4980 (F, US). Colfax Co., vicinity <strong>of</strong> Ute<br />
Park, 2200-2900 m, 11 Sep 1916, fr, P. C. St<strong>and</strong>ley<br />
14698 (NY, US). Lincoln Co., Capitan Mountains,<br />
Pine Lodge, 29 Jun 1979, floral buds, J. P. Hubbard<br />
s.n. (UNM); gravel loam, vicinity <strong>of</strong> Rio Bonito,<br />
west <strong>of</strong> Bonito Lake, in the White Mountains, 7700<br />
ft, 17 Jul 1971, fl, Hutchins 3492 (UNM); along<br />
shady stream in Sierra Blanca Wilderness area, 27<br />
Jul 1974, young fr, W. Wagner 264 (UNM); South<br />
Fork <strong>of</strong> Rio Bonito, ca. 0.2 mi S <strong>of</strong> South Fork<br />
trailhead, Lincoln National Forest, along Rio<br />
Bonito, N33 o 26.806’, W105 o 45.523’, 2325 m,<br />
associated with Urtica, Monarda, Pinus,<br />
Pseudotsuga, Ranunculus, <strong>and</strong> Rubus, uncommon,<br />
16 Jul 2000, fl, J. Wen 4983 (F, US); White Mts.,<br />
Eagle Creek Canyon above (1 mi W <strong>of</strong>) the summer<br />
resort cabins, 2500 m, 1 Aug 1984, fl, R. D.<br />
Worthington 12276 (NY). Los Alamos Co., Apache<br />
Spring, B<strong>and</strong>elier National Monument, 7900 ft, 27<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
Jul 1975, fl, R. Halley 49 (UNM); Water Canyon,<br />
7100 ft, riparian habitat, 6 Sep 1978, fr, G. Tierney<br />
& T. Foxx 17 (UNM). Rio Arriba Co., vicinity <strong>of</strong><br />
Brazos Canyon, swamp, 21 Aug 1914, fl, P. C.<br />
St<strong>and</strong>ley & H. C. Bollman 10700 (US, 2 sheets);<br />
N. Paleo Cr., Camp Zia, San Pedro Parks Wild<br />
Area, San Pedro Mts., 22 Aug 1965, young fr, A.<br />
Fleck s.n. (UNM). S<strong>and</strong>oval Co., Jemez<br />
Mountains, in low ground near stream at recreation<br />
ground north <strong>of</strong> Jemez Springs, 25 Aug 1936, fr,<br />
M. Marcelline 2269 1/2 (F). San Miguel Co.,<br />
vicinity <strong>of</strong> Las Vegas, Aug 1923, fr, Anect 135<br />
(US); Las Vegas, Porvenior Creek, 2400-2800 m,<br />
6 Sep 1926, fr, G. Arsène 17733 (US); Las Vegas<br />
Hot Springs, Aug 1901, fl, H. S. Barber 151 (US).<br />
Santa Fe Co., Santa Clara Canyon, in shady place<br />
near stream at Rancher’s Cabin Transition, 1 Jul<br />
1936, fl, M. Marcelline 1905 (F). Socorro Co.,<br />
gravel loam <strong>of</strong> stream area in South Canyon, in<br />
the Magdelena Mountains, 8000 ft, 28 Jul 1973,<br />
floral buds, Hutchins 4597 (UNM); Mogollon Mts.,<br />
on Mogollon Creek, 23 Jul 1903, fl, 7500 ft., O. B.<br />
Metcalfe 303 (G, NY, P, RSA, US). Torrance Co.,<br />
Manzano Mts., Trigo Canyon, ¼ mile E <strong>of</strong><br />
Kennedy Camp, 2 Sep 1963, fr, E. J. Bedker 1543<br />
(UNM). Union Co., on <strong>and</strong> near the Sierra Gr<strong>and</strong>e,<br />
2100-2925 m, moist thickets, 19 Jun 1911, sterile,<br />
P. C. St<strong>and</strong>ley 6136 (US). Valencia Co., Seboyeta<br />
Canyon bottom, riparian habitat near head <strong>of</strong><br />
canyon, growing in fine moist soil in shade, 7200<br />
ft, 25 Sep 1977, fr, G. A. Marley 912 (UNM).<br />
TEXAS: Jeff Davis Co., Mt. Livermore, 21 Sep<br />
1935, L. C. Hinckley s.n. (SRSC); North top Mt.<br />
Livermore, 21 Aug 1935, fl, with larger fruited<br />
specimen mounted on the same sheet which was<br />
noted to be collected 21 Sep 1935, fr, L. C. Hinckley<br />
405 (F, NY, 2 sheets); Davis Mountains Preserve,<br />
N side, Mt. Livermore, base <strong>of</strong> Palisades <strong>of</strong><br />
“Laura’s Rock,” N30.63898, W104.17350, 7900<br />
ft., apen/Gambel oak/talus, 9 Aug 2002, in fr., J. P.<br />
Karges 2002-08095-6 (SRSC); Mt. Livermore,<br />
above Bridge Gap, along road, below radio shack,<br />
31 Jul 1997, in fr., K. Lund et al. 263 (SRSC); Talus<br />
<strong>and</strong> slopes below high north-facing bluffs <strong>of</strong> Mt.<br />
Livermore, 2200 m, 3 Jun 1928, sterile, E. J.<br />
Palmer 34334 (NY); Mt. Livermore, Davis Mts.,<br />
upper slopes <strong>and</strong> near the peak, lower margin <strong>of</strong><br />
talus, E <strong>of</strong> peak, under Gambel oak, 4 Sep 1982,<br />
in fr., A. M. Powell & S. Powell 3921 (SRSC); Mt.<br />
Livermore, 19 Oct 1935, fr, O. E. Sperry T198 (US,<br />
2 sheets); infrequent low shrub above upper spring,
Madera Canyon, S slope <strong>of</strong> Mt. Livermore, igneous<br />
soil, 13 Sep 1947, in fl., 7800 ft., B. H. Warnock<br />
7511 (SRSC); infrequent herb, high on N slopes<br />
<strong>of</strong> Mt. Livermore, rare, only one plant seen,<br />
igneous soil beneath oaks, Don McIvor Ranch, 25<br />
Aug 1977, in fr., B. H. Warnock 21518 (SRSC);<br />
Mt. Livermore, near the top, N30 o 38.153’,<br />
W104 o 10.456’, 8232 ft, in rocky area at the edge<br />
<strong>of</strong> forest, associated with Quercus depressipes,<br />
Quercus spp., Prunus serotina, Holodiscus<br />
discolor, Astragalus giganteus, <strong>and</strong> Ptelea sp., 22<br />
Aug 2003, fr, fruits dark purple, J. Wen 7250 (F,<br />
US). UTAH: Kane Co., Water Canyon drainage<br />
area, ca. 7.5 miles west <strong>of</strong> Kanab, 6100 ft, 2 Jul<br />
1985, floral buds, J. S. Tuhy 2326 (NY, 2 sheets,<br />
one sterile, one with floral buds). Washington Co.,<br />
Zion Cañon, 25 Jun 1933, floral buds, A. Eastwood<br />
& J. T. Howell 1173 (CAS); Zion Cañon, 18 Jul<br />
1922, fl, M. E. Jones s.n. (DS, RSA, US); Zion<br />
Narrows trail head vicinity, 4500 ft, 9 Jul 1987, fl<br />
& young fr, S. L. Welsh 23829 (NY); Zion grotto,<br />
25 Jun 1928, floral buds, A. M. Woodbury 27 (US).<br />
This species has <strong>of</strong>ten been treated as part <strong>of</strong><br />
<strong>Aralia</strong> racemosa, either as a synonym (Smith<br />
1944), or as a subspecies (Welsh <strong>and</strong> Atwood<br />
1975). Kearney <strong>and</strong> Peebles (1951) also listed A.<br />
bicrenata as a synonym <strong>of</strong> A. racemosa; yet they<br />
also noted morphological differences between the<br />
two. In particular, their inflorescence architecture<br />
is highly distinct with A. bicrenata having the<br />
corymbose architecture <strong>and</strong> A. racemosa with the<br />
racemose overall structure. The stipule <strong>of</strong> A.<br />
bicrenata is <strong>of</strong>ten leafy, whereas that <strong>of</strong> A.<br />
racemosa is small, nonleafy <strong>and</strong> triangular. The<br />
stem <strong>of</strong> A. bicrenata is green to purplish green<br />
whereas that <strong>of</strong> A. racemosa is dark purple.<br />
Furthermore the leaflets <strong>of</strong> A. bicrenata are <strong>of</strong>ten<br />
truncate to subcordate at base, <strong>and</strong> those <strong>of</strong> A.<br />
racemosa are cordate to deeply so.<br />
Kearney <strong>and</strong> Peebles (1951) documented that<br />
the fruits <strong>of</strong> A. bicrenata are eaten by various birds.<br />
4. <strong>Aralia</strong> cordata Thunb. — Fig. 22; color plate<br />
4: A-C.<br />
<strong>Aralia</strong> cordata Thunb., Fl. Jap. 127. 1784. TYPE:<br />
Japan. C. P. Thunberg 7577 (lectotype: UPS-<br />
THUNB, here designated).<br />
<strong>Aralia</strong> nudicaulis Blume, Bijdr.: 870. 1826, nom.<br />
illeg.<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
73<br />
<strong>Aralia</strong> edulis Siebold & Zucc., Fl. Jap. 1: 57, t. 25.<br />
1837. Dimorphanthus edulis (Siebold & Zucc.)<br />
Miq., Comm. Phytogr. 3: 96. 1840. TYPE:<br />
Japan. L 0326346 (L-898,125-0047), “folia<br />
radices edulis,” Siebold 107 (lectotype: L!, here<br />
designated; this specimen bears the h<strong>and</strong> <strong>of</strong><br />
Zuccarini, “<strong>Aralia</strong> edulis S & Z. Japonica”;<br />
isolectotype: GH!).<br />
<strong>Aralia</strong> nutans Franch. & Sav., Enum. Pl. Jap. 2:<br />
376. 1878. TYPE: Japan. Hokkaido, Savatier<br />
3835 (holotype: P!; isotype: P!).<br />
Perennial herb 1.5-2 m tall, <strong>and</strong>romonoecious.<br />
Lower part <strong>of</strong> stem with stiff hairs. Rhizomes stout<br />
<strong>and</strong> horizontal. Leaves 60-100 cm long <strong>and</strong> wide,<br />
tripinnately to bipinnately ternate; upper leaves<br />
pinnate to bipinnate or trifoliolate; stipules 2-2.3<br />
cm long, 0.7-1 cm wide, narrowly triangular, <strong>of</strong>ten<br />
becoming leaf-like <strong>and</strong> enlarged, especially for<br />
those on the lower part <strong>of</strong> the stem; petioles 12-22<br />
cm long, pubescent with s<strong>of</strong>t <strong>and</strong> scabrous hairs;<br />
leaflets 5.5-22 cm long, 4-10.5 cm wide, papery,<br />
basal leaflets ovate, other leaflets ovate to narrowly<br />
so, acuminate to acute at apex, cordate, subcordate<br />
to rounded at base, <strong>of</strong>ten oblique on lateral leaflets,<br />
serrate to serrulate at margin; lateral veins 8-10 on<br />
each side, conspicuous on both surfaces, sparsely<br />
scabrous on adaxial surface, pilose along veins <strong>and</strong><br />
veinlets on abaxial surface, petiolules 0-2 cm long,<br />
pilose. Inflorescence 30-65 cm long, a terminal<br />
panicle with the lower 2-5 primary branches<br />
subtended by leaves rather than bracts, pilose,<br />
consisting <strong>of</strong> 20-50 primary branches, mostly<br />
racemosely or sometimes panicunately arranged<br />
on a main axis, <strong>of</strong>ten 2-8 forming a circle on the<br />
upper half <strong>of</strong> the main inflorescence axis, each<br />
primary branch 3-22 cm long, consisting <strong>of</strong> 1-10<br />
umbels; terminal umbels mostly 35-95-flowered,<br />
pedicels 9-14 mm long, pubescent, tip <strong>of</strong> pedicels<br />
enlarged <strong>and</strong> with a cluster <strong>of</strong> scabrous hairs; lateral<br />
umbels 18-35-flowered, pedicels 7-9 mm long;<br />
bracts <strong>of</strong> primary branches 4-9 mm long, 0.6-0.8<br />
mm wide, linear, pilose; bracteoles 1.6-2 mm long,<br />
0.1-0.2 mm wide, linear, pilose. Sepals 0.3-0.4<br />
mm long, 0.2-0.3 mm wide, narrowly triangular;<br />
petals 1.5-1.8 mm long, 0.9-1 mm wide, ovate, with<br />
a conspicuous vascular bundle in the middle;<br />
stamens 5, 2.2-2.3 mm long, filaments 1.7-2 mm<br />
long, anthers 0.7-0.8 mm long, 0.4-0.5 mm wide,<br />
ovate; ovaries 5-locular, styles 0.6-0.8 mm long at<br />
anthesis, base <strong>of</strong> ovary slightly pilose with a few
74<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
Fig. 22. <strong>Aralia</strong> cordata Thunb. A. Habit with leaf <strong>and</strong> inflorescences. B. Leafy stipule. C. Close-up <strong>of</strong> leaflet margin <strong>and</strong><br />
veinlets on lower surface. D. Flower. E. Young fruit. F. Fruit.
scattered hairs, the top part <strong>of</strong> the ovary<br />
conspicuously exposed <strong>and</strong> appearing like a<br />
stylopodium. Fruits 3.2-3.5 mm long <strong>and</strong> wide,<br />
globose, persistent styles 5-divided, connate at the<br />
base.<br />
Common names: udo.<br />
Uses: young shoots as vegetables.<br />
Phenology: flowering in July to September;<br />
fruiting in August to November.<br />
Distribution: widely distributed in Japan (Fig.<br />
23), also in Ullung-do <strong>of</strong> Korea.<br />
Ecology: deciduous forests, or mixed broadleaf<br />
<strong>and</strong> coniferous forests, edges <strong>of</strong> forests, roadsides<br />
near forests; 50-1500 (-2000) m.<br />
Additional specimens examined: Additional<br />
specimens examined: Japan. Hokkaido: Insula<br />
Jesso, circa Hakodate, 1861, Albrecht s.n. (K, NY,<br />
part 2 <strong>of</strong> the specimen only, part 1 is <strong>Aralia</strong> elata<br />
var. elata); west side <strong>of</strong> Shibetsu city, ca 6 km E <strong>of</strong><br />
Soeushinai, Hokkaido Temporary Game Preserve,<br />
280 m, mixed forests <strong>of</strong> Quercus, Acer <strong>and</strong> Betula<br />
<strong>and</strong> some gymnosperms, 1 Sep 1977, fl, D. E.<br />
Boufford & E. W. Wood 19842 (RSA); Suto-gun,<br />
Kuromatsunai-cho, along a small tributary <strong>of</strong> the<br />
Higashi-gawa River, 230 m, mixed deciduous –<br />
Abies forest with Sasa, 28 Jul 1980, fl, D. E.<br />
Boufford & M. Kato 22429 (RSA); Hakodadi,<br />
1860, C. P. Hoolpson s.n. (K); Yeso, Lingriff 788<br />
(WU); Hokkaido, Mt. Shiribeshi (Yotei-zan),<br />
Makkari course, 4 Aug 1982, fl, H. Takahashi 2853<br />
(RSA); Prov. Kitami, Esashi-gun, Utanobori-cho,<br />
Pankenai-sawa – Futamata-sawa, c. 200 m, 5 Aug<br />
1986, fl, H. Takahashi et al. 6599 (RSA); Hakodadi<br />
[Hakodate], 1859, C. Wilford s.n. (K); Hokadodi<br />
[Hakodate], Jul 1859, C. Wilford s.n. (K). Honshu:<br />
Aomori Pref., Aomori, Oct 1899, Faurie 3322<br />
(WU). Gunma Pref., Tomioka, 1866-1876,<br />
Savatier s.n. (K). Iwate Pref., Kesen-gun, Sanrikumachi,<br />
Okkirai — Koobezaki, 25 Jul 1978, J.<br />
Murata et al. 5993 (TI); Iwate Pref., Hayachine<br />
National Park, on upper slopes above 1300 m on<br />
Mt. Hayachine, serpentine rock, 24 Jul 1984, fl,<br />
D. Sather & G. Caddell 4914 (NCU, 2 sheets).<br />
Kanagawa Pref., Hayama-machi, 2 km from the<br />
coast, 19 Sep 1969, fl, 50 m, beside road along<br />
Shimoyama River, perennial ca. 5 feet high, R. M.<br />
Beauchamp 1033 (RSA, 2 sheets); Yokohama,<br />
1862, Maximowicz s.n. (K, W); Yokohama, Sep<br />
1876, fl, J. P. Bisset s.n. (BM, K); Yokohama, Milne<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
75<br />
s.n. (BM); Nippon media, circa Yokoska<br />
[Yokosuka], Savatier 523 (K, 2 sheets). Kyoto<br />
Pref., SSW slope <strong>of</strong> the Mt. Mikunidake, Miyamacho,<br />
Kitakuwada-gun, 650 m, 27 Aug 1992, fl, S.<br />
Tsugaru & T. Takahashi 16775 (MO, 2 sheets);<br />
Yamashiro, Mt. Hieizan, 14 Aug 1931, floral buds,<br />
M. Tagawa 224 (RSA). Miyagi Pref., Mts. Zao,<br />
Seikei-daichi, 1500 m, 16-19 Jul 1963, H. Ohashi<br />
s.n. (TI); circa Sendai, Oct 1903, U. Faurie s.n.<br />
(BM). Nagano Pref., Karuizawa, 7 Aug 1912, fl,<br />
H. E. Fox s.n. (BM); Nagano Pref., Prov. Shinano,<br />
Shimoinagun, Iida, Mt. Nenjo, Kamimatsukawa,<br />
1500 m, 12 Aug 1963, T. Yamazaki & K. Asano<br />
7518 (TI); Nagano Pref., Prov. Shinano, Hondo,<br />
Suga-daira Osa-son (new Suga-daira Sanada-choo<br />
Chiisagata-gun) Chiisagata-gun, 1300 m, 15 Aug<br />
1953, fl & young fr, M. Furuse 26619 (K, 3 sheets<br />
<strong>of</strong> an entire plant); Nagano Pref., Norikura Heights,<br />
30 km SW <strong>of</strong> Matsumoto, margin <strong>of</strong> deciduous<br />
woodl<strong>and</strong>, 1500 m, 5 Sep 1993, fl & young fr, R.<br />
K. Brummitt 19027 (K); Nagano Pref., North Alps<br />
National Park, along trail between Kamikochi <strong>and</strong><br />
Mt. Hodakka, 20 Sep 1983, young fr, D. Sather &<br />
G. Caddell 2749 (NCU). Niigata Pref., Isl. Sado,<br />
Aikawa-Cho, Bunadaira-yama, Ookuragawa, 600<br />
m, 20 Aug 1985, T. & F. Yamazaki 5069 (KUN);<br />
Basi Mt., Echigo-Koma, 4-6 Oct 1949, J. Ohwi<br />
s.n. (K, 2 sheets). Shizuoka Pref., Ihara-gun,<br />
Fujigawa-machi, along the Arinase River, 150-350<br />
m, 21 Oct 1978, Y. Tateishi et al. 4489 (U);<br />
Shizuoka Pref., Shizuoka-shi, Umegashima,<br />
Ohyakuzure, 35 o 18’N, 138 o 18’E, 1300-1400 m, 5<br />
Oct 1990, fr, Y. Tateishi et al. 16089 (MO). Tochigi<br />
Pref., N side <strong>of</strong> Kansei Pass Hwy, at rest area below<br />
Yumoto Toll gates, Nikko National Park, 15 Sep<br />
1969, fl, 2000 m, R. M. Beauchamp 1003 (RSA);<br />
Prov. Shimotsuke, Hondo, Nokado Kuriyama-son<br />
Shiwoya-gun, 10 Sep 1967, fr, M. Furuse 45807<br />
(K, 2 sheets); Nikko, woods near Nikko Boranical<br />
Garden, Aug 1995, J. Wen 2456 (US). Tokyo,<br />
Oizumi, Nerima-ku, 4 Oct 1926, young fr, T.<br />
Makino s.n. (CAS); Oizumi, Nerima-ku, 19 Sep<br />
1937, fl, T. Makino s.n. (CAS); Tokyo, Sengenone,<br />
from Fussawa to Koiwa, Hinohara-mura,<br />
Nishitama, 22 Sep 1968, H. Ohba 680914 (TI);<br />
woods near Tokyo, J. Wen 2476 (US); Chiba-Ken,<br />
Narita-Shi, 500 Tokko, in secondary woods behind<br />
Hotel Nikko Narita, 21 May 2005, sterile, J. Wen<br />
8542 (US). Toyama Pref.: Arimine, Kaminiikawagun,<br />
1000 m, 4 Aug 1965, T. Yamazaki 9779 (TI).<br />
Wakayama Pref., along the Komori-rindo, Ryujin-
76<br />
mura, Hidaka-gun, sunny steep slope by roadside<br />
in deciduous forest, 1200 m, 15 Oct 1989, late fl,<br />
T. Takahashi et al. 1393 (MO); Wakayama Pref.,<br />
Higashimuro, Kumanogawa-cho, Shikiya, foot <strong>of</strong><br />
the mountain, 50 m, 20 Sep 1964, young fr, young<br />
stem edible, H. Matsushita 196 (RSA). Yamanashi<br />
Pref.: Oshino-mura, NW foot <strong>of</strong> Mt. Fuji, 900-1000<br />
m, 23 Oct 1966, M. Togashi et al. s.n. (TI); W side<br />
<strong>of</strong> Mt. Fuji, 1500 m, 30 Aug 1969, fl, H. Ohba<br />
69828 (BM, 2 sheets). Kyushu: Kumamoto<br />
Prefecture, Mount Aso National Park, in grassl<strong>and</strong><br />
at 700 m, slopes <strong>of</strong> Mt. Aso, 11 Aug 1984, fl, D.<br />
Sather & G. Caddell 5544 (NCU, 2 sheets).<br />
Nagasaki Pref., Nagasaki, 1862, R. Oldham s.n.<br />
(K); Nagasaki, Aug 1862, floral buds, cultivated<br />
as eatable vegetable, R. Oldham 837 (K); Nagasaki,<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
Fig. 23. Map <strong>of</strong> Japan showing the distribution <strong>of</strong> <strong>Aralia</strong> cordata Thunb.<br />
1862, R. Oldham 310 (K, 2 sheets); circa Nagasaki,<br />
Sep 1864, fl, C. J. Maximowicz 10960 (BM). Saga<br />
Pref., Mt. Seburi, at 800 m on road to summit <strong>of</strong><br />
Mt. Seburi, 10 Oct 1984, fl, D. Sather & G. Caddell<br />
6605 (NCU, 2 sheets). Shikoku: Ehime Pref.:<br />
Kamiukena-gun, Yanagidani-mura, Odamiyamakokuyuurin,<br />
850 m, 20 Oct 1980, J. Murata 10106<br />
(TI, 2 sheets); Ehime-Ken, Kamiukena-gun,<br />
Kuma-cho, Higasimyojin, 600-700 m, 26 Oct 1995,<br />
late fl & young fr, Y. Tateishi et al. 40570 (TNM).<br />
Japan, location unknown: 11 Aug, part <strong>of</strong> the<br />
Siebold collection distributed from L, Herb. Bürger<br />
s.n. (K). Japan, Noeji, 4 Aug 1889, floral buds, P.<br />
Faurie 806 (K); Montagun, Ittomori, 11 Sep 1889,<br />
P. Faurie 1159 (K); Japan, Aug 1886, P. Faurie<br />
1284 (K); 10 Aug 1905, U. Faurie 6849 (BM).
Korea. Ullung-do, path along ridge to NE <strong>of</strong> Nam<br />
Yang, infrequent perennial shrub, height & spread<br />
3.5 m, growing in acid loam in full sun, 2 Oct 1982,<br />
fr, Beyer et al. 119 (K, 2 sheets).<br />
Specimens <strong>of</strong> the taxon examined in the Siebold<br />
collection from Japan in the Leiden Herbarium (L):<br />
L 0171191 (L-898,125-0048), Keuy, “specimen<br />
authenticum ad florum japonicum conficiendam<br />
adribitum,” Siebold s.n. [in fl.]; L 0171192 (L-<br />
898,125-0052), 1829, “specimena authentica ad<br />
florum jap. conficiendam adribita,” Siebold s.n. [in<br />
fl. buds]; L 0326339 (L-898,125-0049), Siebold<br />
s.n. [in fl.]; L 0326340 (L-898,125-0051), “fol.<br />
composita!” Siebold s.n. [leaf only]; L 0326341<br />
(L-898,125-0060), specimen on the right side <strong>of</strong><br />
the sheet only, Burger s.n. [in fl.]; L 0326342 (L-<br />
898,125-0061), Burger s.n. [in fl. & floral buds];<br />
L 0326343 (L-898,125-0063), Burger s.n. [in fl.];<br />
L 0326344 (L-898,125-0062), Burger s.n. [in fl.];<br />
L 0326345 (L-898,125-0059), Burger s.n. [in fl.];<br />
L 0423961 (L-898,125-0057), Dosen – Shaw,<br />
Mohnike s.n. [in fl.]; L 0423962 (L-898,125-0055),<br />
Textor s.n. [in fl.]; L 0423963 (L-898,125-0058),<br />
Dosen, Mohnike 51 [in fl.]; L 0423964 (L-898,125-<br />
0050), Siebold 1055 [leaf only]; L 0423965 (L-<br />
898,125-0056), Textor s.n. [infl.].<br />
Cultivated material: Japan. Botanic Garden<br />
<strong>of</strong> the Faculty <strong>of</strong> Science, the University <strong>of</strong><br />
Kanazawa, 12 Sep 1966, S. Yoshitake & S. Kaneda<br />
s.n. (TI). Meguro, Meguro-ku, Tokyo Pref., 14 Aug<br />
1900, fl, T. Makino s.n. (CAS). Oizumi, Nerimaku,<br />
Tokyo Pref., 2 Nov 1931, sterile, T. Makino<br />
s.n. (CAS). Netherl<strong>and</strong>s. Rotterdam, Hortus<br />
Ramlehstraat, 10 Aug 1950, B. K. Boom B3058<br />
(L). Also cultivated in the Great Britain (see cover<br />
<strong>of</strong> Frodin <strong>and</strong> Govaerts 2003).<br />
The isolectotype <strong>of</strong> <strong>Aralia</strong> edulis Siebold &<br />
Zucc. at GH (with leaves <strong>and</strong> flowers) bears the<br />
h<strong>and</strong>writing <strong>of</strong> Siebold “<strong>Aralia</strong> edulis S. & Z.” It<br />
was a specimen originally from Herb. Lugd. Batav.<br />
(L).<br />
5. <strong>Aralia</strong> schmidtii Pojark. — Fig. 24; color plate<br />
4: D-E.<br />
<strong>Aralia</strong> schmidtii Pojark., Fl. URSS 16: 588. 1950.<br />
TYPE: Russia. Sakhalin, near Due, 2 Aug<br />
1860, P. Glehn s.n. (holotype: LE!; isotypes:<br />
G!, K!, NY!).<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
77<br />
<strong>Aralia</strong> racemosa L. var. sachalinensis Regel, Index<br />
Seminum Hort. Bot. Petr. 1864: 22. 1864.<br />
<strong>Aralia</strong> cordata Thunb. var. sachalinensis<br />
(Regel) Nakai, J. Arnold Arbor. 1924. TYPE:<br />
Russia. Sachalin, 1860, F. Schmidt s.n.<br />
(lectotype: LE!, here designated; isolectotypes:<br />
G!, GH!, LE!, NY!).<br />
<strong>Aralia</strong> sachalinensis hort. ex Sieb. et Voss in Vilm.<br />
Blumengärten, 3 Aufl. I, 403. 1896, nom. nud.<br />
Perennial herb 1-3 m tall. Leaves ternately<br />
compound, upper leaves pinnate to bipinnate or<br />
trifoliolate; stipules 5-13 mm long, 1.6-4.3 mm<br />
wide, lanceolate, sparsely covered with a few hairs,<br />
ciliate at margin; leaflets 6-21 cm long, 3-12.5 cm<br />
wide, papery, basal leaflets <strong>of</strong> the pinnae ovate,<br />
other leaflets ovate to narrowly so, acuminate at<br />
apex, cordate to subcordate at base, <strong>of</strong>ten oblique<br />
on lateral leaflets, serrulate at margin; lateral veins<br />
9-11 on each side, conspicuous on both surfaces,<br />
sparsely scabrous on adaxial surface, <strong>of</strong>ten with<br />
scattered hairs on veins <strong>and</strong> veinlets, pubescent<br />
with scabrous hairs on abaxial surface, petiolules<br />
0.1-2.4 cm long, pilose. Inflorescence 30-50 cm<br />
long, a terminal panicle with the lower 3-4 primary<br />
branches subtended by leaves rather than bracts,<br />
pilose, consisting <strong>of</strong> 20-30 primary inflorescence<br />
branches, mostly racemosely or sometimes<br />
panicunately arranged on a main axis, <strong>of</strong>ten 3-5<br />
forming a circle on the upper part <strong>of</strong> the main<br />
inflorescence axis, each primary branch 6-12 cm<br />
long, consisting <strong>of</strong> 1-3 umbels, usually only the<br />
terminal umbel <strong>of</strong> each branch bearing fruits, others<br />
becoming aborted or functionally male; terminal<br />
umbels mostly (25-) 35-60-flowered, pedicels 15-<br />
22 mm long, pubescent, tip <strong>of</strong> pedicels enlarged<br />
<strong>and</strong> with a cluster <strong>of</strong> hairs; lateral umbels 18-25flowered,<br />
pedicels 9-15 mm long; bracts <strong>of</strong> primary<br />
branches 5-10 mm long, 0.7-0.9 mm wide, linear,<br />
somewhat pilose; bracteoles 1.7-2.2 mm long, 0.2-<br />
0.3 mm wide, linear, pilose. Sepals 0.3-0.4 mm<br />
long, 0.2-0.3 mm wide, narrowly triangular; petals<br />
1.6-2 mm long, 1-1.1 mm wide, ovate, with a<br />
conspicuous vascular bundle in the middle;<br />
stamens 5, 2-2.3 mm long, filaments 1.8-2 mm<br />
long, anthers 0.6-0.8 mm long, ca. 0.5 mm wide,<br />
ovate; ovaries 5-locular, styles 0.6-0.8 mm long at<br />
anthesis, connate at base, base <strong>of</strong> ovary slightly<br />
pilose with a few scattered hairs, the top part <strong>of</strong><br />
the ovary conspicuously exposed <strong>and</strong> appearing<br />
like a stylopodium. Fruits 3-3.2 mm long <strong>and</strong> wide,
78<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
Fig. 24. <strong>Aralia</strong> schimdtii Pojark. A. Branch showing upper leaf <strong>and</strong> axillary inflorescence. B. Lanceolate stipule. C. Leaflet<br />
from a lower leaf. D. Leaflet lower surface showing pubescence. E. Umbel. F. Floral bud with pedicel. G. Flower with<br />
pedicel. H. Flower. I. Young fruit (A, D, & F-H – Gage 2512, NY; B, C, E & I – Gage 2632, NY).
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
Fig. 25. Map <strong>of</strong> eastern Asia showing the distribution <strong>of</strong> <strong>Aralia</strong> continentalis Kitag. in China, far eastern Russia, <strong>and</strong> Korea, <strong>and</strong><br />
A. schmidtii Pojark. in Sakhalin <strong>and</strong> Kuril isl<strong>and</strong>s.<br />
globose, persistent styles connate at the base<br />
(divided roughly to the middle).<br />
Phenology: flowering in July-August; fruiting<br />
in August to September.<br />
Distribution: Sakhalin <strong>and</strong> Kuril isl<strong>and</strong>s, Russia<br />
(Fig. 25).<br />
Ecology: Deciduous forests near sea shore.<br />
Additional specimens examined: Russia.<br />
Kuril Isl<strong>and</strong>s, Kurilsk Isl<strong>and</strong>, 20 Jul 1982, floral<br />
buds, V. Barkalov s.n. (MW); Kuril Archipelago,<br />
Iturup, 5 Aug 1988, fl, V. Barkalov s.n. (MW); Kuril<br />
Archipelago, Iturup, Konservnaya Bay, steep slope<br />
above beach, dominated by Sasa, Filipendula with<br />
scattered trees <strong>of</strong> Salix, 45 o 20’7"N, 147 o 59’32"E,<br />
herb 2-3 m tall with huge stipulate compound<br />
leaves, 19 Aug 1996, fl, S. Gage 2512 (NY); Iturup,<br />
inl<strong>and</strong> <strong>of</strong> Dobroye Nachalo Bay, at Lake Natasha,<br />
forested area near lake, with Abies, Prunus,<br />
79<br />
Kalopanax, 44 o 46’11’N, 147 o 11’4"E, 22 Aug 1996,<br />
fl, S. Gage 2632 (NY); Kuril Isl<strong>and</strong>s, Kunashir<br />
Isl<strong>and</strong>, 19 Jul 1985, floral buds, N. Shvedtchikova<br />
s.n. (MW); Kuril Isl<strong>and</strong>s, Kunashir Isl<strong>and</strong>, 26 Aug<br />
1986, young fr, N. Shvedtchikova s.n. (MW).<br />
Sakhalin, Cholmsk, 10 Aug 1978, fl, M. Ignotov<br />
s.n. (MW); Sachalin, western shore, near the mouth<br />
<strong>of</strong> River Axzegu, 31 Aug 1924, A. Krysht<strong>of</strong>ovich<br />
s.n. (LE, 3 sheets); Sachalin, western shore,<br />
Agnevo, 14 Jul 1925, A. Krysht<strong>of</strong>ovich 171 (LE);<br />
Sachalin, Dolinsk, 11 Aug 1952, fl, E. Motorina<br />
(MW). Southern Sachalin, on W side <strong>of</strong> the<br />
mountain slope on the SW shore, Valley Rantomar<br />
or Apple Valley, 20 Aug 1950, M. G. Popov s.n.<br />
(LE); Insula Sachalin, Kasanai, Aug 1860, F.<br />
Schmidt s.n. (LE); Sachalin, 1861, F. Schmidt<br />
18202 (BM); Sachalin, Korsakov, 20 Aug 1986,<br />
N. Shvedtchikova s.n. (MW); Sachalin, Kuril<br />
Isl<strong>and</strong>s, Kunashir Islet, 24 Jul 1986, floral buds,<br />
M. Stolarskaya & M. Maschkova s.n. (MW);
80<br />
Sachalin, Augustinowiz 71 (K); Sagalien, Aug<br />
1908, Faurie 398 (BM).<br />
Nakai (1924) cited <strong>Aralia</strong> racemosa var.<br />
sachalinensis Regel as originally published by<br />
Regel (1864b; in Gartenfl. 13: 100. t. 432. 1864).<br />
Regel (1864a) also provided a brief description <strong>of</strong><br />
<strong>Aralia</strong> racemosa var. sachalinensis (in Index<br />
Seminum Hort. Bot. Petr. 1864: 22). Regel (1864a)<br />
actually has nomenclatural priority, as it was<br />
published on March 8, whereas Regel (1864b)<br />
appeared in April. Regel (1864b) is a detailed<br />
description with an excellent illustration.<br />
When Pojarkova (1950) originally published<br />
<strong>Aralia</strong> schmidtii, she clearly indicated her <strong>Aralia</strong><br />
schmidtii as a new species (not a new name), <strong>and</strong><br />
treated <strong>Aralia</strong> racemosa var. sachalinensis Regel<br />
as a synonym. She used a different type, as A.<br />
racemosa var. sachalinensis was described based<br />
on a plant raised from seeds collected by Schmidt.<br />
<strong>Aralia</strong> schmidtii differs from the closely related<br />
A. cordata by the former’s longer pedicels <strong>and</strong><br />
more numerous flowers per terminal umbel.<br />
6. <strong>Aralia</strong> taiwaniana Y. C. Liu & F. Y. Lu — Color<br />
plate 5: A-H.<br />
<strong>Aralia</strong> taiwaniana Y. C. Liu & F. Y. Lu in Y. C.<br />
Liu, Quart. J. Chinese Forest. 9: 136, t. 8. 1976.<br />
TYPE: China. TAIWAN: Nantou Hsien, F. Y.<br />
Lu & C. H. Ou 1507 (holotype: NCUF!).<br />
Perennial spreading herb 1-2 m tall,<br />
<strong>and</strong>romonoecious. Stem pubescent. Rhizomes<br />
thick. Leaves ternately compound, lower leaves<br />
75-90 cm long <strong>and</strong> wide, tripinnately ternate, upper<br />
leaves pinnate to bipinnate; stipules 4-10 mm long,<br />
2-4 mm wide, narrowly triangular to lanceolate,<br />
pilose, sparsely ciliate at margin; leaflets 5.5-15<br />
cm long, 3-9.5 cm wide, papery, the basal leaflets<br />
<strong>of</strong> the pinnae ovate, other leaflets ovate to narrowly<br />
so, acuminate to abruptly so at apex, truncate,<br />
rounded to subcordate or sometimes cordate at<br />
base, <strong>of</strong>ten oblique on lateral leaflets, finely serrate<br />
to doubly so at margin; lateral veins 8-10 on each<br />
side, conspicuous on both surfaces, scabrous on<br />
adaxial surface, <strong>of</strong>ten with scattered hairs on veins<br />
<strong>and</strong> veinlets, pilose on abaxial surface, petiolules<br />
<strong>of</strong> lateral leaflets 0.3-2.5 cm long, pilose.<br />
Inflorescence a terminal or axillary panicle, pilose,<br />
terminal inflorescence 20-55 cm long, consisting<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
<strong>of</strong> 10-30 primary inflorescence branches, more or<br />
less racemosely arranged on a main axis,<br />
sometimes 6-12 forming a circle on the upper part<br />
<strong>of</strong> the main inflorescence axis, or forming 2-3<br />
circles on the inflorescence axis, each primary<br />
branch 6-18 cm long, consisting <strong>of</strong> 2-7 umbels,<br />
terminal umbels mostly (25-) 35-65-flowered,<br />
pedicels 15-20 mm long, slender <strong>and</strong> pilose, tip <strong>of</strong><br />
pedicels enlarged <strong>and</strong> with a cluster <strong>of</strong> hairs; lateral<br />
umbels not well developed, usually <strong>of</strong> male<br />
flowers, 15-25-flowered, pedicels 3-12 mm long;<br />
bracts <strong>of</strong> primary branches 5-20 mm long, 0.7-2<br />
mm wide, linear, pilose, sometimes becoming<br />
leafy; bracteoles 1.8-2 mm long, 0.3-0.4 mm wide,<br />
linear, pilose. Sepals 0.3-0.5 mm long, 0.4-0.5 mm<br />
wide, triangular; petals 1.8-2 mm long, 0.9-1.1 mm<br />
wide, ovate, with a conspicuous vascular bundle<br />
in the middle, tip slightly enlarged; stamens 5, 1.7-<br />
2.2 mm long, filaments 1.5-2.1 mm long, anthers<br />
0.6-0.7 mm long, 0.4-0.5 mm wide, ovate, light,<br />
yellow; ovaries 5-locular, styles 0.5-0.6 mm long<br />
at anthesis, base <strong>of</strong> ovary nearly glabrous, top part<br />
<strong>of</strong> ovary projected, disk pale yellow turning<br />
magenta. Fruits 3.2-4 mm long <strong>and</strong> wide, globose,<br />
dark purple at maturity, persistent styles divided<br />
from the middle or nearly to the base. Seeds 1.5-<br />
1.8 mm long, 0.9-1 mm wide, 0.4-0.6 mm thick,<br />
kidney-shaped, greenish white.<br />
Common names: Taiwan tu-dang-gui; <strong>and</strong> tudang-gui.<br />
Phenology: flowering from July to September;<br />
fruiting from August to November.<br />
Distribution: endemic to Taiwan (Chiayi,<br />
Hsinchu, Hualien, Ilan, Miaoli, Nantou <strong>and</strong><br />
Taichung) (Fig. 26).<br />
Ecology: in mixed coniferous <strong>and</strong> broadleaf<br />
forests, or in broadleaf forests; (1250) 1700-2900<br />
m.<br />
Additional specimens examined: China.<br />
TAIWAN: Chiayi Co., Alishan, 24 Aug 1990, fl,<br />
C. H. Ou et al. s.n. (TNM). Hsinchu Co., Kuanwu,<br />
20 Jun 1990, fl, C. H. Ou et al. s.n. (TNM). Hualien<br />
Co., Mt. Muhkwa, a shrub near roadside, 24 Jul<br />
1961, M. T. Kao 4153 (A, MSC, TAI); Hualien,<br />
Xiugu, Luanshan, 19 Jul 1973, in fl., F. Y. Lu & C.<br />
H. Ou 1836 (NCUF); Shoufeng District, Mt.<br />
Mukwashan Forest, Halun Station, 2000-2100 m,<br />
chromosome number 2n = 48 from this plant, 23<br />
Sep 1984, young fr, C.-I. Peng 7263 (HAST, 2
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
Fig. 26. Map <strong>of</strong> Taiwan showing the distribution <strong>of</strong> <strong>Aralia</strong> taiwaniana Y. C. Liu & F. Y. Lu.<br />
81
82<br />
sheets). Ilan Hsien, Tatung Hsiang, Szuchi, along<br />
Chiaping Forest Road, 121 o 27’50"E, 24 o 27’01"N,<br />
2300 m, mixed coniferous-broadleaf forest on<br />
mountain slope, 2 m tall, 21 Aug 2000, fl, C.-H.<br />
Lin 617 (HAST, MO); Heping Forest Road, 50K,<br />
2000 m, 3 Aug 1983, sterile, S. Y. Lu 12676 (TAIF).<br />
Miaoli Co., Talu logging tract 28-35K, 2000-2300<br />
m, 26 Jul 1987, late fl & young fr, J. C. Wang & K.<br />
C. Yang 4895 (TAI, 2 sheets); Taian Hsiang, near<br />
36 km marker on 230 Forest Road, 121 o 05’41"E,<br />
24 o 22’08"N, 2400 m, 17 Sep 1995, fr, C. M. Wang<br />
01714 (TNM); Hsishihshan, 24 o 23’00"N,<br />
121 o 06’00"E, 2300 m, 27 Oct 1976, fr, S.-Y. Lu<br />
5971 (TAIF, 2 sheets); Hsishihshan, 24 o 23’N,<br />
121 o 06’E, 2500-2700 m, 27 Oct 1976, fr, B.-P. Yang<br />
74 (TAIF). Nantou Co., Chingching, 24 o 04’0"N,<br />
121 o 10’0"E, 1800-2000 m, 4 Aug 1970, fl, K.-S.<br />
Shyu 635 (TAIF, 2 sheets); Jenai Hsiang, Center<br />
Cross-isl<strong>and</strong> highway, Mayfeng to Tsuifeng,<br />
original broadleaf forest, 2100-2350 m, on wet<br />
place beside paved road, common, plant ca. 2 m<br />
tall, 11 Jul 1995, fl, W.-H. Hu 2821 (HAST, 3<br />
sheets; PE, 3 sheets); May-Feng, 2100 m, 26 Jun<br />
1991, late floral buds, W.-H. Hu 1435 (HAST);<br />
Mayfeng watershed, along dirt road from<br />
Provincial Rd 14-a to Mayfeng Water Source,<br />
broadleaf evergreen forest, 2150-2200 m, 9 Sep<br />
1991, C.-I. Peng 14460 (HAST); Jenai Hsiang,<br />
Sanchia<strong>of</strong>eng, Highl<strong>and</strong>s Experiment Farm,<br />
National Taiwan University, 121 o 11’05"E,<br />
24 o 06’07"N, 2300 m, on mountain slope, waste<br />
farml<strong>and</strong>, common, 13 Jul 2004, fl, Y.-N. Ko et al.<br />
1382 (HAST); Mashan, along Central Cross-Isl<strong>and</strong><br />
Hwy, near road mileage sign 19 km, 121 o 11’30"E,<br />
24 o 6’43"N, 2315 m, mixed coniferous-broadleaf<br />
forest, roadside, semiexposed habitat, occasional,<br />
30 Nov 2005, late fl, C.-I. Huang 2343 (HAST, 2<br />
sheets); Jenai Hsiang, on the way from Tianchi<br />
Preserve Station to Tianchi, trailside, under forest,<br />
forest margin, 4 Sep 1998, fl, T. Y. A. Yang 11188<br />
(HAST, PE); Jenai Hsiang, Kunyang, 2400-2450<br />
m, 8 Jul 1991, fl, C.-H. Chen 1033 (PE, TAIF);<br />
Jenai Hsiang, en route from Yunhai to Tienchih,<br />
24 o 03’N, 121 o 16’E, broadleaf forest phasing to<br />
mixed broadleaf <strong>and</strong> coniferous forest, 2360-2860<br />
m, herb ca. 60 cm tall, fruits purplish black, 13<br />
Oct 1992, H.-L. Ho 561 (MO); Yunhai to Tienchih,<br />
2850 m, trailside, forest margin, 6 Sep 2001, fr, T.<br />
Y. A. Yang et al. 13875 (TNM); Jenai Hsiang,<br />
Juiyenchi Forests Nature Protected Area,<br />
121 o 11’59"E, 24 o 07’01’N, 2250 m, mixed<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
Cryptomeria <strong>and</strong> broadleaf forest, ca. 2 m tall, 8<br />
Sep 2000, young fr, Y.-Y. Huang 99 (MO); Tsuifeng<br />
– Meifeng, 2100-2350 m, 16 Aug 1984, fl & young<br />
fr, Y. Tateishi et al. 17783 (MO); Tsuifeng –<br />
Meifeng, roadside, 16 Aug 1984, W. S. Wang 652<br />
(TAI); Tsuifeng, 8 Dec 1974, fr, C. H. Ou 2909<br />
(TAI); Mayfong, 13 Jun 1980, floral buds, M. T.<br />
Kao 9477 (TAI, 2 sheets); Tsue-fong to Mei-fong,<br />
roadside, 2200-2000 m, 16 Aug 1984, fl, T. Y. Yang<br />
89 (TAI); Meifeng – Chingching Farm, roadside,<br />
1700 – 2100 m, 25 Jul 1987, floral buds, S. F.<br />
Huang & S. Y. Yang 3765 (TAI); Nantou Hsien,<br />
Jenai Hsiang, at km 23 on Hwy 14A, disturbed<br />
vegetation along roadside on steep slope,<br />
24 o 06’32"N, 121 o 13’38"E, 2680 m, ca. 2 m tall,<br />
23 Sep 1997, fr, P. P. Lowry II et al. 4969 (HAST,<br />
MO, 4 sheets); Juiyenhsi, 24 o 08’0"N, 121 o 12’0"E,<br />
1800 m, 29 Jun 1999, floral buds, Y.-P. Cheng 2664<br />
(TAIF, 4 sheets); Meifeng, 24 o 06’0"N,<br />
121 o 10’56"E, M.-J. Lin 285 (TAIF); Meifeng, 16<br />
Jul 1996, M.-J. Lin 154 (TAIF); Mei-feng, roadside,<br />
broadleaf forest, 2000-2100 m, 30 Jun 1996, floral<br />
buds, J. C. Wang et al. 10093 (HAST); Tsuifeng,<br />
24 o 06’N, 121 o 11’, 1700 m, 4 Sep 1973, fr, K.-Y.<br />
Wang s.n. (TAIF, 2 sheets); Tsuifeng, 121 o 11’52"E,<br />
24 o 06’30"N, 2200 m, roadside, 19 Aug 2005,<br />
young fr, C. M. Wang 08185 (TNM); Nantou Co.,<br />
on the way from Chu-Feng to Hong-Xiang, just<br />
below Chui-Feng, at Rueiyan River area,<br />
N24 o 06’32", E121 o 11’21", 2265 m, 5 Nov 2006,<br />
fl, J. Wen 9424 (US). Pingtung Co., Gangkou, 19<br />
Jul 1919, Matuda-Eizi s.n. (TAIF, 2 sheets, 18416<br />
&18418) [maybe cultivated as udo]; Kosyun,<br />
Kankau, 19 Jul 1919, fl, S. Sasaki s.n. (TAI,<br />
082497) [maybe cultivated as udo]. Taichung Co.,<br />
Tahsuehshan Forest Road, 24 o 20’N, 121 o 07’E,<br />
1960 m, 19 Jul 1984, young fr, S.-Y. Lu 14919<br />
(HAST, TAIF, 2 sheets); Hoping Hsiang,<br />
Tashueishan forest track, 1700 m, roadside, 10 Oct<br />
1995, post fr, T. Y. A. Yang & Y. B. Cheng 06356<br />
(HAST, TNM). Kwarenko-Tyo, between Be<br />
Derikku <strong>and</strong> Be Padahu, 2900 m, monte<br />
Tyuosenzan, 30 Jul 1936, fl, N. Fukuyama &<br />
Suzuki-Tokio ST 15202 (PE, TAI).<br />
<strong>Aralia</strong> taiwaniana usually occurs from 1250-<br />
2900 m in altitudes in Taiwan. It is interesting to<br />
note that a few specimens from Pingtung <strong>of</strong><br />
southern Taiwan at very low altitudes are<br />
morphologically abnormal with smaller leaflets,<br />
<strong>and</strong> shorter <strong>and</strong> fewer pedicels. The Pingtung
collections (from Gangkou) were most likely from<br />
cultivated material. The Government <strong>of</strong> Formosa<br />
Nursury formerly stood on this site <strong>and</strong> the<br />
Japanese like udo as vegetables. Gangkou is near<br />
the seashore <strong>and</strong> appears to low in altitude for<br />
natural populations <strong>of</strong> A. taiwaniana.<br />
<strong>Aralia</strong> taiwaniana differs from A. cordata in<br />
that A. taiwaniana has pubescent (vs. hirsute lower<br />
stem surface) <strong>and</strong> does not have leafy stipules (cf.<br />
color plates 4C <strong>and</strong> 5D). Compared to A. cordata,<br />
A. taiwaniana tends to have leafy <strong>and</strong> smaller<br />
inflorescence.<br />
In comparison with <strong>Aralia</strong> continentalis, A.<br />
taiwaniana has finer leaflet margin <strong>and</strong> its pedicels<br />
are statistically longer. It is interesting to note that<br />
the few specimens from Pingtung <strong>of</strong> southern<br />
Taiwan at very low altitudes are morphological<br />
abnormal with smaller leaflets, <strong>and</strong> shorter <strong>and</strong><br />
fewer pedicels. The Pingtung collections (from<br />
Gangkou) were most likely from cultivated<br />
material. The Government <strong>of</strong> Formosa Nursury<br />
was there <strong>and</strong> the Japanese folks like udo as<br />
vegetable. Gangkou appears to be too low in<br />
altitude for <strong>Aralia</strong> taiwaniana, as it is near the<br />
seashore.<br />
<strong>Aralia</strong> taiwaniana resembles A. schmidtii that<br />
both taxa have more than 35 flowers in each<br />
terminal umbel. Yet they can be differentiated as<br />
follows:<br />
1. Leaflet margin finely serrulate, base <strong>of</strong> leaflets<br />
cordate, stipules lanceolate; endemic to the<br />
Sakhalin & Kuril isl<strong>and</strong>s <strong>of</strong> far eastern Russia<br />
........................................................ A. schmidtii<br />
1. Leaflet margin doubly serrate, base <strong>of</strong> leaflets<br />
obliquely truncate to subcordate, rarely cordate,<br />
stipules narrowly triangular; endemic to Taiwan<br />
..................................................... A. taiwaniana<br />
7. <strong>Aralia</strong> continentalis Kitag. — Fig. 27; color<br />
plate 6: A-E.<br />
<strong>Aralia</strong> continentalis Kitag., Bot. Mag. (Tokyo) 49:<br />
228, t. 3. 1935. <strong>Aralia</strong> cordata Thunb. var.<br />
continentalis (Kitag.) Y. C. Zhu in Z. Y. Chang<br />
(ed.), Pl. Medic. Chinae Bor.-Orient.: 787.<br />
1989. TYPE: China. LIAONING: southern<br />
Manchuria, Fengtien Prov. [Liaoning Prov.],<br />
Jalu [Yalu] River valley, near Schildagou, 13<br />
Aug 1897, fl, V. Komarov 1152 (lectotype: TI!,<br />
here designated; isolectotype: BM!, LE!).<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
83<br />
Perennial spreading herb 1-3.3 m tall. Stem<br />
pubescent, lower part sometimes with stiff hairs.<br />
Rhizomes thick. Leaves 50-100 cm long <strong>and</strong> wide,<br />
ternately compound, lower leaves tripinnately<br />
ternate, upper leaves pinnate to trifoliolate or<br />
tripinnately ternate; stipules 5-20 mm long, 3.5-<br />
15 mm wide, pilose; leaflets 5-16.5 cm long, 3-9.5<br />
cm wide, papery, the basal leaflets <strong>of</strong> the pinnae<br />
ovate, other leaflets ovate to narrowly so,<br />
acuminate to acute at apex, cordate to subcordate<br />
or sometimes rounded at base, <strong>of</strong>ten oblique on<br />
lateral leaflets, serrate to doubly so at margin;<br />
lateral veins 7-9 on each side, conspicuous on both<br />
surfaces, scabrous on adaxial surface, <strong>of</strong>ten with<br />
scattered hairs on veins <strong>and</strong> veinlets, pubescent<br />
with scabrous hairs on abaxial surface, petiolules<br />
<strong>of</strong> lateral leaflets 0.1-2.2 cm long, more or less<br />
pilose. Inflorescence a terminal or axillary panicle,<br />
pilose, terminal inflorescence 30-65 cm long,<br />
consisting <strong>of</strong> 20-35 primary branches, panicunately<br />
or racemosely arranged on a main axis,<br />
occasionally 3-5 forming a circle on the upper part<br />
<strong>of</strong> the main inflorescence axis, each primary branch<br />
5-15 cm long, consisting <strong>of</strong> 2-18 umbels, <strong>of</strong>ten the<br />
primary branches further divided into a panicle;<br />
terminal umbels mostly 16-35 (-50)-flowered,<br />
pedicels 6-10 mm long, pubescent, tip <strong>of</strong> pedicels<br />
enlarged <strong>and</strong> with a cluster <strong>of</strong> hairs; lateral umbels<br />
8-13-flowered, pedicels 4-7 mm long; bracts <strong>of</strong><br />
primary branches 5-9 mm long, 0.6-0.8 mm wide,<br />
linear to lanceolate, somewhat pilose; bracteoles<br />
1.2-2 mm long, 0.2-0.3 mm wide, lanceolate,<br />
pilose. Sepals 0.4-0.5 mm long, 0.6-0.7 mm wide,<br />
triangular to broadly so; petals 1.8-2 mm long, 0.9-<br />
1.1 mm wide, ovate, with a conspicuous vascular<br />
bundle in the middle, tip slightly enlarged; stamens<br />
5, 2.5-2.7 mm long, filaments 2-2.4 mm long,<br />
anthers 0.6-0.8 mm long, 0.4-0.5 mm wide, ovate;<br />
ovaries 5-locular, styles 0.6-0.7 mm long at<br />
anthesis, base <strong>of</strong> ovary nearly glabrous, the top part<br />
<strong>of</strong> ovary slightly projected. Fruits 3.5-4 mm long<br />
<strong>and</strong> wide, globose, persistent styles divided to the<br />
middle. Seeds 2-3 mm long, 1.2-1.4 mm wide,<br />
0.8-1 mm thick, kidney-shaped.<br />
Common names: tu dang gui <strong>and</strong> duhe (China),<br />
<strong>and</strong> Dok-whal (Korea).<br />
Uses: leaves as vegetables; roots as bloodregulating<br />
medicine.<br />
Phenology: flowering in July <strong>and</strong> August;<br />
fruiting in September to October.
84<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
Fig. 27. <strong>Aralia</strong> continentalis Kitag. A. Habit showing leaves <strong>and</strong> inflorescences. B. Part <strong>of</strong> a lower leaf showing the terminal<br />
pinna. C. Axillary inflorescence. D. Flower after anthesis. E. Young fruit. F. Umbel with fruits. G. Floral buds. H. Flower.<br />
I. Leaf-like stipule (A, C & G – Ye 2002, MO; B, D & H – Biao 1104, MO; E & F – Wang 60812, A; I – Ching 2957, A).
Distribution: eastern Russia, eastern, central,<br />
south-central, <strong>and</strong> northern part <strong>of</strong> China (to<br />
northern Guangxi province in the southern part <strong>of</strong><br />
China), <strong>and</strong> Korea (Fig. 25).<br />
Additional specimens examined: China.<br />
ANHUI: Huangshan, 28 Jul 1926, S. S. Chien 1172<br />
(PE, W); Whang Shan, S. Anhwei, 3000 ft, shrub<br />
10 ft, flowers greenish, terminal, 12 Jul 1925, fl,<br />
R. C. Ching 2957 (A, K); Shizhilin, near the Guest<br />
House, 30 Jul 1957, L. K. Fu 0776 (WUK);<br />
Huangshan, near the Hot Spring, 14 Jul 1975, floral<br />
buds, K. J. Guan 75598 (PE); Hwangshan,<br />
Yunkusze to Shihszelin, 12 Aug 1935, T. N. Liou<br />
& P. C. Tsoong 2616 (KUN, PE, WUK);<br />
Huangshan, below Beihai, 1700 m, 14 Aug 1973,<br />
fl, G. S. Zhou et al. 469 (PE, 2 sheets); Huangshan,<br />
near Guang-ming-ding, common between 1400-<br />
1700 m, in Picea-Carpinus-Pinus-Acer forest, 17<br />
Jul 1999, J. Wen 4885 (US). FUJIAN: Chongan<br />
Xian, 700 m, 13 Aug 1979, fl, Wuyi Expedition<br />
Team 00210 (PE); Tongmuguan, 800 m, 16 Aug<br />
1979, fl, Wuyi Expedition Team 00482 (PE).<br />
Wuyishan, on the way from Shangang to Gaoqiao,<br />
roadside in sparse forest, 3 m tall, common, 19 Jul<br />
1980, floral buds, Wuyishan Team 80-0118 (MO,<br />
WUK). GANSU: Wen Xian, Baishujiang Nature<br />
Preserve, Qiu Jia Ba, Second Work Section, 2400-<br />
2700 m, at the edge <strong>of</strong> forest, in forest, fruits<br />
reddish, 5 Sep 2007, fr, Baishuijiang Expedition<br />
Team 5728 (PE). GUANGXI: Damiaoshan Xian,<br />
Shanfangqu, Pingshixiang, Jiuwangdashan,<br />
Yuantoutun, Yangmeijie, 700-1000 m, valley,<br />
sparse forest, ca. 0.6 m, 21 Jul 1958, S. Q. Chen<br />
15792 (KUN, PE, WUK); Yangmeijie, 1300 m, 2<br />
Jul 1957, roadside woods, 1.8 m tall, 2 Jul 1957,<br />
floral buds, D. C. Chen 910 (KUN);<br />
Jiuwangdashan, Jiuyang, common, 1280 m, 30 Aug<br />
1958, 0.5 m, S. Q. Chen 15429 (KUN, WUK).<br />
Longsheng, Dadi Xiang, Hongyashan, 1660 m, 29<br />
Jul 1955, fl, Guang-Fu Forestry District Team<br />
00928 (PE); 1630 m, 19 Oct 1955, fr, Guang-Fu<br />
Forestry District Team 01076 (PE). Xingan, Miaoer-shan,<br />
Longtangjiang, 1000 m, 26 Jul 1997, fl,<br />
G. Z. Li 15425 (PE); Mao-Er-Shan, N25 o 54.492’,<br />
E110 o 27.833’, 1526 m, 7 May 2010, J. Wen 11550<br />
(US). Zhiyuan Xian, Wupa, 8 Jul 1959, D.-A.<br />
Huang 61322 (MO). Guangxi, R. C. Ching 7151<br />
(A, NY, PE); T. S. Tsoong 83497 (A). HEBEI:<br />
Dongling, Jul 1935, fl, Y. Liu 11808 (PE); Aug<br />
1935, fl, Y. Liu 11836 (PE); Aug 1930, H. T. Tsai<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
85<br />
50227 (A, PE); Shangpanpo, Tungling, 25 Aug<br />
1930, fr, H. F. Chow 40973 (PE); Liulipingshan,<br />
Tung-ling, Yu 222 (PE). Fuping-hsien, 1200 m, 5<br />
Sep 1934, fr, K. M. Liou 3607 (PE, WUK). Tanghsien,<br />
29 Jul 1934, young fr, K. M. Liou 3112 (PE,<br />
WUK). Tienchiaoshan, Chahar, stream side, 16<br />
Aug 1933, young fr, C. W. Wang 60812 (PE).<br />
Northern Hebei, 10 Sep 1951, fr, W. C. Wang 2473<br />
(PE). HENAN: Lanchuan Xian, 1959, Pucha Team<br />
20325 (PE). Lushi Xian, Shizhiping,<br />
Nanlianpanshan, in sparse forest, valley, shady<br />
area, 1350 m, 22 Aug 1973, fr, K. J. Fu 17097<br />
(WUK); Lushih-hsien, Laochunshan, 1140 m, 12<br />
Aug 1935, young fr, K. M. Liou 5019 (PE, WUK).<br />
HUBEI: Enshi, Taishanmiao Tree Farm, 31 Aug<br />
1957, young fr, G. X. Fu & K. S. Zhang 1397 (PE,<br />
3 sheets, WUK). Hefeng Xian, Huping Farm,<br />
1350-1450 m, 21 Aug 1958, fl, H. J. Li 5744 (MO,<br />
PE, WUK). Lichuan Xian, Xiao He, Honghexi,<br />
Zhaojiayan, Oct 1956, fr, X. B. Wan 283 (WUK);<br />
Lichuan Shi, Moudao Township, Dazhuang<br />
Village, Jingzhulin (Gold Bamboo Forest),<br />
N30 o 29’06", E108 o 43’22", 1350 m, 1 Sep 2001, at<br />
the edge <strong>of</strong> bamboo woods, J. Wen 8138 (US).<br />
Tongshan Xian, Jiugongshan, 18 Oct 1996, fr, C.<br />
S. Ye 3310 (MO, TNM). Xuanen Xian, Gongshan,<br />
in front <strong>of</strong> the Medicine Factory, in shrubby area,<br />
1.5 m tall, 7 Jul 1958, buds, H. J. Li 3577 (PE,<br />
WUK); Dagongshan, 1300-1400 m, 20 Jul 1958,<br />
floral buds, H. J. Li 5264 (PE, WUK). HUNAN:<br />
Jiangyong Xian, Datian, 750 m, P.-X. Tan 63652<br />
(MO). Shangzhi Xian, Badagongshan, 1989, Plant<br />
Chemistry Voucher 683 (PE, 2 sheets). Xingning<br />
Xian, Wanfengshan Tree Farm, Huashijiang, 1200<br />
m, 28 Sep 1996, fr, Z. C. Luo 1661 (PE, TAIF, 2<br />
sheets); on top <strong>of</strong> Dayunshan, 1700 m, 29 Jul 1985,<br />
floral buds, Y. B. Luo 2769 (PE); Shunyao,<br />
Tongzhichong, 29 Jun 1985, floral buds, Y. B. Luo<br />
2487 (PE); Shunhuangshan Tree Farm, Lizhu<br />
Village, Luodanjiang Kou, N26 o 27.505’,<br />
E110 o 58.955’, 682 m, occurring locally in upper<br />
areas, planted near homestead, 5 Aug 2006, fl, J.<br />
Wen 9300 (US); Xinning Xian, Wanfeng Tree<br />
Farm, Ga<strong>of</strong>eng Rd., N26 o 30.252’, E110 o 37.480’,<br />
1316 m, 7 Aug 2006, fl, herb 70-120 cm tall, fls<br />
greenish white, J. Wen 9330 (US). JIANGXI:<br />
Jingan Xian, Beigang Tree Farm, Shizhiyan, 1000<br />
m, near stream in forest, 29 Oct 1982, S. S. Lai et<br />
al. 01450 (PE). Jinggangshan, Liangzhushan,<br />
Goupazhai, 1100 m, near stream, 2 Oct 1958, fr,<br />
fruits purplish black, J. Xiong 2873 (PE, 2 sheets).
86<br />
Qianshan Xian, Wuyishan, Tongmuguan, 1400 m,<br />
22 Aug 1958, fl, S. S. Lai & M. X. Nie 4419 (PE,<br />
WUK); Qianshan Xian, Wuyishan Nature Preserve,<br />
on the way from the Preserve Headquarter to<br />
Zhumukeng, ca. 1500 m, 9 Jul 2001, fl, J. Wen<br />
5545 (US). Shangyou Xian, Wuguaifeng<br />
Commune, Qiyunshan, the First Peak, 20 Aug<br />
1971, fl, Jiangxi Team 537 (PE). Shuichuan Xian,<br />
Dafeng Qu, Linyang, Xiaoshui, in forest, moist<br />
area, 1200 m, 24 Sep 1963, fr, fruits purplish black,<br />
J. S. Yue et al. 4253 (KUN, WUK). Tongmuguan,<br />
800 m, 11 Aug 1958, fl, P. X. Qiu 1873 (PE, 3<br />
sheets). Wuling Xian, Luoxi, 1500 m, in the valley<br />
near stream, herb ca. 2 m tall, 28 Jul 2000, fl &<br />
young fr, J.-H. Zhang 1056 (TAIF). Xiushui Xian,<br />
Tangpai Xiang, roadside, 1100 m, 0.5-1 m tall, 5<br />
Aug 1996, fl, C. S. Ye 2002 (TNM). JILIN: Antu<br />
Xian, riverside vegetation <strong>of</strong> left bank <strong>of</strong> Er Dao<br />
Bai He, 930 m, river alluvions, 1 Aug 1986, D.-Y.<br />
Hong et al. 33.318 (RSA); primary but partly<br />
degraded broadleaf forests N <strong>of</strong> Er Dao Bai He,<br />
800 m, moderately moist to drier road side with<br />
ditches, dark brown forest soils <strong>and</strong> clay, 25 Aug<br />
1986 (fr), D.-Y. Hong et al. 34.433 (RSA); on the<br />
way from Naitoushan to Laolangbao, edge <strong>of</strong><br />
forest, K. M. Liou 3850 (PE); Antu Xian, Erdao<br />
Township, Hanchun Gou, just west <strong>of</strong> Erdao, 5 Aug<br />
1997, J. Wen 3129A (US). Fusong Xian, along the<br />
road from Changbai to Songjianghe, in forest SE<br />
<strong>of</strong> the 50 km marker, 18 Jun 2001, J. Wen 5426<br />
(US). Helong Xian, Shajinggou, in mixed forest,<br />
18 Sep 1959, Yanbian Group II 962 (PE). Journey<br />
from Mukden to Kirin, Tang-ho-ko, Sungari River<br />
to Hui Fa River, May to Aug 1886, fl, H. E. M.<br />
James s.n. (K); Changbaishan, 800 m, under<br />
woods, 1981, fl, B. B. Wan & K. S. Chow 81106<br />
(BM, CAS, K, MO, NY); Changbaishan, north<br />
slope, 1200 m, at the edge <strong>of</strong> mixed forest, 24 Aug<br />
1959, Yanbian Group I 416 (PE, 2 sheets);<br />
Changbaishan, 800 m, in forest, 18 Aug 1981,<br />
young fr, G. S. Zhou et al. 106 (PE). LIAONING:<br />
Benxi Xian, Chaohezhang Commune, 13 Aug<br />
1965, fl, C. Q. Lin 961 (PE). Fengcheng Xian,<br />
Dongxingbaozhi, in forest on slope, C. Q. Lin 1048<br />
(PE). Huairen Xian, Huanggou, in valley near a<br />
stream, 14 Aug 1959, C. S. Wang et al. 3858 (PE),<br />
3859 (WUK). SHAANXI: Fuping Xian, riverside<br />
in the valley, 1300 m, herb 1.5 m, 28 Jul 1989, fl,<br />
J.-S. Ying et al. 0135 (MO). Yang Xian, Huayang,<br />
Jiuchiba, 1400 m, J. X. Yang 01603 (PE, WUK, 2<br />
sheets). SHANXI: Hengqu Xian, Tongshan<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
Commune, southern slope, 1100 m, 13 Jul 1984,<br />
floral buds, Shanxi Team 02243 (Ny, WUK, 2<br />
sheets). SICHUAN: Baoxing Xian, as Pao-hsinghsien<br />
on this specimen, 2400 m, 1 Sep 1936, fl, K.<br />
L. Chu 3780 (BM, PE); as Pao-hsin-hsien, formerly<br />
Mupin, 1954, T.-P. Soong 38937 (KUN, WUK),<br />
39067 (KUN); Shizhishan, slope, grassy area, 2700<br />
m, 16 Aug 1933, fl, T. T. Yü 2368 (PE, 2 sheets).<br />
Emeishan, Biology Department <strong>of</strong> Sichuan<br />
University Expedition Team 51545 (WUK).<br />
Hanyuan Xian, Xianglingshan, 2800 m, 9 Sep<br />
1938, fr, T. P. Wang 9674 (WUK). Meigu Xian,<br />
Liangshan, Houpu, Letuo Qu, 1-1.5 m, 18 Aug<br />
1959, Sichuan Economic Botany Liangshan Team<br />
1674 (KUN). Mianning Xian, Tuowu Qu, Yele,<br />
Shanchahe, in dense forest, 2600 m, 3 m, 20 Jul<br />
1959, S. G. Wu 2061 (KUN, PE, 2 sheets). Shimian,<br />
X. X. Kong 41889 (WUK); Shimian, X. X. Kong<br />
42009 (WUK). YUNNAN: Yiliang Xian:<br />
Chaotianma Tree Farm, below Doukouzhi,<br />
limestone slope, rare, 1 m tall, Northeast Yunnan<br />
Team 677 (KUN, 2 sheets). Manchuria austroorientalis,<br />
1860, Maximowicz s.n. (K, 2 sheets).<br />
Korea. Korea septentrionalis, 6 Jul 1897, fl, V.<br />
Komarov 1152 (K); Hallaisan, Oct 1907, fl &<br />
young fr, T. Taquet 138 (BM); Hallaisan, 1300 m,<br />
13 Aug 1908, fl, Taquet 891 (K); Kang Kai, 25 Jul<br />
1911, fl, R. C. Mills s.n. (PE, 2 sheets); Kang Kai,<br />
3 Aug 1911, young fr, R. C. Mills s.n. (PE). Russia.<br />
Amur, Primorskaya region, Askold Isl<strong>and</strong>, N. A.<br />
Palczewsky s.n. (A, LE); Primorye Territory,<br />
Ussuriisk Town, basin Borisovka river, slope, 16<br />
Aug 1973 (fl), P. Gorovoy s.n. (RSA). Far East,<br />
Telyakovsky Inlet, Gamov Peninsula, N42 o 34’,<br />
E131 o 12’, 290 m, herbaceous perennial to 1.3 x<br />
1.3 m, occasional, growing in dense shade with<br />
Acer pseudosieboldianum <strong>and</strong> Diervilla praecox<br />
in Quercus mongolica, Betula schmidtii, Betula<br />
davurica, Kalopanax pictus, Tilia amurensis<br />
woodl<strong>and</strong>, on gentle NW facing slope in rich loamy<br />
soil with high organic matter content, 20 Sep 1994,<br />
fr, M. Flanagan & A. Kirkham ESUS 130 (K, 2<br />
sheets); Russian Far East, Peninsula Galiv, dense<br />
oak forest, 20 Aug 1931, V. Petrov s.n. (MW, 2<br />
sheets); southern Ussuri region, Pos’etckiy, near<br />
Harbor Seedema, in oak decidous forest, 18 Sep<br />
1933, A. Poretzky 279 (LE); in the mountain <strong>of</strong><br />
Moetzita, Pos’etckiy region, 11 Aug 1926, A.<br />
Savernin 107 (LE); Primorskaya region,<br />
Vladivostoksky District, 4 Aug 1984, fl, P. U.<br />
Zhmylev s.n. (MW, 3 sheets).
Specimens from cultivated source: China.<br />
JIANGXI: Lushan Botanical Garden, 4 Sep 1988,<br />
fr, J. Wen 610 (OS).<br />
Kitagawa (1935) cited five specimens<br />
(syntypes) in the original description <strong>of</strong> <strong>Aralia</strong><br />
continentalis. V. Komarov 1152 is representative<br />
<strong>of</strong> the morphology <strong>of</strong> the Manchurian “A.<br />
continentalis” <strong>and</strong> was distributed to different<br />
herbaria (BM, LE, <strong>and</strong> TI). It is thus selected as<br />
the lectotype for <strong>Aralia</strong> continentalis. It is noted<br />
that Komarov 1152 consists <strong>of</strong> mixed collections<br />
which were made in the summer <strong>of</strong> 1897. The<br />
lectotype was collected on August 13, 1897. Two<br />
other specimens <strong>of</strong> Komarov 1152 were seen, but<br />
were not considered as isolectotypes: (Korea, 6<br />
July 1897, V. Komarov 1152, K; <strong>and</strong> China, Jilin,<br />
Mu-dan-dsian, 10 Jul 1897, V. Komarov 1152, A,<br />
W).<br />
8. <strong>Aralia</strong> cachemirica Decne. — Fig. 28.<br />
<strong>Aralia</strong> cachemirica Decne. in Jacquem. Voy. Inde<br />
4: 72. 1844. TYPE: India. Kashmir, Mt.<br />
Vestervonne, 3087 m, 12 Jun 1831, V.<br />
Jacquemont 718 (lectotype, P!, here<br />
designated).<br />
<strong>Aralia</strong> macrophylla Lindl., Edward’s Bot. Reg. 30:<br />
73. 1844. TYPE: India. North part (type not<br />
seen).<br />
Robust, spreading, <strong>and</strong> unarmed perennial herb<br />
1.5-2.3 m tall; stem green to greenish purple.<br />
Leaves 1.7-2.3 cm long, 0.8-1.2 cm wide, lower<br />
leaves ternately compound, upper leaves pinnate<br />
to bipinnate; stipules narrowly triangular,<br />
membranaceous, ciliate at margin, pilose;<br />
uppermost petioles 4-19 cm long, greenish,<br />
puberulent; leaflets (4)-8-21 cm long, (1.5) 3-8.5<br />
(11) cm wide, thick papery, the pair <strong>of</strong> leaflets<br />
below the terminal leaflet narrowly ovate, other<br />
leaflets ovate to narrowly so, rarely elliptic, acute<br />
to acuminate at apex, rounded to subcordate, rarely<br />
cordate at base, commonly oblique on lateral<br />
leaflets, serrulate to serrate, sometimes doubly so<br />
at margin; lateral veins (8)-10-12-(19) on each<br />
side, conspicuous on both surfaces, nearly<br />
glabrescent to slightly scabrous on adaxial surface,<br />
<strong>of</strong>ten with scattered hairs on veins <strong>and</strong> veinlets,<br />
glabrescent to slightly puberulent on abaxial<br />
surface, petiolules 0.2-2.2 cm long, glabrescent to<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
87<br />
slightly pilose. Inflorescence 15-45 cm long,<br />
terminal <strong>and</strong> axillary, paniculate in overall<br />
structure, puberulent, consisting <strong>of</strong> numerous<br />
umbels, primary inflorescence branches 15-30,<br />
mostly racemosely or sometimes panicunately<br />
arranged, <strong>of</strong>ten 3-8 forming a circle on the main<br />
inflorescence axis, each primary branch 3-10 cm<br />
long, consisting <strong>of</strong> 1-4 umbels, usually only the<br />
terminal umbel <strong>of</strong> each branch bearing fruits, others<br />
becoming aborted; terminal umbels mostly 35-70-<br />
(95)-flowered, pedicels 7-11 (-16) mm long,<br />
pubescent; lateral umbels 15-25-flowered, pedicels<br />
5-8 mm long; bracts <strong>of</strong> primary branches 3.5-8 mm<br />
long, 0.5-1 mm wide, linear, somewhat pilose,<br />
sometimes leafy at the lower part <strong>of</strong> the<br />
inflorescence; bracteoles 1.2-2.2 mm long, linear,<br />
more or less ciliate at margin. Sepals 0.3-0.4 mm<br />
long, broadly to narrowly triangular, persistent on<br />
fruits; petals 1.7-2.2 mm long, ca. 1 mm wide,<br />
ovate, greenish white to white, with a conspicuous<br />
vascular bundle in the middle; stamens 5, 1.8-2<br />
mm long; ovaries 5-locular, styles ca. 0.5 mm long<br />
at anthesis, base <strong>of</strong> styles conspicuously enlarged,<br />
appearing like a stylopodium. Fruits 3-4 mm long,<br />
3.8-4 mm wide, globose, persistent styles ca. 1 mm<br />
long, divided to the middle. Seeds 2-2.2 mm long,<br />
1 mm wide, 0.2-0.25 mm thick, kidney-shaped,<br />
whitish gray, smooth.<br />
Phenology: flowering in late June to<br />
September; fruiting in September <strong>and</strong> October.<br />
Distribution: Northwestern India, northern<br />
Pakistan, <strong>and</strong> western Nepal, most <strong>of</strong>ten collected<br />
from the Kashmir region (Fig. 29).<br />
Ecology: river valleys, streamsides, <strong>and</strong> mixed<br />
forests; 2100-3650 m.<br />
Additional specimens examined. India.<br />
Humid woods above Sendjigam, 2600 m,<br />
V.Jacquemont 913 (P, 2 sheets); V. Jacquemont s.n.<br />
(P). 8-9000 ft, 1884, J. F. Duthie 595 (LE); NW<br />
India, J. F. Royle s.n. (K, LE, 2 sheets); NW India,<br />
1871, J. L. Stewart s.n. (LE, 2 sheets).<br />
HIMACHAL PRADESH: Chamba, NW India,<br />
Aug 1880, young fr, R. Ellis 428 (K); Panji,<br />
Chamba State, Sanch Valley, 9000 ft, 10 Aug 1899,<br />
fl, J. F. Duthie 19618 (K); NW Himalayas, Chamba<br />
State, Dharwas, 9,000 ft, 28 Jun 1896, young fr &<br />
fl, J. H. Lace 1476 (E); Simla, 7-8000 ft, 28 Aug<br />
1849, T. Thomson s.n. (K); Simla, Matiyana, 9000<br />
ft, Sep 1878, in fl, H. Collett 6568 (K, 2 sheets);
88<br />
Theog, 8000 ft, 1 Sep 1886, fl, H. Collett 584 (K,<br />
2 sheets). JAMMU & KASHMIR: V. Jacquemont<br />
717 (P); Astor Valley near Dashkin, 26 Jul 1892,<br />
J. F. Duthie 12254 (BM); Erin Valley, near<br />
B<strong>and</strong>apur, 8,000 ft, 25 Jul 1940, on edge <strong>of</strong><br />
cultivation, F. Ludlow & G. Sherriff 7831 (E);<br />
Kulewan, 7750 ft, 22 Jul 1876, C. B. Clarke 29470<br />
(LE); Sonamurg, 8500 ft, 1 Sep 1876, fl, C. B.<br />
Clarke 30906 (BM, K, 2 sheets, LE, WU); 1 Sep<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
Fig. 28. <strong>Aralia</strong> cachemirica Decne. A. Habit showing leaves <strong>and</strong> inflorescences. B. Leaf base showing stipule morphology. C.<br />
Close-up showing leaflet margin. D. Flower. E. Young fruit.<br />
1876, C. B. Clarke 30918 (K); Kashmir, 7-9500<br />
ft, 28 Sep 1848, fl., T. Thomson s.n. (K, LE, U,<br />
W); Gulmarg, Kashmir, 8500 ft., 11 Aug 1919, H.<br />
H. Rich 1239 (K, 2 sheets); Rajparyan Sanctuary,<br />
10,500 ft, 24 Aug 1943, floral buds, F. Ludlow &<br />
G. Sherriff 9366 (BM, E); below Zaiwan, 9,500 ft,<br />
30 Aug 1956, O. Polunin 56/606 (BM, 2 sheets,<br />
E); Nara Nag, Wanqar Valley, among debris <strong>of</strong> Nara<br />
Nag Ruins, 12 Aug 1840, fl, P. M. Pinfold 252
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
Fig. 29. Map <strong>of</strong> India, Pakistan <strong>and</strong> neighboring regions showing the distribution <strong>of</strong> <strong>Aralia</strong> cachemirica Decne.<br />
89
90<br />
(BM); Prov. Kishtvar, Kishtvar to the Pir Patsaski<br />
or Kishtvar Pass, 1-4 Aug 1856, Schlagintweit 3796<br />
(BM, K); Dunga Fali, Aug 1880, A. P. Young s.n.<br />
(BM, 2 sheets). PUNJAB: J. R. Drummond 24236<br />
(K). SIKKIM. 10-12000 ft, J. D. Hooker s.n. (LE).<br />
UTTARANCHAL: Garhwal, 8000 ft, T. Thomson<br />
s.n. (K, LE); NW India, Tihri - Garhwal, Ganges<br />
Valley, 9-10000 ft, Oct 1881, fr, J. F. Duthie s.n.<br />
(BM, WU). UTTAR PRADESH: Deota, 8000 ft,<br />
17 Jun 1906, H. H. Haines 2245 (K, 2 sheets);<br />
Jaunsar, Hajawa, 8000 ft, Oct 1890, fr, J. S. Gamble<br />
25231 (K); Jaunsar, Mendali, 7500 ft, Oct 1890, J.<br />
S. Gamble 25054 (K); Mendali, 8000 ft, Sep 1898,<br />
fr, J. S. Gamble 27311 (K); NW Himalayas,<br />
Jaunsar, Konain, 19 Sep 1936, young fr, M. B.<br />
Raizada 7263 (E). Nepal. Garjigoth, 9500 ft, 9<br />
Aug 1952, O. Polunin et al. 5045 (BM, 2 sheets,<br />
E). Pakistan. 7000 ft., Gilgit Expedition, Giles<br />
544 (K). Hazara Distr., Kaghan, 7500 ft, 3 Oct<br />
1966, M. A. Giddiqi 4052 (W); Hazara, between<br />
Naran <strong>and</strong> Kaghan, 6 Sep 1988, fl, S. Omer & M.<br />
Qaiser 2748 (K); NW “India”, near Dana, Bulsun<br />
[now in Pakistan], 8000 ft, Sep 1877, H. Collett<br />
5542A (K). PUNJAB: NW Frontier Province,<br />
Murree Hills, Hazaza, Aug 1907, in fl., H. Deane<br />
s.n. (K); Kishtivar, Murree, Mt. Chumbi, 10 Aug<br />
1851, fl., 7-8000 ft., T. Thomson s.n. (K).<br />
Himalaya, Nagk<strong>and</strong>a, 7-10000 ft, 1844, M. P.<br />
Edgeworth 187 (K, 2 sheets).<br />
Cultivated material: Austria. Botanischen<br />
Garten in Wien, 6 Aug 1920, H. Zerny s.n. (W).<br />
Germany. Berlin, Botanischen Garten Berlin-<br />
Dahlem, 13 Aug 1976, Schwerdtfeger 3644 (B, 2<br />
sheets); 18 Jul 1983, Schwerdtfeger 14783 (B, 3<br />
sheets)<br />
Four specimens <strong>of</strong> <strong>Aralia</strong> cachemirica are<br />
preserved at P, which were collected during Victor<br />
Jacquemont’s voyage to India [V. Jacquemont 717,<br />
718, 913 (2 sheets), <strong>and</strong> a specimen without<br />
number]. V. Jacquemont 718 bears both leaves <strong>and</strong><br />
flowers <strong>and</strong> is representaive for the species. It is<br />
thus selected as the lectotype.<br />
Two specimens (J. D. Hooker s.n., LE, from<br />
Sikkim, 10-12000 ft; <strong>and</strong> O. Polunin et al. 5045,<br />
BM, E, from Nepal, below Garjigoth, 9500 ft, 9<br />
Aug 1952) appear to be morphologically<br />
intermediate beteween A. cachemirica <strong>and</strong> A.<br />
tibetana. They are herein cited as A. cachemirica<br />
on a tentative basis. Field studies are required in<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
these areas to ascertain the taxonomic status <strong>of</strong> the<br />
populations represented by these specimens.<br />
The type <strong>of</strong> <strong>Aralia</strong> macrophylla has not been<br />
located at present. Its author, Lindley (1844, p.<br />
73), noted the species as follows: “This is an<br />
herbaceous plant from the North <strong>of</strong> India, looking<br />
like A. racemosa, but much larger in all its parts.<br />
The flowers, which are greenish-yellow, have been<br />
produced in the garden <strong>of</strong> the Horticultural Society,<br />
where the plant has been raised from seeds<br />
presented by the East India Company.”<br />
9. <strong>Aralia</strong> tibetana G. Hoo — Fig. 30; color plate<br />
7: A-F.<br />
<strong>Aralia</strong> tibetana G. Hoo, Acta Phytotax. Sin., Add.<br />
1: 175. 1965. TYPE: China. XIZANG: Tibet<br />
[Xizang Prov.], Rung-sa, Mt. Jolmo Lungma,<br />
under forests <strong>of</strong> Tsuga, 3100 m, 2 Aug 1959,<br />
Mt. Jolmo Lungma Exped. Team 758 (holotype:<br />
PE!).<br />
Panax tripinnatum Wall. ex Don, Gen. Hist. 3: 384.<br />
1834 (non <strong>Aralia</strong> tripinnata Blanco, Fl. Filip.<br />
ed. 1: 223). TYPE: Emodi [Himalaya], at<br />
Gosain Than [now in Nepal), Aug 1821, fl, N.<br />
Wallich 4934 (holotype: BM!; isotypes: G!, K!,<br />
K-W!).<br />
Perennial herb, up to ca. 2 m tall, stem purple<br />
to dark purple, nearly glabrous, bearing 4-5 leaves;<br />
rhizomes thick. Leaves 50-90 cm long, 40-80 cm<br />
wide, tripinnately or bipinnately ternate; uppermost<br />
leaves 12-20 cm long, 10-15 cm wide, bipinnate<br />
to pinnate, or occassionally tripinnate; stipules <strong>of</strong><br />
lower leaves 1.5-2.5 cm long, 1-1.2 cm wide,<br />
membranaceous, narrowly triangular to ovate,<br />
pilose, somewhat divided or toothed at the tip,<br />
enclosing the emerging young shoot, stipules <strong>of</strong><br />
upper leaves 2-3 mm long, triangular in shape;<br />
petioles <strong>of</strong> uppermost leaves 3-7 cm long, pilose;<br />
leaflets 3.5-11.5 cm long, 2.3-5.5 cm wide,<br />
membranaceous, ovate, or occasionally narrowly<br />
so to elliptic, sharply long acuminate or sometimes<br />
acuminate at apex, rounded to subcordate at base,<br />
doubly serrulate to sometimes doubly serrate at<br />
margin, lateral veins 7-9 on each side, sunken on<br />
adaxial surface, conspicuous on both surfaces,<br />
adaxial surface green, scabrous when young, pilose<br />
along veins to glabrescent when mature, abaxial<br />
surface whitish green, pilose, or at least along the<br />
veins <strong>and</strong> veinlets, with conspicuously visible fine
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
Fig. 30. <strong>Aralia</strong> tibetana G. Hoo. A. Leaf with axillary inflorescence. B. Stipule. C. Umbel also showing bracts. D. Inflorescence.<br />
E. Floral bud. F. Flower. G. Flower showing anthers <strong>and</strong> gynoecium after breaking <strong>of</strong>f two petals. H. Fruit. I. Seed, left<br />
showing face view, <strong>and</strong> right showing side view (A, D, H & I - Qinghai-Xizang Team 74-2440, KUN; B, C & E-G - Qinghai-<br />
Xizang Team 6340, KUN).<br />
91
92<br />
veinlets; petiolules 0.1-2 cm long, slightly pilose.<br />
Inflorescence 20-40 cm long, terminal <strong>and</strong> axillary,<br />
consisting <strong>of</strong> numerous umbels, primary<br />
inflorescence branches 12-30, mostly racemosely<br />
arranged, <strong>of</strong>ten 3-7 forming a circle on the main<br />
inflorescence axis, each primary branch 4-8.5 cm<br />
long, consisting <strong>of</strong> 1-4 umbels, usually only the<br />
terminal umbel <strong>of</strong> each branch bearing fruits, others<br />
becoming aborted; terminal umbels mostly 30-70flowered,<br />
pedicels 6.5-11 mm long, pubescent,<br />
enlarged <strong>and</strong> with tufts <strong>of</strong> hairs at the tip, the hairs<br />
<strong>and</strong> the enlarged tip forming a somewhat disk-like<br />
structure at the base <strong>of</strong> the flower; lateral umbels<br />
15-25-flowered, pedicels 4.5-9.5 mm long; bracts<br />
<strong>of</strong> primary branches 5-7 mm long, 1-2 mm wide,<br />
lanceolate to narrowly triangular, ciliate at margin,<br />
sometimes bracts becoming leafy <strong>and</strong> enlarged;<br />
bracteoles 1.6-2.2 mm long, 0.4-0.6 mm wide,<br />
lanceolate, glabrous or nearly so. Sepals 0.4-0.6<br />
mm long, 0.4-0.5 mm wide, narrowly triangular,<br />
persistent on fruits; petals 1.5-2.0 mm long, 1-1.2<br />
mm wide, ovate, pale purplish, with a conspicuous<br />
vascular bundle in the middle; stamens 5, 2.2-2.8<br />
mm long, filaments 2-2.5 mm long, anthers 0.7-<br />
0.8 mm long, 0.4-0.5 mm wide, ovate; ovaries 5locular,<br />
styles 0.7-0.8 mm long at anthesis, tip <strong>of</strong><br />
ovary enlarged <strong>and</strong> projected. Fruits 2.7-3.5 mm<br />
long, 3-3.4 mm wide, globose, purplish black,<br />
persistent styles divided to the middle, 1.1-1.3 mm<br />
long. Seeds 2-2.5 mm long, 1-1.2 mm wide, 0.6-<br />
0.7 mm thick, kidney-shaped, whitish gray,<br />
smooth.<br />
Common name: Xizang tu dang gui.<br />
Phenology: flowering in July <strong>and</strong> August;<br />
fruiting in September to October.<br />
Distribution: in the eastern Himalaya, including<br />
eastern <strong>and</strong> central Nepal, Sikkim <strong>of</strong> India, western<br />
Bhutan, <strong>and</strong> southern Tibet <strong>and</strong> western Sichuan<br />
<strong>of</strong> China (Fig. 31).<br />
Ecology: in shady forests, edge <strong>of</strong> woods,<br />
grassy slopes, among rocks <strong>and</strong> shrubs, especially<br />
in mixed Rhododendron forests; 2400-4250 m<br />
(8,000-14,000 ft).<br />
Additional specimens examined: Bhutan.<br />
Bhutan. Mindook La, 31 Jul 1884, in fl., Dungboo<br />
2 (P). Tilaging Timpu, 8000 ft, 17 Aug 1914, R. E.<br />
Cooper & A. K. Bulley 2567 (BM); Dotura +<br />
Timpu, 1 Oct 1914, fr, 10,000 ft, R. E. Cooper 2983<br />
(BM); west Bhutan, Damthaug, 30 Sep 1933, F.<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
Ludlow & G. Sherriff 535 (BM); Belierea<br />
Deucheng, 7000 ft, 20 Jul 1949, fl, in bamboo <strong>and</strong><br />
Tsuga forest, 3-4 ft high, F. Ludlow et al. 21302<br />
(BM). China. SICHUAN: Kangding, 11 Aug<br />
1930, young fr, Z. P. Huang et al. 1743 (WUK);<br />
Kangding, 8 Aug 1930, young fr, Z. P. Huang et<br />
al. 1854 (PE, WUK). XIZANG: Chola Xian, Jiba<br />
Xiang, at the edge <strong>of</strong> secondary forest, 3500 m, 1-<br />
2 m tall, 19 Jul 1975, fl, C. Y. Wu et al. 75-1043<br />
(KUN, 2 sheets, PE, 2 sheets); Lebu Qu, 3200 m,<br />
25 Aug 1975, young fr, Qinghai-Xizang Additional<br />
Collections Team 751563 (KUN, PE); Lebu Qu,<br />
Deyinggou, shrubby area, roadside, 3200 m, 1-1.5<br />
m tall, 8 Sep 1975, fr, Qinghai-Xizang Additional<br />
Collections Team 751842 (KUN, 2 sheets, PE, 2<br />
sheets); Tibet, Cuona, Mama Xiang, 3062 m, 19<br />
Sep 2009, Z.-L. Nie et al. 788 (KUN). Jinong Xian,<br />
Jinong zhen, Rukacun 4277 m, 28 Sep 2009, Z.-L.<br />
Nie et al. 1023 (KUN); Jinong Qu, Luka, 2960 m,<br />
13 Oct 1975, fr, plant 2.5 m, fruit black, Qinghai-<br />
Xizang Team 75-34 (PE); Tuodan, 3450 m, 5 Jul<br />
1975, Zhong Kao Hui 75-512 (PE, 3 sheets); from<br />
Jinong to Bangxing, in Pinus forest on slopes, 2850<br />
m, herb 1.5 m tall, 13 Jul 1975, floral buds,<br />
Qinghai-Xizang Team 6340 (KUN); Tuodang,<br />
Dongshan, in Abies-Quercus mixed forest, at the<br />
edge, 3300-3400 m, plant 0.5-0.8 m, C. Y. Wu et<br />
al. 75-531 (KUN, 2 sheets). Medog Xian,<br />
Kanongla Mountain, South side, edge <strong>of</strong> forest,<br />
common, 50-60 cm tall, 20 Aug 1982, fl, S. Z. Chen<br />
& B. S. Li 00296 (PE, 2 sheets). Yadong Xian,<br />
Yadong, Xiaoyadong xiang, Lilasha, 3187 m, 23<br />
Sep 2009, Z.-L. Nie et al. 938 (KUN); Qinbeigou,<br />
in mixed forests, 3350 m, Zhong Kao Hui 75-896<br />
(PE, 3 sheets); Nanguolashan, in Tsuga forest, 14<br />
Aug 1975, young fr, Zhong Kao Hui 75-1051 (PE,<br />
3 sheets); Kailingang, shrubby area near fields,<br />
3200 m, 24 Jul 1960, fl, G. X. Fu 403 (PE);<br />
Hongqingang Qu, at the edge <strong>of</strong> Pinus forest, 2800<br />
m, 9 Sep 1974, fr, Qinghai-Xizang Team 74-2149<br />
(PE); near Xiashimazheng, shrubby area on slopes,<br />
2900 m, 13 Sep 1974, fr, Qinghai-Xizang Team<br />
74-2440 (KUN, PE, 2 sheets); near Yadong, 23 Jul<br />
1953, fl, in forest, B. Q. Zhong 5929 (PE, 2 sheets).<br />
Tibet, Chumbi, 9 Jul 1913, R. E. Cooper & A. K.<br />
Bulley 270 (BM, E); Chumbi, 8,000 ft, 25 Aug<br />
1913, R. E. Cooper 650 (BM); Chumbi, 10 Aug<br />
1857, A. Fleming 331 (E); Yatung, 27 o 51’N,<br />
88 o 35’E, H. E. Hobson s.n. (K). Sikong, Nan Yuen,<br />
near Chang Lou-pine, Ta-hsiang lin, 2200 m, 28<br />
Aug 1939, fr, C. Y. Chiao 2069 (A). India.
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
Fig. 31. Map <strong>of</strong> eastern Asia showing the distribution <strong>of</strong> <strong>Aralia</strong> tibetana G. Hoo.<br />
SIKKIM: Lachung, 9500 ft, 10 Aug 1892, G. A.<br />
Gammie 712 (P); Ichu Zeu, 8 Aug 1877, fl, G. King<br />
4436 (BM); East District, Karponong, 8 Aug 1980,<br />
P. K. Hajra 525 (BSHC); Changu, rare, B. Krishna<br />
2238 (BSHC); East Sikkim, Karponang – Chhangu,<br />
4 Aug 1985, D. C. S. Raju 4128 (BSHC, 2 sheets);<br />
North District, on way from Lachen to Thangu,<br />
3500 m, ca. 1.5 m tall, 26 Aug 1982, P. Chakraborty<br />
2397 (BSHC, 2 sheets); North Sikkim, Lachung –<br />
Zakophyak, 3000 m, 25 Jul 1990, S. Kumar & P.<br />
Singh 9562 (BSHC, 3 sheets); North Sikkim,<br />
Lachung to Dambergang, shrubs to 2 m, flowers<br />
in umbel, greenish, 27 Jul 1998, S. S. Dash & A.<br />
Maih 18555 (BSHC, 3 sheets); North Sikkim,<br />
between Phuni <strong>and</strong> Yakche, 3230 m, 14 Jul 1996,<br />
G. P. Singh & D. G. Long 17851 (BSHC). Nepal.<br />
C Nepal, Bagmati Zone, Rasuwa Distr., Chyauche<br />
Kharka (3600 m) – a bridge (2010 m) – Lingju<br />
(2040 m), 28 o 14’N, 85 o 07’E, 12 Aug 1994, fl, F.<br />
Miyamoto et al. 10271 (BM); C Nepal, Bagmati<br />
Zone, Rasuwa Distr., Pati Kharka (3760 m) – cross<br />
a river (2800 m) – a Kharka (near Pabil Kharka,<br />
2860 m), 28 o 14’N, 85 o 09’E, 5 Aug 1994, fl, F.<br />
Miyamoto et al. 20169 (BM), 10207 (BM, 2<br />
sheets); Kasuwa Khola, 11,000 ft, mixed<br />
Rhododendron forest, 19 Aug 1975, L. W. Beer<br />
93<br />
25333 (BM); East Nepal, Baroya Khimty-Thakma<br />
Khola, 16 Nov 1963, sterile, H. Hara et al.<br />
6301046 (BM, K); Langtang forest area, 1 Aug<br />
1949, O. Polunin 1639 (BM, 2 sheets, E); near<br />
Dhorpatan, 9,500 ft, 12 Jul 1954, Stainton et al.<br />
3474 (BM, E); central Nepal, Langtang, 28 o 15N,<br />
85 o 30E, 8000 ft, 22 Sep 1965, J. D. A. Stainton<br />
5147 (BM); Lete, S <strong>of</strong> Tukucha, Kali G<strong>and</strong>aki,<br />
among shrubs <strong>and</strong> tall herbs on grass slopes, height<br />
5 ft, berries black, 10,000 ft, 17 Sep 1954, Stainton<br />
et al. 7904 (BM); Simbua Khola, 27 o 30’N,<br />
87 o 57’E, 10,000 ft, 24 Jun 1969, floral buds, L. H.<br />
J. Williams 903 (BM); Toketey, 13,000-14,000 ft,<br />
L. Dhevoj 0493 (BM, 2 sheets, E).<br />
Panax decompositum Wall. ex DC. (de<br />
C<strong>and</strong>olle 1830) was previously treated as a<br />
synonym <strong>of</strong> <strong>Aralia</strong> cachemirica by Clarke (1879).<br />
Wen et al. (2002) placed it as the synonym <strong>of</strong> <strong>Aralia</strong><br />
tibetana. My recent examination <strong>of</strong> N. Wallich<br />
4935 at K-W has shown that this specimen belongs<br />
to <strong>Aralia</strong> armata (Wall. ex G. Don) Seem.<br />
I noted a few specimens from Sichuan, China<br />
which are similar to <strong>Aralia</strong> tibetana. But they differ<br />
from the latter that the lateral leaflets immediately
94<br />
below the terminal leaflets have relatively long<br />
petiolules (14-18 mm long); <strong>and</strong> that the teeth at<br />
the leaflet margin are short <strong>and</strong> triangular with a<br />
slightly gl<strong>and</strong>ular tip. These specimens are cited<br />
as below: China. SICHUAN: Baoxin: Bao-Hsin<br />
(Moupin), in shadow or damp places, creeping<br />
thick roots used as asprine, Jul-Aug 1939, young<br />
fr, S. Y. Hu 1242 (A, 2 sheets). Dujiangyan Shi,<br />
west <strong>of</strong> Kuan Hsien, woodl<strong>and</strong>s, Pan-lan-shan,<br />
8000 ft, Oct 1910, fr, E. H. Wilson 4285 (A, 2<br />
sheets). Heishui, Wabo, 2750 m, 8 Jun 1959, floral<br />
buds, Chuan-Jing-A (Sichuan Economic Plants<br />
Aba) 1333 (KUN). Hongqi, Lema, herb 1 m tall,<br />
31 Jul 1959, fl, Sichuan Economic Plants<br />
Liangshan Team 1510 (KUN). Leibo Xian,<br />
Shanlingang, Dagudui Xiang, Huangmaolang,<br />
2700 m, herb 20-30 cm, 19 Jun 1959, Sichuan<br />
Economic Plants Liangshan Team 0801 (KUN).<br />
Lieng Ho Kou, Aug 1938, Wang & Wen 694 (A).<br />
Western China, locality unknown, 8500 ft., 2-3 ft,<br />
Jul 1903, in fl., E. H. Wilson 3710 (A, BM, K, P).<br />
Further collections <strong>and</strong> field studies are<br />
required to determine whether these collections<br />
represent a new species or just <strong>Aralia</strong> tibetana.<br />
10. <strong>Aralia</strong> fargesii Franch. — Color plate 8: A-F.<br />
<strong>Aralia</strong> fargesii Franch., J. Bot. (Morot) 10: 302.<br />
1896. TYPE: China. CHONGQING: Sutchuen<br />
oriental [now Chongqing Shi], District<br />
<strong>of</strong> Tchen-keou-tin [Chengkou], R. P. Farges s.n.<br />
(holotype: P!, photo [A]!; isotypes: P(4)! photo<br />
[A]!, leaves only).<br />
<strong>Aralia</strong> kansuensis G. Hoo, Acta Phytotax. Sin.,<br />
Addit. 1: 174. 1965. TYPE: China. GANSU:<br />
Xigu (now Zhouqu Xian), Heiyugou,<br />
Shanshengyeliang, on a Quercus tree, 3100 m,<br />
19 Jul 1951, young fr, Z. B. Wang 14444<br />
(holotype: WUK!; isotype: PE!).<br />
Perennial spreading herb 0.7-1.5 m tall. Stem<br />
purple to purplish green; rhizomes thick <strong>and</strong><br />
horizontal in older plants, younger ones well well<br />
developed tap roots. Leaves 27-45 cm long, 20-<br />
35 cm wide, 3-4 on each plant, tripinnately or<br />
bipinnately ternate; stipules 8-10 mm long, 2-3 mm<br />
wide, narrowly triangular, membranaceous;<br />
petioles <strong>of</strong> uppermost leaves 7-12.5 cm long,<br />
pilose; leaflets (3-) 5.5-11 cm long, (1.5-) 3-8.5<br />
cm wide, membranaceous to chartaceous, ovate to<br />
broadly or narrowly so, acuminate to sharply so at<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
apex, cordate, subcordate, to rarely truncate at base,<br />
doubly serrate at margin, lateral veins 7-9 on each<br />
side, conspicuous on both surfaces, adaxial surface<br />
green, sparsely scabrous, abaxial surface light<br />
green, pilose along the veins <strong>and</strong> veinlets,<br />
petiolules 4-20 mm long, pilose. Inflorescence 15-<br />
43 cm long, terminal <strong>and</strong> axillary, a loose panicle<br />
consisting <strong>of</strong> an umbellate structure at the top <strong>and</strong><br />
2-3 branches below, pilose, the top umbellate<br />
structure with 4-6 primary branches, each primary<br />
inflorescence branch 7-14 cm long, mostly with<br />
1-6 umbels; terminal umbels 15-25-flowered,<br />
pedicels 5-6 mm long, pubescent; lateral umbels<br />
7-15-flowered, pedicels 4-5 mm long; bracts <strong>of</strong><br />
primary branches leafy to lanceolate, variously<br />
sized; bracteoles 1.3-1.5 mm long, 0.1-0.2 mm<br />
wide, linear to lanceolate. Sepals 0.5-0.6 mm long,<br />
0.6-0.7 mm wide, triangular, persistent on fruits;<br />
petals 1.6-1.8 mm long, 1-1.1 mm wide, ovate, with<br />
a conspicuous vascular bundle in the middle;<br />
stamens 5, filaments 1.9-2.1 mm long, anthers ca.<br />
1 mm long, ovate; ovaries 5-3-locular, styles 5-3<br />
lobed to the middle. Fruits 4-5 mm in diameter,<br />
globose, persistent styles divided to the middle.<br />
Seeds 3-3.2 mm long, 1.3-1.4 mm wide, ca. 0.2<br />
mm thick, kidney-shaped, whitish, smooth.<br />
Common name: long yan du he.<br />
Uses: roots as blood-regulating medicine, also<br />
for arthritis.<br />
Phenology: flowering in July; fruiting in August<br />
to September.<br />
Distribution: Hubei, Chongqing, Gansu,<br />
Sichuan, Shaanxi <strong>and</strong> Qinghai provinces <strong>of</strong> China<br />
(Fig. 32).<br />
Ecology: in forests or thickets, relatively rich<br />
habitats; 1750-3400 m.<br />
Additional specimens examined: China.<br />
CHONGQING: Western China, Wu Shan (Mt.<br />
Wu), 9000 ft., 5 Jul 1903, in fl., E. H. Wilson 3698<br />
(BM, K, 2 sheets, P, 2 sheets, PE). GANSU:<br />
Tianshui, Pianlegou, 2300 m, 8 Jul 1963, fl, Q. X.<br />
Li 422 (PE). Xigu (now Zhuqu Xian), Shatan,<br />
Dahaigou, 2500 m, 29 Jul 1951, fl, Z. B. Wang<br />
14625 (WUK). Zhouqu Xian, Boyu, Qumagou,<br />
in gully mixed forest, 2600 m, 11 Aug 1993, fr, Y.<br />
S. Lian et al. 93-339 (MO). HUBEI: Fang, A.<br />
Henry 6785 (K, 2 sheets, NY). Shennongjia Forest<br />
Dist., Xiao Nong Tan, J. Wen 1442 (US); Xiao<br />
Nong Tan, in Abies forest, N31 o 28.73’,
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
Fig. 32. Map <strong>of</strong> China showing the distributions <strong>of</strong> <strong>Aralia</strong> fargesii Franch. <strong>and</strong> A. atropurpurea Franch.<br />
E110 o 18.11’, 2180 m, 28 Jun 2001, J. Wen 5446<br />
(US); Shennongjia Forest Dist., on the road to Xiao<br />
Nong Tan, N31 o 29.91’, E110 o 22.27’, 1640 m, 28<br />
Jun 2001, J. Wen 5455 (US). QINGHAI: Xunhua,<br />
Tianchi, shrubby area on slope, 4 Aug 1985, Huang<br />
Tu Team 5939 (WUK, 2 sheets); Xunhua, Mengda,<br />
Muchanggou, in forest, 2900 m, 29 Jun 1964, fl,<br />
Z. F. Wei 390 (WUK). SHAANXI: Mei Xian,<br />
Taibeishan, Tangyu, Ketianguan, forests, Zhu et al.<br />
2435 (PE). Xitaibaishan, near Wafangzhi, 1750<br />
m, slope, 7 Jul 1959, fl, J. X. Yang 681 (WUK);<br />
Taibaishan, ravine to the east <strong>of</strong> Ta-T’ien, 15 Jul<br />
1938, fl, T. N. Liou & P. C. Tsoong 2175 (PE).<br />
SICHUAN: Baoxing Xian, Pao-hsing-hsien, 25<br />
Jul 1936, young fr, K. L. Chu 3300 (BM, PE); Paohsin-hsien,<br />
formerly known as Mupin,<br />
Dachangpeng, near the 52th Peng, 7 Aug 1954,<br />
young fr, T.-P. Soong 38957 (KUN, PE, WUK).<br />
Dajing Xian, Wanli Xiang, Wanlicheng Gou, in<br />
95<br />
forest near the stream, 3400 m, 19 Jul 1958, young<br />
fr, Q. Li 78132 (PE, WUK). Emei Shan, Baiyunshi,<br />
30 Jul 1952, fl, W. P. Fang et al. 31839 (PE); Emei<br />
Shan, Baiyunshi, T. Y. Zhou & G. J. Xu 466 (PE);<br />
Emei Shan, Luohanpo, 2200 m, 8 Jul 1996, young<br />
fr, Y.-L. Xu 960329 (MO); Emei Shan,<br />
Leidongping, 2900 m, 29 Jul 1997, J. Wen 3079<br />
(US). Ganle, Haitang, Yue-er-gou, 22 Jul 1959,<br />
2200 m, Sichuan Economic Plants 4304 (PE, 2<br />
sheets). Heishui, Luhua, 2850 m, 31 Jul 1957,<br />
young fr, Q. Li 73940 (PE); Guwanshi, Muozhigou,<br />
15 Aug 1957, fr, 2750 m, Q. Li 73998 (PE). Lifan,<br />
Mung-twin-ko, Aug 1941, young fr, S. Y. Hu 1933<br />
(A), 2075 (A). Mou Xian & Li Xian (formerly<br />
Lifan Xian), 1952, Z. He & Z. L. Zhou 12948 (PE).<br />
Wenchuan Co., Ts’ao Po, Yin-Chang-Kao, 5 Aug<br />
1942, fr, S. Y. Hu 2591 (A, 2 sheets). Sichuan,<br />
location unknown, 1930, T. Tang 23346a (PE).
96<br />
<strong>Aralia</strong> fargesii differs from A. atropurpurea in<br />
its styles connate to the middle (vs. completely<br />
distinct in A. atropurpurea), well developed thick<br />
rhizomes (vs. well developed tap roots) in older<br />
plants, inflorescence a loose panicle with an<br />
umbellate structure on top <strong>and</strong> 2-3 additional<br />
primary branches below (vs. an umbellate<br />
inflorescence in A. atropurpurea), <strong>and</strong> shorter<br />
pedicels (5-6 mm vs. 8-12 mm in terminal umbels).<br />
In general, <strong>Aralia</strong> fargesii is a bigger <strong>and</strong> a more<br />
robust plant ca. 1-1.5 m tall,<br />
<strong>Aralia</strong> kansuensis G. Hoo is now treated as the<br />
synonym <strong>of</strong> A. fargesii. The type <strong>of</strong> <strong>Aralia</strong><br />
kansuensis bears smaller leaflets which are 3-5 cm<br />
long <strong>and</strong> 1-2.5 cm wide. There are two<br />
inflorescences on the type specimen; one is an<br />
umbellate structure with 7 primary branches, <strong>and</strong><br />
the other bears two whorls <strong>of</strong> primary branches<br />
(6-7 branches each). The type resembles closely<br />
A. fargesii except that the leaflets are smaller. I<br />
interpret the morphotype in “A. kansuensis” as the<br />
extreme condition at the higher altitudes <strong>and</strong> its<br />
westernmost distribution. Another specimen (Z.<br />
B. Wang 14625) was also collected in Xigu <strong>of</strong><br />
Gansu, but at a lower altitude (2500 m). This<br />
specimen shows the typical morphology <strong>of</strong> A.<br />
fargesii. Huang Tu Team 5939 was collected from<br />
Xunhua <strong>of</strong> Qinghai province at the altitude <strong>of</strong> 2800<br />
m. One <strong>of</strong> the two sheets has smaller leaflets (3-5<br />
cm long, 1.5-2 cm wide), closely resembling the<br />
type specimen <strong>of</strong> A. kansuensis, <strong>and</strong> the other sheet<br />
has much larger leaflets (7-8.5 cm long, 3-4.7 cm<br />
wide).<br />
11. <strong>Aralia</strong> atropurpurea Franch. — Fig. 33.; color<br />
plate 9: A-H.<br />
<strong>Aralia</strong> atropurpurea Franch., J. Bot. (Morot)<br />
10:301. 1896. Panax atropurpureus (Franch.)<br />
H<strong>and</strong>.-Mazz., Vegetationsbilder 22 (8): 9.<br />
1932. TYPE: China. YUNNAN: in woods <strong>of</strong><br />
Ta-long-tan near Tapin-tze, 10 Jul 1888, P. J.<br />
M. Delavay 4530 (holotype: P!; isotypes: P, 2<br />
sheets!).<br />
<strong>Aralia</strong> yunnanensis Franch., J. Bot. (Morot) 10:<br />
303. 1896. <strong>Aralia</strong> fargesii Franch. var.<br />
yunnanensis (Franch.) H. L. Li, Sargentia 2:<br />
103. 1942. TYPE: China. YUNNAN: [Dali<br />
Shi], Kichan [Jizhushan or Jishan], Tapintze,<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
21 Aug 1889, P. J. M. Delavay 4027 (lectotype,<br />
designated by Li, 1942, P!; isolectotype: P!).<br />
<strong>Aralia</strong> melanocarpa (H. Lév.) Lauener, Notes Roy.<br />
Bot. Gard. Edinburgh 32: 94. 1972.<br />
Eleutherococcus melanocarpus H. Lév., Bull.<br />
Acad. Int. Géogr. Bot. 24: 282. 1914. TYPE:<br />
China. YUNNAN: near Ma-li-ouan, 2550 m,<br />
Oct 1913, E. E. Maire s.n. (holotype: P!;<br />
isotypes: E, cited in Lauener 1972, P!, W!).<br />
<strong>Aralia</strong> dumetorum H<strong>and</strong>.-Mazz., Symb. Sin. 7:<br />
701. 1933. TYPE: China. YUNNAN: near<br />
Ma-li-ouan, 2600 m, Jul 1913, E. E. Maire s.n.<br />
(holotype: W!).<br />
Perennial herb 0.5-1 m tall, <strong>and</strong>romonoecious.<br />
Main roots 15-40 cm long, deep, forked <strong>of</strong>ten<br />
unequally. Stem purplish green, with 2-4 leaves.<br />
Lower leaves 60-75 cm long, 50-70 cm wide,<br />
tripinnately ternately compound; upper leaves 17-<br />
25 cm long, 13-22 cm wide, bipinnately ternate to<br />
pinnate or sometimes trifoliate; stipules 3-4 mm<br />
long, 2-3 mm wide, narrowly triangular, more or<br />
less ciliate at margin; petioles 2.3-14.5 cm long,<br />
slightly pilose to glabrescent, occasionally more<br />
densely pilose, especially in Muli area <strong>of</strong> Sichuan<br />
province; leaflets 2.5-7.8 cm long, 1.5-5 cm wide,<br />
1-5 per pinnule, chartaceous to sometimes<br />
membranaceous, ovate to narrowly ovate, or<br />
sometimes broadly so, acuminate to sharply so at<br />
apex, cordate, obliquely cordate, subcordate,<br />
truncate to rounded at base, finely doubly serrate<br />
at margin, lateral veins 7-9, conspicuous on both<br />
sides, adaxial surface green, scabrid, abaxial<br />
surface grayish green, pilose along the veins <strong>and</strong><br />
veinlets, petiolues more or less pilose, those <strong>of</strong><br />
lateral leaflets (0) 0.3-2 cm long, gradually<br />
becoming shorter toward the tip <strong>of</strong> the pinnae,<br />
those <strong>of</strong> terminal leaflets 1.2-3.5 cm long.<br />
Inflorescence 10-25 cm long, loose, terminal <strong>and</strong><br />
axillary, more or less umbellate in outline,<br />
sometimes with one or two umbels below the<br />
primary umbellate structure, glabrous, primary<br />
branches 2-4, each with 1-5 umbels, each primary<br />
branch 8-20 cm long, subtended by a leafy bract<br />
or a lanceolate bract, sizes <strong>of</strong> bracts varied greatly,<br />
terminal umbels 15-25-flowered, pedicels 8-12 mm<br />
long, pilose to nearly glabrous, lateral umbels 8-<br />
14-flowered, pedicels 5-11 mm long, pilose to<br />
nearly glabrous, bracteoles 1.5-1.8 mm long, 0.2-<br />
0.35 mm wide, lanceolate. Flowers dark purple,
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
Fig. 33. <strong>Aralia</strong> atropurpurea Franch. A. Habit showing a leaf <strong>and</strong> inflorescences. B. Rhizome <strong>and</strong> root. C. Base <strong>of</strong> petiole<br />
showing stipule morphology. D. Leaf on a leaf on the lower part <strong>of</strong> the stem. E. Close-up showing leaflet margin <strong>and</strong><br />
pubescence on lower surface. F. Flower after anthesis. G. Young fruit. H. Floral bud. I. Flower (A-I - Wen 3057, US).<br />
97
98<br />
floral buds globose, with several flowers aborted,<br />
perhaps functionally male, on each terminal umbel,<br />
more so on lateral umbels at fruiting stage; sepals<br />
minute, 0.5-0.6 mm long, 0.4-0.6 mm wide,<br />
triangular or narrowly so, spreading at anthesis,<br />
recurved afterwards, persistent into fruiting stage;<br />
petals 1.5-2.0 mm long, 1.2-1.3 mm wide, ovate;<br />
stamens 1.5-1.7 mm long, filaments 1.2-1.5 mm<br />
long, anthers 0.4-0.5 mm long, 0.3-0.4 mm wide,<br />
oblong; gynoecium conical at base, styles 5, 0.7-<br />
0.9 mm long, completely divided after anthesis,<br />
the apical portion <strong>of</strong> the ovary somewhat projected.<br />
Fruits 5-6 mm in diameter, globose, 5-ridged when<br />
dry, with 5 persistent recurved styles. Seeds 3.5-<br />
3.7 mm long, 2.1-2.2 mm wide, ca. 0.2 mm thick,<br />
kidney-shaped, whitish.<br />
Common name: nong zhi long yan du he.<br />
Uses: roots used as blood-regulating medicine<br />
<strong>and</strong> for correcting irregular menstrual period.<br />
Phenology: flowering in June to September;<br />
fruiting from July to October.<br />
Distribution: Sichuan <strong>and</strong> Yunnan <strong>of</strong> China<br />
(Fig. 32).<br />
Ecology: in broadleaf or mixed conifer <strong>and</strong><br />
evergreen broadleaf forests; 1800-3300 m.<br />
Additional specimens examined: China.<br />
SICHUAN: Huidong, Baisha Xiang, 3100 m, 19<br />
Jun 1959, fl, S. G. Wu 771 (KUN). Muli Xian,<br />
Muli, Consinliang near Ngerya, on the border <strong>of</strong><br />
Chungtien, 2400 m, in woods near stream, 24 Aug<br />
1939, fl, K. M. Feng 2791 (KUN, 3 sheets); Muli,<br />
near Benxi, 2600 m, T. T. Yü 14050 (A, KUN, 2<br />
sheets, PE); Muli Xian, Xiamaidi Xiang, Mianbu<br />
Village, 41 Km S <strong>of</strong> Muli along rt S216, between<br />
km markers 495 & 496, N27 o 44’53.6",<br />
E101 o 12’46.3", 2560 m, 6 Aug 2007, fr, J. Wen et<br />
al. (Tibet-MacArthur Expedition) 1917 (F, KUN,<br />
US). YUNNAN: locality unknown, specimen<br />
received on 10 Aug 1921, R. P. Maire s.n. (P);<br />
woods <strong>of</strong> Ta Long Tan near Ta Pin Tze, 10 Jul 1888,<br />
P. J. M. Delavay 4536 (P, 2 sheets); woods near<br />
Che Tcho Tze <strong>of</strong> Ta pin Tze, 2000 m, 27 Aug 1885,<br />
in late fl., M. Delavay s.n. (P); Ta Pin Tze, 22 Aug<br />
1885, P. J. M. Delavay s.n. (P); Ta Long Tan, Ta<br />
Pin Tze, 26 Jul 1885, in fl., P. J. M. Delavay s.n.<br />
(P). Gorges near San Ictrang Kiou (Ho Kin, =<br />
Heqin), 6 Aug 1885, P. J. M. Delavay s.n. (P).<br />
Woods in the mountains near Ma-li-ouan, 2600 m,<br />
flowers green <strong>and</strong> fruits black, August, R.-P. Maire<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
s.n. (P); woods near Ma-li-ouan, 2600 m, Aug, E.<br />
E. Maire s.n. (P, 2 sheets). Tchong-chan, 23 Aug<br />
1905, in fl & young fr., F. Ducloux 3941 (P, 3<br />
sheets). Bingchuan Xian, Jizhushan, Zhushengshi,<br />
near the creek in front <strong>of</strong> the temple, 2280 m, plant<br />
70 cm tall, 26 Jul 1988, young fr, J. Wen 518 (OS);<br />
Jizhushan, on the way from Zhushengshi to<br />
Jingdian, plant 50-65 cm tall, petals dark purple,<br />
anther white at first, turning purple, young fruits<br />
green, turning purple, common, 27 Jul 1988, young<br />
fr & fl, J. Wen 522 (OS); Bingchuan Xian,<br />
Jizhushan, 2500 m, 24 Jul 1997, J. Wen 3048 (US).<br />
Dali Xian, between km marker 12 <strong>and</strong> 13 on road<br />
from Xiaoguan to Dacang, 2400-2500 m, 25 o 31’N,<br />
100 o 12’E, 9 Jul 1984, floral buds, 1984 Sino-Amer.<br />
Bot. Exped. 928 (A, CAS, KUN, PE, US). Dayao<br />
Xian, Shantai Qu, Shantai, 2000 m, herb 0.6 m, 9<br />
Jul 1965, fl, Woody Oil Plants Team 65-0329<br />
(KUN); Shantai Qu, Bakou Chun, Duodihe, Buzha,<br />
Yangwozhi, in pine forest, 12 Jul 1965, fl, Woody<br />
Oil Plants Team 65-0429 (KUN, 2 sheets). Deqin<br />
Xian, near Chang Jiang, on the way from Chizhong<br />
to Xila, 2600 m, in broadleaf forest, herb ca. 1 m<br />
tall, 5 Oct 1959, fl & fr, Kunming Work Station<br />
23918 (KUN, 3 sheets). Eshan Xian, Shi Qu, NW<br />
<strong>of</strong> the Qu Office, 1800 m, 1 Oct 1958, S. G. Wu<br />
151 (KUN, 2 sheets). Fumin Xian, Yongding,<br />
Tuodan, Gangkuanggou, 2600 m, in pine forest,<br />
herb 80 cm tall, B.-Y. Qiu 596355 (KUN). Heqin<br />
Xian, Huangping Qu, Jundeng Commune, Upper<br />
Dapingzhi, 2450 m, 16 Aug 1963, Northwest<br />
Yunnan Jingshajiang Team 6546 (KUN); Heqin,<br />
Lianping, Beitouqin, 7 Aug 1940, R. C. Ching<br />
23567 (KUN, 2 sheets); Heqin, Baiyan, Shaxi,<br />
2700 m, in woods, 8 Sep 1940, fr, R. C. Ching<br />
24437 (KUN, 3 sheets). Jingdong Xian,<br />
Chingtung, Ta-Mai-Ti Hou Shan, 1900 m, in mixed<br />
forest, 19 Oct 1939, fl & fr, M. K. Li 0720 (KUN,<br />
WUK, 2 sheets). Kunming, Xishan, Huatingshi,<br />
2000 m, in sparse forest, herb 0.8 m, 21 Aug 1953,<br />
fr, B. Y. Qiu 50212 (KUN); Xishan, Taihuashi, 5<br />
Nov 1938, K. M. Feng 116 (KUN); Xishan, behind<br />
Taihuashi, at the edge <strong>of</strong> pine forest, plant 60 cm<br />
tall, 27 Aug 1988, fr, J. Wen & X. P. Li 599 (OS);<br />
Xishan, just behind Taihuashi, in mixed pine <strong>and</strong><br />
evergreen broadleaf forest, associated with Pinus,<br />
Castanopsis, Rhododendron, wild ginger,<br />
Metapanax delavayi, <strong>and</strong> Smilax, 2250 m, plant<br />
35-60 cm tall, stem purple <strong>and</strong> zigzag, 14 Jun 1992,<br />
fl, J. Wen 1216 (A, F, US). Lanping Xian, Jingding<br />
Qu, Gaoping, 29 Jul 1965, fl, J.-S. Yang 5042
(KUN). Lijiang Xian, Yulong, Dahuoshan, 24 Aug<br />
1961, fl, Y.-F. Qi 612694 (KUN); Lidjiang [Lijiang<br />
Xian], 3000-3300 m, 11 Jul 1915, H. F. Hanel-<br />
Mazzetti 7025 (W, WU); Li-Kiang-Hsien, 2800 m,<br />
woods, Jun 1935, floral buds, C. W. Wang 71583<br />
(PE); Yuhu Xiang, Xuesongchun, on the way from<br />
Xuesong to Wenhai, Halagu area, 0.5-1.2 m tall, 7<br />
Aug 1988, fr, J. Wen 548 (OS); Lijiang Xian,<br />
Baisha Xiang, Yuhu Village, in mountainous forest<br />
near Shanchahe (three river junction) <strong>of</strong> the Yulong<br />
Snow Mountain, 26 Jul 1997, J. Wen 3057<br />
(US)Yuhu Xiang, Xue Song Village, N27 o 01’27.5",<br />
E100 o 12’12.5", 2952 m, 20 Jul 2007, young fr &<br />
fl, J. Wen et al. (Tibet-MacArthur Expedition) 926<br />
(F, KUN, US). Luquan Xian, Lu-Chieh, Tien Wei,<br />
2600 m, rocky place, 25 Oct 1940, fr, Y. P. Chang<br />
346 (KUN). Songming Xian, Guodong, 22 Aug<br />
1957, young fr, 2380 m, B. Y. Qiu 54837 (KUN, 2<br />
sheets); 24 Aug 1957, fl, B. Y. Qiu 54965 (KUN);<br />
Songming, Shaodian Qu, Queying Chun, 1800 m,<br />
streamside, in bushes, herb 0.5 m, 14 Oct 1953, fr,<br />
B. Y. Qiu 50428 (KUN, 2 sheets); Guodong,<br />
Laojishan, 2200 m, 8 Jul 1960, fl, B. Y. Qiu 57877<br />
(WUK); Kan-Ho, 2200 m, in a valley, 13 Oct 1950,<br />
fr, P. I. Mao 207 (KUN, PE). Tuodian Xian<br />
(formerly in Shuangbai Xian), Chayeqing, 5 Oct<br />
1958, fr, Z. Q. Huang 164 (KUN, 2 sheets). Weixi<br />
Xian, Pantiange Xiang, Zhigezi Village, ca 3.7 km<br />
from the junction <strong>of</strong> the road to Pantiange with Rt<br />
Shangrila-Weixi, N27 o 20’43.8", E99 o 15’31.7",<br />
2767 m, 30 Jul 2007, fr, J. Wen et al. (Tibet-<br />
MacArthur Expedition) 1487 (F, KUN, US).<br />
Wuding Xian, Shi Shan, 2200 m, in Quercus forest,<br />
moist habitat, 1 m, 15 Oct 1960, fl, Yunnan Tropical<br />
Biological Resources Team 60-160 (KUN, 2 sheets,<br />
WUK). Zhongdian, Baidi, 2500-2700 m, 1-1.5 m,<br />
18 Aug 1962, fl, Zhongdian Team 1018 (KUN, 2<br />
sheets); Baidi, 18 Aug 1962, fl, 2500 m, limestone,<br />
0.8 tall, Zhongdian Team 2289 (KUN); Haba,<br />
Zhazhamu, Longwang, Shaba, 2650 m, herb 0.8<br />
m, Zhongdian Team 63-3193 (KUN). Yunnan, S.<br />
Ten s.n. (WU). Western China, 1914, E. E. Maire<br />
294 (BM).<br />
When describing <strong>Aralia</strong> yunnanensis, Franchet<br />
(1896) cited two specimens (Delavay 4027 <strong>and</strong><br />
Delavay 4581). The two specimens are different<br />
in habits (Delavay 4027 herbaceous, <strong>and</strong> Delavay<br />
4581 woody) <strong>and</strong> they apparently represent two<br />
distinct species. Li (1942) typified <strong>Aralia</strong><br />
yunnanensis with the herbaceous Delavay 4027,<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
99<br />
then treated it as a variety <strong>of</strong> A. fargesii. Li’s<br />
definition <strong>of</strong> <strong>Aralia</strong> fargesii was broad. The type<br />
<strong>of</strong> <strong>Aralia</strong> fargesii was collected near Chengkou <strong>of</strong><br />
the now Chongqing Shi. <strong>Aralia</strong> fargesii has<br />
connate styles to the middle <strong>and</strong> much larger<br />
leaves, <strong>and</strong> more robust inflorescences with more<br />
flowers in each umbel.<br />
<strong>Aralia</strong> yunnanensis is herein treated as a<br />
synonym <strong>of</strong> <strong>Aralia</strong> atropurpurea. At the type<br />
locality <strong>of</strong> A. yunnanensis in Jishan or also known<br />
as Jizhushan, typical A. atropurpurea also occurs.<br />
Many intermediates were found between the two<br />
“species.” In general, plants <strong>of</strong> “A. yunnanensis”<br />
grow at somewhat higher altitudes. Both “species”<br />
can co-occur <strong>and</strong> there is a lack <strong>of</strong> consistent<br />
differences between the two. I thus recognize them<br />
as one species. Both names were published in the<br />
same paper by Franchet in 1896, with A.<br />
atropurpurea on p. 301, <strong>and</strong> A. yunnanensis on p.<br />
303. Also A. yunnanensis was associated with a<br />
species in <strong>Aralia</strong> sect. Pentapanax. I thus prefer<br />
the use <strong>of</strong> <strong>Aralia</strong> atropurpurea, which describes<br />
the <strong>of</strong>ten purple stem <strong>of</strong> the species.<br />
The holotype <strong>of</strong> <strong>Aralia</strong> dumetorum H<strong>and</strong>.-<br />
Mazz. from W bears the h<strong>and</strong>writing <strong>of</strong> H<strong>and</strong>el-<br />
Mazzetti “<strong>Aralia</strong> dumetorum H<strong>and</strong>.-Mazz., sp. n.”<br />
This specimen was collected by E. E. Maire in July<br />
1913. Interestingly, the herbarium label said “fl.<br />
gries - fruits noirs” (flowers greyish <strong>and</strong> fruits<br />
black). The specimen had flowers, which appears<br />
to be collected in July, but no fruits were seen on<br />
the specimen.<br />
Examination <strong>of</strong> the types <strong>of</strong> <strong>Aralia</strong> dumetorum<br />
<strong>and</strong> A. melanocarpa suggests that these two “taxa”<br />
are identical. The type localities <strong>of</strong> both<br />
Eleutherococcus melanocarpa H. Lév. [= <strong>Aralia</strong><br />
melanocarpa (H. Lév.) Lauener] <strong>and</strong> A. dumetorum<br />
were Ma-li-ouan <strong>of</strong> Yunnan. But the type <strong>of</strong><br />
Eleutherococcus melanocarpa was collected in<br />
October in the same year by E. E. Maire when the<br />
plants were bearing fruits, whereas that <strong>of</strong> <strong>Aralia</strong><br />
dumetorum was collected in July. Furthermore,<br />
some E. E. Maire (Edward Ernest Maire)<br />
specimens from Yunnan at P were labeled with the<br />
collector R.-P. Maire. “R.-P. Maire” represented<br />
Révérend Pére Maire because Edward Ernest<br />
Maire was a missionary priest in Yunnan then.<br />
I herein treat both <strong>Aralia</strong> melanocarpa<br />
<strong>and</strong> A. dumetorum as synonyms <strong>of</strong> A. atropurpurea.<br />
Li (1942) cited Wilson 4185 (A) as A. dumetorum,<br />
<strong>and</strong> noted that A. dumetorum possessed terminal
100<br />
paniculate inflorescence. Apparently Li did not<br />
have access to the holotype <strong>of</strong> A. dumetorum during<br />
WWII, when his treatment <strong>of</strong> <strong>Aralia</strong>ceae <strong>of</strong> China<br />
was completed. H<strong>and</strong>el-Mazzetti (1933) described<br />
the inflorescence <strong>of</strong> A. dumetorum as a terminal<br />
panicle, <strong>and</strong> this taxon was described based on one<br />
specimen (the type). The holotype <strong>of</strong> A.<br />
dumetorum actually bears an inflorescence with<br />
three main branches originating from one point,<br />
two <strong>of</strong> them having two umbels <strong>and</strong> one with a<br />
single umbel. This is here interpreted as an<br />
umbellate structure, just like that in A.<br />
atropurpurea, which usually has 2-4 primary<br />
branches originating from one point.<br />
E. E. Maire noted on the label <strong>of</strong> the type<br />
specimens <strong>of</strong> <strong>Aralia</strong> melanocarpa that the flowers<br />
were greenish white <strong>and</strong> fruits black. The holotype<br />
has fruits with purple tints. All <strong>Aralia</strong> atropurpurea<br />
has dark purple fruits.<br />
12. <strong>Aralia</strong> apioides H<strong>and</strong>.-Mazz. — Fig. 34; color<br />
plate 10: A-G.<br />
<strong>Aralia</strong> apioides H<strong>and</strong>.-Mazz., Symb. Sin. 7: 701.<br />
1933. TYPE: China. YUNNAN: in region <strong>of</strong><br />
L<strong>and</strong>sang-djiang (Mekong) <strong>and</strong> Lu-djiang<br />
(Salween) divide, Saoa-lumba, 28 o , 3500-3600<br />
m, 18 Jun 1916, H. H<strong>and</strong>el-Mazzetti 8979<br />
(holotype: W!; isotypes: WU!, 3 sheets).<br />
Perennial herb ca. 1 m tall. Rhizomes thick<br />
<strong>and</strong> horizontally oriented. Tap roots forked.<br />
Leaves 40-55 cm long, 30-40 cm wide, ternately<br />
compound; stipules 0.5-1.5 cm long, 0.4-1.2 cm<br />
wide, triangular to narrowly so; petioles 15-18 cm<br />
long, glabrescent; leaflets 2-4.5 cm long, 1.2-2.2<br />
cm wide, 3-5 per pinnule, membranaceous, ovate,<br />
acuminate at apex, obliquely truncate to sometimes<br />
acute or slightly subcordate at base, irregularly<br />
doubly serrate at margin with relatively deep teeth,<br />
sometimes lobed, lateral veins 6-8, conspicuous<br />
on both sides, adaxial surface green, scabrid,<br />
abaxial surface grayish green, pilose along the<br />
veins <strong>and</strong> veinlets, petiolues 2-11 mm long,<br />
gradually becoming shorter toward the tip <strong>of</strong> the<br />
pinnae, more or less pilose. Inflorescence 28-40<br />
cm long, with a long stalk 19-30 cm long, with 6-<br />
14 primary branches umbellately arranged at the<br />
upper part, sometimes with one or two branches<br />
below the top umbellate structure, slightly pilose,<br />
each primary branch subtended by a lanceolate to<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
linear bract or a reduced simple leaf or a trifoliate<br />
structure, variously sized, each branch with 1-5<br />
umbels, 7-9 cm long, terminal umbels 9-15flowered,<br />
pedicels 7.5-13 mm long, pilose, lateral<br />
umbels 7-10-flowered, pedicels 5-7 mm long,<br />
pilose, bracteoles 1.3-1.5 mm long, 0.4-0.7 mm<br />
wide, lanceolate to narrowly triangular. Floral buds<br />
somewhat globose. Sepals minute, 0.6-0.7 mm<br />
long, 0.4-0.5 mm wide, narrowly triangular to<br />
triangular; petals 1.3-1.5 mm long, 0.7-1 mm wide,<br />
ovate; stamens 1.3-1.6 mm long, filaments 1.1-1.3<br />
mm long, anthers 0.5-0.6 mm long <strong>and</strong> wide,<br />
broadly ovate; gynoecium conical at base, styles<br />
5, 0.8-1 mm long, completely divided after<br />
anthesis, the apical portion <strong>of</strong> the ovary slightly<br />
projected. Fruits 4.5-5.5 mm long, 4-5 mm wide,<br />
globose, styles 1.2-1.5 mm long, distinct <strong>and</strong><br />
persistent, purple.<br />
Common name: qing ye long yan du he.<br />
Phenology: flowering in June-July; fruiting in<br />
July-August.<br />
Distribution: in northwestern Yunnan,<br />
southeastern Tibet, <strong>and</strong> western Sichuan, China<br />
(Fig. 35).<br />
Ecology: common in Picea forests, or mixed<br />
conifer <strong>and</strong> broadleaf forests, shady <strong>and</strong> moist<br />
habitats; 3000-3600 m.<br />
Additional specimens examined: China.<br />
SICHUAN: Baoxin Xian, formerly Mupin, 1954,<br />
T.-P. Soong s.n. (KUN, no. 0563335). Miyaluo,<br />
Xiamuozhigou, 2900 m, 8 Aug 1956, fr, D. P. He<br />
45641 (WUK). Western Szechuan, woods around<br />
Tachiew-lu [Kangting], 8-9,000 ft, E. H. Wilson<br />
982 (BM). Eastern Tibet, Fsukou, R. P. Soulié 1143<br />
(P). YUNNAN: Deqin Xian, Shi-mian-chang,<br />
1981, Picea-Quercus forest, 3200 m, Kunming<br />
Institute <strong>of</strong> Botany Team 130 (KUN); Deqin,<br />
Benzhilan, Yongluobu, in Picea forest, streamside,<br />
common, 3300 m, 6 Jul 1981, Kunming Vegetation<br />
Team s.n. (KUN); Deqin, Qianlelongle, 3240 m,<br />
29 Aug 1937, fr, T. T. Yü 9913 (KUN); Dokerla,<br />
A-tun-tze, 3000 m, ravine, streamside, 3 May 1935,<br />
fl, C. W. Wang 64935 (PE, 2 sheets); Deqin, divide<br />
<strong>of</strong> Chang-Lu rivers, i.e., Lanchangjiang (Mekong)<br />
<strong>and</strong> Lujiang (Salween) divide, near Dokerla, 3000-<br />
3300 m, frequent, herb 3-4 ft, fr black, mixed forest<br />
near stream, 3 Aug 1940, fr, K. M. Feng 5909<br />
(KUN, 3 sheets); Deqin, Benzhilan Commune,<br />
Yongle Teng, 3200 m, in Picea forest, herb 30-50
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
Fig. 34. <strong>Aralia</strong> apioides H<strong>and</strong>.-Mazz. A. Habit with leaves <strong>and</strong> inflorescence. B. Fruit. C. Flower after anthesis. D. Floral bud.<br />
E. Flower. F. Upper part <strong>of</strong> rhizome. G. Seed. H. Umbel (A&F – Yü 22458, A; B & G – Wilson 4158, A; C, D, E & H – Wang<br />
63870, A).<br />
101
102<br />
cm, 3 Jul 1981, fl, Qinghai-Xizang Team 1850<br />
(KUN); on the way from Deqin to Benzhilan, near<br />
122 Daoban, 3700 m, near the edge <strong>of</strong> Picea forest<br />
on slopes, herb 80-100 cm tall, rare, 12 Jul 1981,<br />
fl, Qinghai-Xizang Team 2663 (KUN); Deqin Xian,<br />
Meili Snow Mountains, on the way from Xidan<br />
Hot Spring to the pass, near Bayi, N28 o 24’06.4",<br />
E98 o 49’22.8", 3581 m, 28 Jul 2007, fr, J. Wen et<br />
al. (Tibet-MacArthur Expedition) 1446 (F, KUN,<br />
US). Gongshan Xian, Sewalongba (type locality<br />
<strong>of</strong> <strong>Aralia</strong> apioides), 3400 m, on grassy slope, 25<br />
Aug 1938, fr, T. T. Yü 22458 (KUN, 2 sheets);<br />
divide <strong>of</strong> Chang-Lu rivers, i.e., Lanchangjiang<br />
(Mekong) <strong>and</strong> Lujiang (Salween) divide, near<br />
Cizhong, on the way to Sila, 3000-3300 m, herb 1<br />
m, mixed forests, 17 Jul 1940, young fr, K. M. Feng<br />
5534 (KUN, 2 sheets); on the way from Gongshan<br />
to Dulongjiang, between the 12 th Bridge to<br />
Dongsha<strong>of</strong>ang, in Abies forest on slope, herb 1-<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
Fig. 35. Map <strong>of</strong> China showing the distribution <strong>of</strong> <strong>Aralia</strong> apioides H<strong>and</strong>.-Mazz.<br />
1.5 m, 25 Jul 1982, young fr, Qinghai-Xizang Team<br />
8574 (KUN, 2 sheets); Binzhongle, Yongshongta,<br />
in Tsuga forest on slopes, 3200 m, herb 1.5 m, 25<br />
Jun 1982, fl & young fr, Qinghai-Xizang Team<br />
7597 (KUN, 2 sheets); First District, on the way<br />
from Alulaka to Dengzhong, 2880 m, in valley<br />
under mixed forest, 0.8 m tall, 5 Jun 1960, floral<br />
buds, Nan Shui Bei Diao Team 9245 (PE). Lanping<br />
Xian, Hexi Qu, 1965, Lanping Herbal Medicine<br />
Co. s.n. (KUN). Weixi Xian, 3000 m, Jun 1935,<br />
floral buds, C. W. Wang 63731 (KUN); Weixi, 3500<br />
m, near ravine, Jun 1935, fl, C. W. Wang 63870<br />
(PE, 2 sheets); Weixi, Weideng Commune, from<br />
Xinghua to Haizhi, on slopes in mixed forest, herb<br />
1.5 m, 26 May 1982, floral buds, Qinghai-Xizang<br />
Team 6856 (KUN). Zhongdian Xian [= Shangrila<br />
Xian], Xiao Zhongdian, Jisha, in Picea forest,<br />
common, 14 Jun 1981, Kunming Vegetation Team<br />
s.n. (KUN); Zhongdian, Xundong, 3200 m, 17 Jun
1963, fl, in Picea forest, Yunnan Tropical<br />
Biological Resources Team s.n. (KUN); N flank<br />
<strong>of</strong> Haba Snow Range, plant 3 ft, by stream in<br />
wooded valley in slopes, 22 Jun 1939, fl, K. M.<br />
Feng 1351 (KUN, 2 sheets); Shigao Snow<br />
Mountain, near Dege Cow Farm, in forest, 8 Jul<br />
1939, fl, K. M. Feng 1566 (KUN, 2 sheets, PE);<br />
Wuchun, Lanong, Shita, 3200 m, in Picea forests<br />
near stream, 6 Sep 1959, fr, fruit black, rare, K. M.<br />
Feng 23499 (KUN); Zhongdian, 3200 m, in Picea<br />
forest, 17 Jun 1963, fl, Zhongdian Team 632539<br />
(KUN); Tianbao Snow Range, in forest, 3300 m,<br />
herb 1 m tall, 14 Jun 1981, fl, Qinghai-Xizang Team<br />
1071A (KUN); Xiao Zhongdian, on the way to<br />
Tianchi Lake, N27 o 29’04.5", E99 o 41’44", 3700 m,<br />
24 Jul 2007, fr, herb 1-1.5 m, J. Wen et al. (Tibet-<br />
MacArthur Expedition) 1241 (F, KUN, US); Xiao<br />
Zhongdian, on the road to Tianchi Lake,<br />
N27 o 36’52.6", E99 o 42’45.6", 3354 m, 16 Jun 2009,<br />
fl, J. Wen et al. (Tibet-MacArthur Expedition) 2283<br />
(F, KUN, US); Xiao Zhongdian, on the road to<br />
Tianchi Lake, N27 o 36’52.6", E99 o 42’45.6", 3450<br />
m, 17 Jun 2009, fl, J. Wen et al. (Tibet-MacArthur<br />
Expedition) 2382 (F, KUN, US). XIZANG: Bomi<br />
Xian, on the way from Bomi to Gawalong Lake,<br />
N29 o 50.984’, E95 o 43.476’, 3046 m, 23 Jun 2009,<br />
young fr, J. Wen et al. (Tibet-MacArthur<br />
Expedition) 2680 (F, KUN, US). Chayu [Tsayu]<br />
Xian, Ridong Xiang, in Picea forest on slopes,<br />
3500 m, 18 Sep 1982, fr, black, Qinghai-Xizang<br />
Team 10555 (KUN).<br />
The type specimens <strong>of</strong> <strong>Aralia</strong> apioides have<br />
young leaves with unopened floral buds. One <strong>of</strong><br />
the isotypes has somewhat more mature leaves,<br />
with leaflets up to 3.5 cm long <strong>and</strong> 2.3 cm wide,<br />
<strong>and</strong> margin irregularly doubly serrate.<br />
This species differs from <strong>Aralia</strong> atropurpurea<br />
by its irregularly doubly serrate leaflet margin with<br />
relatively deep teeth or sometimes lobed (vs. finely<br />
doubly serrate at margin in A. atropurpurea).<br />
13. <strong>Aralia</strong> glabra Matsum. — Fig. 36.<br />
<strong>Aralia</strong> glabra Matsum., Bot. Mag. (Tokyo) 12: 17.<br />
1899. TYPE: Japan. Nikko, 4 Oct 1895, fl &<br />
immature fr, Matsumura s.n. (holotype: TI!).<br />
<strong>Aralia</strong> glabra Matsum., Bot. Mag. (Tokyo) 11: 441.<br />
1897. nom. nud.<br />
Glabrous perennial herb 0.6-0.9 m tall. Stem<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
103<br />
purplish green. Rhizome thick. Lower leaves 45-<br />
75 cm long, 30-65 cm wide, tripinnately ternate;<br />
upper leaves 17-31 cm long, 13-22 cm wide, bi- or<br />
tripinnately ternate; stipules 7-10 mm long, 4-5 mm<br />
wide, narrowly triangular, membranaceous, ciliate<br />
at the upper margin; petioles 3-9 cm long; leaflets<br />
3.5-11 cm long, 2-6.5 cm wide, (2) 3-5 (6) per<br />
pinnule, membranaceous, ovate to narrowly ovate,<br />
occasionally broadly so, acuminate to long<br />
acuminate at apex, cordate, obliquely so or rounded<br />
at base, doubly serrate at margin, lateral veins 7-9<br />
on each side, conspicuous on both sides, adaxial<br />
surface green, sparsely scabrid, abaxial surface<br />
grayish green, slightly pilose along the veins <strong>and</strong><br />
veinlets, petiolues 0-2.5 (-4) cm long, gradually<br />
becoming shorter toward the tip <strong>of</strong> the pinnae,<br />
glabrous. Inflorescence 30-48 cm long, loose,<br />
more or less umbellate in outline, glabrous, primary<br />
branches 4-8, with 3-5 primary branches<br />
umbellately arranged at the top, 1-3 scattered along<br />
the main axis, each primary branch 8-25 cm long,<br />
with 1-5 umbels; terminal umbels 15-42-flowered,<br />
pedicels 12-33 mm long, glabrous or nearly so;<br />
lateral umbels 8-14-flowered, pedicels 6-11 mm<br />
long, glabrous or nearly so; bracts subtending the<br />
primary branches 7-9 mm long, 0.9-1.3 mm wide,<br />
lanceolate to linear; bracteoles 1.2-1.7 mm long,<br />
0.8-1.2 mm wide, lanceolate to narrowly triangular.<br />
Flowers dark purple, floral buds somewhat<br />
globose, with several flowers aborted on each<br />
terminal umbel, more so on lateral umbels at<br />
fruiting stage; sepals minute, 0.4-0.6 mm long, 0.5-<br />
0.7 mm wide, triangular to broadly so, spreading<br />
at anthesis, recurved afterwards, persistent into<br />
fruiting stage; petals 1.8-2.0 mm long, 1.2-1.3 mm<br />
wide, ovate, sometimes forming a calyptra <strong>and</strong><br />
falling <strong>of</strong>f at anthesis as a unit; stamens ca. 1.8<br />
mm long, filaments ca. 1.5 mm long; gynoecium<br />
conical at base, styles 5, completely divided after<br />
anthesis, ca. 0.8 mm long, the apical portion <strong>of</strong> the<br />
ovary somewhat projected. Fruits 4.1-4.5 mm long,<br />
4.2-4.5 mm wide, globose, 5-ridged when dry.<br />
Seeds 3.2-3.5 mm long, 1.5-1.6 mm wide, 0.4-0.5<br />
mm thick, kidney-shaped.<br />
Common name: Miyama-udo.<br />
Phenology: flowering in July to October;<br />
fruiting in August to October.<br />
Distribution: in central Honshu, Japan (Fig.<br />
37).<br />
Ecology: in Abies or Tsuga forests, shady
104<br />
habitats; 1430-1810 m.<br />
Additional specimens examined: Japan.<br />
Honshu. Pref. Nagano, Owiki to Denge, 1430-<br />
1470 m, 25 Aug 1961, Chien-Chang Hsu 3190 (TI);<br />
Azumi-mura, Minamiazumi-gun, Abo Pass, 1810<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
Fig. 36. <strong>Aralia</strong> glabra Matrum. A. Habit showing a leaf <strong>and</strong> inflorescences. B. Base <strong>of</strong> petiole showing stipule morphology. C.<br />
Close-up <strong>of</strong> leaflet margin. D. Leaflet <strong>of</strong> a lower leaf. E. Floral bud. F. Flower. G. Flower after anthesis. H. Fruit (A-G –<br />
Murata 16727, US; H – Asano 5060, TI).<br />
m, in Tsuga diversifolia forest, flowers dark purple,<br />
27 Jul 1964, in fl., Kana & T. Yamashita 8963<br />
(MSC, TI). Shinano Prov. Sanpuku Pass, the South<br />
Alps, 6 Aug 1964, K. Asano 5060 (TI). Yamanashi<br />
Pref., Prov. Kai, Seitetsu-machi Nirazaki-shi, Mt.<br />
Hoowoo, half way up from the foot, 1800 m, forest,
20 Aug 1949, young fr, M. Furuse 21403 (K, 2<br />
sheets); Prov. Kai, en route from Sakeishi to Mt.<br />
Daibosatsu, 1800 m, 15 Jul 1962, G. Murata 16727<br />
(US); Mt. Hoozan, 1700 m, 27 Jul 1956, T.<br />
Yamazaki 3740 (A).<br />
<strong>Aralia</strong> glabra is a close relative <strong>of</strong> A. fargesii<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
Fig. 37. Map <strong>of</strong> Japan showing the distribution <strong>of</strong> <strong>Aralia</strong> glabra Matrum.<br />
105<br />
in central China. It differs from the latter by its<br />
longer pedicels <strong>and</strong> its glabrous stems <strong>and</strong><br />
inflorescences. The two species have similar<br />
inflorescence architecture, with most primary<br />
branches umbellately arranged on the upper part<br />
<strong>of</strong> the main axis, <strong>and</strong> 1-3 additional primary<br />
branches at the lower part.
106<br />
14. <strong>Aralia</strong> henryi Harms — Fig. 38; color plate<br />
16: A-B.<br />
<strong>Aralia</strong> henryi Harms, Bot. Jahrb. Syst. 23: 12. 1896<br />
(15 Sep). TYPE: China. HUBEI: Hupeh,<br />
1885-88, in fr, A. Henry 6655 (lectotype, here<br />
designated, K!, isolectotypes: BM!, E!, G!, 3<br />
sheets, GH!, 2 sheets, photo <strong>of</strong> one GH<br />
isolectotype at P, NY!, also its photo with a<br />
packet containing a leaflet at A!, P!, US!, 2<br />
sheets, US Herb. #s 801790 & 801791).<br />
<strong>Aralia</strong> pilosa Franch., J. Bot. (Morot) 10: 302.<br />
1896 (16 Sep). TYPE: China. CHONGQING:<br />
Su-tchuen oriental [now Chongqing Shi],<br />
Tchen-keou-tin [Chengkou], R. P. Farges s.n.<br />
(lectotype: P!, here designated; isolectotypes:<br />
P!, 5 sheets).<br />
Perennial herb 0.25-1 m tall. Rhizomes<br />
horizontal, elongated, with or without internodes.<br />
Stem purplish green, glabrous to pubescent with<br />
long hairs. Leaves ternately compound, decreasing<br />
in size upwards, lower leaves 20-45 cm long, 18-<br />
32 cm wide, upper leaves 6-12 cm long, 6-8 cm<br />
wide, ternately palmately compound; stipules 7-<br />
23 mm long, 1.5-4 mm wide, membranaceous,<br />
glabrous, lanceolate, adnate to the base <strong>of</strong> petiole;<br />
petioles 2-13 cm long, gradually becoming shorter<br />
toward the tip <strong>of</strong> the stem, sparsely pubescent with<br />
long hairs; leaflets 3.5-10 cm long, 2-6 cm wide,<br />
1-3 per pinnule, membranaceous, elliptic or<br />
broadly so, sometimes ovate, acuminate at apex,<br />
cordate, acute or occasionally rounded at base,<br />
doubly serrate at the upper 2/3 <strong>of</strong> the margin, the<br />
lower 1/3 <strong>of</strong> margin mostly serrate, lateral veins<br />
8-10 on each side, conspicuous on both surfaces,<br />
adaxial surface green, sparsely pubescent with long<br />
s<strong>of</strong>t hairs along the veins or glabrescent, abaxial<br />
surface light green, pubescent along the veins <strong>and</strong><br />
veinlets, hairs 0.5-2 mm long, petiolules 0-20 mm<br />
long, pilose. Inflorescence 5-20 cm long, terminal<br />
<strong>and</strong> axillary, relatively small, loose paniculate in<br />
outline, glabrous or nearly so, primary branches<br />
1-3, scattered along the main axis, each primary<br />
branch subtended by a triangular bract, with 1<br />
umbel, 1-3-flowered, pedicels 3-5.5 mm long,<br />
glabrous; bracts subtending primary branches 1.5-<br />
2 mm long, 0.9-1.3 mm wide, narrowly triangular<br />
to lanceolate, glabrous; bracteoles 1.5-1.8 mm<br />
long, 0.5-0.8 mm wide, narrowly triangular,<br />
glabrous. Sepals 0.4-0.5 mm long, 0.35-0.45 mm<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
wide, rounded, spreading at anthesis; petals 1.2-<br />
1.4 mm long, 0.8-1.0 mm wide, ovate; stamens ca.<br />
1.3 mm long; gynoecium 3-5-locular, styles 3-5,<br />
ca. 0.5 mm long, almost divided to the base. Fruits<br />
4.5-5 mm long, 4-4.5 mm wide, red, globose, 5seeded.<br />
Seeds 3.6-4.3 mm long, 2-2.2 mm wide,<br />
0.3-0.4 mm thick, kidney-shaped.<br />
Common name: rou mao long yan du he.<br />
Phenology: flowering in June to July; fruiting<br />
in July to late August.<br />
Distribution: in Anhui, Chongqing, Gansu,<br />
Guizhou, Hubei, Hunan, Shaanxi, Sichuan, <strong>and</strong><br />
Yunnan provinces <strong>of</strong> China (Fig. 39).<br />
Ecology: in mesic forests, shady <strong>and</strong> moist<br />
habitats; 1000-2300 m.<br />
Additional specimens examined: China.<br />
ANHUI: in ravine below Mt. Shu Shin, 1500 m,<br />
18 Jul 1936, P. G. Tsoong 3538 (PE, 2 sheets,<br />
WUK). CHONGQING: Nanchuan, Jishanming,<br />
1750 m, 23 Jul 1957, fl, J. H. Xiong et al. 42236<br />
(WUK); Jingfushan, Mafanzhi, 1500 m, 13 Jul<br />
1986, Jing Fu Shan Team 1741 (PE); Jingfushan,<br />
Dayan, 2000 m, 21 Jul 1957, J. H. Xiong & Z. L.<br />
Zhou 92161 (PE); Jingfushan, Fenghuangshi, 1600<br />
m, 20 Jul 1957, fr, G. F. Li 62967 (PE); Nanchuan,<br />
Jingfushan, below the cable line, along the old trail<br />
toward the north entrance <strong>of</strong> Gufodong,<br />
N29 o 01’56", E107 o 11’06", 2085 m, 4 Sep 2004,<br />
herbs 35-50 cm tall, in bamboo thickets, J. Wen<br />
8196 (US). Wushan, A. Henry 6655A (K). Wulong<br />
Xian, Baimashan, Huangying, 1460 m, 4 Jun 2005,<br />
fl, S.-R. Yi et al. 0234 (PE). Wuxi Xian, 6 Aug<br />
1958, G. H. Yang 59114 (PE, WUK). GANSU:<br />
Bikou Xian, Heidonggou Shui Songpingshan, 1000<br />
m, on the slope in forest, 9 Jul 2006, young fr,<br />
Baishuijiang Expedition 1661 (PE). Wen Xian,<br />
Liujiaping, Qixinggou, Toudaotaizi, in forest, 1756<br />
m, 21 Aug 2006, young fr, Baishuijiang Expedition<br />
2477 (PE). GUIZHOU: Dafang Xian, Baila Qu,<br />
Jiulongshan, 1950 m, 16 Aug 1959, Bi Jie Team<br />
873 (KUN, PE, 2 sheets); Jiulongshan, 1950 m,<br />
16 Aug 1959, fr, Bi Jie Team 862 (PE). HUBEI:<br />
A. Henry 6088 (CAL, K). Badagongshan, near the<br />
Medicine Factory, 5 th branch, moist soil, 1400 m,<br />
herb 80-100 cm, 8 Jul 1958, H. J. Li 3446 (WUK).<br />
Hefeng Xian, Huping Farm, Huangbaipeng, in<br />
forest near streamside, 1300-1400 m, 21 Aug 1958,<br />
fr, purple, common, H. J. Li 5754 (PE, WUK).<br />
Shennongjia Forestry Dist., near Gangqianyan, in
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
Fig. 38. <strong>Aralia</strong> henryi Harms. A. Entire plant. B. Close-up to show leaflet margin <strong>and</strong> pubescence on lower leaf surface. C.<br />
Flower. D. Fruit <strong>and</strong> flowers on two umbels (Henry 6655, E).<br />
107
108<br />
dense broadleaf forest, associated with Dryopteris,<br />
Phellodendron chinense, Chloranthus chinense,<br />
Phoebe, Veratrum, Pilea, <strong>and</strong> Lindera, 1260 m, 14<br />
Jul 1992, young fr, J. Wen 1362 (A, F, OS, US).<br />
HUNAN: Shangzhi Xian, Bamaoxi Commune,<br />
Tianpingshan, Xiaozhuangping, 1650 m, 10 Jul<br />
1975, fr, fruits bright red, B. G. Li & S. B. Wan<br />
750219 (PE). Shunwangshan, 22 Jul 1982, fr, Z.<br />
C. Luo 07 (PE). Xingning Xian, Zhi-yun-shan,<br />
Sep 1984, fr, Zhi Yun Shan Team 360 (PE, 2 sheets),<br />
388 (PE), 920 (PE), 929 (PE), 931 (PE, 2 sheets),<br />
931A (PE), 933 (PE), 962 (PE, 2 sheets).<br />
SHAANXI: Eryang Xian, Guan-shan-liang-gou,<br />
1600 m, 20 Jul 1952, young fr, K. J. Fu 5140 (PE).<br />
Ningshan Xian, Chaizhiping, Banyanggou, 22 Jul<br />
1987, Z. H. Wu 90-808 (WUK). SICHUAN:<br />
Emeishan, 14 Jun 1955, fl, China-Soviet Union<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
Fig. 39. Map <strong>of</strong> China showing the distribution <strong>of</strong> <strong>Aralia</strong> henryi Harms.<br />
Expedition Team 2039 (PE). Hongqi, Lewu, 31<br />
Jul 1959, fr, 2300 m, herb 1 m, Sichuan Economic<br />
Plants Liangshan 1526 (KUN, PE). Leibo, 1934,<br />
T. T. Yü 3977 (PE). YUNNAN: Malipo Xian,<br />
Jinchang, Zhongzhai, Yanyang Cave, herb 0.6 m<br />
high, fruit red, 1800 m, 23 Aug 2004, fr, Y.-M. Shui<br />
et al. 43014 (KUN, US); Malipo Xian,<br />
Xiajingchang Xiang, Zhongzhai Chun,<br />
Hongshuiba, Yanyang Cave (Ram Cave),<br />
N23 o 10’18.1", E104 o 47’47.9", 1895 m, 25 Sep<br />
2008, perennial ca 60-90 cm tall, J. Wen 10640<br />
(US).<br />
The type specimens <strong>of</strong> A. henryi bear flowers,<br />
young fruits, <strong>and</strong> mature fruits on the same plant.<br />
The reproductive biology <strong>of</strong> the species is poorly<br />
known. The flowers appear to be bisexual.
<strong>Aralia</strong> henryi is highly distinctive in <strong>Aralia</strong> in<br />
that it bears red fruits. Other <strong>Aralia</strong> species have<br />
dark purple fruits. Furthermore, its ovaries are 3-<br />
5 locular, rather than 5-locular in the section. I<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
109<br />
considered the option <strong>of</strong> recognizing it as a distinct<br />
section. Nevertheless, it seems most closely allied<br />
with members <strong>of</strong> <strong>Aralia</strong> sect. <strong>Aralia</strong>, <strong>and</strong> I herein<br />
place it in this section.<br />
ARALIA SECT. NANAE HARMS<br />
<strong>Aralia</strong> sect. Nanae Harms, Bot. Jahrb. 23: 12, 1896.<br />
Dioecious herb with long horizontal rhizomes,<br />
acaulescent, with one ternately compound leaf <strong>and</strong><br />
one inflorescence arising from the base, trichome<br />
simple <strong>and</strong> slender, occasionally branching at the<br />
tip. Umbels (2) 3 (7) in a cluster with a long scape<br />
(1018 cm). Flowers 5merous, styles 5, free. Fruits<br />
blackish putple.<br />
One species widely distributed in North<br />
America, especially common in eastern North<br />
America.<br />
15. <strong>Aralia</strong> nudicaulis L. — Fig. 40; color plate<br />
11: A-G.<br />
<strong>Aralia</strong> nudicaulis L., Sp. Pl.: 274. 1753. TYPE:<br />
Clifford herbarium, sheet 113, <strong>Aralia</strong> No. 2<br />
(lectotype: BM!, designated by Wen <strong>and</strong> Reveal<br />
1992).<br />
<strong>Aralia</strong> nudicaulis var. prolifera Apgar, Bull. Torrey<br />
Bot. Club 14: 166. 1887. <strong>Aralia</strong> nudicaulis f.<br />
prolifera (Apgar) Britton, Bull. Torrey Bot.<br />
Club 17: 124. 1890. TYPE: U.S.A. NEW<br />
JERSEY: near Lambertville, 1887, A. G. Apgar<br />
s.n. (holotype: NY!).<br />
<strong>Aralia</strong> nudicaulis var. elongata Nash, Bull. Torrey<br />
Bot. Club 20: 374. 1893. TYPE: U.S.A. NEW<br />
YORK: Greene Co., in the Catskill Mountains,<br />
on the top <strong>of</strong> Cairo Round Top, ca. 2 miles from<br />
Cairo, rocky woods, 11 Jul 1893, fr, with<br />
collector’s notes “several sterile plants seen,<br />
but only one in fruit,” 2000 ft, G. V. Nash s.n.<br />
(holotype, NY!).<br />
<strong>Aralia</strong> nudicaulis f. depauperata Marie-Vict., Nat.<br />
Canad. 71: 206. 1944. TYPE: Canada.<br />
QUÉBEC: Longueuil, comté de Chambly, bois,<br />
11 Jun 1941, F. F. Marie-Victorin & Roll<strong>and</strong>-<br />
Germain 56602 (holotype: MT!; isotypes: GH!,<br />
NY!, US!, all three isotypes are vegetative].<br />
Acaulescent perennial herb, 0.35-0.7 m tall,<br />
highly clonal, dioecious. Rhizome long, branched<br />
<strong>and</strong> horizontal; upright stem 3-10 cm long,<br />
depending upon the age <strong>of</strong> the individual. Leaves<br />
30-60 cm long <strong>and</strong> wide, ternately compound,<br />
usually single or occasionally two arising from the<br />
tip <strong>of</strong> the upright stem (rhizome in essence),<br />
purplish at nodes; stipules absent; petiole 10-45<br />
cm long; leaves usually with 3 (-4) primary<br />
segments, each segment with 4-7 leaflets when<br />
leaves are bipinnately ternate, sometimes leaves<br />
tripinnately ternate; leaflets 6-13 cm long, 4-7.5<br />
cm wide, papery, ovate to oblong, acute to<br />
acuminate at apex, acute to broadly so at base,<br />
commonly oblique on lateral leaflets, doubly<br />
serrate at margin; lateral veins 8-9 on each side,<br />
conspicuous on both surfaces, leaflets green,<br />
glabrous to nearly glabrescent on upper surface,<br />
glabrous to slightly pilose on lower surface,<br />
petiolule 0-1.8 cm long, pilose with long hairs<br />
when young, becoming glabrescent to nearly so.<br />
Inflorescence acaulescent, glabrous, appearing at<br />
the same time as the young leaves, purplish<br />
initially, then becoming green, usually with 3 or<br />
variously 2-7 umbels at the apex <strong>of</strong> a long stalk,<br />
occasionally the peduncle <strong>of</strong> 1-2 umbels shortened<br />
drastically with the flowers clustering at the base<br />
<strong>of</strong> the peduncle <strong>of</strong> the well-developed umbel;<br />
inflorescence stalk 15-30 cm long, glabrous,<br />
peduncle <strong>of</strong> the umbels 2.5-9 cm long, glabrous;<br />
each umbel 25-100-flowered, male umbels usually<br />
having more flowers per umbel; pedicels 9-18 mm<br />
long, slightly pilose with long hairs to glabrous,<br />
tip slightly enlarged at anthesis; bracts at the base<br />
<strong>of</strong> the inflorescence stalk 7-25 mm long, 7-11 mm<br />
wide, usually 3-5 embracing the tip <strong>of</strong> the stem to<br />
protect the buds, ovate to oblong, glabrous; bracts<br />
at the base <strong>of</strong> the peduncle 3-5 mm long, 0.8-1<br />
mm wide, lanceolate, glabrous; bracteoles 0.4-0.8<br />
mm long, 0.25-0.4 mm wide, narrowly triangular,<br />
glabrous <strong>and</strong> caducous. Sepals 0.5-0.6 mm long
110<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
Fig. 40. <strong>Aralia</strong> nudicaulis L. A. Habit with leaf, inflorescence <strong>and</strong> rhizome. B. Young inflorescence appearing at the same time<br />
as the leaf opens. C. Horizonal rhizome, upright rhizome, bract at the base <strong>of</strong> leaf <strong>and</strong> inflorescence. D. Leaflet. E. Leaf<br />
segment. F. Leaflet margin. G. Female umbel. H&I. Umbels with fruits. J. Male floral bud. K. Opening male flower. L.<br />
Male flower. M. Male flower after anthesis. N. Female floral bud. O. Female flower. P. Female flower after anthesis. Q.<br />
Fruit.
<strong>and</strong> wide, triangular, glabrous, persistent on fruits;<br />
petals 2.2-2.8 mm long, 1.1-1.2 mm wide, ovate,<br />
white to purplish white, with a conspicuous<br />
vascular bundle in the middle, spreading to<br />
recurved at anthesis; stamens 5, erect at anthesis,<br />
filaments 2.5-3.5 mm long; anthers 1-1.2 mm long,<br />
0.7-0.8 mm wide, oblong, whitish yellow; ovaries<br />
5- or occasionally 6-locular; styles 1.2-1.3 mm long<br />
at anthesis, completely distinct to the base, base<br />
not projected. Fruits 6-8 mm long, 5-7 mm wide,<br />
globose, purplish black, persistent styles 3-3.5 mm<br />
long, completely distinct to the base. Seeds 4.5-<br />
5.5 mm long, 2.5-3 mm wide, 0.9-1.1 mm thick,<br />
kidney-shaped, whitish gray, smooth.<br />
Common names: wild sarsaparilla, aralia, false<br />
sarsaparilla, American sarsaparilla, shotbush, small<br />
spikenard, wild licorice, rabbitroot, <strong>and</strong><br />
salsepareille (Quebec).<br />
Uses: roots <strong>and</strong> rhizomes used as an alterative,<br />
tonic, or antisyphilitic, <strong>of</strong>ten used by the American<br />
Indians in decoction.<br />
Phenology: flowering from May to July;<br />
fruiting in June to August.<br />
Distribution: widely distributed in North<br />
America from Newfoundl<strong>and</strong> to northern Alberta<br />
<strong>and</strong> north-central British Columbia, southward to<br />
northern Georgia in the eastern range <strong>and</strong> Colorado<br />
<strong>and</strong> eastern Washington in the western range (Fig.<br />
41).<br />
Ecology: moist or dry woods, thickets, riparian<br />
areas, margins <strong>of</strong> prairies, or bog edges; 50-2700<br />
m.<br />
Representative specimens examined: Canada.<br />
ALBERTA: Stettler District, N <strong>of</strong> Battle River,<br />
woods, 3 Jun 1926, fl, A. H. Brinkman 2072 (US).<br />
Rosedale, vicinity <strong>of</strong> Calgary, 30 May & 5 Jun<br />
1914, fl, M. E. Moodie 2 (F); Rosedale, wooded<br />
hill, 30 May 1914, fl, M. E. Moodie 36 (US);<br />
vicinity <strong>of</strong> Rosedale, 2200-2500 ft, 1 Jun 1915, fl,<br />
M. E. Moodie 930 (F, NY, US). S<strong>and</strong> Pt., north<br />
shore <strong>of</strong> Lake Athabaska, about 58 o 56’N,<br />
110 o 42’W, 700 ft, 4 Sep 1932, fr, H. M. Raup & E.<br />
C. Abbe 4561 (NY). District de Jasper-Edson, a<br />
l’ouest du lac Iosegun, Piceetum, 16 Jul 1958, fl,<br />
B. Boivin & J.-M. Perron 12535 (NY). Near<br />
Rockie River, Athabaska Delta, rich woods, 29 Jun<br />
1920, fl, H. M. Laing 102 (US). Jasper, 3472 ft,<br />
Jasper National Park, Aug 1943, sterile, E.<br />
Scamman 3082 (US). Athabasca River, Athabasca<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
111<br />
(Long Rapid), [on the herbarium sheet, this<br />
specimen was labeled from Northwest Territories,<br />
but the locality is in the present-day Alberta], 17<br />
Aug 1903, fr, A. E. Preble & M. Cary 127 (US).<br />
BRITISH COLUMBIA: Liard Hot Springs, woods,<br />
21 Jun 1946, young fr, J. P. Anderson & R. G.<br />
Brown 9881 (US). Vicinity <strong>of</strong> Field, Burgess Trail,<br />
5000-6000 ft, 29 Jun 1906, fl, S. Brown 357 (US).<br />
12 miles east southeast <strong>of</strong> New Hazelton along road<br />
to Smithers, occasional in open, rocky slope, 1100<br />
ft, 15 Jul 1954, young fr, J. A. Calder et al. 13026<br />
(US). About 6 miles SSE <strong>of</strong> Savona along road to<br />
Tunkwa Lake, common in deciduous woods by<br />
creek, flowers white, 2500 ft, 30 May 1956, fl, J.<br />
A. Calder et al. 16828 (US). Along Deadman River<br />
about 18 miles NNW <strong>of</strong> Savona, common in<br />
deciduous woods by river, 2400 ft, 31 May 1956,<br />
fl, J. A. Calder et al. 16865 (US). 5.5 miles on<br />
road from Bridge Lake to Littlefort, common on<br />
grassy slope in open, mixed woods at edge <strong>of</strong> lake,<br />
4700 ft, 13 Jun 1956, fl, J. A. Calder et al. 17415<br />
(NY). Alaska Highway, Mile 270 Alaska Highway<br />
south <strong>of</strong> Ft. Nelson, 58 o 31’N, 122 o 48’W, 15 Jun<br />
1960, fl, J. A. Calder & J. M. Gillett 25404 (US).<br />
Mackenzie District, Fort Simpson, disturbed<br />
ground along road through woods, in shade,<br />
61 o 52’N, 121 o 22’W, 15 Jun 1955, fl, W. J. Cody &<br />
J. M. Matte 8242 (NY). Southeastern British<br />
Columbia, underbrush growth in larch <strong>and</strong> alder<br />
forest 5 miles north <strong>of</strong> Kingsgate, 14 Jul 1948, fr,<br />
E. Y. Dawson 4991 (RSA). Glacier National Park,<br />
Bald Mountain Trail to Grizzly Creek, 51 o 21’N,<br />
117 o 26’W, mossy woods beside bulldozed trail,<br />
3250 ft, 27 Jul 1972, fl, E. Haber & M. J.<br />
Shchepanek 1613 (NY). Vicinity <strong>of</strong> Alaska<br />
Highway, Mile 300, Fort Nelson, 58 o 48’N,<br />
122 o 43’W, common in a Populus tremuloides-Rosa<br />
acicularis forest on hill about 0.5 km north <strong>of</strong> the<br />
Highway, along the Simpson Trail, 458 m, 26 Jun<br />
1959, fl, S. G. Shetler & K. J. Stone 3061 (US). At<br />
mile 497 on the Alaskan Highway at Laird Hot<br />
Springs, north side <strong>of</strong> river, 6 Jun 1975, fl, J. & C.<br />
Taylor 18659 (NY). Alpine slopes <strong>of</strong> Green Mt.,<br />
near Haylmore, mostly 7000 ft, 28 Jul 1938, J. W.<br />
& E. M. Thompson 661 (F, NY). MANITOBA:<br />
District de MacDonald, Coteau de Prairie,<br />
Montagne Pembina, entre Saint-Lupicin et<br />
Roseisle, Reviere la Graisse, bois d’alluvions, 29<br />
Aug 1960, fr, B. Boivin 13927 (NY). NEW<br />
BRUNSWICK: Campobello Isl<strong>and</strong>, Jul-Aug 1888,<br />
fl, J. Donnell Smith 779 (US). Gr<strong>and</strong> Manan, J. T.
112<br />
Rothrock s.n. (F). NEWFOUNDLAND: Valley <strong>of</strong><br />
Exploits River, Gr<strong>and</strong> Falls, dry rocky clearings,<br />
10 Jul 1911, fl, M. L. Fernald & K. M. Wieg<strong>and</strong><br />
5948 (US). Near Topsail, Conception Bay, 12-19<br />
Aug 1901, damp woods, C. D. Howe & W. F. Lang<br />
1307 (NY); Topsail, dry rocky soil, hillside, 19 Aug<br />
1901, C. D. Howe 1338 (F, NY). Frenchman’s<br />
Cove, 19 Jul 1921, fr, K. K. Mackenzie s.n. (NY).<br />
NOVA SCOTIA: Picton Co., Bay View, 6 Aug<br />
1906, fr, C. B. Robinson 221 (NY). ONTARIO:<br />
Vicinity <strong>of</strong> S<strong>and</strong>y Lake, 21 Jul 1957, common <strong>and</strong><br />
widespread through aspen <strong>and</strong> coniferous forest<br />
<strong>of</strong> the area, E end <strong>of</strong> lake, D. R. Moir 4221 (F).<br />
Parry Sound Dist., the Georgian Bay Isl<strong>and</strong>s<br />
opposite Shawanaga Township, Hemlock Isl<strong>and</strong>,<br />
6 Jul 1942, young fr, E. D. McDonald Jr. 18 (US).<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
Fig. 41. Map <strong>of</strong> North America showing the distribution <strong>of</strong> <strong>Aralia</strong> nudicaulis L.<br />
Hemlock-white pine-Thuja occidentalis-tamarack-<br />
Betula lutea swamp along SE side <strong>of</strong> highway 2,<br />
NE <strong>of</strong> Wardsville, 8 Sep 1952, sterile, J. A.<br />
Steyermark 74610 (F). In river bottom woods<br />
under birch <strong>and</strong> balsam, densely shaded, near<br />
mouth <strong>of</strong> Wolf River, 2 mi NE <strong>of</strong> Dorion P. O.,<br />
Dorion Twp., 29 Jul 1963, fr, C. E. Garton 9786<br />
(F). Carleton Co., Ottawa District, Wright’s Cove,<br />
Prescott Hwy, Nepean Twp., W. H. Minshall 106<br />
(NY). Frontenac Co., Kingston, 1897, A. Boyd s.n.<br />
(F); Kingston, Sharbot Lake, 16 Jul 1898, fr, J.<br />
Fowler s.n. (US). Leeds Co., Rideau Park, Oliver’s<br />
Ferry, 19 Jun 1898, young fr, T. W. Edmondson<br />
s.n. (NY). Timiskaming Co., 15 mi NW <strong>of</strong><br />
Kirkl<strong>and</strong> Lake, in mixed second growth forest, 4-<br />
11 Aug 1935, fr, E. C. & T. G. Yuncker 5440 (F,
NY). PRINCE EDWARD ISLAND: Kings Co.,<br />
about 2 miles north <strong>of</strong> Souris, in mape – beech<br />
woods with ferns, 16 Jun 1953, fl, A. J. Smith 80<br />
(NY). QUEBEC: Matamek River Dist., North<br />
Shore, 28 Jun 1927, fl, P. W. Bowman 70 (US).<br />
Longue Pointe de Mingan, 50 o 18’N, 64 o 01’W,<br />
roadsides <strong>and</strong> forests, abundant, 15 Jun 1950, fl,<br />
J. M. Gillett 4985 (US). Woods by Lac Chicobi,<br />
NW <strong>of</strong> Amos, fruits purplish black, 6 Aug 1959,<br />
fr, C. V. Morton 11608 (US). Lac Mistassini,<br />
Territoire de Mistassini, pres de la Baie<br />
Yawatagami (dans la Baie Abatagouche), dans le<br />
bois, 73 o 47’W, 50 o 32’N, 365-415 m, 11 Jul 1944,<br />
fl, J. Rousseau & E. Rouleau 5 (US). Archipel de<br />
Mingan, Ile a la Proie, dans les bois, F. F. Marie-<br />
Victorin & Roll<strong>and</strong>-Germain 18507 (NY). Comté<br />
D’Argenteuil, Saint-Adolphe, forêt d’arbres à<br />
feuilles décidues, Lac Vingt-Sous, 19 Jul 1942, F.<br />
F. Marie-Victorin & Roll<strong>and</strong>-Germain 55285 (NY,<br />
US). Bonaventure Co., Port-Daniel Prov. Preserve,<br />
in woods near river, under Picea-Betula-Populus<br />
forest, abundant with variable leaf <strong>and</strong><br />
inflorescence architecture, 1 Jul 1999, J. Wen 4735<br />
(CS. F, US). Gatineau Co., Wakefield, 1 Jun 1939,<br />
fl, H. A. Senn 1385 (NY). La Haute-Gaspésie<br />
Regional County Municipality, Parc (Quebec) de<br />
la Gaspesie, Mont-Albert, on the way from the<br />
visitor’s center to the top <strong>of</strong> Mont-Albert,<br />
N48 o 56.198’, W66 o 10.376’, 250-1000 m,<br />
abundant, 8 Aug 2007, fl, plant 30-45 cm tall,<br />
rhizomatous perennial, J. Wen 10440 (US). St.-<br />
Jean-Chrysostôme, Comté Lévis, 7 Jun 1966, G.<br />
Lemieux 9837 (BKF). Rimouski Co., Bic, Parc du<br />
Bic, near the Farm House, on hillside woods near<br />
the ocean, in forests <strong>of</strong> Populus-Acer-Abies, 2 Jul<br />
1999, J. Wen 4739 (F, US); Rimouski area, Bic,<br />
Parc du Bic, near Ferme Rioux, in Populus<br />
tremuloides (aspen) forest, N48 o 21.420’,<br />
W68 o 47.808’, 10 m, 9 Aug 2008, fr, herb ca 30-40<br />
cm tall, fruits globose, blackish purple, J. Wen<br />
10466 (US). Sainte-Rose-du-Dégelé, Comté<br />
Témiscouata, 22 Jun 1964, J. L. Blouín et al. 7198<br />
(BKF). Pontiac Co., Onslow Twp., 24 May 1941,<br />
fl, H. A. Senn 1786 (F). Oka, SW <strong>of</strong> Montreal,<br />
Parc d’Oka, in Acer-Quercus-Populus forests, fruit<br />
dark purple to blackish, globose mostly with 5<br />
persistent styles, 5-8 mm in diameter, with<br />
persistent narrowly triangular calyx lobes ca. 1 mm<br />
long, 2 Jul 1999, J. Wen 4743 (F, US).<br />
SASKATCHEWAN: Lake Athabaska, Charlot Pt.,<br />
59 o 36’N, 109 o 13’W, 15 Jun 1935, H. M. Raup 6142<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
113<br />
(F). La Loche, mile 64.5 N Buffalo Narrows – La<br />
Loche Road, 56 o 27’N, 109 o 22’W, under Populus<br />
tremuloides woods with scattered smaller Picea<br />
glauca, common understory herb, 29 Jun 1971, fr,<br />
V. L. Harms 17640 (NY). District de Melfort, Lac<br />
La Ronge, 35 miles au sud, Pinède sablonneuse et<br />
ouverte, 20 Jul 1954, young fr, B. Boivin & D.<br />
Dunbar 10360 (NY). U.S.A. COLORADO:<br />
Boulder Co., N St. Vrain Creek, Rocky Mountain<br />
National Park, 3 Sep 1937, sterile, 8600 ft, R. A.<br />
Schneider 758 (F). Douglas Co., Pike National<br />
Forest, S <strong>of</strong> Sedalia on road to Devil’s Head<br />
campground, 7500 ft, ponderosa pine forest with<br />
some Douglas fir, locally common along stream,<br />
13-14 Aug 1983, fr, D. Keil 17760 (TEX);<br />
occasional in steep wooded ravine <strong>of</strong> West Plum<br />
Creek near the Haystack Ranch (Perry Park South<br />
Ranch), 2 Jul 1981, fl, G. J. Goodman & C. A.<br />
Lawson 8672 (NY); Platte Canõn, H. N. Patterson<br />
37 (F). El Paso Co., vicinity <strong>of</strong> Pikes Peak, 8000<br />
ft, 23 Jun 1896, fl, Bilmore 1301 (US); shaded spot,<br />
Ute Pass Trail, Pike National Forest, foothills, 15<br />
Jun 1980, fl, L. J. Dorr 1604 (TEX); Colorado<br />
Springs, Engelmann’s Canyon, 31 May 1878, fl,<br />
noted as new to Colorado, M. E. Jones 139 (NY,<br />
US). Jefferson Co., Pike National Forest, Tarryall<br />
Mts., Lost Creek Wildness Area, along Goose<br />
Creek in mixed forest <strong>of</strong> Pseudotsuga menziesii<br />
<strong>and</strong> Pinus ponderosa with Alnus, Betula, Acer <strong>and</strong><br />
Cornus along stream, 8200 ft, 5 Jun 2002, fl, T.<br />
Hogan 3835 (NY). Larimer Co., mountains, 20<br />
Aug 1895, fr, C. S. Cr<strong>and</strong>all s.n. (NY); Young’s<br />
Gulch, 8 miles NW <strong>of</strong> Poudre Park, 6500 ft, 15<br />
Jun 1996, J. Wen 2487 (CS, US). Jack Brook, 2700<br />
m, 26 Jun 1901, fr, F. E. & E. S. G. Clements 230<br />
(US). CONNECTICUT: Hartford Co., mixed<br />
mesic woods, junction <strong>of</strong> the new Int. 291 & 84<br />
(between Hartford <strong>and</strong> New Britain), 31 May 1967,<br />
fl, H. E. Ahles 66006 (US). Litchfield Co., moist<br />
woods near Deer Isl<strong>and</strong>, Bantam Lake, Morris, 29<br />
Aug 1919, G. E. Nichols s.n. (YU); Meriden, 2 Jun<br />
1843, fl, B. Livinston s.n. (YU); Middlesex Co.,<br />
Chester, 22 May 1932, fl, R. H. Burton s.n. (YU).<br />
New Haven Co., New Haven, Jun 1856, fl, D. C.<br />
Eaton s.n. (YU); Milford, 26 May 1906, fl, Eames<br />
& Godfrey s.n. (YU); New Haven, 30 May 1884,<br />
fl, W. E. Safford 94 (US); Orange, Maltby Lakes,<br />
21 May 1884, fl, A. L. Winton, Jr. s.n. (YU).<br />
DELAWARE: Nordl<strong>and</strong>s, Jun 1880, W. M. Canby<br />
s.n. (F). DISTRICT OF COLUMBIA: Jun 1893,<br />
E. S. Burgess s.n. (NY); Windy Gap, 17 May 1888,
114<br />
fl, E. S. Burgess s.n. (NY). GEORGIA: Rabun<br />
Co., on north slopes <strong>of</strong> Rabun Bald NW <strong>of</strong> its<br />
summit, in rich loam soil <strong>of</strong> deciduous woods, 3850<br />
ft, 18 May 1947, fl, W. H. Duncan 7560 (NY, US).<br />
IDAHO: Bonner Co., Colburn, in alder swamp, 14<br />
Aug 1941, fr, J. H. Christ 12837 (NY). Kootenai<br />
Co., Lake Peud d’Orville, 28 Jul 1892, fr, J. H.<br />
S<strong>and</strong>berg 772 (US). Hope, in rock slide, 5 May<br />
1914, fl, M. B. Dunkle 402 (NY). Upper Priest<br />
River, 3000 ft, 8 Jul 1925, young fr, C. Epling 7179<br />
(F). ILLINOIS: Glencoe, 30 May 1905, fl, F. C.<br />
Gates 344 (F). Roger’s Park, 20 May 1898, fl, L.<br />
M. Umbach s.n. (F). Chicago, north branch<br />
Chicago River, open woods, M. Bradley 7877 (F).<br />
Cook Co., Jurgensen’s Woods, 183 rd <strong>and</strong> Cottage<br />
Grove, mixed deciduous forest, 1 Jul 1967, fr, S.<br />
F. Glassman 8380 (F). Kankakee Co., in shaded<br />
ground near Rock Creek in Kankakee River State<br />
Park south <strong>of</strong> Deselm, 31 May 1953, fl, F. A. Swink<br />
2083 (F). La Salle Co., wooded hillside, Starved<br />
Rock, 1-7 Jun 1909, fl, J. M. Greenman et al. 32<br />
(F). Ogle Co., Castle Rock, 4 miles SW <strong>of</strong> Oregon,<br />
shaded s<strong>and</strong> at base <strong>of</strong> s<strong>and</strong>stone bluff, 21 Jul 1940,<br />
sterile, R. A. Schneider 1648 (F). Peoria Co.,<br />
Springdale Cemetery, 1866, J. T. Stewart s.n. (F).<br />
INDIANA: Clarke, open woods, 4 Jun 1900, young<br />
fr, O. E. Lansing Jr. 891 (F). Kosciusko Co.,<br />
Mineral Springs, 21 Jul 1928, fr, mixed woods, N.<br />
V. Hayne 958 (F). Lake Co., North Clark Road,<br />
Gary, mesophytic woods, 22 May 1954, fl, R. M.<br />
Eiseman 21 (F). Porter Co., rich acid soil in white<br />
oak - Cornus florida – red maple forest on hills <strong>of</strong><br />
Valparaiso Moraine, along hwy 6, about 1 mi E <strong>of</strong><br />
Woodville junction, 23 May 1946, fl, J. A.<br />
Steyermark 63558 (F). Steuben Co., in low border<br />
<strong>of</strong> Graveyard Lake among tamarack, 28 May 1905,<br />
fl, C. C. Deam s.n. (US). Warren Co., Pine, 15<br />
May 1880, fl, M. Bross s.n. (F). IOWA: Boone<br />
Co., Ledges, 17 Aug 1897, fl, L. H. Pammel & C.<br />
R. Ball 740 (US). Clayton Co., upl<strong>and</strong> woods,<br />
McGregor Heights, 21 Aug 1920, B. Shimek s.n.<br />
(NY). Fayette Co., 15 Jun 1894, fl, B. Fink s.n.<br />
(US). Johnson Co., 4 May 1895, fl, woods,<br />
frequent, T. J. Fitzpatrick s.n. (F). Louisa Co.,<br />
along runs in richly wooded bluff overlooking<br />
south part <strong>of</strong> Lake Odessa, Jefferson Twp., 15 May<br />
1954, fl, R. A. Davidson 1978 (TEX, US).<br />
KENTUCKY: 1840, C. W. Short s.n. (NY). Lewis<br />
Co., Big Sulphur Creek, mesophytic woods, 9 May<br />
1942, fl, E. L. Braun 4508 (US). Rowan Co., Perry<br />
Branch, 21 May 1939, sterile, E. L. Braun 2494<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
(US). MAINE: Cumberl<strong>and</strong> Co., Cape Elizabeth,<br />
May 1895, young fr, E. E. Gayle 714 (US). Knox<br />
Co., Camden Hills State Park, Mt. Battie, 1.5 mi<br />
N <strong>of</strong> Camden, 069 04.27W, 044 13.58N, granite<br />
outcrop, exposed, with Arctostaphylos, Quercus,<br />
Prunus, Myrica, Vaccinium, Juniperus, 14 Jul<br />
1999, fr, W. Hess et al. 8809 (F, 2 sheets, NY).<br />
Penobscot Co., valley <strong>of</strong> Wassataquoik River, from<br />
McLeod’s to City Camp, 16 Jul 1900, young fr, M.<br />
L. Fernald s.n. (US). Washington Co., bank along<br />
Narraguagus River, Cherryfield, 5 Jul 1912, young<br />
fr, S. F. Blake 4041 (LL). MARYLAND: Allegany<br />
Co., dry woods with clearings on Dan’s Rock area<br />
on Conemaugh formation, about 2 miles eastsoutheast<br />
<strong>of</strong> Midl<strong>and</strong>, 12 Sep 1969, sterile, R. M.<br />
Downs 8765 (NCU). Baltimore Co., Gunpowder<br />
Falls State Park, area between end <strong>of</strong> Schroeder<br />
Ave. <strong>and</strong> Perry Hall Rd., Perry Hall, north-facing<br />
slope, shaded rocky habitat, near a river, locally<br />
rare, 22 May 1991, J. Wen 818 (A). Frederick Co.,<br />
Catoctin Mountain Park, moist rocky area near<br />
stream, with Tsuga canadensis, <strong>and</strong> Kalmia<br />
latifolia, 5 May 1973, fl, C. J. Hickey II 276 (NCU).<br />
Garrett Co., Backbone Mountain, 23 Jun 1931, fr,<br />
E. L. Core s.n. (NY); rich damp woods, Garrett<br />
Co., 3 Jul 1882, fl, J. Donnell Smith 194 (US).<br />
MASSACHUSETTS: Bristol Co., South<br />
Dartmouth, Salters Point, 15 Jun 1917, fl, A. King<br />
22 (F). Franklin Co., Northfield Twp., Lachang<br />
Brook just upstream from jct <strong>of</strong> rts. 63 <strong>and</strong> 10 in N<br />
part <strong>of</strong> town, 65 m in elevation, 1 May 1991, J.<br />
Wen et al. 787 (A). Hampshire Co., Haydenville<br />
(Williamsburg), pine woods, Mountain Street<br />
Reservoir, 1 Jun 1975, H. E. Ahles 79975 (BKF, 2<br />
sheets); South Amherst, 26 May 1881, fl, P.<br />
Blatchford s.n. (F); Amherst, 25 May 1880, C. W.<br />
Minott s.n. (US); Mt. Holyoke, 20 Jul 1895, sterile,<br />
A. F. Storms s.n. (US). Plymouth Co., edge <strong>of</strong> dry<br />
woods, Plymouth, 22 Jun 1912, fruits <strong>of</strong>f, S. F.<br />
Blake 3827 (LL). MICHIGAN: Baraga Co., near<br />
Otter Lake area, at the Access site <strong>of</strong> DNR <strong>of</strong> the<br />
Otter River Dam, in maple-Tilia-Fraxinus mixed<br />
deciduous forest, associated with Tilia americana,<br />
Acer sp., <strong>Aralia</strong> racemosa, Corylus cornuta,<br />
Prunus serotina, Prunus sp., Fraxinus sp.,<br />
Osmorhiza claytonii, O. longistylis, Parthenocissus<br />
quinquefolia, Trillium sp., <strong>and</strong> Cornus sp., 27 Jun<br />
1999, J. Wen 4717 (F, US). Clinton Co., Essex<br />
Twp., 2.5 miles SE <strong>of</strong> Maple Rapids, slight northfacing<br />
slope, 20 Jun 1952, sterile, G. W. Parmelee<br />
& P. A. Hyypio 155 (MSC). Crawford Co., SE
side <strong>of</strong> K P Lake, west facing slope, 10 m from<br />
shoreline, mesic northern forest, mixed hardwood<br />
forest, shade, dry loamy s<strong>and</strong>, common, perennial<br />
herb to 4 dm, flowers white with fragrance <strong>of</strong><br />
honey, mostly vegetative, assoc. with Acer<br />
saccharum, Quercus rubra, Pteridium, Vaccinium<br />
spp., Maianthemum, 1 Jun 1992, in fl., E. M.<br />
Chittenden & A. J. Peil 481 (MSC). Delta Co.,<br />
1.3 miles E <strong>of</strong> Isabella, in conifer-mixed hardwood<br />
forest on s<strong>and</strong>, pine needle litter, 19 Jun 1959, in<br />
young fr., W. T. Gillis 2964 (MSC). Houghton Co.,<br />
moist shaded edge <strong>of</strong> logging trail on Limestone<br />
Mountain, 13 Jun 1957, in young fr., P. A. Hyypio<br />
596 (MSC). Ingham Co., thicket along railroad<br />
tracks, white flowers, 6 Jun 1953, in fl. & young<br />
fr., D. H. Norris 72 (MSC). Keweenaw Co., in the<br />
upper peninsula, dry coniferous forest, in Agate<br />
Harbor, 8 Jul 1936, fr, F. J. Hermann 7879 (US).<br />
Mackinac Co., Hiawatha National Forest, near<br />
Three Lakes Campground, in pine-maple-birch<br />
mixed forest, 26 Jun 1999, J. Wen 4704 (CS, F,<br />
US). Mason Co., Grant Twp., southern end <strong>of</strong><br />
vegetation isl<strong>and</strong>, berries black, 6 Aug 1979, in<br />
mature fr., B. T. Hazlett 712 (MSC); 3/4 mile SW<br />
<strong>of</strong> Rock Harbor Lodge, Isle Royle National Park,<br />
moist woods, 22 Jun 1960, in fl., R. A. Janke 3815<br />
(MSC). Oceana Co., in cedar swamp on<br />
Roscommon mucky s<strong>and</strong>, 2 mi E <strong>of</strong> Crystal Valley,<br />
infrequent in cedar swamps <strong>of</strong> the Manistee<br />
National Forest, assc. with Adiantum pedatum, 14<br />
May 1977, fl., T. S. Mustard 319 (MSC).<br />
MINNESOTA: Becker Co., about 2.5 miles E <strong>of</strong><br />
White Earth, 47 o 5’23"N, 95 o 46’54"W, 19 May<br />
1993, fl, W. R. Smith 22148 (F). Clearwater Co.,<br />
Itasca Park, NE <strong>of</strong> campus, 13 Jul 1929, young fr,<br />
M. L. Grant 2769 (US); west shore <strong>of</strong> Lake Mary,<br />
wooded area, 17 Jun 1973, young fr, D. J. Kitz 6<br />
(F). Lake Co., Two Harbors, Jun 1893, fl, E. P.<br />
Sheldon s.n. (US). Lake <strong>of</strong> the Woods Co., mixed<br />
forest, one mile up from the mouth <strong>of</strong> Pine Creek,<br />
Angle Inlet ¾ <strong>of</strong> a mile west <strong>of</strong> the creek, 21 Jul<br />
1939, fr, J. W. & M. F. Moore 11032 (TEX). Saint<br />
Louis Co., Duluth, 11 Jun 1936, fl, O. Lakela 1407<br />
(F, NY, US). MISSOURI: Clark Co., NE-facing<br />
slope along Des Moines River, 0.5-1.5 miles NW<br />
<strong>of</strong> Athens, 30 May 1941, fl, J. A. Steyermark 28689<br />
(F). Pike Co., 0.75-1.5 miles SE <strong>of</strong> Ashburn,<br />
wooded slopes just at base <strong>of</strong> Limestone Bluffs,<br />
13 May 1939, J. A. Steyermark 22371 (F).<br />
MONTANA: Bigfork, 12 Aug 1902, fr, conifer<br />
woods, 3100 ft, H. N. Whitford 210 (F). Flathead<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
115<br />
Lake <strong>and</strong> vicinity, Yellow Bay, 11 Aug 1908, fr, J.<br />
Clemens s.n. (F). Flathead Co., E bank <strong>of</strong> South<br />
Fork <strong>of</strong> Kootenai River, 0.5 mi south <strong>of</strong> Hwy #2,<br />
in rich moist woods, 11 Jul 1965, fr, C. L. Hitchcock<br />
24019 (NY). Glacier National Park, 13 Jul 1914,<br />
young fr, A. S. Hitchcock 12017 (US); Glacier<br />
National Park, vicinity <strong>of</strong> Glacier Hotel<br />
(“Lewis’s”), at the head <strong>of</strong> Lake McDonald, 960-<br />
1050 m, 27 Aug 1919, P. C. St<strong>and</strong>ley 18277 (US);<br />
Glacier National Park, on rich forest floor along<br />
trail to Avalanche Lake, 3500-3800 ft, 7 Jul 1937,<br />
fl, T. G. & E. C. Yuncker 6709 (F, NY). Lake<br />
McDonald, 28 Aug 1903, L. M. Umbach 776 (F,<br />
US). Lake Co., East Shore <strong>of</strong> Flathead Lake, 1<br />
mile south <strong>of</strong> Yellow Bay, in fairly deep woods,<br />
27 Jun 1948, fr, C. L. Hitchcock 17716 (US); small<br />
tributary <strong>of</strong> Goat Creek, birch <strong>and</strong> poplar, 12 Jun<br />
1968, fl, M. Mooar 8003 (NY). NEBRASKA:<br />
Brown Co., Fairfield Creek, 3 May 1890, fl, C.<br />
Rutter s.n. (US). Cheery Co., 9.5 miles east <strong>of</strong><br />
Valentine on gravel hwy 12, plant common at edge<br />
<strong>of</strong> woods along trail at base <strong>of</strong> north-facing<br />
Quercus-Betula bluff, along Niobrara River, 9 Jun<br />
1975, fl, S. P. Churchill 5625 (NY). NEW<br />
HAMPSHIRE: Chesire Co., along SR 123, 1.2<br />
miles S <strong>of</strong> jct with SR 9, hemlock woods, 10 Aug<br />
1986, J. Wen et al. 136 (OS). NEW YORK: Clinton<br />
Co., Clintonville S<strong>and</strong>plain, wooded hillside, 16<br />
Jun 1953, young fr, V. E. Rudd 802 (US), 807 (US).<br />
Dutchess Co., vicinity <strong>of</strong> Clove, rich woods, 18<br />
Aug 1915, P. C. St<strong>and</strong>ley & H. C. Bollman 11875<br />
(US). Madison Co., Pecksport, 9 Jun 1916, fl, W.<br />
R. Maxon 6181 (US). Orange Co., vicinity <strong>of</strong><br />
Highl<strong>and</strong> Falls, in the Hudson Highl<strong>and</strong>s, altitudes<br />
ca. 50 m, 7 Jul 1910, fr, E. A. Mearns 23 (US). St.<br />
Lawrence Co., Waddington, 5 Jun 1914, fl, O. P.<br />
Phelps 721 (US). Sullivan Co., 3.5 mi NE <strong>of</strong><br />
Westbrookville, Bashakill State Wildlife Area, <strong>of</strong>f<br />
NY 209, 074 30.65W, 041 32.25N, 5 Jul 1999,<br />
sterile, W. Hess et al. 8684 (F). Tompkins Co.,<br />
Danby, 6 Aug 1885, fr, F. V. Coville s.n. (US);<br />
woodl<strong>and</strong> 6-mile creek gorge, near Ithaca, 15 May<br />
1941, fl, D. Iseley 1262 (F); Ithaca, path by Baher<br />
Laboratory <strong>of</strong> Chemistry, 4 May 1942, late fl, E.<br />
D. McDonald Jr. s.n. (US). NORTH CAROLINA:<br />
Avery Co., rich woods on slope, just W <strong>of</strong> jct. NC<br />
105 & 185, common, 22 May 1968, J. W. Hardin<br />
13220 (NCSC). Ashe Co., Bluff Mtn., oak-heath<br />
dominated by Quercus borealis, Q. alba,<br />
Rhododendron, Menziesia, Vaccinium <strong>and</strong><br />
Gaylussacia, 25 May 1974, in fl., R. L. Kologiski
116<br />
& C. H. Perino 190 (NCSC). Buncombe Co.,<br />
slopes <strong>of</strong> Craggy Mountain, 25 May 1897, fl,<br />
Biltmore 28a (US). Haywood Co., roadside, some<br />
20 mi from Lake Junaluska, on Max Patch Rd, 17<br />
Jun 1934, fl, H. J. Oosting 34303 (F); near summit<br />
<strong>of</strong> Cold Mtn, 27 Jun 1958, G. S. Ramseur 4655<br />
(NY). Macon Co., moist hemlock forest, Chestnut<br />
Ridge, E <strong>of</strong> Highl<strong>and</strong>s, 4 Aug 1951, R. K. Godfrey<br />
et al. 51656 (NCSC); along Rt US 28, ca 7 miles<br />
SE <strong>of</strong> jct with Rt 441, 29 Apr 2002, fl, J. Wen<br />
6269 (US). Orange Co., wooded slope, 2 miles<br />
SW <strong>of</strong> Hillsboro, 20 May 1961, fl, H. E. Ahles<br />
53987 (NCU). Swain Co., wooded slope, 0.3 miles<br />
south <strong>and</strong> 2.7 miles east <strong>of</strong> Alarka, 6 Jun 1956,<br />
young fr, H. E. Ahles 14228 (NCU). Transylvania<br />
Co., Devil’s Courthouse on the Blue Ridge<br />
Parkway northeast <strong>of</strong> NC 215, heath bald <strong>and</strong> rock<br />
outcrop, 14 Jun 1974, fl, D. E. Boufford et al. 14256<br />
(NCU); near <strong>and</strong> on rocky summit <strong>of</strong> Devil’s<br />
Courthouse, S <strong>of</strong> Sunburst (Lake Logan), 5750 ft,<br />
12 Jun 1955, fl, J. W. Hardin 712a (US). Watauga<br />
Co., Rich Mountain N <strong>of</strong> Boone, 27 May 1991, J.<br />
Wen 849 (A); Rich Mountain area, along Rich<br />
Mountain Rd., 4 miles E <strong>of</strong> jct with Rt 421, 1 Jun<br />
1994, J. Wen 1526 (US); Rich Mountain area, along<br />
Rich Mountain Rd, 5.5 miles from jct with Rt 460,<br />
on the E side <strong>of</strong> Rt 709, near a ravine, 1 Jun 1994,<br />
J. Wen 1535 (US). NORTH DAKOTA: Neche, 23<br />
Jul 1892, Lee 284 (NY). Benson Co., in woods,<br />
Devil’s Lake, 1 Jul 1905, fl, J. Lundell s.n. (US).<br />
Bottineau Co., in woods near Lake Metigoshe,<br />
Turtle Mts., 9 Jun 1949, fl, V. E. Rudd 685 (US).<br />
Ramsey Co., Devils Lake, 1 Jul 1905, fl, in shaded<br />
forests, J. Lunell s.n. (NY). OHIO: Ashtabula Co.,<br />
along Rt 45, near Rock Creek, in Quercus-Acer-<br />
Fraxinus forest, on a dead tree, 26 Jul 1986, J. Wen<br />
100 (OS). Fairfield Co., Sugar Grove, 20 May<br />
1916, sterile, E. C. Leonard 3973 (US); Sugar<br />
Grove, 25 May 1901, fl, C. S. Mead 2628 (F); Ohio<br />
State University Property, Clear Creek, 10 May<br />
1986, fl, J. Wen 83 (OS), 85 (OS); Clear Creek, 25<br />
Sep 1988, J. Wen 626 (OS, US). Highl<strong>and</strong> Co.,<br />
Fort Hill, mesic woods, 31 Aug 1955, E. L. Braun<br />
s.n. (US). Lawrence Co., woods, head <strong>of</strong> Rock<br />
Hollow, 30 Jul 1951, sterile, F. Bartley 1377 (US).<br />
Portage Co., frequent, marshy woods on E side <strong>of</strong><br />
Twp. Rd. 272, 500 ft N <strong>of</strong> the intersection with<br />
Twp. Rd. 275, Hiran Twp., 26 May 1978, fl, B. A.<br />
Andreas 1827 (US). PENNSYLVANIA: vicinity<br />
<strong>of</strong> Philadelphia, 24 May 1892, fl, without leaves,<br />
W. S. Hall s.n. (F). Bradford Co., Kellogg<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
Mountain, 1.5 mile W <strong>of</strong> Kellogg, open rocky oak<br />
woods, near fire tower at 2200 ft, 19 Jun 1942,<br />
young fr, R. W. Bohl 4193 (NY). Clearfield Co.,<br />
Burnside Twp., along Solley Rd., 1.4 miles E <strong>of</strong><br />
its jct with US Hwy 219, 1.4 miles S <strong>of</strong> Burnside,<br />
common on dry rocky slope with mixed hemlockpine<br />
woods, 7 Jun 1997, fl, T. G. Lammers & M. A.<br />
Vincint 9927 (F). Crawford Co., moist woods, Blair<br />
Bridge Rd., near Linesville, 3 May 1943, fl, H. H.<br />
Harrison s.n. (F). Lackawanna Co., on west<br />
slope,<strong>of</strong> Moosic Mts, 1 mi SE <strong>of</strong> Dunmore, 22 May<br />
1946, fl, S. L. Glowenke 5192 (NY). Lancaster<br />
Co., vicinity <strong>of</strong> Smithville, 21 May 1890, fl, J. K.<br />
Small s.n. (F). Lycoming Co., Deer Hole<br />
Mountain, South Williamsport to summit, 200-400<br />
m, 21 Sep 1928, sterile, W. W. Eggelston & J. W.<br />
Kelly 22752 (US). Snyder Co., Selinsgrove, in<br />
thicket along Middle Creek, 16 May 1925, H. N.<br />
Moldenke 2395 (NY). Sullivan Co., Hillsgrove<br />
<strong>and</strong> vicinity, Hoagl<strong>and</strong> Branch, affluent <strong>of</strong> Elk<br />
Creek, 18 Jun 1981, fr, T. Plowman 10536 (F).<br />
Warren Co., Sheffield, in deep woods, 19 Jun 1948,<br />
fl, H. N. Moldenke 19455 (WU). York Co., vicinity<br />
<strong>of</strong> McCalls Ferry, 5-7 Jul 1904, sterile, J. N. Rose<br />
& J. H. Painter 8145 (US). SOUTH DAKOTA:<br />
Deadwood, Rocky hillside, 28 Jul 1913, fr, W. P.<br />
Carr 131 (F, US); Black Hills, Rapid Creek & Dark<br />
Canyon, H. E. Hayward 840 (F); Custer State Park,<br />
head <strong>of</strong> French Creek, 1927, H. E. Hayward 868<br />
(F); canyon W <strong>of</strong> Whitewood, N-facing slope, H.<br />
E. Hayward 1203 (F); Black Hills, Spearfish<br />
Canyon, 1927, H. E. Hayward 1348 (F), 1439 (F);<br />
Black Hills, Harney Peak, E-facing slope, 1927,<br />
H. E. Hayward 1716 (F); Black Hills, Mystic, 23<br />
Aug 1908, fr, S. S. Visher s.n. (F); Savoy,<br />
mesophytic canyon, associated with yellow pine,<br />
spruce, 8 Aug 1926, fr, H. E. Hayward 290 (F);<br />
Savoy, limestone, 5300 ft, 23 Jun 1910, fl, J.<br />
Murdoch Jr. 4143 (F); Terry Peak, H. E. Hayward<br />
1922 (F). Deadwood, near Pinecrest, mesophytic<br />
valley, 5 Aug 1926, H. E. Hayward 240 (F, 2<br />
sheets); 1131 (F); Black Hills, in dense woods, 22<br />
Jul 1910, young fr, A. Nelson 9456 (US); Piedmont<br />
<strong>and</strong> Little Elk Creek, 28 Jun 1892, fl, 4000 ft, P. A.<br />
Rydberg 732 (US); Piedmont <strong>and</strong> Little Elk Creek,<br />
28 Jun 1892, 4000 ft, P. A. Rydberg 733 (US).<br />
Custer Co., Mayo, shady slope, 13 Jun 1914, young<br />
fr, W. H. Over 1790 (US). Lawrence Co., 5 mi W<br />
<strong>of</strong> US 385, Black Hills National Forest, 103<br />
44.09W, 44 14.65N, 2000 m, locally common, 14<br />
Jul 1998, sterile, W. Hess et al. 8260 (F); Spearfish
Canyon, ca. 10 mi SW <strong>of</strong> Spearfish along hwy 14-<br />
A on canyon floor, 4 Aug 1965, fr, J. M. Kane 2481<br />
(LL). VERMONT: Bennington Co., Manchester,<br />
29 Jun 1898, fr, M. A. Day 78 (F, US, 2 sheets);<br />
along SR 9, opposite to the entrance to Woodford<br />
State Park, moist woods, a large population <strong>of</strong> over<br />
1000 individuals, 9 Aug 1986, J. Wen et al. 131<br />
(OS); Woodford State Park, at the edge <strong>of</strong> Quercus-<br />
Larix forest, 9 Aug 1986, J. Wen et al. 133 (OS).<br />
Caledonia Co., Peacham, 7 Jun 1892, fl, A. F.<br />
Stevens s.n. (F). Windham Co., on rocky cliff on<br />
mountainside, Jamaica, 16 May 1937, fl, H. N.<br />
Moldenke 9558 (NY). VIRGINIA: Amherst Co.,<br />
Bluff Mt., 21 Aug 1964, R. S. Freer 2782 (SRSC).<br />
Augusta Co., St. Marys River, upper valley, 30 May<br />
1966, young fr, Freer & Hooks 3856 (NCU). Bath<br />
Co., Flag Rock, in woods, 9 Jun 1930, fl, C. V.<br />
Morton 1903 (US). Botetourt Co., ca 10 miles NE<br />
Catawba in rich bluff woods, 17 May 1960, fl, R.<br />
Kral 10274 (NCU). Floyd Co., Blue Ridge<br />
Parkway, 2.5 mi N jct US 58, E <strong>of</strong> road, 30 Jun<br />
1978, M. J. Warnock 926 (TEX). Giles Co., Salt<br />
Pond Mt, 2 Jun 1890, fl, H. A. Brown et al. s.n.<br />
(NY); near Mountain Lake along Rt 700, 9 Sep<br />
1986, J. Wen et al. 179 (OS); near Mountain Lake<br />
Biological Station, 9 Sep 1986, J. Wen et al. 186<br />
(OS); 5.3 miles from jct. <strong>of</strong> Rt 460 <strong>and</strong> Rt 700 on<br />
Rt 700 toward Mountain Lake, rocky slope, in oak<br />
<strong>and</strong> maple forest, 25 May 1991, J. Wen 833 (A);<br />
Mountain Lake area, along Rt 613, ca 3.2 miles<br />
SE <strong>of</strong> jct with Rt 668, 27 Apr 2002, J. Wen 6234<br />
(US). Grant Co., mixed mesophytic woods with<br />
clearing on northwest slope facing north branch<br />
<strong>of</strong> Potomac River on s<strong>and</strong>y <strong>and</strong> carbonaceous<br />
shales <strong>and</strong> s<strong>and</strong>stones, ca. 1 mile northeast <strong>of</strong><br />
Bayard, 27 Aug 1969, sterile, R. M. Downs 7243<br />
(NCU). Greene Co., Shen<strong>and</strong>oah National Park,<br />
near Pocosin Shelter, in open light woods or bushy<br />
area, 18 May 1940, fl, E. H. Walker 2746 (US).<br />
Rappahannock Co., Shen<strong>and</strong>oah National Park, by<br />
stump in wet woods, 15 May 1938, fl, E. H. Walker<br />
2301 (US). Roanoke Co., 2.2 mi S <strong>of</strong> jct Co. 696<br />
& US 221, 30 May 1968, young fr, F. C. James<br />
10196 (NCU). Rockingham Co., on US 33 in the<br />
George Wahington National Forest about 5 miles<br />
W <strong>of</strong> West Virginia <strong>and</strong> Virginia State line, 22 Jun<br />
1968, young fr, F. C. James 11071 (NCU). Smyth<br />
Co., Pond Mountain, E Marion, 3000 ft, 25 May<br />
1892, J. K. Small s.n. (F, 2 sheets). Warren Co.,<br />
along Skyline Dr. in Shen<strong>and</strong>oah National Park,<br />
ca 1 mile N <strong>of</strong> Hoagwallow Flats Overlook,<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
117<br />
N38o48’07.1", W78o10’40.6", 756 m, 4 Jul 2007,<br />
fr, herb ca 0.5 m tall, fruits dark purple, J. Wen<br />
9840 (US). WASHINGTON: Pend Orielle Co.,<br />
wooded slope <strong>of</strong> Z* Canyon, 25 Jun 1933, young<br />
fr, J. W. Thompson 9207 (NY, US). Stevens Co.,<br />
in dense woods on South Fork <strong>of</strong> Sheep Creek,<br />
near Frontier, 28 May 1939, fl, C. W. Sharsmith<br />
4038 (NY, US). WEST VIRGINIA: Preston Co.,<br />
Masontown, 7 Jun 1940, fl, M. Maysilles 10 (US).<br />
Wetzel Co., Fish Creek, near Littleton, 2 Jun 1931,<br />
young fr, E. L. Core 2666 (NY). WISCONSIN:<br />
Columbia Co., Wisconsin Bells, 31 May 1961, G.<br />
N. Jones 32369 (F). Door Co., Ephraim, rocky<br />
places, Jul 1909, fl, C. F. Millspaugh 3552 (F, 2<br />
sheets). Douglas Co., at the Lower Falls <strong>of</strong> the<br />
Amnicon Falls, N46o36.660’, W91o53.451’, 240<br />
m, 5 Sep 2007, perennial ca 40-50 cm tall, clonal,<br />
J. Wen 9940 (US). Marathon Co., along 5th Ave.,<br />
0.9 mi S <strong>of</strong> jct. with North Lane or 1.9 mi S <strong>of</strong> jct<br />
with County Rt A, associated with Fraxinus<br />
americana, Onoclea sensibilis, Tilia americana,<br />
Quercus rubra, Acer rubrum, Impatiens sp., Rubus<br />
sp., Hydrophyllum virginicum, <strong>and</strong> Ulmus<br />
americana, roadside, 14 Sep 1994, J. Wen 1810<br />
(US). Oconto Co., E side <strong>of</strong> US Hwy 41, 1.5 miles<br />
N <strong>of</strong> its junction with County Hwy J, scattered in<br />
swampy coniferous forest, 10 Jun 1994, young fr,<br />
T. G. Lammers & M. A. Vincent 9018 (F).<br />
WYOMING: Crook Co., Black Hills, 9.5 air miles<br />
E <strong>of</strong> Sundance, ponderosa pine slopes, stream<br />
bottom, <strong>and</strong> meadows, wooded slopes, 9 Jul 1984,<br />
young fr, 4600 ft, R. L. Hartman 17576 (NY).<br />
Sundance Mt, 3 Jul 1896, fl, A. Nelson 2140 (NY,<br />
US).<br />
This is the most widely distributed species <strong>of</strong><br />
<strong>Aralia</strong> in North America. Although it has been<br />
treated as herbaceous (e.g., Smith 1944), it is<br />
woody in essence, because it has a woody<br />
horizontal rhizome <strong>and</strong> a short upright woody stem.<br />
Flanagan <strong>and</strong> Bain (1988) remarked that the plant<br />
behaves like an “underground or buried shrub.”<br />
Most woody species <strong>of</strong> <strong>Aralia</strong> are distributed in<br />
warmer areas. <strong>Aralia</strong> nudicaulis is the<br />
northernmost species <strong>of</strong> the entire genus. Scoggan<br />
<strong>and</strong> Cody (1979) also recorded the species in the<br />
southern Yukon Territory, although the author has<br />
not examined collections there yet. The herbaceous<br />
habit is apparently secondary or derived in the<br />
genus.<br />
The generic name “<strong>Aralia</strong>” initially coined by<br />
Tournefort (1700) is thought to have derived from
118<br />
the Indian common name “aralie” for <strong>Aralia</strong><br />
nudicaulis (Marie-Victorin 1964; Wen <strong>and</strong> Reveal<br />
1992).<br />
<strong>Aralia</strong> nudicaulis is the only species so far well<br />
documented to be dioecious in <strong>Aralia</strong>. Other<br />
species are primarily <strong>and</strong>romonoecious. Barrett<br />
<strong>and</strong> Helenurm (1981) <strong>and</strong> Bawa et al. (1982) also<br />
noted that in rare cases inflorescences <strong>of</strong> the species<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
may contain perfect flowers or both male <strong>and</strong><br />
female flowers. The male <strong>and</strong> female flowers are<br />
dimorphic. The female flowers have five long<br />
styles <strong>and</strong> five short stamens with non-functional<br />
<strong>and</strong> smaller anthers. The male flowers bear five<br />
long stamens <strong>and</strong> five short styles (also see<br />
Flanagan <strong>and</strong> Bain 1988).<br />
ARALIA SECT. HUMILES HARMS<br />
<strong>Aralia</strong> sect. Humiles Harms, Bot. Jahrb. 23: 13. 1896.<br />
Unarmed branched shrub. Leaves pinnate to<br />
bipinnate, stipulate; trichomes usually branched<br />
when present into dendroids, stomates on both<br />
abaxial <strong>and</strong> adaxial leaf surfaces. Umbels 5-15,<br />
loosely clustered into panicles, inflorescence<br />
terminal. Flowers 5 (-6)-merous. Fruits dark<br />
purple.<br />
This section consists <strong>of</strong> three species distributed<br />
in Central <strong>and</strong> southwestern North America.<br />
Key to species <strong>of</strong> <strong>Aralia</strong> sect. Humiles<br />
1. Leaves <strong>and</strong> leaflets pubescent ..................................................................................................... A. humilis<br />
1. Leaves <strong>and</strong> leaflets glabrous .............................................................................................................. 2<br />
2. Leaves bipinnate; inflorescence glabrous ............................................................................... A. regeliana<br />
2. Leaves pinnate; inflorescence pilose; endemic to Baja California ....................................... A. scopulorum<br />
16. <strong>Aralia</strong> humilis Cav. — Figs. 42-44; color plate<br />
12: A-G.<br />
<strong>Aralia</strong> humilis Cav., Icon. 4: 7, t. 313. 1797.<br />
TYPE: cultivated in the Madrid Botanical<br />
Garden from seeds originally collected from<br />
Mexico during the Sessé <strong>and</strong> Mociño<br />
Expedition, yet the type specimen not found in<br />
the Sessé <strong>and</strong> Mociño herbarium, plate 313 in<br />
Icones et descriptions plantarium <strong>of</strong> Cavanilles<br />
(1797) (lectotype, here designated).<br />
<strong>Aralia</strong> pubescens DC., Cat. Pl. Horti Monsp.: 80.<br />
1813. Type: Herb. J. Gay, donne par M.e Dunal<br />
(en hort Monsp.) en Juin 1817 (lectotype: K!,<br />
here designated) [with the h<strong>and</strong> <strong>of</strong> de C<strong>and</strong>olle,<br />
“<strong>Aralia</strong> pubescens Dec<strong>and</strong>.”<br />
<strong>Aralia</strong> scabra C. Presl ex DC., Prodr. 4: 258. 1830.<br />
TYPE: Mexico. Presl s.n. in Herb. Haukearis<br />
(holotype: G; isotype: BM!).<br />
<strong>Aralia</strong> brevifolia Marchal, Bull. Acad. Roy. Sci.<br />
Belgique, II, 47: 74. 1879. TYPE: Mexico.<br />
Merattan, San Andres, Liebmann 33 (holotype:<br />
C, photo at US).<br />
<strong>Aralia</strong> chilapensis Sessé & Moç., Pl. Nov. Hisp.:<br />
48. 1888. TYPE: Mexico. GUERRERO:<br />
Chilapa, Sessé & Mocino watercolor plate <strong>of</strong><br />
‘’<strong>Aralia</strong> chilapensis Sp. N.’’, accession number<br />
6331.0246 in the Torner Collection, Hunt<br />
Institute for Botanical Documentation,<br />
Pittsburgh (lectotype, here designated).<br />
<strong>Aralia</strong> pinnata Sessé & Moç., Pl. Nov. Hisp.: 48.<br />
1888. TYPE: Mexico. JALISCO, Ahuijullo,<br />
flowering in December, type not seen in the<br />
Sessé & Moçiño herbarium.<br />
Pentapanax mexicanus C. B. Shang & X. P. Li, in<br />
Y. W. Yuan & al. (eds.), Proc. Intern. Symp.<br />
Bot. Gard. 1990: 628, t. 2. 1990. <strong>Aralia</strong><br />
mexicana (C. B. Shang & X. P. Li) Frodin in
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
Fig. 42. <strong>Aralia</strong> humilis Cav. A. Habit with leaves <strong>and</strong> inflorescence. B. Close-up <strong>of</strong> lower leaflet surface showing pubescence<br />
<strong>and</strong> teeth at margin. C. Flower. D. Fruit.<br />
D. Frodin & R. Govaerts, World Checklist<br />
Bibliog. <strong>Aralia</strong>ceae 72. 2003. TYPE: Mexico.<br />
JALISCO: lava beds near Zapotlan, a small<br />
tree, 15-20 ft, 13 May 1893, in fr., C. G. Pringle<br />
4366 (holotype: P!; isotypes: BM!, F!, G!, LE!,<br />
MEXU!, MSC!, P[2]!, US!, WU!).<br />
Shrub to small tree 0.5-10 m tall. Stem grayish<br />
119<br />
to dark brown, branchlet terete. Stipule 1.5-2.5<br />
mm long, 0.9-1.5 mm wide, adnate to petiole base,<br />
the free portion triangular to narrowly so, pilose;<br />
Leaves 9-27 cm long, 6-16 cm wide, usually<br />
pinnate in architecture, sometimes bipinnate,<br />
pinnate leaves with (3-) 5-9 leaflets, accessary<br />
leaflets <strong>of</strong>ten present, especially on bipinnate<br />
leaves; petioles 5-9 cm long, pubescent with many
120<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
Fig. 43. <strong>Aralia</strong> humilis Cav. A. Habit with leaves <strong>and</strong> inflructecence. B. Older flower after falling <strong>of</strong>f <strong>of</strong> petals <strong>and</strong> anthers. C.<br />
Flower. D. Floral buds. E. Fruit. F. Infructescence. G. Base <strong>of</strong> leaves <strong>and</strong> inflorescence showing bracts. H. Lower leaflet<br />
surface showing pubescence with dendroid hairs (A & H – Chiang et al. F-2594, F; B-D & G – Carlson 4076, F; E & F –<br />
Pringle 4366, F).
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
Fig. 44. <strong>Aralia</strong> humilis Cav. A. Stem with a bipinnate leaf. B. Stem <strong>and</strong> lower part <strong>of</strong> petiole showing stipules. C. Inflorescence.<br />
D. Floral buds. E. Umbel with flowers at anthesis.<br />
curved white hairs; leaflets 2.8-7.5 cm long, 1.6-<br />
6.5 cm wide, papery, ovate to broadly or narrowly<br />
ovate, acute to acuminate at apex, rounded, broadly<br />
acute to slightly subcordate at base, symmetrical<br />
or sometimes oblique, serrulateto serrate at margin,<br />
tip <strong>of</strong> teeth gl<strong>and</strong>ular, lateral veins 5-6, conspicuous<br />
above <strong>and</strong> below, adaxial surface pilose with white<br />
121<br />
s<strong>of</strong>t dendroid or branched hairs, green to dark<br />
green, abaxial surface usually densely pilose with<br />
branched hairs, light green, petiolule 1.5-16 mm<br />
long, pilose with <strong>of</strong>ten branched hairs.<br />
Inflorescence 12-30 cm long, 8-20 cm wide,<br />
terminal at the branch apex, more or less pubescent<br />
with branched s<strong>of</strong>t hairs, green to purplish green,
122<br />
consisting <strong>of</strong> 5-10 primary branches paniculately<br />
arranged on a main axis; inflorescence bracts 5-7<br />
arranged in several whorls at the base <strong>of</strong><br />
inflorescence, 3-4 mm long, 4-5 mm wide, semipersistent,<br />
triangular to broadly so in shape, dark<br />
brown in color, glabrous or glabrescent; each<br />
primary branch consisting <strong>of</strong> 1-3 umbels, terminal<br />
umbels with 25-45 flowers, lateral umbels 15-20<br />
flowered, with peduncles 3-10 cm long; pedicels<br />
glabrous to pubescent, slightly enlarged at the tip,<br />
pedicels <strong>of</strong> terminal umbels 12-23 mm long, those<br />
<strong>of</strong> lateral umbels 5-8 mm long; bracts <strong>of</strong> primary<br />
branches 2-3.5 mm long, 1-2 mm wide, narrowly<br />
triangular to lanceolate, pilose, <strong>of</strong>ten caducous;<br />
bracteoles 1.8-3.5 mm long, 0.7-1.2 mm wide,<br />
lanceolate, more or less pilose. Sepals 0.4-0.5 mm<br />
long <strong>and</strong> wide, triangular; petals 2-2.2 mm long,<br />
1.3-1.5 mm wide, ovate, greenish white; stamens<br />
5, filaments 2.2-2.5 mm long, anthers 0.8-1 mm<br />
long, 0.6-0.7 mm wide, oblong; styles 0.9-1.1 mm<br />
long at anthesis, divided; floral disk slightly<br />
projected. Fruits 5-6.5 in diameter, globose, dark<br />
purplish black, bloomy, persistent styles 1.5-2 mm<br />
long, divided up to the base <strong>and</strong> recurved, with the<br />
base <strong>of</strong> the style projected <strong>and</strong> exposed between<br />
the persistent sepals <strong>and</strong> the divided part <strong>of</strong> the<br />
styles. Seeds 4-5 mm long, 2-2.2 mm wide, 1.5-<br />
1.8 mm thick, kidney-shaped.<br />
Common names: c<strong>and</strong>elilla (Durango, Mexico);<br />
cuajilotillo (Sinaloa, Mexico); tacamajaca (Jalapa,<br />
Guatemala).<br />
Uses: in Chihuahua, flowers, leaves <strong>and</strong> bark<br />
used as tea for fever, wood for making rattles <strong>and</strong><br />
violin (see R. A. Bye 5996, MEXU).<br />
Phenology: flowering in August to September,<br />
December, January, <strong>and</strong> February; fruiting in<br />
August, late September, October, December,<br />
February, March, May <strong>and</strong> June.<br />
Distribution: widely distributed in Mexico,<br />
Guatemala, Honduras, south to Nicaragua, <strong>and</strong><br />
north to Arizona <strong>of</strong> U.S.A. (Fig. 45).<br />
Ecology: thickets, pine forests, mixed forests,<br />
rocky outcrops, lava beds, scrub woodl<strong>and</strong>, rocky<br />
hill, <strong>and</strong> selva baja caducifolia; (100) 750-2655<br />
m.<br />
Additional specimens examined: Guatemala.<br />
BAJA VERAPAZ: Mun. San Jeronimo Km 137<br />
Carretera La Cumbre-Salama, 1030 m, 24 Jul 1988,<br />
fl, P. Tenorio L. 14778 (BM). CHIQUIMULA:<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
llanos around Ipala, 900 m, dry slopes, 23 Oct<br />
1939, fr, J. A. Steyermark 30322 (F); along Rio<br />
Taco, between Chiquimula <strong>and</strong> Montana Barriol,<br />
3-15 miles northwest <strong>of</strong> Chiquimula, 500-1200 m,<br />
pine slopes, shrub 5-8 ft tall, fruit wine-colored,<br />
26 Oct 1939, fr, J. A. Steyermark 30647 (F).<br />
HUEHUETENANGO: 23 Aug 1896, fl, C. & E.<br />
Seler 3030 (US); along road between San Rafael<br />
Pétzal <strong>and</strong> Colotenango, 1700-1900 m, 14 Aug<br />
1942, fl, shrub 5 ft tall, J. A. Steyermark 50543<br />
(F); along road 13 km west <strong>of</strong> Huehuetenango, near<br />
Puente de Xinaxó, 1800 m, dry steep oak forest,<br />
shrub 1.5-2 m tall, 30 Dec 1940, fr, P. C. St<strong>and</strong>ley<br />
81503 (F); near crossing <strong>of</strong> Río San Juan Ixtán,<br />
east <strong>of</strong> San Fafael Petzal, 1730 m, dry bushy<br />
limestone slope, naked shrub 2 m, fruit black,<br />
shining, 9 Jan 1941, fr, P. C. St<strong>and</strong>ley 82862 (F);<br />
between Nentón <strong>and</strong> Las Palmas, via Yalisjao,<br />
Rincón Chiquite, Chiaquial, Guaxacaná, in Sierra<br />
de los Cuchumatanes, 800-1200 m, 30 Aug 1942,<br />
fl, J. A. Steyermark 51650 (F); thickets <strong>and</strong> forest<br />
in deep canyon <strong>of</strong> a tributary <strong>of</strong> Río Blanco, about<br />
5 km W <strong>of</strong> Aguacatán, 2000 m, 4 Dec 1962, fl, L.<br />
O. Williams et al. 22369 (F, G, US). JALAPA:<br />
vicinity <strong>of</strong> Jalapa, 1360 m, damp thicket, shrub 2<br />
m, 7-18 Nov 1940, fr, P. C. St<strong>and</strong>ley 76513 (F);<br />
rocky scrub-oak forest on hills northeast <strong>of</strong> Jalapa,<br />
1400-1600 m, shrub 1-3 m, occasional, fruit blackpurple,<br />
10 Nov 1940, fr, P. C. St<strong>and</strong>ley 76837 (F);<br />
mountains along road between Jalapa <strong>and</strong> Monjas,<br />
1500 m, oak forest, shrub or small tree, fruit<br />
blackish purple, “tacamajaca,” 11 Nov 1940, fr, P.<br />
C. St<strong>and</strong>ley 76887 (F); west <strong>of</strong> San Pedro Oinula,<br />
1000 m, brushy hillside, shrub 1 m tall, 12 Nov<br />
1940, sterile, P. C. St<strong>and</strong>ley 77119 (F). JUTIAPA:<br />
Lago Retana, between Ovejero <strong>and</strong> Progreso, 600<br />
m, dry rocky slopes <strong>of</strong> escarpment bordering lake,<br />
26 Nov 1939, fr, J. A. Steyermark 32024 (F).<br />
QUÍCHÉ: 1942, J. I. Aquilar 1421 (F). ZACAPA:<br />
lower slopes <strong>of</strong> Sierra de las Minas, along trail<br />
above Río Hondo, 250-900 m, shrub 10 ft tall,<br />
berries green, turning into wine color, leaves light<br />
green, grassy area, 11 Oct 1939, fr, J. A. Steyermark<br />
29555 (F). Honduras. CHULUTECA: fls verdes,<br />
arbusto 4 m, sobre rocas, bosque de Pinos 2 kms<br />
arriba de San Marcos de Colon, 6 Aug 1955, fl,<br />
1100 m, A. Molina R. 5414 (US); San Marcos, in<br />
pine forest area, 1000 m, 16 Nov 1946, fr, L. O.<br />
Williams & A. Molina R. 10889 (F).<br />
COMAYAGUA: pinel<strong>and</strong> 13 mi SE <strong>of</strong><br />
Siguatepeque, 3100 ft, bushy shrub 1.5-2.5 m high,
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
Fig. 45. Map <strong>of</strong> Central America <strong>and</strong> U.S.A. showing distribution <strong>of</strong> <strong>Aralia</strong> humilis Cav.<br />
locally common, buds purplish, flowers greenish,<br />
1 Aug 1962, fl, G. L. Webster et al. 12725 (F, US);<br />
Siguatepeque, 1050 m, rocky hillside, shrub ca. 6<br />
ft tall, 11 Jul 1936, floral buds, T. G. Yuncker et al.<br />
5800 (F, G). COPÁN: shrub 1.5 m, occasional in<br />
thickets along Yaragua creek, 1 mile west <strong>of</strong> Copán<br />
Ruinas, 500 m, 29 Aug 1975, fl, A. Molina R. & A.<br />
R. Molina 30843 (F). EL PARAÍSO: Colectado<br />
en Guinope, 100 m. de altura, 18 Apr 1981, E.<br />
Izaguirre 131 (BM); open savannah, Las Mesas<br />
region near Yuscaran, shrub to 10 ft, Aug 1960, fl,<br />
H. W. Pfeifer 1531 (US, 2 sheets); near Las Casitas,<br />
in pine-oak forest, 900 m, 10 Aug 1947, fl, P. C.<br />
St<strong>and</strong>ley 12067 (F); Quebrada de El Bosque, east<br />
123<br />
<strong>of</strong> Danlí, wet forested quebrada, about 840 m, shrub<br />
2 m, 18 Feb 1949, sterile, P. C. St<strong>and</strong>ley 16765<br />
(F); tree 6 m, in barranco 8 kms west <strong>of</strong> Ojo de<br />
Agua, 900 m, 19 Oct 1946, fr, L. O. Williams & A.<br />
Molina R. 10659 (F); Las Casitas, 900 m, in oakpine<br />
forest, 10 Aug 1947, fl, L. O. Williams & A.<br />
Mollina R. 13236 (F). FRANCISCO MORAZÁN:<br />
20 km N <strong>of</strong> Talanga along the road to Cedros, 800-<br />
1000 m, Pinus – Quercus forest on hills, 4 Oct<br />
1986, G. Davidse & G. E. Pilz 31617 (BM); fls<br />
verdes, planta 0.5-1 m, Matorrales húmedos de<br />
Quebrada Terragra, entre Tatascán y Maraita,<br />
Drainage <strong>of</strong> the Rio Yeguare, at about 87 o W <strong>and</strong><br />
14 o N, 27 Jul 1951, fl, 1200 m, A. Molina R. 4077
124<br />
(F, US); fls cremas, árbol 2-5 m, frecuente,<br />
Barranco en el pinar abierto entre los Kms 11-12<br />
carretera Suyapa a La Montañita, 11 Sep 1963, fl,<br />
A. Molina R. 12849 (F); fls verdes, arbusto 1-1.5<br />
m, poco frecuente, bosque mixto pino-roble entre<br />
Kms 9 y 13 carretera La Montañita y El Zamorano,<br />
11 Sep 1965, fl, 1300 m, A. Molina R. 15228 (F,<br />
US); fls yellowish, weak shrub 2 m tall, common<br />
in pine forest <strong>and</strong> thickets on way to San Antonio<br />
de Oriente, 900 m, 26 Sep 1973, fr, A. Molina R.<br />
& A. R. Molina 27963 (F, US); flowers yellowish,<br />
weak shrub 3 m, in thickets near San Antonio de<br />
Oriente cemetery, 1300 m, 25 Jul 1979, fl, A.<br />
Molina R. 31748 (F); Cerro Gr<strong>and</strong>e, bosque seco<br />
subtropical, 1000 m, 8 Oct 1983, fl, F. José Padilla<br />
16 (BM); along ridge, 1500 m, by trail to San<br />
Antonio de Oriente, shrub to 10 ft in rocks, Aug<br />
1960, H. W. Pfeifer 1753 (RSA, US); Region <strong>of</strong><br />
Agua Amarilla, above El Zamorano, in pine-oak<br />
forest, 780 m, shrub 1 m, fr purple-black, 22-30<br />
Nov 1946, fr, P. C. St<strong>and</strong>ley & L. O. Williams 435<br />
(F); above El Zamorano, road from Jicarito toward<br />
El Pedregal, in pine-oak-region, 875 m, shrub 2<br />
m, scarce, 14 Aug 1947, early floral buds, P. C.<br />
St<strong>and</strong>ley 12252 (F); along Río Caparrosa, above<br />
El Zamorano, 900 m, Sep-Oct 1943, fl, P. C.<br />
St<strong>and</strong>ley 12621 (F); Región <strong>of</strong> Río de la Orilla,<br />
southeast <strong>of</strong> El Zamorano, rocky hillside oak<br />
thicket, shrub 2 m, 900-950 m, 11 Aug 1949, floral<br />
buds, P. C. St<strong>and</strong>ley 22409 (F); Santa Lucia, 10<br />
km al NE de Tegucigalpa, 24 Sep 1983, J. Torres<br />
45 (BM); Santa Inés, 850 m, Aug 1943, fl & young<br />
fr, J. Valerio Rodriguez 480 (F); Santa Inés, 850<br />
m, 4 Nov 1943, fr, J. Valerio Rodriguez 1531 (F);<br />
Jicarito, in pine-oak forest, shrub to 4 m tall, 1000<br />
m, 21 Oct 1946, fr, L. O. Williams & A. Molina R.<br />
10712 (F); drainage <strong>of</strong> the Río Yeguare, at about<br />
longitude 87 o W <strong>and</strong> latitude 14 o N, 1250 m, near<br />
Tatumbla, rocky hillside, shrub 2 m, 19 Aug 1947,<br />
floral buds, L. O. Williams & A. Molina R. 13291<br />
(F). OCOTEPEQUE: weak shrub 2 m, common,<br />
cut over pine-oak forest El Cerro, vicinity <strong>of</strong> San<br />
Antonio, 1300 m, 30 Aug 1968, fl, A. Molina R.<br />
22478 (F). Mexico. CHIAPAS: steep rocky slope<br />
with Quercus along Mexican Highway 190 in the<br />
Zinacantán paraje <strong>of</strong> Multajoc, municipio <strong>of</strong><br />
Ixtapa, 3500 ft, 17 Aug 1965, fl, D. E. Breedlove<br />
11845 (F). Shrubby slope 4 mi SW <strong>of</strong> Bochil along<br />
road to Soyalo, Minicipio <strong>of</strong> Bochil, 4500 ft., 21<br />
Aug 1965, fl, D. E. Breedlove 12085 (F, LL, US);<br />
steep slope with Quercus along Mexican Highway<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
190, 3 mi S <strong>of</strong> La trinitaria, municipio <strong>of</strong> La<br />
Trinitaria, 5100 ft, 14 Oct 1965, D. E. Breedlove<br />
& P. H. Raven 13230 (F); slopes with Pinus <strong>and</strong><br />
Quercus 6-8 km west <strong>of</strong> Teopisca on the side <strong>of</strong><br />
Cerro Chenek’ultik, Municipio <strong>of</strong> Totolapa, 2150<br />
m, 16 Aug 1972, D. E. Breedlove 27076 (LL);<br />
steep-walled canyon at the head <strong>of</strong> the Rio de la<br />
Venta at the Chorreadero near Derna, tropical<br />
deciduous forest, Hauya, Ceiba, Tabebuia, <strong>and</strong><br />
Capparis, Municipio <strong>of</strong> Ocozocoautla de Espinosa,<br />
800-1000 m, 16 Dec 1972, fr, D. E. Breedlove &<br />
R. F. Thorne 30289 (RSA); slope with Pinus,<br />
Quercus <strong>and</strong> Liquidambar near Pueblo Nuevo<br />
Solistahuacán, Municipio <strong>of</strong> Pueblo Nuevo<br />
Solistahuacán, 5800 feet, 22 Aug 1967, fr, O. F.<br />
Clarke 427 (RSA); Municipio la Trinitaria, 6-7 km<br />
S <strong>of</strong> La Trinitaria, 1400-1500 m, rocky limestone<br />
hills with open deciduous forest, treelet 3 m tall,<br />
leaves bipinnate, 17 Nov 1984, fr, G. Davidse, M.<br />
Sousa, O. Téllez, E. Martínez & J. Davidse 29973<br />
(BM, MO); along road between Teneapa <strong>and</strong><br />
Yajalon, 3000-5000 ft, 13 Oct 1895, fr, E. W.<br />
Nelson 3237 (US); between San Cristobal <strong>and</strong><br />
Teopisca, 6700-8500 ft, 4 Dec 1895, fr, E. W.<br />
Nelson 3451 (US); Hacienda Monserrate, Sep<br />
1923, C. A. Purpus 9160 (BM, US), 9232 (F); slope<br />
with Quercus <strong>and</strong> Pinus along the creek “Chenek’<br />
Ha” near Amatenango, Municipio <strong>of</strong> Amatenango<br />
del Valle, 5700 ft, 28 Jul 1967, fr, A. Shilom T.<br />
2692 (F); steep slope with Quercus, Pinus,<br />
Liquidambar, Podocarpus <strong>and</strong> Magnolia along the<br />
ridge above Pueblo Nuevo Solistahuacán,<br />
municipio <strong>of</strong> Pueblo Nuevo Solistahuacán, 6500<br />
ft, 15 Aug 1967, fl, A. Shilom T. 2889 (F); 3 mi W<br />
<strong>of</strong> Pueblo Nuevo Solistahuacán, 9 Sep 1963, fl, R.<br />
A. Marin 88 (F); Chiapas, along Rt 195, 5 km N <strong>of</strong><br />
Soyalo, Mpio. Bochil, 1483 m, 16 o 54.27’N,<br />
92 o 55.54’W, shrubs ca 1 m tall on shrubby slope,<br />
leaves pinnate to bipinnate, 15 May 2006, fl, J.<br />
Wen & E. Martinez 8707 (US). CHIHUAHUA:<br />
Mpio. De Batopilas, Barranca de Batopilas,<br />
vicinity <strong>of</strong> La Bufa, Arroyo Bakosiachi, Nov 1973,<br />
young fr, R. A. Bye 5952 (NCU); Sierra Charuco,<br />
Rio Fuerte, 22 Jul 1935, fr, upper Sonoran, canyon,<br />
H. S. Gentry 1516 (F); Guayanopa Canyon, Sierra<br />
Madre Mts., 24 Sep 1903, fl, 3600 ft, M. E. Jones<br />
s.n. (US); Majarachic, 7 Sep 1939, floral buds, I.<br />
W. Knobloch 5802 (MSC); Rio Bonito, Dec 1936,<br />
fl, H. LeSueur 1153 (F), 1154 (F); C<strong>and</strong>amena<br />
Barranca transect on road from Cruz Verde to<br />
C<strong>and</strong>amenaRiver, 28 o 06’40"N to 28 o 20’30"N,
108 o 17’W, 5500-3250 feet, from pine-oak woods<br />
at 7000 feet to thorn forest & riparian Platanus-<br />
Sassafridium assn. at 3250 feet, 18 Mar 1986, fr,<br />
but leafless, P. S. Martin et al. 78 (RSA); Mpio.<br />
De Madera, Rio Sirupa, 44 km al Se de Madera,<br />
29 Sep 1982, young fr, P. Tenorio L. 1881 (MEXU);<br />
woods <strong>of</strong> Acacia, Quercus, Garrya, Juniperus et<br />
al., on limestone hills 3 mi NW <strong>of</strong> Comitan, 5700<br />
ft, 10 aug 1962, fl, G. L. Webster et al. 12906 (MO).<br />
COLIMA: Desviación cerca Río Salado, 5 Dec<br />
1959, fl, F. Mir<strong>and</strong>a 9068 (MEXU). DURANGO:<br />
Mpio. El Mezquital, camino de El Mezquital a<br />
Temoaya, 193\0 m, 22 Jan 1987, fl, M. Gonzalez<br />
& S. Acevedo 2163 (MEXU); Sianori, 1924, J. G.<br />
Ortega 5388 (US); San Ramon, 21 Apr – 18 May<br />
1906, fr, E. Palmer 110 (F). GUANAJUATO: el<br />
Baral Blanco, ladera S, municipio de Tarimoro,<br />
orilla de arroyo con vegetacion de matorrales<br />
secundarios, 2300 m, 25 May 1987, fr, H. Díaz-<br />
Barriga 3739 (F). GUERRERO: Llano, 4 m high,<br />
local, Manchon, Mina, 28 Sep 1936, in fl., G. B.<br />
Hinton et al. 9605 (P, RSA, US, 2 sheets); Montes<br />
de Oca, Vallecites, oak woods, 2 m shrub, 30 Sep<br />
1937, fr, G. B. Hinton 11440 (RSA, US); Arroyo,<br />
tree 3 m, fls white, Manchon, Mina, Dec 1937, fr,<br />
but leafless, G. B. Hinton et al. 11295 (RSA, US, 2<br />
sheets); Apaxtla, Sep 1926, late fl, B. P. Reko 4977<br />
(US). JALISCO: Salcillo, 1 Jun 1892, fr, like<br />
elder, a bush, M. E. Jones 25 (US); bluffs <strong>of</strong><br />
Barranca, 20 May 1891, fr, C. G. Pringle 5144<br />
(US); barranca <strong>of</strong> Guadalajara, 28 Sep 1891, sterile,<br />
C. G. Pringle s.n. (US); on road between Bolaños<br />
<strong>and</strong> Guadalajara, 21 Sep 1897, late fl, J. N. Rose<br />
3049 (US); Guadalajara, shrub or small tree<br />
growing on or among rocks, hillsides near the Rio<br />
Gr<strong>and</strong>e de Santiago, 27 km E <strong>of</strong> Guadalajara, 5200<br />
ft, 26 Aug 1941, sterile, W. & M. Leavenworth 1898<br />
(F). MEXICO: District <strong>of</strong> Temascaltepec, Puerto<br />
Salitre, 1300 m, 20 Sep 1932, fl, 3 m high, G. B.<br />
Hinton 1789 (G); Tenayac, Temascaltepec, 1500<br />
m, 15 Aug 1933, fl, rocky hill, 3.5 m high, 1500<br />
m, G. B. Hinton 4430 (BM, F); Tenayac, 1520 m,<br />
24 Nov 1933, fr, G. B. Hinton 5141 (F).<br />
MICHOACAN: oeste de Santa Gertrudis,<br />
municipio de Zacapu, matorral pedregoso, ladera<br />
de cerro, 2000 m, arbusto de 2.5 m de alto, 18 Jan<br />
1989, fl & young fr, no leaves, A. Grimaldo N. 511<br />
(F)). MORELOS: Xochitepec, Nov 1934, fr, J.<br />
Elcoro 1152 (US); Sep 1935, fl, J. Elcoro 1153<br />
(US); hills, Cuernavaca, 5000 ft, Oct – Nov 1895,<br />
in fr., C. G. Pringle 6237 (BM, F, G, LE, MSC, P, 2<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
125<br />
sheets, US, 2 sheets, WU). OAXACA: San Felipe,<br />
village north <strong>of</strong> Oaxaca, valley <strong>of</strong> Rio San Felipe<br />
(also called Toma) on slope <strong>of</strong> San Felipe<br />
Mountain, 5700-7200 ft, 7 Mar 1949, fr, no leaves,<br />
M. C. Carlson 1355 (F); Cerro San Felipe, 2100<br />
m, 7 Mar 1898, fl, C. Couzatti & V. Gouzaley 674<br />
(US); vicinity <strong>of</strong> Cerro Zempoaltepetl, along trail<br />
from Tlahuitoltepec to Santo Domingo Albarradas,<br />
18-25 km west-southwest <strong>of</strong> summit, upright<br />
spreading shrub to 5 m, in lowl<strong>and</strong> Bursera scrub<br />
woodl<strong>and</strong>, 1400 m, 16 Aug 1950, fl, B. Hallberg<br />
984 (US); Oaxaca, 2000 m, Oct 1902, fr, Herrerá<br />
s.n. (G); Distrito Teposcolula, Mpio. Yolomecatl,<br />
4.5 km de Yolomecatl, sobre la carretera de<br />
terraceria a Nican<strong>and</strong>uta, 17 o 29’44.8", 97 o 37’0.5",<br />
2250 m, plant highly pubescent, bosque de Pinus,<br />
Quercus y Juniperus, 10 Apr 2003, fr, J. Ismael C.<br />
23891 (MEXU); valley <strong>of</strong> Oaxaca, 6000 ft, 20 Sep<br />
1894, fl, E. W. Nelson 1435 (US); hills near<br />
Tamazulapam, 7800 ft, 13 Nov 1894, sterile, E. W.<br />
Nelson 1952 (US, 2 sheets); hills near Oaxaca,<br />
6000 ft., 3 Dec 1895, in fl & young fr, leaves<br />
bipinnate, 5-8 ft, C. G. Pringle 6173 (F, G, P, RSA,<br />
US, 2 sheets, WU); Monte Alban, near Oaxaca City,<br />
5500-6000 ft, 23 Oct 1894, young fr, C. L. Smith<br />
898 (US); Oaxaca, San Felipe, Toma de Agua, 1729<br />
m, 17 o 6.916’N, 96 o 42.628’W, in remnant forest <strong>of</strong><br />
Pinus, Taxodium, Anona, Ipomoea, <strong>and</strong> Alnus, 13<br />
May 2006, J. Wen & E. Martínez 8679 (US, 2<br />
sheets). PUEBLA: approx. 4 km al S de<br />
Xochiltepec, 7 km al N de San Luis Atolotitlán,<br />
arbol de 2.5 m, frutos morados, 23 Mar 1982, fr, F.<br />
Chiang C. 2291 (MEXU, RSA); 6 km al E de San<br />
Luis Atolotitlán, por la terraceria rumbo a Caltepec,<br />
18 11’N, 97 25’W, 2200 m, cerro pedregoso rojizo,<br />
con Senecio praecox, Ipomoea murucoides, etc, 8<br />
Jun 1985, fr, F. Chiang et al. F-2594 (F, G); Cerros<br />
calizos al NE Tehuacán, 1750 m, 21 Feb 1986, fr,<br />
no leaves, infl. pubescent, A. Salinas T. et al. F-<br />
3144 (MEXU, MO). SINALOA: Mpio. Sinaloa<br />
de Leyva ceros de alrededor de la finca de adobe y<br />
los laurels, 1350 m, 26 Feb 1988, fl, G. Bojórquez<br />
& H. Aguiar 545 (MEXU); Balboa, Jan 1923,<br />
young fr, J. G. Ortega 5051 (US); Mpio.<br />
Badiraguato a 56 km de Badiraguato rumbo a<br />
Surutato, bosque tropical caducifolio, 1000-1200<br />
m, 8 Dec 1987, fr, R. Vega et al. 2645 (MEXU).<br />
SONORA: San Javier, a 1 km de la Carr. Fed. 16,<br />
28 o 34’48"N, 109 o 44’42", 750 m, 12 May 1993, fl,<br />
A. Búúrquez 93-111 (MEXU); San Javier,<br />
Microondas en el Cerro El Durazno, 28 o 36’43",
126<br />
109 o 45’25", 1075 m, A. Búúrquez & D. Yetman 95-<br />
213 (MEXU); Sierra de Alamos, Canon La Huerta<br />
(eastern tributary <strong>of</strong> Canon Agua Escondida),<br />
26 o 59’N, 108 o 59’W, 1300 m, growing in canyon<br />
bottom in riparian forest with Eysenhardtia,<br />
Lysiloma watsoni, Montanoa rosei, Guardiola<br />
platyphylla, <strong>and</strong> Quercus tuberculata, 8 m tall tree<br />
with gray fissured bark <strong>and</strong> greensish perianth,<br />
uncommon, 19 Mar 1994, fl, M. Fishbein et al.<br />
1617 (MEXU); 5.5 mi E <strong>of</strong> Nacozari International<br />
Airport, 30 o 28’N, 109 o 29’W, rock outcrop,<br />
Quercus-Arctostaphylos, 5000 ft, L. Nash & E.<br />
Lehto L19400 (ASU); vicinity <strong>of</strong> Alamos, high up<br />
in Sierra de Alamos, 19 Mar 1910, young fr & fl,<br />
J. N. Rose et al. 13097 (US); Cerro del Capulin,<br />
northwest <strong>of</strong> Aribabi, 6100 ft, 4 Sep 1939, floral<br />
buds, S. S. White 2728 (MEXU); La Mina Verde<br />
31 kilometers de Cumpas, 23 Sep 1934, fr & fl, I.<br />
L. Wiggins 7415 (US); Sonora, locality unknown,<br />
1890, C. V. Hartman 143 (US). VERACRUZ:<br />
Maltrata, Jan 1883, E. Kerber 266 (LE, US).<br />
Mexico locality unknown, May 1842, Pl. Mexic.<br />
Liebm. 1270, <strong>Aralia</strong>ceae n. 32, leaves very<br />
pubescent, C. L. Felipe 1270 (US); Mexico,<br />
Chapuleo, Dec 1841, fl & young fr, leafless,<br />
Liebmann 12135 (F). Nicaragua. ESTELI: N<br />
slope <strong>of</strong> Cerro Tamabú, 13 o 02’N, 86 o 17’W, 1200-<br />
1400 m, roadside pasture <strong>and</strong> grazed oak forest,<br />
with a few scattered pines above, 16 Oct 1979, W.<br />
D. Stevens 14925 (BM). JINOTEGA: southwest<br />
<strong>of</strong> Jinotega, along road to La Cantera <strong>and</strong> Los<br />
Pinos, in region <strong>of</strong> pine forest, 1050-1350 m, moist<br />
thicket, shrub 2 m, scarce, 25 Jun 1947, sterile, P.<br />
C. St<strong>and</strong>ley 10160 (F); ca. 4.7 km S <strong>of</strong> Jinotega<br />
(square) on old road to Hwy 3 near Matagalpa,<br />
then along road to city dump <strong>and</strong> ab<strong>and</strong>oned mine,<br />
13 o 02’N, 86 o 00’W, 1150-1250 m, open pine-oak<br />
forest on steep rocky slopes, 25 Aug 1978, W. D.<br />
Stevens 10139 (BM). U.S.A. ARIZONA: Cochise<br />
Co., Dragoon Mts., Stronghold Canyon East along<br />
USFS Trail 279, 5000-5200 ft, dry riparian zone<br />
in mixed oak woodl<strong>and</strong> <strong>and</strong> oak chaparral on<br />
gravelly granite slopes, with Quercus, Garrya,<br />
Cupressus, Pinus, Arbutus, <strong>and</strong> Juniperus,<br />
subshrub to 5 dm, infrequent, 7 Sep 1983, fl, T. F.<br />
Daniel 2993 (ASU); Mule Mountains, west side,<br />
14 Sep 1961, fl, L. N. Goodding 258-61 (NCU);<br />
Dragoon Mountains, Stronghold Canyon East, 1/6<br />
mi up first major canyon coming into Stronghold<br />
Canyon from FS Road, just above first waterfall,<br />
1540 m, boulderly canyon, local, rare, shrub 4 ft<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
tall, berries maroon-purple, juicy, 8 Nov 1991,<br />
mature fr, W. Hodgson 6433 (ASU, DES); Dragoon<br />
Mts., Cochise Stronghold, 5000 ft, 18 Sep 1936,<br />
late fl, S. W. Hutchinson 7321 (CAS, RSA);<br />
Huachuca Mts., 4 Sep 1903, 7000 ft, M. E. Jones<br />
s.n. (NY); Texas Canyon, decomposing granite,<br />
5200 ft, E. Lehto 2154 (ASU, NCU); Huachuca<br />
Mts., Aug 1882, Lemmon Herb. 2616 (G), Lemmon<br />
Herb. 2716 (BM, F, G, LE, P, US); southern<br />
Huachuca Mountains, top <strong>of</strong> Montezuma Peak, oak<br />
<strong>and</strong> pinyon, 7000-7600 ft, 16 Jul 1990, sterile, B.<br />
D. Parfitt 4387 (ASU); southern Huachuca<br />
Mountains, canyon above Yaqui Spring, 6000-6300<br />
ft, 16 Sep 1990, fr, B. D. Parfitt & C. M. Christy<br />
4696 (ASU); 16 Sep 1990, fl, B. D. Parfitt & C.<br />
M. Christy 4698 (ASU); Mule Mountains, Banning<br />
Creek Canyon, ca. 2.5 miles northwest <strong>of</strong> the tunnel<br />
at Bisbee on hwy 80, O. F. Clarke property on the<br />
steep rocky slopes below Juniper Flats, oak<br />
woodl<strong>and</strong> with Quercus emoryi, Q. arizonica,<br />
Agave parryi, Dasylirion, Eragrostis, Andropogon,<br />
etc, 5600 ft, 31 Aug 1984, fl, A. C. S<strong>and</strong>ers et al.<br />
5195 (TEX); Texas Canyon, ca. 70 miles E <strong>of</strong><br />
Tucson on I-70, dry habitat with large boulders,<br />
mixed with Quercus sp., 18 Sep 1999, young fr,<br />
shrub ca. 3 m tall, J. Wen 4974 (CS, F, US); near<br />
Fort Huachuca, Aug 1894, fr, T. E. Wilcox 309<br />
(US). Pima Co., shady canyons, W slope <strong>of</strong><br />
Baboquivari Peak, 30 Sep 1944, fl, O. M. Clark<br />
12555 (NY); Toro Canyon, Baboquivari Mts., 29<br />
Aug 1931, fl, M. F. Gilman 39 (NY); on shaded<br />
slope in canyon about 1 mile above guest house,<br />
western side <strong>of</strong> Mt. Baboquivari, 4000 ft, 6 Oct<br />
1944, fl, F. W. Gould et al. 2681 (NY, US);<br />
Sycamore Canyon, Baboquivari, 22 Oct 1945,<br />
young fr, L. N. Gooding 230-45 (NY); Baboquivari<br />
Mts., 19 Sep 1931, M. E. Jones s.n. (BM, DS,<br />
RSA); Baboquivari Mts., Thomas Canyon near<br />
Max Seep & Broken Trough Spring, rare on shaded<br />
slope above canyon bottom, 5200 ft, 22 Sep 1982,<br />
fl, M. Mittleman 550 (ASU); Stone Cabin Canyon,<br />
Santa Rita Mountains, 5000 ft, 12 Sep 1903, fr, J.<br />
J. Thornber 182 (ASU, NY). Santa Cruz Co., on<br />
shaded rocky slopes, 4000 ft, Sycamore Canyon,<br />
near Ruby, 30 Sep 1944, fl, R. A. Darrow & H. S.<br />
Haskell 2023 (NY); Santa Rita Mts., 25 Aug 1903,<br />
4500 ft, M. E. Jones s.n. (BM, NY); AZ 289 S <strong>of</strong><br />
Pena Blanca Rec. area, among big whitish<br />
boulders, grassl<strong>and</strong> with Quercus emoryi, Q.<br />
oblongifolia, Arctostaphylos pungens, Rhus<br />
choriophylla, <strong>and</strong> Mimosa dysocarpa, 8 Sep 1976,
fl, L. McGill & E. Lehto L20373 (ASU, NY, US).<br />
Smith (1944) indicated that <strong>Aralia</strong> humilis<br />
occurs in southern New Mexico <strong>and</strong> Arizona to<br />
Guatemala. Frodin <strong>and</strong> Govaerts (2003) indicates<br />
the distribution from southern New Mexico,<br />
southern Arizona, Mexico to C. America. So far I<br />
have not seen any collections from New Mexico<br />
<strong>and</strong> the species was not recorded in floras <strong>of</strong> New<br />
Mexico by Wooten <strong>and</strong> St<strong>and</strong>ley (1915) <strong>and</strong> Martin<br />
<strong>and</strong> Hutchins (1980). Within the U.S., the species<br />
appears to occur only in Arizona.<br />
There seems to be no specimens in the Sessé &<br />
Mociño herbarium corresponding to the original<br />
material <strong>of</strong> <strong>Aralia</strong> chilapensis. The watercolor<br />
plate <strong>of</strong> ‘’<strong>Aralia</strong> chilapensis Sp. N.’’, accession<br />
number 6331.0246 in the Torner Collection <strong>of</strong> Hunt<br />
Institute for Botanical Documentation, Pittsburgh<br />
is herein selected as the lectotype. The Torner plate<br />
bears the number “346” near the top (also see Field<br />
Museum negative 30658, designated as “S. + M.<br />
Pl. 346”). A print <strong>of</strong> 30658 is now at the US<br />
National Herbarium with the courtesy from the<br />
Field Museum Herbarium.<br />
Pentapanax mexicanus C. B. Shang & X. P. Li<br />
was described based on Pringle 4366 collected<br />
from Zapotlan, Jalisco, Mexico (Xiang <strong>and</strong> Li<br />
1990). The holotype was indicated to be at PE,<br />
<strong>and</strong> no isotypes were cited in the original<br />
description. I examined a specimen <strong>of</strong> Pringle<br />
4366 at P, indicated by the senior author C. B.<br />
Shang as the holotype; <strong>and</strong> I also failed to locate<br />
any specimen <strong>of</strong> Pringle 4366 at PE. In an earlier<br />
draft <strong>of</strong> the paper by Xiang <strong>and</strong> Li (1990)<br />
distributed by the junior author, the holotype <strong>of</strong><br />
the species was indicated to be at P. It seems that<br />
“PE” was a misprint <strong>of</strong> “P” in the protologue when<br />
the holotype was indicated. This raises an<br />
intriguing issue on whether the name Pentapanax<br />
mexicanus is valid, as the holotype was not in the<br />
herbarium as indicated in the original description.<br />
Nevertheless, Pringle 4366 clearly is a specimen<br />
<strong>of</strong> <strong>Aralia</strong> humilis.<br />
My delimitation <strong>of</strong> <strong>Aralia</strong> humilis is the same<br />
as that <strong>of</strong> Smith (1944). St<strong>and</strong>ley (1924)<br />
recognized <strong>Aralia</strong> pubescens as from Sonora to<br />
Oaxaca <strong>of</strong> Mexico based on pedicel pubescence.<br />
Nevertheless, he suggested that A. pubescens may<br />
be a synonym <strong>of</strong> A. humilis (St<strong>and</strong>ley 1924, p.<br />
1081). I examined the collections across Mexico<br />
<strong>and</strong> also made field observations in Mexico <strong>and</strong><br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
127<br />
Arizona. Initially I attempted to recognize three<br />
entities within the current concept <strong>of</strong> <strong>Aralia</strong><br />
humilis. These three entities include <strong>Aralia</strong> humilis<br />
s. s. from southern Arizona <strong>and</strong> northern Mexico<br />
(Chihuahua, Durango, Sinaloa, <strong>and</strong> Sonora), <strong>Aralia</strong><br />
pubescens from Jalisco, Michoacan, Oaxaca,<br />
Puebla <strong>and</strong> Colima <strong>of</strong> Mexico with pubescent <strong>and</strong><br />
more or less shorter pedicels, <strong>and</strong> <strong>Aralia</strong><br />
chilapensis widespread in central to southern<br />
Mexico, Guatemala, Honduras, <strong>and</strong> south to<br />
Nicaragua with glabrous <strong>and</strong> somewhat longer<br />
pedicels. Clearly these distinctions have lots <strong>of</strong><br />
intermediates <strong>and</strong> the narrower species concept will<br />
make it difficult practically to identify taxa in<br />
Oaxaca, Jalisco <strong>and</strong> Puebla where both “<strong>Aralia</strong><br />
chilapensis” <strong>and</strong> “A. pubescens” <strong>and</strong> their<br />
intermediate forms occur. I herein treat <strong>Aralia</strong><br />
humilis as a variable species especially concerning<br />
its pubescence <strong>and</strong> length <strong>of</strong> pedicels <strong>and</strong> leaflet<br />
shape. Most likely <strong>Aralia</strong> humilis may be the<br />
progenitor <strong>of</strong> both <strong>Aralia</strong> regeliana <strong>and</strong> A.<br />
scopulorum. Based on the leaf architecture <strong>and</strong><br />
leaflet morphology as well as geographic<br />
distribution, <strong>Aralia</strong> regeliana may have derived<br />
from the bipinnate forms <strong>of</strong> <strong>Aralia</strong> humilis from<br />
central to southern Mexico; <strong>and</strong> A. scopulorum <strong>of</strong><br />
Baja California may be a derivative <strong>of</strong> A. humilis<br />
from northern Mexico. Detailed phylogeographic<br />
analyses are needed to construct the speciation<br />
history <strong>of</strong> <strong>Aralia</strong> sect. Humiles.<br />
17. <strong>Aralia</strong> regeliana Marchal — Fig. 46; color<br />
plate 13: C-F.<br />
<strong>Aralia</strong> regeliana Marchal, Bull. Acad. Roy. Sci.<br />
Belgique, ser. 2, 47: 73. 1879. TYPE: Mexico.<br />
TAMAULIPAS: Tanque Colonada ad Victoria,<br />
Aug 1842, Karwinsky 777 (holotype: LE!).<br />
Deciduous shrub 1-4 m tall. Stem grayish<br />
brown, branchlets terete. Leaves 10-20 cm long,<br />
7-13 cm wide, bipinnate or mixed with pinnate<br />
ones, with 3-7 leaflets, accessary leaflets absent;<br />
stipule 2.5-3.5 mm long, 1.2-1.5 mm wide, adnate<br />
to petiole base, lanceolate in outline, ciliate at<br />
margin; petioles 3-8.5 cm long, glabrous; leaflets<br />
(2) 3-6 cm long, 2-3.8 cm wide, chartaceous,<br />
broadly ovate to ovate, acuminate at apex,<br />
subcordate to truncate at base, symetrical, sparsely<br />
serrate at margin, teeth <strong>of</strong>ten callose-tipped, lateral<br />
veins 6-7, conspicuous above <strong>and</strong> below, abaxial
128<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
Fig. 46. <strong>Aralia</strong> regeliana Marchal. A. Habit with leaves, stem <strong>and</strong> inflorescence. B. Inflorescence. C. Leaves on upper portion<br />
<strong>of</strong> stem, showing leaf variation in architection. D. Leaf with accessory leaflets. E. Bipinnate leaf with accessory leaflets. F.<br />
Leaf. G. Lower leaf surface <strong>and</strong> teeth at margin. H. Floral buds. I. Flower. J. Flower after anthesis. K. Fruit.
<strong>and</strong> adaxial surface glabrous, or occasionally with<br />
a few scattered hairs on the veins on the upper<br />
furface, petiolules 1.5-8.5 mm long, glabrous.<br />
Inflorescence 6-15 cm long, 5-11 cm wide, terminal<br />
at the branch apex, pilose, consisting <strong>of</strong> 3-10<br />
primary branches arranged on a main axis;<br />
inflorescence bracts 5-8, arranged spirally at the<br />
base, 5-7 mm long, 3-4 mm wide, not persistent,<br />
narrowly triangular in shape, brown in color; each<br />
primary branch with 1-3 umbels with the terminal<br />
umbel well developed <strong>and</strong> the lateral ones poorly<br />
developed or aborted, terminal umbels with 25-35<br />
flowers, lateral umbels 15-25 flowered, peduncles<br />
1.5-3 cm long; pedicels pilose, slightly enlarged<br />
at the tip, pedicels <strong>of</strong> the main terminal umbel at<br />
the tip <strong>of</strong> each inflorescence 12-16 mm long, those<br />
<strong>of</strong> the terminal umbel <strong>of</strong> primary branches 7-12<br />
mm long, those <strong>of</strong> lateral umbels <strong>of</strong> 5-7 mm long;<br />
bracts <strong>of</strong> primary branches 5-7 mm long, 2-3.5 mm<br />
wide, narrowly triangular to lanceolate, ciliate at<br />
margin; bracteoles 2-3 mm long, 0.8-1 mm wide,<br />
lanceolate, ciliate at margin. Sepals 0.3-0.4 mm<br />
long <strong>and</strong> wide, rounded to triangular; petals 2.1-<br />
2.3 mm long, 1.3-1.5 mm wide, ovate; stamens 5-<br />
6, filaments 2.3-2.5 mm long, anthers 0.6-0.7 mm<br />
long, ovate; styles 5-6, ca. 1.0 mm long after falling<br />
<strong>of</strong>f <strong>of</strong> petals, divided; floral disk projected. Fruits<br />
6-6.5 long, 5.2-5.5 mm wide, ovoid globose, black<br />
with purple juice, persistent styles 1.2-1.5 mm long,<br />
divided up to the base. Seeds 4.7-5 mm long, 2-<br />
2.2 mm wide, 1.5-1.6 mm thick, kidney-shaped.<br />
Common name: jamoncillo (Tamaulipas,<br />
Mexico).<br />
Phenology: flowering from February to March;<br />
fruiting in April to June, also in October.<br />
Distribution: endemic to Mexico (Coahuila,<br />
Durango, Guanajuato, Hidalgo, Queretaro, San<br />
Luis Potosi, <strong>and</strong> Tamaulipas) (Fig. 47).<br />
Ecology: in canyons, limestone hill slopes,<br />
open limestone ridges, steep rocky slopes; 1250-<br />
3138 m.<br />
Additional specimens examined: Mexico.<br />
COAHUILA: Sierra de Jimulco <strong>and</strong> up to 3 km N<br />
<strong>of</strong> Mina San Jose which is 8 km NE <strong>of</strong> Estacion<br />
OTTO, 25 o 6’30"-8"30"N, 103 o 13’30"W, 1800-<br />
3138 m, mat. Esp. lat. – chaparral on higher slopes,<br />
steep to very steep slopes <strong>of</strong> limestone in places<br />
highly mineralized calcareous, assoc. with Acacia<br />
berl<strong>and</strong>ieri, A. crassifolia, Fouquieria, higher is<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
129<br />
Quercus spp., 27 Sep 1972, in fr., F. Chiang et al.<br />
9549b (MEXU, TEX); ca. 26 (air) miles SW <strong>of</strong><br />
Torreon in Sierra de Jimulco, ca. 6 (air) mi SSW<br />
<strong>of</strong> La Rosita, along trail to summit, on open<br />
limestone ridge between two canyons, just below<br />
oak forest, with Artemisia, Agave, Daslirion,<br />
Opuntia, Cercocarpus, Croton, Cordia, <strong>and</strong><br />
grasses, small gnarled tree, to 6 inches in diameter,<br />
4 ft tall, 6 ft wide at crown, fls yellowish, 8200 ft,<br />
18 Sep 1973, in fr., 25 o 10’N, 103 o 15’W, J.<br />
Henrickson 13172 (MEXU, RSA, TEX); Sierra<br />
de Jimulco, N-facing cliffs, NW-facing notches,<br />
25 o 11’N, 103 o 12’W, 1600-2150 m, mostly<br />
chaparral, lower some matorral desertico,<br />
limestone, assoc. with Fraxinus greggii, Lindleya,<br />
Agave parrasana, Bonnetiella anomala, <strong>and</strong><br />
Juniperus sp., 27 Jun 1973, in young fr. & late fl.,<br />
M. C. Johnston et al. 11484 (LL, MEXU, MO);<br />
Sierra de Jimulco, Mina San Jose, ca. 10 km al NE<br />
de la Flor de Jimulco, 103 o 13’30’’W y 25 o 6’30"’N,<br />
2150 m, veg. Matorral de Acacia crassifolia, A.<br />
berl<strong>and</strong>ieri, Lindleya mespiloides, Cercocarpus<br />
mojadensis y Vauquelinia californica, 25 Aug<br />
1988, veg., J. A. Villarreal 4377 (TEX); 25 Aug<br />
1988, in fr., J. A. Villarreal 4388 (TEX); Sierra de<br />
Jimulco, mina San Jose, vereda hacia la cima,<br />
25 o 08’N, 103 o 13’W, matorral de Bonetiella, Agave<br />
lechuguilla, Acacia berl<strong>and</strong>ieri, Flourensia,<br />
Hechitia, <strong>and</strong> Spiraea, 1800-1850 m, 10 Aug 1994,<br />
fr., J. A. Villarreal 7807 (MEXU, TEX, 2 sheets).<br />
DURANGO: northwestern third <strong>of</strong> Sierra del<br />
Rosario, 25 o 42’ – 25 o 45’N, 103 o 57’ – 104 o 00’W,<br />
1800-2655 m, chaparral on top, steep limestone<br />
sierra with some zones <strong>of</strong> igneous mineralization,<br />
limestone derived gravel, assoc. with Garrya, Rhus<br />
virens, Arctostaphylos, <strong>and</strong> Agave macroculmis,<br />
rare, 25 Jun 1973, fl. & young fr., M. C. Johnston<br />
et al. 11466 (LL, MO). GUANAJUATO: Puerto<br />
de La Calera, Mpio. De Atarjea, 1900 m, 12 Jul<br />
1990, fr, E. Ventura & E. Lopez 8241 (MEXU).<br />
Cerro de Veracruz, Mpio. Atarjea, 16 May 1990,<br />
fr, 1250 m, E. Ventura & E. Lopez 7998 (MEXU).<br />
HIDALGO: Barranca de Venados, al principio, a<br />
1 km de Paso de León, 1800 m, 3 Jun 1976, fr, A.<br />
Delgado S. & R. Hernández 239 (MEXU, 2 sheets).<br />
Barranca de Venados, mpio de Atotonilco el<br />
Gr<strong>and</strong>e, planta arbustiva, de 1-2 m, 1900 m, 8 May<br />
1981, fr, R. Hernández M. 6058 (MO). 3 km al SE<br />
de Venados, 2 Nov 1975, F. Gonzalez M. 8408<br />
(MEXU); Municipio Cardonal, Barranco<br />
Tolantongo, El Marmo, the third curve from the
130<br />
bottom <strong>of</strong> Grutas de Tolantongo, limestone slope,<br />
rocky, 1550 m, 21 Mar 1988, fl, J. Wen & E.<br />
Martinez S. 384 (OS, US). QUERETARO: ca. 80<br />
km NE <strong>of</strong> Querétaro, lower limit <strong>of</strong> piñon-juniper<br />
belt in dry mountains above Pilón on road to Pinal<br />
de Amoles, 2700 m, occasional on N slopes <strong>of</strong><br />
barranca, 24 Apr 1949, in fl. & young fr., R.<br />
McVaugh 10348 (BM, LL, 2 sheets, MEXU, MO,<br />
TEX, US). SAN LUIS POTOSI: Minas de San<br />
Rafael, May 1911, C. A. Purpus 5010 (BM, E, MO,<br />
US), 5011 (BM, MEXU, MO, US); Guadalcázar,<br />
6 km al W del Crucero carr. Matehuala-Noria de<br />
las Flores, Cañada a mano, izquierda hacia Noria<br />
de Las Flores (frente cerro Chirrion), arbolito de<br />
4-5 m, con fr., 27 Jun 2000, R. Torres Colin 15689<br />
(MEXU, TEX). TAMAULIPAS: Mpio.<br />
Miquihuana, Loc. Rincón de las Vacas, 7 km de<br />
Miquihuana, entre Miquihuana y la Perdina, 1870<br />
m, pinar de P. cembroides en laderas N, arbusto de<br />
4 m, escasa, blanco-verdosas, 23 May 1974, fl, F.<br />
González-Medrano 7029 (RSA); 18 km by winding<br />
road SE <strong>of</strong> Bustamente toward La Presita <strong>and</strong> Tula,<br />
23 o 20’N, 99 o 40’W, 1700 m, Pinar, gypseous<br />
limestone slope, calcareous gypseous gravelly soil,<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
Fig. 47. Map <strong>of</strong> Mexico showing the distributions <strong>of</strong> <strong>Aralia</strong> regeliana Marchal <strong>and</strong> A. scopulorum Br<strong>and</strong>egee.<br />
assoc. with Pinus nelsonii, Arctostaphylos,<br />
Coldenia, Ilex, <strong>and</strong> Lindleya, 20 May 1973, in fl.,<br />
M. C. Johnston et al. 11149 (LL, MO); Jaumave<br />
Valley, ca. 2000 ft, 1 Jun 1898, E. W. Nelson 4463<br />
(US); 5.8 mi N <strong>of</strong> Rte. 101 on road to Bustamante,<br />
23 o 20’N, 99 o 41’, 2100 m, matorral/chaparral,<br />
limestone mountains, associated with Quercus<br />
spp., Dasylirion, Juniperus, Brahea, Agave, Rhus,<br />
Dalea, Cowania, Ephedra, Mortonia, <strong>and</strong> Pinus<br />
nelsonii, 26 May 1974, in fl., T. Wendt & F. Chiang<br />
185 (TEX). Tamaulipas, Felipe Angeles Ojo de<br />
Agua San Jose, 2-3 km al W de Ejido, Municipio.<br />
Bustamante, 1600 m, matorral alto subinerme,<br />
arbol 2 m, 5 Oct 1985, R. Diaz 00494 (MO).<br />
Mexico, La Miquiguana, 1841-42, Karwinsky 1248<br />
(LE).<br />
<strong>Aralia</strong> regeliana is characterized by its glabrous<br />
<strong>and</strong> bipinnate leaves with ovate leaflets having a<br />
long acuminate apex. It is noted that E. Ventura &<br />
E. Lopez 9217 (el Pinalito Redondo, por Carricillo,<br />
Mpio. De Atarjea, Guanajuato, MEXU) appears<br />
to be A. regeliana, yet they have pubescent leaflets.<br />
Because this collection was from transitional zones
<strong>of</strong> <strong>Aralia</strong> regeliana <strong>and</strong> A. humilis, it may represent<br />
a hybrid <strong>of</strong> the two species.<br />
18. <strong>Aralia</strong> scopulorum Br<strong>and</strong>egee — Fig. 48;<br />
color plate 13: A-B.<br />
<strong>Aralia</strong> scopulorum Br<strong>and</strong>egee Proc. Calif. Acad.<br />
Sci. 2:165. f. 8. 1889. Pentapanax scopulorus<br />
(Br<strong>and</strong>egee) C. B. Shang in Y. W. Yuan & al.<br />
(eds.), Proc. Intern. Symp. Bot. Gard. 1990:<br />
630, 1990. TYPE: Mexico. BAJA<br />
CALIFORNIA SUR: Comondu Canon,<br />
common on rocks, 12 Mar 1889, fl & young fr,<br />
T. S. Br<strong>and</strong>egee s.n. (holotype, UC!; isotypes,<br />
A!, GH!, GH photo!, PH!, US!).<br />
Deciduous shrub to small tree 1.5-8 m tall.<br />
Stem grayish brown, branchlet terete. Leaves 10-<br />
20 cm long, 7-13 cm wide, pinnate, with 3-7<br />
leaflets, accessary leaflets absent; stipule 3-4 mm<br />
long, 1-1.5 mm wide, adnate to petiole base,<br />
lanceolate in outline, ciliate at margin; petioles 3-<br />
8.5 cm long, glabrous; leaflets (2) 3-6 cm long, 2-<br />
3.8 cm wide, chartaceous, broadly ovate to ovate,<br />
acuminate at apex, subcordate to truncate at base,<br />
symetrical, sparsely serrate at margin, teeth <strong>of</strong>ten<br />
callose-tipped, lateral veins 6-7 on each side,<br />
conspicuous above <strong>and</strong> below, abaxial <strong>and</strong> adaxial<br />
surface glabrous, or occasionally with a few<br />
scattered hairs on the veins on the upper furface;<br />
petiolules 1.5-8.5 mm long, glabrous.<br />
Inflorescence 6-15 cm long, 5-11 cm wide, terminal<br />
at the branch apex, pilose, consisting <strong>of</strong> 3-10<br />
primary branches arranged on a main axis;<br />
inflorescence bracts 5-8, 5-7 mm long, 3-4 mm<br />
wide, not persistent, arranged spirally at the base,<br />
narrowly triangular in shape, brown in color; each<br />
primary branch with 1-3 umbels with the terminal<br />
umbel well developed <strong>and</strong> the lateral ones poorly<br />
developed or aborted, terminal umbels with 25-35<br />
flowers, lateral umbels 15-25-flowered; peduncles<br />
1.5-3 cm long; pedicels <strong>of</strong> the main terminal umbel<br />
at the tip <strong>of</strong> each inflorescence 12-16 mm long,<br />
those <strong>of</strong> the terminal umbel <strong>of</strong> primary branches<br />
7-12 mm long, those <strong>of</strong> lateral umbels <strong>of</strong> 5-7 mm<br />
long, pilose, slightly enlarged at the tip; bracts <strong>of</strong><br />
primary branches 5-7 mm long, 2-3.5 mm wide,<br />
narrowly triangular to lanceolate, ciliate at margin;<br />
bracteoles 2-3 mm long, 0.8-1 mm wide,<br />
lanceolate, ciliate at margin. Sepals 0.3-0.4 mm<br />
long <strong>and</strong> wide, rounded to triangular; petals 2.1-<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
131<br />
2.3 mm long, 1.3-1.5 mm wide, ovate; stamens 5-<br />
6, filaments 2.3-2.5 mm long, anthers 0.6-0.7 mm<br />
long, ovate; styles 5-6, divided, ca. 1.0 mm long<br />
after falling <strong>of</strong>f <strong>of</strong> petals, floral disk projected.<br />
Fruits 6-6.5 long, 5.2-5.5 mm wide, ovoid globose,<br />
black with purple juice; persistent styles 1.2-1.5<br />
mm long, divided up to the base. Seeds 4.7-5 mm<br />
long, 2-2.2 mm wide, 1.5-1.6 mm thick, kidneyshaped.<br />
Common names: sauco cimarrón, <strong>and</strong> sauchio.<br />
Phenology: flowering from February to May;<br />
fruiting in April to June, also in October.<br />
Distribution: Baja California (primarily in Baja<br />
California Sur), Mexico (Fig. 47).<br />
Ecology: in canyons, deep canyon bottom,<br />
hillsides, among rocks, hill ridge, rocky slopes,<br />
volcanic rock outcrops, <strong>and</strong> also around high arid<br />
rims (in lesser stature); 200-1600 m.<br />
Additional specimens examined: Mexico.<br />
BAJA CALIFORNIA NORTE: ca. 3 on southeast<br />
ridge, Cerro la S<strong>and</strong>ía, 28 o 24’N, 113 o 27.5’W, 1300<br />
m, 24 Jan 1964, sterile, shrub ca. 2 m tall, R. Moran<br />
11545 (RSA, US); ca 10 individuals on north slope<br />
under cliff, ca. 2.5 mi east <strong>of</strong> La S<strong>and</strong>ía, 28 o 24’N,<br />
113 o 25.5’W, 925 m, 25 Jan 1964, R. Moran 11554<br />
(LL). BAJA CALIFORNIA SUR: ca. 2 km SW <strong>of</strong><br />
San Francisco de la Sierra, north side <strong>of</strong> Sierra<br />
Agua Verde, north-running canyon with Dodonaea<br />
viscosa, Nicotiana glauca, Nolina beldingii,<br />
Perezia palmeri, Rhamnus crocea, Rhus<br />
integrifolia, <strong>and</strong> Washingtonia robusta, N27 o 35’,<br />
W113 o 02’, 1150 m, sympodial tree to 6 m tall, as<br />
broad, 20 Feb 1994, fl, M. A. Baker et al. 11308<br />
(ASU); dry, rocky, broad bed <strong>of</strong> Arroyo Carrizal,<br />
east <strong>of</strong> Rancho El Horno (NE <strong>of</strong> San Xavier), 550-<br />
700 m, 25 o 53’N, 111 o 31.5’W, A. Carter & R. Ferris<br />
3816 (MEXU, MO, TEX); Sierra de la Giganta,<br />
La Victoria, 540 m, 25 o 52.5’N, 111 o 25’W, 20 Mar<br />
1960, A. Carter & R. Ferris 3893 (BM, MEXU,<br />
MO, TEX); deep, vertical-walled canyon, Aguaje<br />
de los Encinos (S side <strong>of</strong> Cerro Giganta), 26 o 5’N,<br />
111 o 35’W, 850 m, 27 Mar 1960, in young fr., A.<br />
Carter & R. Ferris 3984 (BM, MEXU, MO);<br />
volcanic rock outcrops, vicinity <strong>of</strong> La Tinaja, Mesa<br />
de San Alejo (west <strong>of</strong> San Javier), 750 m,<br />
25 o 51.5’N, 111 o 34.5’W, 10 Mar 1961, fl, A. Carter<br />
& H. Sharsmith 4165 (MEXU); on steep northfacing<br />
canyon wall, La Esperanza, 25 o 48’N,<br />
111 o 24.5’W, 20 Apr 1962, A. Carter 4393 (BM,
132<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
Fig. 48. <strong>Aralia</strong> scopulorum Br<strong>and</strong>egee. A. Habit with leaves <strong>and</strong> inflorescence. B. Close-up <strong>of</strong> leaf margin <strong>and</strong> lower leaflet<br />
surface. C. Flower. D. Fruit.
MEXU); Canada south <strong>of</strong> Rancho de Los Encinos,<br />
Valle de Los Encinos (south side <strong>of</strong> Cerro Giganta),<br />
26 o 3.5’N, 111 o 35’W, 7 Jun 1963, A. Carter & J.<br />
Reese 4565 (BM, MEXU, MO); south-facing slope<br />
<strong>of</strong> Cerro del Pinto, N <strong>of</strong> Portezuelo de San Antonio,<br />
headwater <strong>of</strong> Arroyo el Coyote (SE <strong>of</strong> La Soledad<br />
<strong>and</strong> N <strong>of</strong> Cerro Mechudo, 850 m, 24 o 50.5’N,<br />
110 o 44’W, 21 Feb 1970, fl, A. Carter 5453<br />
(MEXU); Sierra San Francisco, southwest edge <strong>of</strong><br />
Mesa San Jorge, ca. 8 km southwest <strong>of</strong> San<br />
Francisco <strong>and</strong> 13 km WNW <strong>of</strong> Santa Marta, near<br />
27 o 33.5’N, 113 o 05.75’W, 9 Jun 1984, fr, 800 m, J.<br />
Dice et al. 507 (RSA); Sierra San Francisco, SW<br />
edge <strong>of</strong> Mesa San Jorge, ca. 8 km SW <strong>of</strong> San<br />
Francisco de la Sierra, near 27 o 33.5’N, 113 o 05.75’,<br />
800 m, 24 Jun 1986, young fr, J. C. Dice et al. 680<br />
(MEXU); Volcan de Lar Virgines, 1893, 1500 m,<br />
Diguet s.n. (P); La Champagna, Sierra de las<br />
Palmas, south <strong>of</strong> Santa Rosalia, 27-29 Apr 1952,<br />
in fl. & young fr., Nolina grassl<strong>and</strong> over undulating,<br />
broken terrain <strong>of</strong> volcanic mountain top, 4500-<br />
5000 ft, treelet 4-6 m, with spreading crown,<br />
drought-pauperized leaves, & white flowers, H. S.<br />
Gentry & W. B. Fox 11806 (LL, MEXU); Los<br />
Encinos, Sierra Giganta, deep canyon bottom, 2500<br />
ft, 27 Feb 1939, fl, H. S. Gentry 4262 (K, MEXU,<br />
MO, US); Sierra San Francisco, just southeast <strong>of</strong><br />
town <strong>of</strong> San Francisco de la Sierra, 113 o 00’39.4"W,<br />
27 o 35’23"N, 1200 m, rocky dry slopes <strong>and</strong> narrow<br />
drainages, frequent large shrub, 20 Apr 1994, fl,<br />
W. Hodgson 8121 (ASU); Sierra San Francisco,<br />
Canon Santa Theresa, ca. ½-3/4 mile south <strong>of</strong><br />
Arroyo Solidad, 113 o 04’14.2"W, 27 o 37’51.1"N,<br />
515 m, in canon, north-south orientation, common<br />
shrub, called sauchio by local inhabitants, 21 Apr<br />
1994, fr, W. Hodgson 8148 (ASU, DES); Arroyo<br />
Undo Ranch, Loreto, 26 Oct 1930, M. E. Jones<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
133<br />
27094 (RSA); ca. Cucua “El Raton,” 27 o 35’N,<br />
113 o 03’W, 27 Jan 1989, young fr, 1420 m, J. L.<br />
León 3560 (RSA); summit, Cerro San Juan, rather<br />
scarce, from 950 m, 1250 m, 27 o 58’N, 113 o 00’W,<br />
4 Feb 1964, fl, R. Moran 11586 (RSA); fairly<br />
common on lower north slope <strong>of</strong> Volcan las Tres<br />
Virgenes, seen from 800 m, 1100 m, tree to 8 m<br />
tall, trunk to 8 dm thick, or with several trunks<br />
from a base <strong>of</strong> 1 m, bark gray, 11 Apr 1973, in fl.,<br />
27 o 29’N, 112 o 36’W, R. Moran 20405 (ASU, LL,<br />
MO, NCU); fairly common, Cerro la Laguna,<br />
highest peak <strong>of</strong> Sierra San Francisco, seen 700-<br />
1500 m, 27 o 35’N, 113 o 02’W, 1450 m, 24 Nov 1976,<br />
sterile, R. Moran 23846 (MSC); Aguajidi Santana,<br />
35 miles N <strong>of</strong> San Ignacio, 3400 ft, 4 Oct 1905,<br />
young fr, E. W. Nelson & E. A. Goldman 7189 (US);<br />
rocky slopes San Pablo, Santa Gertridis,1000-2000<br />
ft, Jan-Mar 1898, C. A. Purpus s.n. (K, US, WU);<br />
rocky mountain flats at San Francisco de la Sierra,<br />
27 36’N, 113 01’W, 28 May 1992, fl & young fr,<br />
J. Rebman et al. 1420 (ASU); Sierra Agua Verde,<br />
upper parts <strong>of</strong> the large canyon 1.5 miles to the<br />
west <strong>of</strong> the town <strong>of</strong> San Francisco de la Sierra,<br />
growing with Condalia, Nolina, Polygala<br />
apopetala, <strong>and</strong> palm trees, 113 o 02’W, 27 o 36’N, tree<br />
to 6 m tall, 6 Mar 1994, fl, J. P. Rebman 2372<br />
(ASU, RSA); woods <strong>of</strong> Bursera, Lysiloma, <strong>Aralia</strong><br />
et al., in east-facing ravine, hilly slopes W <strong>of</strong> Volcán<br />
Las Tres Vírgenes, 200 m, 27 o 30’N, 112 o 38’W, 29<br />
Mar 1989, fl, G. L. Webster 26161 (MEXU, 2<br />
sheets); scrub <strong>and</strong> woodl<strong>and</strong>s in the vicinity <strong>of</strong><br />
Cueva de la Ratón, 4 mi SW <strong>of</strong> San Francisco de<br />
la Sierra, 1000 m, 27 o 24’N, 113 o 00’W, 25 Mar<br />
1989, fl, G. L. Webster 26312 (MEXU); Camp <strong>and</strong><br />
hillsides near Comondu, shrub 2 m high, stalks<br />
erect, 26 Apr 1931, young fr, I. L. Wiggins 5489<br />
(RSA, US).<br />
ARALIA SECT. SCIADODENDRON (GRISEB.) J. WEN<br />
<strong>Aralia</strong> sect. Sciadodendron (Griesb.) J. Wen, comb. et stat. nov.<br />
Sciadodendron Griseb., Bonpl<strong>and</strong>ia 6: 7. 1858. Type species: Sciadodendron excelsum Griseb.<br />
Coemansia Marchal, Bull. Acad. Roy. Sci. Belgique 47: 94. 1879. nom. illeg., non Tiegh. & Monnier in<br />
Ann. Sci. Nat. Bot., Ser. 5, 17: 392. 1873.<br />
Coudenbergia Marchal, Bull. Acad. Roy. Sci. Belgique 47: 514, 1879. Type species: Coudenbergia<br />
warmingiana (Marchal) Marchal.<br />
Megalopanax Ekman ex Harms, Notizbl. Bot. Gart. Berlin 9: 122, 1924. Type species: Megalopanax<br />
rex Ekman ex Harms.
134<br />
Unarmed glabrous shrub or tree, highly<br />
branched. Leaves tri- to bipinnate or quadripinnate.<br />
Umbels clustered into panicles, several<br />
inflorescences usually aggregated at tip <strong>of</strong> stem or<br />
a somewhat short branch, with persistant bracts at<br />
the base, pedicels articulated or non-articulated at<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
the base <strong>of</strong> the flower. Flowers 5-, 6- or 8-12merous,<br />
styles connate or at least at the base,<br />
occasionally completely distinct. Fruits dark<br />
purple.<br />
Key to species <strong>of</strong> <strong>Aralia</strong> sect. Sciadodendron<br />
Five species from Central <strong>and</strong> South America.<br />
1. Pedicels not articulated ...................................................................................................................... 2<br />
1. Pedicels articulated ........................................................................................................................... 3<br />
2. Fruits subglobose, 6-7 mm long, peduncles relatively slender, 1-1.5 mm in thickness .......... A. excelsa<br />
2. Fruits ovoid globose to globose, 7-8 mm long, peduncle stout, 2-3 mm in thickness ........... A. bahiana<br />
3. Ovary 5-locular ...................................................................................................................... A. soratensis<br />
3. Ovary 6-10-locular ........................................................................................................................... 4<br />
4. Leaflets entire at margin, subcordate to less <strong>of</strong>ten rounded at base; peduncle <strong>of</strong> umbel thick, 3-5 mm<br />
in diameter; umbels 90-120-flowered; endemic to Cuba ........................................................... A. rex<br />
4. Leaflets sparsely serrate at margin, rounded at base; peduncle <strong>of</strong> umbel 1.5-2 mm in diameter; umbels<br />
25-40-flowered; distributed in South America (Brazil <strong>and</strong> Paraguay, extending to Argentina)<br />
.......................................................................................................................................... A. warmingiana<br />
19. <strong>Aralia</strong> soratensis Marchal — Fig. 49; color<br />
plate 14: A-F.<br />
<strong>Aralia</strong> soratensis Marchal, Bull. Acad. Roy. Sci.<br />
Belgique 47: 75. 1879. TYPE: Bolivia. LA<br />
PAZ: Prov. Larecaja, near Sorata, San Pedro,<br />
2600 m, Mar 1860, fr, G. M<strong>and</strong>on 570<br />
(lectotype, designated by Wen, 1993, P!;<br />
isolectotype, K!, W!).<br />
Pentapanax angelicifolius Griseb., Abh. Konigl.<br />
Ges. Wiss. Göttingen 24: 144. 1879.<br />
Coudenbergia angelicifolium (Griseb.) C. B.<br />
Shang in Y. W. Yuan & al. (eds.), Proc. Intern.<br />
Symp. Bot. Gard. 1990: 632. 1990. TYPE:<br />
Argentina. TUCUMAN: near la Cruz, “Palo<br />
de San Antonia,” hier höherer schlanker Baum,<br />
22-25 Dec 1872, P. G. Lorentz & G. Hieronymus<br />
1173 (holotype, GOET!; isotypes, GOET!, 2<br />
sheets).<br />
<strong>Aralia</strong>? weberbaueri Harms, Repert Spec. Nov.<br />
Regni Veg. 15: 254. 1917. TYPE: Peru.<br />
HUANCAVELICA: Prov. Angaraes, left side<br />
<strong>of</strong> the river Huarpa, 2800-2900 m, Jun 1910,<br />
fr, A. Weberbauer 5668 (lectotype, designated<br />
by Wen, 1993, A!; isolectotypes: G!, GH!, K!,<br />
MO!, NY!, US!, 2 sheets, photo at K).<br />
Tree 4-15 m tall, about 7-15 cm in dbh,<br />
polygamo-monoecious. Bark furrowed, gray.<br />
Leaves 80-125 cm long, 70-120 cm wide, bipinnate<br />
to tripinnate, sometimes pinnate near the<br />
inflorescence; stipule 3-5 mm long, 2-3 mm wide,<br />
adnate to the base <strong>of</strong> petiole, the free portion<br />
triangular to narrowly so, subcoriaceous, glabrous;<br />
petioles purplish green to light green, 20-50 cm<br />
long; rachises green, turning purplish, subtended<br />
by a pair <strong>of</strong> accessory 3-5-foliolate pinnae at the<br />
base <strong>of</strong> the first two lower pinnae <strong>and</strong> a pair <strong>of</strong><br />
accessory leaflets at the upper pinnae; leaflets 5.5-<br />
10.5 cm long, 2.5-6.0 cm wide, glabrous,<br />
acuminate at apex, rounded, obtuse to subcordate,<br />
sometimes truncate at base, sparsely finely serrate<br />
to sometimes crenately serrate at margin, adaxial<br />
surface green, abaxial surface light green, petiolule<br />
6-17 mm long, glabrous. Inflorescence terminal,<br />
consisting <strong>of</strong> an aggregation <strong>of</strong> 2-5 panicles on a<br />
short axis, each panicle 17-30 cm long, 10-20 cm<br />
wide, with 25-40 primary branches arranged<br />
racemosely on a main axis, primary branches 4-9<br />
cm long, glabrous, each with 2-5 umbels, lateral
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
Fig. 49. <strong>Aralia</strong> soratensis Marchal. A. Leaf. B. Inflorescence. C. Close-up <strong>of</strong> lower leaflet surface <strong>and</strong> margin. D. Floral bud.<br />
E. Flower. F. Flower after anthesis. G. Fruit (A & C – Schnell 189, W; B - Vervoorst & Cuezze 7632, W; D-F – Meyer s.n.,<br />
15 Sep 1967, W; G - Lorentz & Hieronymus 214, GOET).<br />
135
136<br />
umbels <strong>of</strong>ten aborted or functionally male, bracts<br />
<strong>and</strong> bracteoles persistent, glabrous, primary bracts<br />
(subtending the individual panicles) 3-5 mm long,<br />
6-8 mm wide, triangular, subcoriaceous, secondary<br />
bracts 3.5-5 mm long, 2.5-4 mm wide, narrowly<br />
triangular, bracteoles 0.6-0.9 mm long, 0.4-0.6 mm<br />
wide, narrowly triangular; umbels 12-25-flowered,<br />
pedicels 5-9 mm long, glabrous, articulated near<br />
the base <strong>of</strong> the flower. Flowers yellow-green,<br />
nearly odorless, appearing at the same time as<br />
leaves; sepals minute, triangular; petals 2.1-2.3 mm<br />
long, 1.2-1.3 mm wide, greenish white, ovate;<br />
filaments 2.1-2.5 mm long, anthers 0.8-1 mm long,<br />
oblong; ovaries 5-locular. Fruits 4-5 mm in<br />
diameter, globose, blackish purple when mature,<br />
persistent styles slightly divided at the tip, connate<br />
4/5 at the lower part, floral disc 2-3 mm in diameter<br />
at the fruiting stage, conspicuous.<br />
Local names: sacha paraiso (Jujuy, Salta <strong>and</strong><br />
Tucumán, Argentina), paraiso (Jujuy, Argentina),<br />
yapicay (by Guarani tribe, Chuquisaca, Bolivia),<br />
<strong>and</strong> mara blanca (Santa Cruz, Bolivia).<br />
Uses: firewood or making utensils (Potosi <strong>of</strong><br />
Bolivia), medicinal (Chuquisaca, Bolivia).<br />
Phenology: flowering in September to October;<br />
fruiting in December.<br />
Distribution: northern Argentina, Bolivia, <strong>and</strong><br />
Peru (Fig. 50).<br />
Ecology: scattered in valley bottom <strong>and</strong> dry<br />
deciduous scrub on hillsides, open dry forests,<br />
premontane moist forest, slopes with semideciduous<br />
forests, <strong>and</strong> dry legume-dominated<br />
vegetation; 380-2900 m.<br />
Additional specimens examined: Argentina.<br />
JUJUY: Jujuy, Dr. Grisebach & Dr. Lorentz 11/77<br />
(K); Dec 1878, fl, Grisebach s.n. (K); S. Larenzo,<br />
Nov 1873, in fl., P. G. Lorentz & G. Hieronymus<br />
214 (GOET, 2 sheets, NY, US); Prov. Jujuy, Depto.<br />
Ledesma, Arroyo la Cantara in the Cerro de<br />
Calilegua, 16 km north <strong>of</strong> Calilegua, overhanging<br />
streambank, 600 m, 19 Oct 1938, fl, W. J. Eyerdam<br />
& A. A. Beetle 22669 (K); Limite Camino, Salta –<br />
Jujuy, 1280 m, 15 Sep 1967, in fl., F. Meyer s.n.<br />
(W). Jujuy, 1913, Schuel 189 (W). San Pedro,<br />
Cuesta de las Lajitas, 12 Nov 1978, fl, A. L.<br />
Cabrera et al. 29806 (MO). Depto. Capital, Cuesta<br />
de Las Lajitas, 18 Nov 1980, fl, A. L. Cabrera et<br />
al. 32084 (MO). SALTA: Dep Orán, Gartagal, 500<br />
m, Oct 1940, fl., Schreiter 11380 (A, U). Dep Orán,<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
Selva Tucumano-Boliviana, Rio los Sosa, ca. 60<br />
km SW <strong>of</strong> Tucuman, N Argentina, subtropical<br />
rainforest, 700 m, 24 Nov 1967, late fl., J. H. de<br />
Haas 886 (U). Dep. Oran, Vade Honde (Rio<br />
Zenta), 380 m, 26 Oct 1970, P. Vervoorst & A. R.<br />
Cuezze 7632C (GH, W). TUCUMÁN: San Pablo,<br />
15 Jul 1888, sterile, Lillo 163 (P). Tucumán, Yerba<br />
Buena, 650 m, tree 10 m tall, 16 Nov 1924, fl, S.<br />
Venturi 1081 (US, 2 sheets) 1081b (MO, US).<br />
Tucuman, Dep Rio Chico, Iseaba, 600 m, 20 Dec<br />
1913, fl, Monetti 1748 (NY). Bolivia.<br />
CHUQUISACA: Prov. Hern<strong>and</strong>o Siles, zona de la<br />
Hacienda Naurenda, 83 km al S de Monteagudo,<br />
alrededor de la Hacienda Ipati, 1 km al S, camino<br />
al Ingre, 1150 m, bosque boliviano-tucumano,<br />
suelo franco-arenoso, moderadamente inclinado,<br />
sub-humedo, 20 Nov 1991, fl., L. Rea et al. 74<br />
(LPB). Prov. Tamina/H. Siles, por el camino de<br />
Padilla a Monteagudo, 80 km, 1650 m, 20 Sep<br />
1980, in fl., Mühlbauer 01 (LPB).<br />
COCHABAMBA: Ayopaya, Sailapata, 1200 m,<br />
tree 6-8 m tall, dry slopes, Oct 1935, fr, M.<br />
Cárdenas 3361 (US). Prov. Campero quebrada de<br />
Mataral a Yuraj Pampa, 2120 m, bosque seco de<br />
Anadenanthera macrocarpa, arbolito 7 m, 23 Mar<br />
1993, sterile, C. Antezana 650 (MO). Prov.<br />
Mizque, desvio de la carretera a Tucma Alta, ca. 5<br />
km de Mizque, 2300 m, 19 Nov 1990, in fl, E.<br />
Saravia et al. 35A (LPB). LA PAZ: Prov. Inquisivi,<br />
comunidad Khora-Vilabarranco, ladera izquierda<br />
del rio Miguillas, 21 km de Choquetanga, 16 o 40’S,<br />
67 o 20’W, 1470 m, bosque semideciduo, alterado<br />
en generacion con predominio de Cactacees, 14<br />
Jun 1994, sterile, N. Salinas 3471 (LPB). Prov.<br />
Inquisivi, 10 km al N de Inquisivi por el camino a<br />
Suri, alrededores del Puente sobre el Río Kato,<br />
bosque seco con Prosopis, Acacia, Schinopsis y<br />
Pereskia, 16 o 48’S, 67 o 11’W, 2100-2200 m, 12 Mar<br />
1988, sterile, J. C. Solomon & M. Nee 18122 (MO).<br />
POTOSÍ: Prov. Chayanta, Localidad Cruz Kasa,<br />
2535 m, Sep 1996, in fr., V. H. Zamora 194 (LPB).<br />
SANTA CRUZ: Prov. Caballero, 1-15 km E <strong>of</strong><br />
Comarapa (17 o 54’S, 64 o 29’W), dry legumedominated<br />
vegetation, ca. 1940 m, tree 5-6 m tall,<br />
15-16 Jan 1990, in fl., L. J. Dorr & L. C. Barnett<br />
7059 (LPB); 8.7 km (by road) E <strong>of</strong> Saipina on<br />
gravel road to Pulquina, 18 o 06’05"S, 64 o 31’05"W,<br />
1710 m, narrow canyon with arid thorn scrub<br />
vegetation with many cacti, small tree 7 m tall, 15<br />
cm in dbn, with 3 trunks, inflorescence hanging<br />
with fruit, a nearby tree had only flowers <strong>and</strong> the
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
Fig. 50. Map <strong>of</strong> South America showing the distribution <strong>of</strong> <strong>Aralia</strong> soratensis Marchal <strong>and</strong> A. bahiana J. Wen.<br />
137
138<br />
inflorescences were more or less erect, flowers<br />
yellow-green, fruit green with purple tinge, 9 Dec<br />
2005, in fr (also in fl in this season), M. Nee et al.<br />
53714 (NY, 2 sheets). Prov. Florida, between<br />
Huerba Buena <strong>and</strong> Aqua Clasa on road from<br />
Mairana to Matasal, steep sided valley with<br />
irrigated cultivation in valley bottom <strong>and</strong> dry<br />
deciduous scrub on hillsides, scattered trees, in dry<br />
bushl<strong>and</strong> above the river, 1400 m, 28 Sep 1996, fl,<br />
leafless, J. R. I. Wood 11444 (K, 2 sheets); 5 km<br />
(by road) SE <strong>of</strong> Bermejo, along highway from<br />
Santa Cruz to Samaipata, gorge <strong>of</strong> Río Piraí, steep<br />
slopes with semi-deciduous subtropical forest with<br />
many Mimosoid legumes, 18 o 09’53"S,<br />
63 o 36’20"W, 840 m, 17 Jan 2006, sterile, M. Nee<br />
& J. Wen 53807 (LPB, NY, US, USZ); 6.4 km S <strong>of</strong><br />
Mataral on road to Vallegr<strong>and</strong>e, 18 o 10’05"S,<br />
64 o 11’58"W, 1460 m, dry forest on slopes, with<br />
Capparis speciosa, Carica quercifolia,<br />
Aspidosperma quebracho-blanco, Prosopis<br />
kuntzei, Cereus comarapanus, Harrisia<br />
tetracantha, <strong>and</strong> Jodina rhombifolia, tree ca. 8 m<br />
tall, 18 cm dbh, flowers yellow-green, anthers<br />
nearly white, fruit green, older ones tinged with<br />
light purple, none yet ripe, 20 Jan 2006, young fr,<br />
M. Nee & J. Wen 53878 (LPB, NY, US, USZ); 0.6<br />
km NE <strong>of</strong> central square <strong>of</strong> Mairana, on dirt road<br />
to Campamento La Younga <strong>of</strong> Parque Nacional<br />
Amboro, 18 o 07’04"S, 63 o 56’05"W, 1380 m,<br />
narrow valley with dry forest <strong>of</strong> Schinopsis<br />
haenkena, Anadenanthera macrocarpa,<br />
Cnidoscolus cnicodendron, Cereus comarapanus,<br />
<strong>and</strong> Porlieria microphylla, tree ca. 8 m tall, 18 cm<br />
dbh, bark furrowed, fruit turning very datk purple<br />
<strong>and</strong> globose with truncate top, 19 Jan 2006, fr, M.<br />
Nee & J. Wen 53845 (LPB, NY, US, USZ). Prov.<br />
Vallegr<strong>and</strong>e, 0.5 km N <strong>of</strong> highest point on road<br />
from El Trigal to Mataral, 12 km (by air) NNW <strong>of</strong><br />
El Trigal, steep slopes with semi-deciduous forest,<br />
18 o 12’S, 64 o 12’W, 1700 m, 15 Dec 1990, in fl &<br />
fr., M. Nee 40304 (GH, 2 sheets, LPB, 2 sheets,<br />
MO, 2 sheets, NY, 2 sheets, US); Vallegr<strong>and</strong>e, on<br />
decent from Pucara to Santa Rosa in Rio Gr<strong>and</strong>e<br />
valley, open dry well-developed forest on steep<br />
slopes, scattered trees, constituents <strong>of</strong> open dry<br />
forest, 1900 m, 12 Feb 1996, J. R. I. Wood 10637<br />
(K). Prov. Warnes, E side <strong>of</strong> Okinawa No. 1,<br />
17 o 14’S, 62 o 53’W, 250 m, brushy areas, very flat,<br />
28 Jan 1987, sterile, M. Nee 33832 (NY). Manuel<br />
María Caballero, Saipina, Estancia Buena Vista, 6<br />
km NW del pueblo, vegetación en transición de<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
monte seco y sub<strong>and</strong>ino con presencia de<br />
Dodoneae viscose y Tipuana spp., 2400 m,<br />
18 o 03’18"S, 64 o 39’44"W, 2 Jan 1994, fl, J.<br />
Balcazar 11 (MO, 2 sheets). Peru. 1839-1840, in<br />
floral buds, M. Cl. Gay 486 (P). Entre Cusco y<br />
Abancay, Departamentos de Cusco y Apurímac,<br />
20 Nov 1947, fl, R. Ferreyra 2751 (US).<br />
APURÍMAC: 2600 m, 1909-1914, A. Weberbauer<br />
5840 (US). CUSCO: Prov. Anta, Río Apurimac<br />
drainage, dry inter-Andecan Valley, 2400-2500 m,<br />
below Limatamba, mostly along side road to<br />
Mollepata, 13 o 32’S, 72 o 30’W, tree 10 m, fls green,<br />
10 Jan 1984, fl, A. Gentry et al. 44106 (MO); Prov.<br />
Anta, Limatambo, Sisal-Cunyacc, stony habitat,<br />
2300 m, tree 6-8 m tall, 14 Mar 1963, past fr, C.<br />
Vargas C. 14342 (US). HUANCAVELICA: Prov.<br />
Tayacaja, road from Pampas to Rio Huanchuy,<br />
between milestone 19 km <strong>and</strong> 20 km from Pampas,<br />
S12 o 18’15", W74 o 50’78", 2669 m, dry slope with<br />
s<strong>and</strong>y soil, tall entirely leafless tree, in full fl, very<br />
rare, 20 Sep 2001, M. Wiegend et al. 5841 (BM).<br />
Central Electrica Mantaro, 4 Feb 1968, sterile, J.<br />
Soukup 5417 (US). TUMBES: Cerros de Amotape<br />
15-25 km SE <strong>of</strong> Cherrelique, premontane moist<br />
forest, along Quebrada Los Conejos, 04 o 09’S,<br />
80 o 37’W, 600-800 m, treelet 2 m, sterile, 9 Jun<br />
1987, A. Gentry & C. Diaz 58212 (MO).<br />
20. <strong>Aralia</strong> warmingiana (Marchal) J. Wen — Fig.<br />
51.<br />
<strong>Aralia</strong> warmingiana (Marchal) J. Wen, Brittonia<br />
45: 54. 1993. Coemansia warmingiana<br />
Marchal, Bull. Acad. Roy. Sci. Belgique 47:95.<br />
1879. Coudenbergia warmingiana (Marchal)<br />
Marchal, Bull. Acad. Roy. Sci. Belgique<br />
47:514. 1879. Pentapanax warmingianus<br />
(Marchal) Harms in Engler & Prantl, Nat.<br />
Pflanzenfam. 3(8): 56. 1898. TYPE: Brazil.<br />
MINAS GERAES: Near LagoaSanta, 22 Aug,<br />
E. Warming s.n. (holotype: BR).<br />
Pentapanax ulei Harms, Bot. Jahrb. 42:237. 1908.<br />
Coudenbergia ulei (Harms) C. B. Shang in Y.<br />
W. Yuan & al. (eds.), Proc. Intern. Symp. Bot.<br />
Gard. 1990: 632. 1990. TYPE: Brazil.<br />
BAHIA: Catinga near Tambury, Oct 1906, in<br />
flower, largely unopen, pedicels clearly<br />
articulated, E. Ule 7073 (lectotype, K!, here<br />
designated) [the lectotype at K bears Harms’<br />
h<strong>and</strong>writing “Pentapanax ulei Harms, n. sp.”
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
Fig. 51. <strong>Aralia</strong> warmingiana (Marchal) J. Wen. A. Leaf. B. inflorescence. C. Infructescence. D. Stem with the base <strong>of</strong> an<br />
inflorescence. E. Close-up <strong>of</strong> lower leaflet surface <strong>and</strong> margin. F. Leaf segment. G. Floral bud. H. Flower. I. Flower after<br />
anthesis. J. Fruit. K. Fruit, apical view (A, B, E & G-I – Stutz 2280, US; C, F, J & K – Smith 14120, US; D - Filho, Martins<br />
& Stubblebine 13280, E).<br />
139
140<br />
Ule’s main set went to Berlin <strong>and</strong> was destroyed<br />
in the World War II.]<br />
<strong>Aralia</strong> fluminensis Glaziou, Bull. Soc. Bot. France<br />
56 (3): 332. 1909. TYPE: Brazil: RIO<br />
JANEIRO: Rio Janeiro, Feb 1874, A. Glaziou<br />
6559 (lectotype, P!, here designated;<br />
isolectotype, K!)<br />
Tree 5-30 m tall, dbh to 40 cm, hermaphrodite.<br />
Leaves 45-100 cm long, 40-90 cm wide, tripinnate,<br />
with 4-6 pairs <strong>of</strong> pinnae; stipules 5-10 mm long,<br />
4-6 mm wide, adnate to the base <strong>of</strong> petiole, the<br />
free portion triangular, subcoriaceous, glabrous;<br />
petioles 15-45 cm long, purplish green; rachises<br />
green, turning purplish, subtended by a pair <strong>of</strong><br />
accessory 3-5-foliolate pinnae at the base <strong>of</strong> the<br />
first two lower pinnae <strong>and</strong> a pair <strong>of</strong> accessory<br />
leaflets at the upper pinnae; leaflets 3-6.5 cm long,<br />
1.5-3.5 cm wide, glabrous, acuminate at apex,<br />
rounded at base, sparsely serrate at margin, adaxial<br />
surface green, abaxial surface light green, petiolule<br />
articulated below the blade, that <strong>of</strong> lateral leaflets<br />
4-8 mm long, that <strong>of</strong> terminal leaflets 13-20 mm<br />
long, glabrous. Inflorescence terminal, consisting<br />
<strong>of</strong> an aggregation <strong>of</strong> 2-5 panicles on a short axis,<br />
each panicle 13-23 cm long, 10-15 cm wide, with<br />
8-20 umbels racemosely arranged on a main axis,<br />
main axis with light brownish scales, appearing<br />
rough, each umbel with a thick peduncle, peduncle<br />
4-7 cm long, 1.5-2 mm thick, with light brownish<br />
scales, thus appearing rough, bracts <strong>and</strong> bracteoles<br />
persistent, glabrous, primary bracts subtending the<br />
individual umbels 4-8 mm long, 5-8 mm wide,<br />
triangular, coriaceous, bracteoles 1.7-3 mm long,<br />
1.2-1.5 mm wide, narrowly triangular, glabrous.<br />
Flowers appearing at the same time as young leaves<br />
open; umbels 25-40-flowered, pedicels 4-7 mm<br />
long at anthesis, 10-13 mm long when fruiting,<br />
glabrous, stout, articulated near the base <strong>of</strong> the<br />
flower; sepals 6-8, minute, triangular; corolla<br />
cream, petals 7-8, 2.4-2.6 mm long, 1.1-1.2 mm<br />
wide, narrowly ovate; filaments 2.1-2.5 mm long,<br />
anthers 0.8-1 mm long, oblong; ovaries 6-8-locular.<br />
Fruits 5-6 mm long, 4-5 mm wide, ovoid globose,<br />
blackish purple when mature, persistent styles 1.8-<br />
2.1 mm long at the fruiting stage, stout, slightly<br />
divided at the tip, connate 4/5 at the lower part.<br />
Common names: salbugueiro falso <strong>and</strong><br />
salbugueirão (Minas Gerais, Brazil), <strong>and</strong> caroba<br />
(Paraguay).<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
Phenology: flowering from March to<br />
December; fruiting from October to January.<br />
Distribution: Brazil, Paraguay, <strong>and</strong> Argentina<br />
(Fig. 52).<br />
Ecology: in dry deciduous caatinga forest on<br />
calcareous hills; 150-300 m.<br />
Additional specimens examined: Argentina.<br />
FORMOSA: Territurro de Formosa, Guayenlec,<br />
arbol giganterio, fr, P. Jórgensen 3254 (MO, US).<br />
Brazil. MINAS GERAIS: Vicosa, 11 Jan 1935,<br />
sterile, J. G. Kuhlmann 54822 (NY); Sagarana,<br />
Projeto Integrato de Colonizacao (INCRA), NW<br />
Minas Gerais, c. 16 o S, 47 o W, wood very weak <strong>and</strong><br />
useless, no good for firewood or charcoal, flowers<br />
with same smell as Hedera helix, 19 Sep 1985, fl,<br />
J. A. Ratter et al. 5047 (K, 2 sheets, NY, 2 sheets);<br />
Pasmado (mun. Itaobim), Minas Gerais, 200 m,<br />
14 Jun 1986, fl, G. Hatschbach & J. M. Silva 50381<br />
(K, 2 sheets, MEXU, NY, US). APA Carste de<br />
Lagoa Santa, Lagoa Santa, 3 Mar 1996, A. E. Brina<br />
& L. V. Costa 32.691 (K, 2 sheets). PARAÍBA:<br />
4.5 km de Mata Gr<strong>and</strong>e, na Estrada para Bonito de<br />
Santa Fé, contacto floresta-caatinga, 30 Nov 1971,<br />
fl, Acad. Bras. de Ciências 1101 (MEXU).<br />
PARANÁ: Londrina, Floresta Godoy, proximo<br />
parc. 5 e 14 da porcao norte, 18 Mar 1994, fl, tree<br />
30 m tall, F. Chagas et al. 01682 (K); Parque<br />
Estadual Mata dos Godoy, 3 Mar 1989, fl, F.<br />
Chagas & Silva 00000, F UEL No. 10742 (K). RIO<br />
JANEIRO: Rio Janeiro, 28 Oct 1878, vegetative,<br />
Glaziou 10895 (K, LE, P, 2 sheets); Rio Janeiro,<br />
Engenho Noro, 8 Aug 1873, Glaziou s.n. (P); Rio<br />
Janeiro, Imbolica, 2 Aug 1891, Glaziou 19416a<br />
(P). SANTA CATARINA: Mun. Itapiranga, forest<br />
above Rio Uruguai, Barra Macaco Branco, ca.<br />
27 o 10’S, 53 o 46’W, 150-250 m, 18 Dec 1964, fl.,<br />
ovary 6-locular, 20 m tall, L. B. Smith & R. M.<br />
Klein 14120 (MO, NY, P, US). SÃO PAULO: Est.<br />
de S. Paulo, Mun. de S. Simão, faz. Aritozina, 29<br />
Jan 1982, in fl., H. F. Leitao Filho, F. R. Martins<br />
& W. H. Stubblebine 13280 (E). Paraguay. Chino,<br />
tree 15 m tall, Oct 1879, in fr., B. Balansa 3758<br />
(BM, K, P, 2 sheets). ALTO PARANÁ: Para<br />
Parayguazu route, C.F.A.P. (Ciudad del Este, Alto<br />
Paraná), Puerto Stroessner, Km 12, 2 Jan 1985, fr,<br />
arbre 23 m, L. C. Stutz O. 2280 (US). SAN<br />
PEDRO: Primavera, small sterile tree, 3 Jun 1961,<br />
A. L. Woolston 1260 (K, NY). On the Parana 26 o -<br />
27 o S lat., D. Parodi 4/83 (K).
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
Fig. 52. Map <strong>of</strong> South America showing the distribution <strong>of</strong> <strong>Aralia</strong> warmingiana (Marchal) J. Wen.<br />
141
142<br />
Three syntypes were cited by Glaziou when he<br />
described <strong>Aralia</strong> fluminensis (Glaziou 1909, p.<br />
332) <strong>and</strong> they are Glaziou 6559, 1895, <strong>and</strong><br />
109416a. I have seen the following Glaziou<br />
collections: Glaziou 6559 (K, P), Glaziou 10895<br />
(K, LE, P, 2 sheets), Glaziou 19416a (P), <strong>and</strong><br />
Glaziou s.n. (P) collected from Engenho Noro <strong>of</strong><br />
Rio Janeiro, 8 Aug 1873 (see specimens cited for<br />
detailed information <strong>of</strong> the four collections). All<br />
<strong>of</strong> these specimens bear the h<strong>and</strong>writing <strong>of</strong> Glaziou<br />
“<strong>Aralia</strong> fluminensis Glaz.” The specimens clearly<br />
represent <strong>Aralia</strong> warmingiana <strong>and</strong> they are all<br />
vegetative b,earing no flowers nor fruits, as the<br />
original description says. I suspect Glaziou 1895<br />
<strong>and</strong> 109416a in the original description (Glaziou<br />
1909) were misprinted <strong>and</strong> they actually represent<br />
Glaziou 10895 <strong>and</strong> 19416a. Because <strong>of</strong> the<br />
confusion concerning two <strong>of</strong> the three syntypes, I<br />
herein select Glaziou 6559 as the lectotype for<br />
<strong>Aralia</strong> fluminensis.<br />
21. <strong>Aralia</strong> rex (Ekman ex Harms) J. Wen — Fig.<br />
53.<br />
<strong>Aralia</strong> rex (Ekman ex Harms) J. Wen, Brittonia<br />
45: 53. 1993<br />
Megalopanax rex Ekman ex Harms, Notizbl. Bot.<br />
Gart. Berlin 9: 122. 1924. TYPE: Cuba.<br />
HABANA: Lomas de Camoa, 25 km from<br />
Habana, 29 Sep 1923, fl, E. L. Ekman 17551<br />
(lectotype, K!, here designated; isolecotypes,<br />
BM!, MO!, NY!, 2 sheets, US!).<br />
Tree 5-13 m tall, hermaphrodite. Leaves ca.<br />
100 cm long, tripinnate, to pinnate near the<br />
inflorescence, with 3-4 pairs <strong>of</strong> pinnae; stipules<br />
adnate to the base <strong>of</strong> petiole, lanceolate,<br />
subcoriaceous, glabrous; petioles 10-20 cm long;<br />
the primary pinnae subtended by a pair <strong>of</strong> accessory<br />
3-5-foliolate pinnae or a small bipinnate structure<br />
on the at the base <strong>of</strong> the lowermost pinnae; leaflets<br />
4.8-8 cm long, 2.4-4.3 cm wide, glabrous,<br />
membranaceous to chartaceous, acuminate at apex,<br />
subcordate to rounded at base, entire at margin,<br />
adaxial surface green, abaxial surface light green,<br />
petiolule articulated below the blade, that <strong>of</strong> lateral<br />
leaflets 2-8 mm long, that <strong>of</strong> terminal leaflets 10-<br />
30 mm long, glabrous. Inflorescence paniculate<br />
with a thick main axis, main axis 13-20 cm long,<br />
glabrous, brown, conspicuously lenticellate, with<br />
ca. 2 whorls <strong>of</strong> umbels, each whorl with 10-20<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
umbels; each umbel with a thick peduncle,<br />
peduncle 7-14 cm long, 2.5-5 mm thick, articulated<br />
near the middle, rough <strong>and</strong> lenticellate; bracts <strong>and</strong><br />
bracteoles persistent, glabrous, primary bracts<br />
subtending the individual umbels 4-6 mm long, 3-<br />
5 mm wide, triangular, coriaceous, bracteoles 1-<br />
1.5 mm long, 0.7-1 mm wide, narrowly triangular,<br />
glabrous. Flowers appearing after the opening <strong>of</strong><br />
young leaves; umbels 90-120-flowered, pedicels<br />
9-13 mm long at flowering stage, glabrous, slender,<br />
articulated at the base <strong>of</strong> the flower. Sepals minute,<br />
triangular, 4-7-toothed on the calyx ring; petals 7-<br />
10, 3.5-5 mm long, 1.5-2 mm wide, narrowly<br />
oblong to lanceolate; filaments 4-5 mm long,<br />
anthers 2-3 mm long, oblong; ovaries 7-10-locular.<br />
Fruits 6-7 mm in diameter, subglobose, with 6-8<br />
ribs when dry, styles stout, persistent stigmas<br />
divergent.<br />
Phenology: flowering in September; fruiting in<br />
November.<br />
Distribution: endemic to Cuba (Havana <strong>and</strong><br />
Cienfuegos) (Fig. 54).<br />
Ecology: on limestone rocks, mountain forests,<br />
thickets or low forests; ca. 450 m.<br />
Conservation: a rare species in need <strong>of</strong><br />
conservation.<br />
Additional specimens examined: Cuba.<br />
CIENFUEGOS: Topaz de Colantes, W80 o 15’,<br />
N21 o 55’, 16 Nov 1988, sterile, 450 m, mountain<br />
forest, trees 5-13 m tall, R. Dechamps & R. Carrera<br />
12190 (MO). HABANA: Lomas de Camoa, on<br />
naked limestone rocks, in thickets or low forests,<br />
6 Nov 1921, E. L. Ekman 13451 (NY); 22 Nov<br />
1921, E. L. Ekman 13523 (BM, MO).<br />
Harms (1924) cited three specimens when<br />
describing Ekman’s Megalopanax rex (Ekman<br />
17551, 13451 <strong>and</strong> 13523). The original specimens<br />
were most likely at B <strong>and</strong> were destroyed during<br />
WWII. Ekman 17551 bears many flowers <strong>and</strong> is<br />
most widely distributed. The Kew specimen <strong>of</strong><br />
Ekman 17551 bears Harms’ h<strong>and</strong>writing <strong>and</strong> is thus<br />
selected as the lectotype here.<br />
22. <strong>Aralia</strong> excelsa (Griseb.) J. Wen — Fig. 55;<br />
color plate 15: A-F.<br />
<strong>Aralia</strong> excelsa (Griseb.) J. Wen, Cathaya 13/14:<br />
96. 2002, non Linden, Cat. Pl. Exot. 11: 34.
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
Fig. 53. <strong>Aralia</strong> rex (Ekman ex Harms) J. Wen. A. Leaf. B. Inflorescence. C. Lower leaflet surface. D. Tip <strong>of</strong> peduncle<br />
showing the thickened joint area <strong>of</strong> the pedicels <strong>and</strong> the peduncle. E. Floral bud. F. Floral bud. G. Flower. I. Flower. J.<br />
Flower after anthesis.<br />
143
144<br />
Fig. 54. Map <strong>of</strong> Cuba showing the distribution <strong>of</strong> <strong>Aralia</strong> rex.<br />
1856, nom. nud. Sciadodendron excelsum<br />
Griseb,, Bonpl<strong>and</strong>ia 6: 7. 1858. TYPE:<br />
Panama. Duchassaing s.n. (lectotype: GOET!,<br />
here designated).<br />
Reynoldsia americana Donn.Sm., Bot. Gaz. 49:<br />
455. 1910. TYPE: Costa Rica. Peninsula<br />
Nicoya, tree <strong>of</strong> medium height, nomen<br />
vernaculare “Chile”, Jan 1900, A. Tonduz<br />
13823 (holotype: US!, photo at NY; isotype:<br />
US!).<br />
Pentapanax granatensis Rusby, Desc. S. Amer. Pl.<br />
72. 1920. TYPE: Colombia. MAGDALENA:<br />
Santa Marta, near Masinga, 250 ft, a tree to 40<br />
ft or more, occasional in forest below 1500 ft,<br />
flowers March-April, white, flowers terminal<br />
10-20 short branchlets crowded together<br />
forming a large cluster, leaves deciduous about<br />
the time flowering, leaves 3-pinnate, 22 Mar<br />
1898-99, H. H. Smith 1595 (holotype: NY!;<br />
isotypes: E!, K!, P!, 2 sheets, PH!, U!, US!, 2<br />
sheets).<br />
Highly branched <strong>and</strong> unarmed tree (4) 8-30 m<br />
tall, dbh 20-50 cm, hermaphrodite, bark furrowed.<br />
Leaves 60-125 cm long, 50-100 cm wide,<br />
quadripinnately to tripinnately compound, or<br />
bipinnately compound when young, with 5-6 pairs<br />
<strong>of</strong> primary pinnae; stipules 10-13 mm long, 1.5-<br />
2.5 mm wide, adnate to the base <strong>of</strong> petiole,<br />
lanceolate, subcoriaceous, glabrous <strong>and</strong> more or<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
less ciliate at margin; petioles 20-50 cm long,<br />
purplish green; rachises green, turning purplish,<br />
subtended by a pair <strong>of</strong> accessory 3-5-foliolate<br />
pinnae at the base <strong>of</strong> each primary pinna; leaflets<br />
4-7 cm long, 1.5-4 cm wide, glabrous, apex<br />
acuminate, base rounded, margin sparsely serrate,<br />
adaxial surface green, abaxial surface light green,<br />
petiolule more or less articulated below the blade,<br />
that <strong>of</strong> lateral leaflets 0-5 mm long, that <strong>of</strong> terminal<br />
leaflets 10-20 mm long, glabrous. Inflorescence<br />
consisting <strong>of</strong> 1-6 panicles on a short branch, each<br />
panicle 7-15 cm long, 5-12 cm wide, with 10-20<br />
umbels racemosely arranged on a main axis, main<br />
axis glabrous, lower portion appearing rough <strong>and</strong><br />
with scales, each umbel with a peduncle 3.5-10<br />
cm long, 1-1.5 mm thick, glabrous, each subtended<br />
by a few coriaceious broadly triangular bracts,<br />
bracts <strong>and</strong> bracteoles persistent, glabrous, primary<br />
bracts subtending the individual umbels 3-6 mm<br />
long, 3.5-7 mm wide, triangular, coriaceous,<br />
bracteoles 1-1.4 mm long, 0.8-1.1 mm wide,<br />
narrowly triangular, glabrous. Flowers appearing<br />
before or at the same time as young leaves open;<br />
umbels 15-35-flowered, pedicels 4.5-8 mm long<br />
at flowering stage, 5-9 mm long when fruiting, nonarticulated<br />
below the flower, glabrous, stout.<br />
Sepals 9-10, minute, semicircular or<br />
inconspicuous; petals 9-10, 2.6-2.8 mm long, 1-<br />
1.5 mm wide, narrowly ovate, <strong>of</strong>ten calyptrate <strong>and</strong><br />
falling together, yellowish white; filaments 3-4 mm
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
Fig. 55. <strong>Aralia</strong> excelsa (Griseb.) J. Wen. A. Leaf with accessory pinna. B. Stem with inflorescences. C. Inflorescence. D.<br />
Floral bud. E. Flower after anthesis. F. Fruit. (A – Steyermark et al. 109907, NY; B - Moreno 25344, BM; C-E - Smith<br />
1595, NY; F – Folsom 3436, BM).<br />
145
146<br />
long, anthers 1.5-1.8 mm long, ca. 1 mm wide,<br />
narrowly oblong; ovaries 9-10-locular. Fruits 6-7<br />
mm long, 7-8 mm wide, subglobose, dark purple<br />
when mature, persistent styles slightly divided at<br />
the tip, largely connate, stylar column 1.2-1.5 mm<br />
long at the fruiting stage, stout.<br />
Vernacular names: calenturo (Colombia),<br />
cebratano, lagarto, <strong>and</strong> corroncho de lagarto (El<br />
Salvador), lagarto <strong>and</strong> palo de lagarto (Nicaragua),<br />
cedro macho (Chamela, Jalisco, Mexico), jobo de<br />
lagarto (Alligator Jobo), <strong>and</strong> mangabé (Panama),<br />
<strong>and</strong> “Chile” (Costa Rica).<br />
Uses: in Panama, the species was cultivated as<br />
a street tree; <strong>and</strong> the petioles were employed for<br />
making bird cages (St<strong>and</strong>ley 1928). The natives<br />
<strong>of</strong> Panama apply the macerated leaves with<br />
beneficial effect to ulcers. In Nicaragua, it was<br />
used as fence post; <strong>and</strong> the ashes <strong>of</strong> the wood are<br />
used for making soap.<br />
Phenology: flowering in January to July;<br />
fruiting in May to August.<br />
Distribution: Colombia, El Salvador, Haiti,<br />
Mexico (Jalisco, Colima, <strong>and</strong> Islas Marías), Costa<br />
Rica, Honduras, Nicaragua, Panama, <strong>and</strong><br />
Venezuela (Fig. 56).<br />
Ecology: Tropical lowl<strong>and</strong> dry forests, or dry<br />
forest – moist forest transitional areas, densely<br />
forested floodplains to more open hillsides (selva<br />
mediana); 0-860 m.<br />
Additional specimens examined: Colombia.<br />
Municipio Fonseca: Corregimiento Distraccion,<br />
sitio Las Casitas-El Socorro, bosque a 2 km, sitio<br />
modera damente pendiente con abundantes rocas,<br />
bien drenado, algo perturbado, parcela, 500 m,<br />
72 o 59.4’W, 10 o 9.1’N, 31 Aug 1990, sterile, O.<br />
Marul<strong>and</strong>a & J. Betancur 2128 (K, 2 sheets).<br />
Costa Rica. ALAJUELA: entre Guachipelin y<br />
Valean de La Vieja, 26 May 1932, sterile, A. M.<br />
Brenes 15535 (NY); along quebrada, simple shrub<br />
6 ft, El Coyolar, 240 m, Apr 1924, sterile, P. C.<br />
St<strong>and</strong>ley 40062 (US). GUANACASTE: Parque<br />
Nacional Santa Rosa Sendero Indio Desnudo,<br />
10 o 50’20"N, 85 o 37’00"W, 320 m, arbol de 16 m x<br />
40 cm DAP, 22 Jul 1992, sterile, Q. Jiménez et al.<br />
1085 (NY); pasture l<strong>and</strong> <strong>and</strong> deciduous forest now<br />
in full leaf in the areas <strong>of</strong> Bahia El Coco, Bahia<br />
Playa, Hermosa, <strong>and</strong> Sardinal, 0-150 m, 10 o 32’N,<br />
85 o 40’W, 29-31 Jul 1971, floral buds, W. C. Burger<br />
& M. Burger 7742 (NY); Santa Rosa, Parque<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
Nacional Santa Rosa, Dry Tropical Forest Research<br />
Center, near the cross <strong>of</strong> the big house to the<br />
administration, 10 o 50’17"N, 85 o 36’52"W, 310 m,<br />
20 Sep 2002, J. Wen & R. Aguilar 6779 (F, US).<br />
PUNTARENAS: Cantón de Garabito, Cuenca del<br />
Río Gr<strong>and</strong>e de Tárcoles, camino a Playa Guacalillo,<br />
09 o 51’30"N, 84 o 37’30"W, 100 m, arbol de 10 m x<br />
35 cm DAP, 28 Sep 1993, sterile, Q. Jiménez et al.<br />
1358 (NY, 2 sheets). El Salvador. Depto.<br />
Ahuachapán, San Francisco Menéndez, El Corozo,<br />
Mariposario, zona baja “Mariposario”, 200 m,<br />
13 o 49’N, 89 o 59’W, 18 May 2000, fr, J. M. Rosales<br />
787 (BM, MEXU); zona baja “Los Peralta”, 175<br />
m, 13 o 49’N, 89 o 59’W, 7 Jun 2000, fr, J. M. Rosales<br />
1270 (BM, MEXU). Region NE <strong>of</strong> Chalatenango,<br />
near Comalapa, Dept. Chalatenango, 1500 ft, 16<br />
Nov 1958, tree 60 ft, bark brittle, thick with<br />
conspicuous longitudinal corrugations, P. H. Allen<br />
& M. L. van Severen 7093 (NY, US). Laguna de<br />
Maquique, Departamento de la Unión, 60 m in<br />
altitude, 18 Feb 1922, sterile, P. C. St<strong>and</strong>ley 20932<br />
(NY, US); vicinity la Unión, 150 m, Feb 1922,<br />
sterile, P. C. St<strong>and</strong>ley 20829 (NY, US). Near Los<br />
Apoyos, Dept. Santa Ana, shrub or a small tree <strong>of</strong><br />
very h<strong>and</strong>some aspect, trunk straight, corky, not<br />
branched, 1924, sterile, S. Calderón 2171 (US).<br />
Haiti. 5 Jul 1927, fl, E. L. Ekman H7966 (K, NY);<br />
Plain Centrale, Hinche, ravine, Papaye, 225 m, 6<br />
May 1926, fr, E. L. Ekman H6009 (US, 2 sheets,<br />
one fr, one leaves). Massif de la Hotte, gr. Mome<br />
Rochclois, Miragoane, 30 Jul 1926, sterile, E. L.<br />
Ekman H6545 (US). Massif des Mathiux,<br />
l’Arcahaie, Montagne Terrible, northern slope, ca.<br />
700 m, limestone, 7 Feb 1926, fl, E. L. Ekman<br />
H5524 (US). Honduras. DEPT. COMAYAGUA:<br />
Matorrales humedos y cafetales cerca de Quebrada<br />
la Jutera, 3 km de la Libertad, 20 May 1956, 600<br />
m, A. Molina R. 7039 (NY). Mexico. COLIMA:<br />
selva camino Camotlán, 28 Jan 1958, sterile,<br />
Mir<strong>and</strong>a 8820 (MEXU). JALISCO: Mpio. La<br />
Huerta, Rancho Cuixmala, Cumbres 1, Arroyo<br />
Cajones, ca. 3 km inl<strong>and</strong> from the Puerto Vallarta<br />
– Barra de Navidad hwy, tropical deciduous forest<br />
with Lonchocarpus, Ruprechtia, Bursera, near lat.<br />
19 o 2 in arroyo, uncommon tree 15 m, 19 Aug 1991,<br />
fr, E. L. Lott 3802 (K, MEXU, RSA); Mpio. La<br />
Huerta, Estación de Biología Chamela (UNAM),<br />
19 o 30’N, 105 o 03’W, 26 May 1982, fl, arbol 8 m,<br />
corteza gris, fissurada, flores amarillas, E. J. Lott<br />
1055 (MEXU, 2 sheets); Mpio. La Huerta, Rancho<br />
Cuixmala, along the road into the Cumbres via the
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
Fig. 56. Map <strong>of</strong> Central America <strong>and</strong> northern South America showing the distribution <strong>of</strong> <strong>Aralia</strong> excelsa (Griseb.) J. Wen.<br />
Rio Cuitzmala, 9 km above the Hwy 200 bridge<br />
<strong>and</strong> 5 km above the river, in a heavily wooded<br />
canyon, 104 o 55’W, 19 o 30’N, 150 m, tropical semideciduous<br />
forest, a scarce 10-15 m high tree, 50<br />
cm dbh, on the canyon bottom, bark gray, deeply<br />
furrowed <strong>and</strong> somewhat corky, 13 Sep 1991, fr, A.<br />
C. S<strong>and</strong>ers 11212 (RSA); Camino Antiguo a 100<br />
m de la Vereda Tejón, arbol 4 m, botones verdes, 1<br />
Apr 1982, floral buds, E. J. Lott 964 (MEXU);<br />
Cumbres de Cuixmala, el 45 camino a Cumbres I,<br />
Municipio La Huerta, 19 o 26’00"N, 104 o 58’45"W,<br />
100 m, selva mediana subcaducifolia, primaria,<br />
suelo pardo arenoso, arbol 25 m, abundante, fruto<br />
seco, 25 Aug 1988, fr, R. Acevedo R. & J. L.<br />
Martinez 965 (MEXU); Estación de Biología<br />
UNAM, Chamela, Mpio. La Huerta Camino<br />
Antiguo Norte 150, 13 Dec 1989, sterile, S. H.<br />
Bullock 2080 (K, MEXU); 2 Jul 1978, fl, J. A. S.<br />
Magallanes 1110 (MEXU); arbol 18 m, 28 Apr<br />
1974, fl, L. A. Pérez J. & M. Pérez G. 834 (MEXU);<br />
L. A. Pérez J. 111 (MEXU); arbol 17 m, 8 Aug<br />
1974, fr, L. A. Pérez J. 958 (MEXU); ca.<br />
Campamento, arbol 15 m, botones Amarillo-<br />
147<br />
verdosas, 2 Jul 1978, fl, L. A. Pérez J. 1770<br />
(MEXU); selva mediana sub-caducifolia, suelo<br />
metamorfico, 16 Aug 1977, fr, J. A. S. Magallanes<br />
777 (MEXU). NAYARIT: Islas Marías, Isla Ma.<br />
Magdalena, hacia el SW (centro de la isla),<br />
021 o 27’N, 106 o 27’W, arbol 3 m, selva baja<br />
caducifolia, 27 Nov 1986, sterile, F. Chiang C.<br />
1085 (RSA). Nicaragua. U.S. North Pacific<br />
Exploring Expedition, 1853-56, fl, C. Wright 8 (P,<br />
US). CARAZO: area de la Estación Biológica de<br />
Chacocentro, en el limite departamental Carazo-<br />
Rivas, bosque seco tropical, 0-100 m, 22-23 Jun<br />
1984, fr, D. Soza, A. Grijalva & M. Ar<strong>and</strong>a 81<br />
(BM). CHONTALES: 6.3 km from Hwy on road<br />
to Cuapa, 12 o 11’N, 85 o 25’W, 170 m, dry rocky<br />
hills, 13 Jun 1984, fr, W. D. Stevens 22963 (BM,<br />
MEXU, NY). MANAGUA: along a dry wash near<br />
km 8, Carretera Sur, outskirts <strong>of</strong> Managua, tropical<br />
dry forest – moist transition, 200 m, 19 Oct 1976,<br />
D. Neill 1082 (BM, 2 sheets); Carretera a<br />
Montelimar, comarca Aduana, al Norte del rio<br />
Aduana, 80-100 m, 21 Jul 1980, in fl., M. Guzman,<br />
D. Castro & A. Montiel 398 (BM); ca. 5 km NNW
148<br />
<strong>of</strong> Hwy 12 along road on ridge <strong>of</strong> Sierra de<br />
Mateare, 12 o 07’N, 86 o 23’W, 420 m, roadside <strong>and</strong><br />
slope, fence-post tree, 28 Jan 1978, W. D. Stevens<br />
6192 (BM, MEXU, NY). MASAYA: Jardín<br />
Botánico UCA, km 17.5 carretera Managua-<br />
Masaya, bosque seco, 2 Sep 1982, J. C. S<strong>and</strong>ino<br />
3492 (BM); Parque Nacional Volcan Masaya, dry<br />
forest near west shore <strong>of</strong> Laguna de Masaya, 140<br />
m, 10 Apr 1978, fl, D. Neill 3458 (BM).<br />
MATAGALPA: entre “Sta. Juana” y el “Rincon<br />
del Diablo” (camino a “Puertas Viejas”), 12 o 32-<br />
34’N, 85 o 57-59’W, 860 m, 6 Apr 1983, in young<br />
fr, M. Araquistain 3478 (BM); “La Mojada,” 10<br />
km al SO de Esquipulas, 12 o 37’N, 85 o 51’W, 400<br />
m, 24 Jan 1985, floral buds, P. P. Moreno 25344<br />
(BM). Panama. At Turbaco, Apr 1862, fl, S.<br />
Hayes 734 (BM, K). CANAL ZONE, 1923, sterile,<br />
H. Johansen 15 (US), H. Johansen 43 (US);<br />
Balboa, P. C. St<strong>and</strong>ley 25481 (US), 25558 (US),<br />
32104 (US, 2 sheets). CHIRIQUI: Km 103 W <strong>of</strong><br />
Panama City, tree 15 m in fence row, mature leaves<br />
to 75 cm, 20 Feb 1978, fl, B. Hammel 1604 (BM,<br />
NY). COCLE: Rio Hato, street trees, 27 May<br />
1977, sterile, J. F. Folsom 3436 (BM); pasture<br />
along the Rio Hato at Pan. Am. Hwy bridge,<br />
scattered trees <strong>and</strong> fringe <strong>of</strong> forest along rio, alt.<br />
20 m, tree 10 m tall, 30 cm dbh, twigs thick, s<strong>of</strong>t,<br />
one branch with huge compound leaves, tree<br />
otherwise leafless with new leaves beginning to<br />
come out from tips <strong>of</strong> twigs, flowers yellow green,<br />
with musty odor, 19 Mar 1974, late fl, M. Nee<br />
10784 (US). PANAMÁ: Río Las Lajas, alt. ca. 20<br />
m, tree 12 m, flowers greenish white, 5 Feb 1939,<br />
fl, no leaves, P. H. Allen 1604 (NY, US); along<br />
Interamerican Hwy 2 miles west <strong>of</strong> turn<strong>of</strong>f to El<br />
Valle, tree 17 m, flowers cream, fruits in huge<br />
globular clusters, becoming purple, sweet <strong>and</strong><br />
fleshy, 16 May 1971, fl & fr, T. B. Croat 14629<br />
(RSA, 2 sheets); Río Tecumen, moist thicket, shrub<br />
or tree 10-20 ft, frequent, 3 Jan 1924, sterile, P. C.<br />
St<strong>and</strong>ley 29415 (US); Las Sabanas, 4 Dec 1923,<br />
sterile, “mangabé”, open slope, tree 15-40 ft,<br />
common, petioles used to make bird cages, P. C.<br />
St<strong>and</strong>ley 25892 (US); Río Tapia, moist forest, shrub<br />
or tree 10-20 ft, P. C. St<strong>and</strong>ley 28218 (US); Taboga<br />
Isl<strong>and</strong>, tree 20 ft, frequent, brushy slope, “Jobo de<br />
lagarto”, Dec 1923, sterile, P. C. St<strong>and</strong>ley 27858<br />
(US); St<strong>and</strong>ley 27023 (US); near Matías<br />
Hernández, moist thicket, shrub 8-12 ft, 30 Dec<br />
1923, sterile, P. C. St<strong>and</strong>ley 28941 (US); Nuevo<br />
San Francisco, 12 Jan 1924, sterile, P. C. St<strong>and</strong>ley<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
30706 (US); between Las Sabanas <strong>and</strong> Matías<br />
Hernández, in thicket, shrub or tree 10-20 ft,<br />
common, 21 Jan 1924, sterile, P. C. St<strong>and</strong>ley 31813<br />
(US). Chepo, trunk 35 ft, 24 inches in diameter,<br />
“Jobo de lagarto”, 1924, fl, no leaves, H. C. Kluge<br />
50 (US). Venezuela. ARAGUA: Parque Nacional<br />
Pittier, along road between El Limon <strong>and</strong> Guamita,<br />
1 km below Guamita, 700 m, 20 Oct 1961, sterile,<br />
J. A. Steyermark 89765 (NY, US). GUÁRICO:<br />
Morros de San Juan, Jun 1960, sterile, L.<br />
Aristeguieta 4232 (US); Altagracia de Orituco-<br />
Aimará, Mar 1966, fl, L. Aristeguieta 6047 (NY,<br />
US); Hacia San Francisco de Macaira, entre<br />
Altagracia de Orituco y Tamanaco, Jun 1966, fr,<br />
arbol de unos 8 m, frutos esféricos, de color verde<br />
oscuro cu<strong>and</strong>o maduros, con pulpa azucarada,<br />
comido por los páraros, L. Aristeguieta 6157 (NY,<br />
U, US). LARA: Districto Palavecino, terreno<br />
plano en sabanas altern<strong>and</strong>o con matorrales y selva<br />
tropófila, entre Sarare y El Altar, 350 m, 2 Jun<br />
1974, J. A. Steyermark, R. Smith, S. Nehlin & M.<br />
Lobo 109907 (NY, U, 2 sheets). YARACUY: a 5<br />
km este de Yaritagua, 1 Apr 1980, fl, N. Romírez<br />
367 (K).<br />
N. Romírez 367 (K) from Venezuela shows<br />
transitional pedicel articulation. The tip <strong>of</strong> pedicel<br />
has a slight constriction, but is not articulated.<br />
Grisebach (1858) cited that Duchassaing’s<br />
name “Arbor excelsa,” which apparently represents<br />
a misprint. The lectotype from GOET bears the<br />
name “<strong>Aralia</strong> excelsa” below Grisebach’ writing<br />
“Sciadodendron excelsum.” The type is a<br />
collection made after the flowering with petals<br />
already falling <strong>of</strong>f. The ovaries on the type are<br />
clearly 10-12-locular, as also indicated on the type<br />
specimen by Grisebach.<br />
I am aware that Linden (1856) used <strong>Aralia</strong><br />
excelsa as a nomen nudum. Hooker (1862)<br />
mentioned that the name <strong>Aralia</strong> excelsa was used<br />
for the plant Leea coccinea Planch. (Leeaceae) in<br />
the horticulture trade with no authority. As the<br />
nomen nudum by Linden has no nomenclatural<br />
st<strong>and</strong>ing, the new combination <strong>of</strong> <strong>Aralia</strong> excelsa<br />
(Griseb.) J. Wen is the correct name for this widely<br />
distributed taxon <strong>of</strong> <strong>Aralia</strong>ceae in Central America<br />
extending to South America.<br />
The specimen Bernardi 7382 (K, NY) collected<br />
from Bolívar, Venezuela clearly has an articulation<br />
below the flower. It was collected in floral bud<br />
condition <strong>and</strong> appeared to be 5-merous in floral
parts, but each petal seemed to split into two. Like<br />
A. warmingiana, the speciemen has an articulation<br />
at the pedicel tip, yet the flowers seem to be smaller<br />
than those <strong>of</strong> <strong>Aralia</strong> warmingiana. Geographically<br />
it is out <strong>of</strong> the normal distributional range <strong>of</strong> A.<br />
warmingiana. The inflorescence <strong>and</strong> floral<br />
morphology are similar to those <strong>of</strong> A. excelsa,<br />
except that they differ in the pedicel articulation.<br />
This collection may represent a new species, but<br />
more collections are needed to confirm its identity.<br />
Its collection data are as below: Venezuela, bosque<br />
de San Mateo, El Cristo-La Paragua, Estado<br />
Bolivar, 300 m, 7 Mar 1959, floral buds, no leaves<br />
yet, A. L. Bernardi 7382 (K, NY).<br />
23. <strong>Aralia</strong> bahiana J. Wen, sp. nov. — Fig. 57.<br />
<strong>Aralia</strong> bahiana J. Wen. TYPE: Brazil. BAHIA:<br />
Mun. de Itiuba, Fazenda Experimental da<br />
Epaba, arvore de approximadamente 6 m de<br />
altura, fuste retilineo com casca lembr<strong>and</strong>o o<br />
pau-d’arco-roxo, madeira mole, estepe arborea<br />
densa e floresta estacional decidual,<br />
afroamentos rochosos, 10 o 43’S, 39 o 50’W, 26<br />
May 1983, young fr, G. C. P. Pinto & H.<br />
Bautista 91/83 (holotype: US!; isotype: NY!).<br />
Arbor. Folia ampla tripinnata, usque 50-70 cm<br />
longa, foliola glabra, ovata, apice acuminata, basi<br />
rotundata, margine parce serrulata. Umbellae 15-<br />
35-florae, pedicellis inarticulatis. Fructus oblongoglobosus,<br />
stylis 8-9.<br />
Branched <strong>and</strong> unarmed tree 6-7 m tall. Leaves<br />
50-70 cm long, 50 cm wide, tripinnately<br />
compound; stipules 6-7 mm long, 1.2-1.5 mm<br />
wide, adnate to the base <strong>of</strong> petiole, lanceolate,<br />
subcoriaceous, glabrous; petioles 15-20 cm long;<br />
each pinna subtended by a pair <strong>of</strong> accessory 3-5foliolate<br />
pinnae; leaflets 3-5 cm long, 1.5-2.5 cm<br />
wide, glabrous, ovate, acuminate at apex, rounded<br />
at base, sparsely <strong>and</strong> inconspicuously serrulate at<br />
margin with a few teeth, adaxial surface green,<br />
abaxial surface light green, petiolule more or less<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
149<br />
articulated below the blade, that <strong>of</strong> lateral leaflets<br />
0-2 mm long, that <strong>of</strong> terminal leaflets 10-15 mm<br />
long, glabrous. Inflorescence consisting <strong>of</strong> 4-5<br />
umbellate units on a short branch, each unit with<br />
5-10 umbels racemosely arranged on a main axis,<br />
main axis 4-6 cm long, glabrous, appearing rough<br />
<strong>and</strong> with scales, each umbel with a peduncle 4-6<br />
cm long, 2.5-3 mm thick, glabrous <strong>and</strong> more or<br />
less rough, each subtended by a few bracts at the<br />
base <strong>and</strong> ca. 0.5-1 cm from the base, bracts <strong>and</strong><br />
bracteoles persistent, glabrous, primary bracts<br />
subtending the individual umbels 4-6 mm long, 5-<br />
7 mm wide, triangular, coriaceous, bracteoles 1.8-<br />
2.2 mm long, 1.2-1.5 mm wide, narrowly<br />
triangular, glabrous. Umbels 15-35-flowered,<br />
pedicels 6-13 mm long, glabrous, stout, nonarticulated<br />
below the flower. Sepals 8-9, minute,<br />
inconspicuous; petals 8-9, 4-4.5 mm long, 1.2-1.3<br />
mm wide, narrowly ovate, seemingly noncalyptrate,<br />
white; filaments 3.5-4 mm long, anthers<br />
1.6-1.9 mm long, 0.9-1 mm wide, narrowly oblong;<br />
ovaries 8-9-locular, persistent sepals 8-9, minute,<br />
inconspicuous. Fruits 7-8 mm long, 6-8 mm wide,<br />
ovoid globose to globose, persistent styles slightly<br />
divided at the tip, largely connate, stylar column<br />
1-1.5 mm long.<br />
Vernacular name: tingui-bravo.<br />
Phenology: with a few flowers in late April,<br />
<strong>and</strong> in young fruits from late April to late May.<br />
Distribution: Bahia, Brazil (Fig. 50).<br />
Ecology: Estepe Arborea Densa e Floresta<br />
Estacional Decidual.<br />
Additional specimens examined: Brazil.<br />
Bahia: Município de Santa Terezinha, ca. 5 km W<br />
da Estrada Santa Terezinha-Itatim, em uma Estrada<br />
vicinal dist<strong>and</strong>o ca. 14 km E de Itatim, 26 Apr 1994,<br />
in fruit & with a few flowers, L. P. de Quiroz & N.<br />
S. Nascimento 3853 (K, 3 sheets).<br />
<strong>Aralia</strong> bahiana is closely related to A. excelsa,<br />
sharing the character <strong>of</strong> the non-articulated pedicel.<br />
It is distinguished from the latter by the very stout<br />
peduncle <strong>and</strong> the ovoid globose fruits <strong>of</strong> the new<br />
species.
150<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
Fig. 57. <strong>Aralia</strong> bahiana J. Wen. A. Branch with leaves showing leaf architecture. B. Branch showing young inflorescence on<br />
old branch. C. Young infructescence. D. Young fruit. E. Older fruit (A-E - Pinto & Bautista 91/83, US).
<strong>Aralia</strong> sololensis Donnell-Smith, Bot. Gaz. 56: 58.<br />
1913. TYPE: Guatemala. Prope Patulul, Dept.<br />
Sololá, Feb 1906, fl, W. A. Kellerman 5828<br />
(holotype: US!; isotype: OS!)<br />
= Sambucus simpsonii Rehder (Adoxaceae)This<br />
This study was supported by the National<br />
Science Foundation (DEB-0196051, <strong>and</strong> DEB-<br />
0108536), the Institute <strong>of</strong> Botany <strong>of</strong> the Chinese<br />
Academy <strong>of</strong> Sciences, the Field Museum, the<br />
Smithsonian Institution (the Small Grants Program<br />
<strong>of</strong> the National Museum <strong>of</strong> Natural History, the<br />
Botany Collections Fund, the Walcott Fund, <strong>and</strong><br />
the Laboratory <strong>of</strong> Analytical Biology), <strong>and</strong> the John<br />
D. <strong>and</strong> Catherine T. MacArthur Foundation. Field<br />
<strong>and</strong>/or herbarium assistance was kindly provided<br />
by many colleagues <strong>and</strong> friends: Susan Agnew,<br />
Reinaldo Aguilar, Qing Ban, Manzhu Bao, John<br />
Beaman, Stephan Beck, John Boggan, Zhiyu Cao,<br />
A. S. Chauhan, Martin Cheek, Hongfeng Chen,<br />
Longqin Chen, Yansheng Chen, Zhiyuan Chen,<br />
Fern<strong>and</strong>o Chiang, Nancy Coile, A. P. Das, J. I.<br />
Davis, Yunfei Deng, Michael Dillon, Hanmin<br />
Dong, James Dong, Michael Dong, Larry Dorr,<br />
Barbara Ertter, Kuomei Feng, David Frodin, Dezhi<br />
Fu, Song Ge, Gang Hao, James W. Hardin, M.<br />
Hasebe, De-yuan Hong, Chiming Hu, Pu-hua<br />
Huang, Sovanmoly Hul, Stefanie Ickert-Bond,<br />
Hugh H. Iltis, M. Ito, Yunsheng Jiang, Samuel B.<br />
Jones, Jr., Shusheng Lai, Thomas G. Lammers, Dezhu<br />
Li, Heng Li, Liangqian Li, Na Li, Rong Li,<br />
Xiangping Li, Xuedong Li, Qi Lin, Yourun Ling,<br />
Ho-Yih Liu, Yan Liu, Zhengyu Liu, Pete Lowry,<br />
Zhengwei Lu, Xiaozhong Luo, Yibo Luo, A. A.<br />
Mao, Esteban Martinez, Lucinda McDade, Ying<br />
Meng, Michael Nee, Bing Ni, Zelong Nie,<br />
Christine Niezgoda, Simon Owens, Ze-hui Pan,<br />
Arun K. P<strong>and</strong>ey, Jose Panero, M. K. Pathak, Ching-<br />
I Peng, Mike Powell, Alan Prather, John Pruski,<br />
Haining Qin, Jim Reveal, Rusty Russell, M.<br />
Sanjappa, Ed Schilling, Suhua Shi, N. P. Singh,<br />
Larry Skog, Jim Soloman, David M. Spooner, Tod<br />
Stuessy, Hang Sun, Hui Tang, Guangfu Tao, Roy<br />
Vickery, Jose L. Villasenor R., Huilin Wan,<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
EXCLUDED NAMES<br />
ACKNOWLEDGEMENTS<br />
151<br />
species is clearly not a member <strong>of</strong> <strong>Aralia</strong>ceae<br />
<strong>and</strong> was excluded from the New World <strong>Aralia</strong><br />
(see Smith 1936; St<strong>and</strong>ley <strong>and</strong> Williams 1966;<br />
Frodin <strong>and</strong> Govaerts 2003).<br />
Chaoliang Wang, Jinghua Wang, Meila Wang, Xiao<br />
Wei, Tom Wendt, Carol Wilson, Emily Wood, C.-<br />
Y. Wu, Fuwu Xin, Kexue Xu, Aleck Yang, Gan<br />
Yao, Tingshuang Yi, Nelson Zamora, Mingli<br />
Zhang, Qifa Zhang, Zhiyun Zhang, Jim Zech, <strong>and</strong><br />
Shiliang Zhou. The curators <strong>of</strong> A, ASU, B, BKF,<br />
BM, BSHC, CAS, DES, DS, E, F, G, GH, GOET,<br />
HAST, K, K-W, KUN, L, LE, LL, LPB, MEXU,<br />
MO, MSC, MT, MU, NCSC, NCUF, NY, OS, P,<br />
PE, PH, RSA, SRSC, TAI, TAIF, TEX, TI, TNM,<br />
U, UC, UNM, US, W, WU, WUK, <strong>and</strong> YU are<br />
gratefully acknowledged for permitting the<br />
examination <strong>of</strong> their specimens. I thank Jeff Martz,<br />
Nancy Klaud, Zorica Dabich <strong>and</strong> Alice Tangerini<br />
for providing the botanical illustrations, Larry Dorr,<br />
Dan Nicolson, Fred Barrie, Pedro Acevedo <strong>and</strong><br />
Warren Wagner for helpful discussions <strong>and</strong> advice<br />
on nomenclature, historical references, localities<br />
<strong>and</strong>/or collectors, Jimmy Triplett <strong>and</strong> Akiko<br />
Soejima for checking locality names <strong>of</strong> <strong>Aralia</strong><br />
cordata from Japan, Christian Feuillet for<br />
interpreting references in French, <strong>and</strong> Paul<br />
Peterson for valuable advice during the preparation<br />
<strong>of</strong> the manuscript. Zelong Nie, Hanmin Dong <strong>and</strong><br />
Michael Dong helped plot the distributional maps,<br />
Stefanie Ickert-Bond <strong>and</strong> Sue Lutz put together the<br />
plates on pollen <strong>and</strong> trichome morphology, Tracy<br />
Belaski made the 14 color plates with InDesign,<br />
<strong>and</strong> Sue Lutz <strong>and</strong> Stanley Yankowski helped with<br />
SEM work on trichomes <strong>and</strong> pollen morphology.<br />
John Boggan helped draft distributional maps,<br />
provided valuable comments on the earlier version<br />
<strong>of</strong> the manuscript, <strong>and</strong> helped with obtaining<br />
interlibrary loans. Finally, without the love <strong>and</strong><br />
support from my family, Hanmin, Michael, James,<br />
<strong>and</strong> Gr<strong>and</strong>ma <strong>and</strong> Gr<strong>and</strong>pa Dong, this monograph<br />
series <strong>of</strong> <strong>Aralia</strong> would never have been completed!
152<br />
Backer, C. A. <strong>and</strong> R. C. Bakhuizen van den Brink.<br />
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Barrett, S. C. H. <strong>and</strong> K. Helenurm. 1981. Floral<br />
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(<strong>Aralia</strong>ceae). Evolution 35: 752-762.<br />
Barrett, S. C. H. <strong>and</strong> J. D. Thomson. 1982. Spatial<br />
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Can. J. Bot. 60: 1662-1670.<br />
Bawa, K. S., C. R. Keegan <strong>and</strong> R. H. Voss. 1982.<br />
Sexual dimorphism in <strong>Aralia</strong> nudicaulis L.<br />
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Blair, A. 1975. Karyotype <strong>of</strong> five plant species<br />
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Blanco, M. 1877. Flora de Filipinas, 3 rd edition.<br />
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Batavia: L<strong>and</strong>s Drukkerij.<br />
Boufford, D. E. <strong>and</strong> S. A. Spongberg. 1983.<br />
Eastern AsianEastern North American<br />
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Bowden, W. M. 1945. A list <strong>of</strong> chromosome<br />
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Cambodge du Laos et du Viet-Nam genre<br />
<strong>Aralia</strong> L. Adansonia 4: 461-465.<br />
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Cavanilles, A. J. 1797. Icones et descriptions<br />
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observationum circa species novas aut non satis<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
LITERATURE CITED<br />
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158<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
ALPHABETICAL INDEX TO COLLECTORS<br />
Abrams 1954 (1)<br />
Acad. Bras. de Ciências 1101 (20)<br />
Acevedo &. Martínez 965 (22)<br />
Adams , J. E. 21(1)<br />
Adams, J. W. & Hopkins, G. 897 (2)<br />
Ahart, L. 2483 (1); 5106 (1)<br />
Ahart, L. & P. 4214 (1)<br />
Ahles 14228 (15); 40851 (2); 46779 (2); 49417<br />
(2); 49872 (2); 53987 (15); 61512 (2); 66006<br />
(15); 79975 (15); 84804 (2)<br />
Ailts 234 (2)<br />
Albrecht s.n. (4), 1861<br />
Alex<strong>and</strong>er et al. 5480 (1); 5587 (1)<br />
Allard 3158 (2); 11526 (2)<br />
Allen 1604 (22)<br />
Allen & van Severen 7093 (22)<br />
Anderson 3041 (1)<br />
Anderson & Brown 9881 (15)<br />
Andreas 1827 (15)<br />
Andreas, B.A. & T. S. Cooperrider 2186 (2)<br />
Anect 135 (3)<br />
Antezana 650 (19)<br />
Apgar s.n. (15)<br />
Applegate 2703 (1); 6528 (1)<br />
Aquilar 1421 (16)<br />
Araquistain 3478 (22)<br />
Aristeguieta 4232 (22); 6047 (22); 6157 (22)<br />
Arsène 17733 (3)<br />
Asano 5060 (13)<br />
Auclair & Walther 57 (2)<br />
Augustinowiz 71 (5)<br />
Austin s.n. (1); 512 (1)<br />
Baishuijiang Expedition 1661 (14); 2477 (14)<br />
Baishuijiang Expedition Team 5728 (7)<br />
Baker et al. 11308 (18)<br />
Balansa 3758 (20)<br />
Balcazar 11 (19)<br />
Ball s.n. (1)<br />
Ballard s.n. (2)<br />
Barber 151 (3)<br />
Barclay et al. 1456 (1)<br />
Barkalov s.n. (5), 20 Jul 1982; s.n. (5), 5 Aug<br />
1988<br />
Bartley 1377 (15)<br />
Batchelder s.n. (2)<br />
Beauchamp 1003 (4); 1033 (4)<br />
Beckwith 128 (1)<br />
Bedker 1543 (3)<br />
Beer 25333 (9)<br />
Benítez P. 2930 (3)<br />
Benke 362 (2)<br />
Benson 2196 (1)<br />
Bernardi 7382 (20)<br />
Beyer et al. 119 (4)<br />
Bigelow s.n. (1)<br />
Bi Jie Team 862 (14); 873 (14)<br />
Biltmore 28a (16); 1301 (15)<br />
Bioletti s.n. (1)<br />
Biology Department <strong>of</strong> Sichuan University<br />
Expedition Team 51545 (7)<br />
Bisset s.n. (4)<br />
Blake 837 (1); 2329 (2); 3827 (15); 4041 (15);<br />
6725 (2)<br />
Blanchard s.n. (2), 10 Sep 1883; s.n. (2), 29 Jul<br />
1892<br />
Blankinship s.n. (1), 7 Sep 1929<br />
Blatchford s.n. (15)<br />
Blewitt 2587 (2)<br />
Blouín et al. 7198 (15)<br />
Blumer 1625 (3); 1625a (3)<br />
Bohrer 1263 (3)<br />
Bohl 4193 (15)<br />
Boivin, B. 13927 (15)<br />
Boivin, B. & D. Dunbar 10360 (15)<br />
Boivin, B. & J.-M. Perron 12535 (15)<br />
Bojórquez & Aguiar 545 (16)<br />
Bol<strong>and</strong>er s.n. (1)<br />
Boom B3058 (4)<br />
Boufford, D.E. & M. Kato 22429 (4)<br />
Boufford, D.E. & E.W.Wood 19842 (4)<br />
Boufford, D.E. et al. 14256 (15); 15452 (2)<br />
Bowerman 233 (1)<br />
Bowman 70 (15)<br />
Boyd s.n. (15)<br />
Bracelin, H. P. 1092 (1)<br />
Bracelin, N. F. 1439 (1); 2290 (1)<br />
Bradley 7877 (15)<br />
Bradshaw s.n. (2)<br />
Br<strong>and</strong>egee s.n. (1)<br />
Br<strong>and</strong>egee, K. s.n. (1), Jun 1905<br />
Br<strong>and</strong>egee, T. S. s.n. (18), 12 Mar 1889<br />
Braun, E. L. 2494 (15); 3579 (2); s.n. (2), 7 Jul<br />
1903; 4508 (15); s.n. (15), 31 Aug 1955<br />
Braun, L. s.n. (2), 3 Aug 1910
Bravo Bolaños 1314 (3)<br />
Breedlove, D.E. 11845 (16); 12085 (16); 27076<br />
(16)<br />
Breedlove, D.E. & P.H. Raven 13230 (16)<br />
Breedlove, D.E. & R. F. Thorne 30289 (16)<br />
Brenes 15535 (22)<br />
Brewer 747(1)<br />
Brina & Costa 32.691 (20)<br />
Brinkman 2072 (15)<br />
Brinton s.n. (2)<br />
Britton et al. s.n. (2)<br />
Bross s.n. (15)<br />
Brown, H. A. et al. s.n. (15), 2 Jun 1890<br />
Brown, H. E. 903 (1)<br />
Brown, S. 357 (15)<br />
Browne & Browne Jr 4417 (2)<br />
Brumbach 8414 (2)<br />
Brummitt 19027 (4)<br />
Bullock 2080 (22)<br />
Bürger s.n. (4)<br />
Burger & Burger 7742 (22)<br />
Burgess s.n. (2); s.n. (15), Jun 1983; s.n. (15)<br />
Burk 505 (2)<br />
Burton s.n. (2); s.n. (15)<br />
Búúrquez, A. 93-111 (16)<br />
Búúrquez, A. & D. Yetman 95-213 (16)<br />
Bye 5952 (16); 5996 (16); 6628 (3)<br />
Cabrera et al. 29806 (19); 32084 (19)<br />
Calder, J. A. & J. M. Gillett 25404 (15)<br />
Calder, J. A. et al. 13026 (15); 16828 (15);<br />
16865 (15); 17415 (15)<br />
Calderón 2171 (22)<br />
Camp 38-3 (3)<br />
Canby s.n. (2); s.n. (15)<br />
Canright 327a (2)<br />
Cárdenas 3361 (19)<br />
Carlson 1355 (16)<br />
Carpenter s.n. (2)<br />
Carr 131 (15)<br />
Carter, A. 4393 (18); 5453 (18)<br />
Carter, A. & R. Ferris 3816 (18); 3893 (18);<br />
3984 (18)<br />
Carter, A. & J. Reese 4565 (18)<br />
Carter, A. & H. Sharsmith 4165 (18)<br />
Carter, E. 331 (1)<br />
Chagas, F. & Silva 00000, FUEL No. 10742<br />
(20)<br />
Chagas, F. et al. 01682 (20)<br />
Chakraborty 2397 (9)<br />
Chamberl<strong>and</strong> 950 (2)<br />
Ch<strong>and</strong>ler 816 (2)<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
159<br />
Chaney 187 (2)<br />
Chang 346 (11)<br />
Chapman s.n. (2)<br />
Chase, A. 1949 (2)<br />
Chase, V. H. 3455 (2); 10781 (2); 13596 (2)<br />
Chen, C.-H. 1033 (6)<br />
Chen, D. C. 910 (7)<br />
Chen, S. Q. 15429 (7); 15792 (7)<br />
Chen, S.Z. & B.S. Li 00296 (9)<br />
Cheng 2664 (6)<br />
Chestnut 220 (1)<br />
Chiang, F. 1085 (22); 2291 (16)<br />
Chiang, F. et al. F-2594 (16); 9549b (17)<br />
Chiao 2069 (9)<br />
Chien 1172 (7)<br />
China-Soviet Union Expedition Team 2039 (14)<br />
Ching 2957 (7); 7151 (7); 23567 (11); 24437<br />
(11)<br />
Chittenden & Peil 481 (15)<br />
Chow 40973 (7)<br />
Christ 12837 (15)<br />
Chu 3780 (7); 3300 (10)<br />
Chuan-Jing-A (Sichuan Economic Plants Aba)<br />
1333 (see discussion in 9)<br />
Churchill, J. R. s.n. (2), 18 Aug 1886 ; s.n. (2), 3<br />
Sep 1921; s.n. (2), 26 Jul 1915<br />
Churchill, S. P. 5625 (15)<br />
Clark, H. S. s.n. (2)<br />
Clark, O. M. 12555 (16)<br />
Clark, R. C. 1165 (2)<br />
Clarke, C. B. 29470 (8); 30906 (8); 30918 (8)<br />
Clarke, O. F. 427 (16)<br />
Clausen 7904 (2)<br />
Clemens, J. s.n. (2), 28 Jul 1909; s.n. (15), 11<br />
Aug 1908<br />
Clements 230 (15)<br />
Cody & Matte 8242 (15)<br />
Coile 3376 (2)<br />
Colin 15689 (17)<br />
Collector unknown s.n. (3)<br />
Collett 6568 (9); 584 (9); 5542A (8)<br />
Collins & Pease s.n. (2)<br />
Congdon s.n. (1)<br />
Constance s.n. (1)<br />
Cooper, R.E. 650 (9); 2983 (9)<br />
Cooper, R. E. & A. K. Bulley 270 (9); 2567 (9)<br />
Core s.n. (19); 2666 (15)<br />
Couzatti & Gouzaley 674 (16)<br />
Coville s.n. (15)<br />
Cr<strong>and</strong>all s.n. (15)<br />
Croat 14629 (22)
160<br />
Curtiss s.n. (2)<br />
Daniel, T. F. 2993 (16)<br />
Daniel, T. F. & S. Nelson 3572 (3)<br />
Darrow & Haskell 2023 (16)<br />
Dash & Maih 18555 (9)<br />
Davidse, G. & G. E. Pilz 31617 (16)<br />
Davidse, G., M. Sousa, O. Téllez, E. Martínez &<br />
J. Davidse 29973 (16)<br />
Davidson 1978 (15)<br />
Davis 4853 (2)<br />
Davy s.n. (1)<br />
Dawson 4991 (15)<br />
Day 77 (2); 78 (15)<br />
Deam s.n. (2), 25 Aug 1897; 495 (2); s.n. (15),<br />
28 May 1905<br />
Deane s.n. (8)<br />
Dechamps & Carrera 12190 (21)<br />
de Haas 886 (19)<br />
Delavay, M. s.n. (11), 27 Aug 1885<br />
Delavay, P. J. M. 4027 (11); 4530 (11); 4536<br />
(11); s.n. (11), 22 Aug 1885; s.n. (11), 26 Jul<br />
1885; s.n. (11), 6 Aug 1885<br />
Delevoryas s.n. (2)<br />
Delgado & Hernández 239 (17)<br />
de Quiroz & N. S. Nascimento 3853 (23)<br />
Detling 9040 (1)<br />
Dhevoj 0493 (9)<br />
Diamond 1297 (2)<br />
Díaz-Barriga 3739 (16)<br />
Diaz 00494 (17)<br />
Dice et al. 507 (18); 680 (18)<br />
Diguet s.n. (18)<br />
Dodge s.n. (2)<br />
Donnell Smith 194 (15); 779 (15)<br />
Dorr, L. 1604 (15)<br />
Dorr, L. & L. Barnett 7059 (19)<br />
Downs 1580 (2); 7243 (15); 8124 (2); 8765 (15)<br />
Drummond 24236 (8)<br />
Duchassaing s.n (22)<br />
Ducloux 3941 (11)<br />
Dudley s.n. (1)<br />
DuMond 1307 (2); 1809 (2)<br />
Duncan, D. s.n. (1)<br />
Duncan, W.H. 7560 (15)<br />
Dungboo 2 (9)<br />
Dunkle 402 (15)<br />
Duthie 595 (8); 12254 (8); 19618 (8); s.n. (8)<br />
Dutton s.n. (2)<br />
Eames, A.W. s.n .(2)<br />
Eames & Godfrey 269 (2); s.n. (15)<br />
Earle 4596 (1)<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
Eastwood, A. 13922 (1)<br />
Eastwood, A. & J. T. Howell 1173 (3)<br />
Eaton s.n. (2), Meriden, Aug 1856; s.n. (2), New<br />
Haven; s.n. (15), New Haven, Jun 1856<br />
Ebinger 857 (2)<br />
Edgeworth 187 (8)<br />
Edmondson 1333 (2); s.n. (15)<br />
Edwards s.n. (1)<br />
Eggleston & Kelly 22748 (2); 22752 (15)<br />
Eiseman 21 (15)<br />
Ekman 13451 (21); 13523 (21); 17551 (21);<br />
H5524 (22); H6009 (22); H6545 (22); H7966<br />
(22)<br />
Elcoro 1152 (16); 1153 (16)<br />
Ellis 428 (8)<br />
Elmer 4987 (1)<br />
Engelmann s.n. (1)<br />
Epling 7179 (15)<br />
Everett et al. 23942 (1)<br />
Ewan 9426 (1); 9433 (1)<br />
Eyerdam & Beetle 22669 (19)<br />
Fang et al. 31839 (10)<br />
Farges s.n. (10); s.n. (14)<br />
Faurie 398 (5); 3322 (4)<br />
Faurie,P. 806 (4); 1159 (4); 1284 (4)<br />
Faurie, U. s.n. (4); 6849 (4)<br />
Felipe 1270 (16)<br />
Feng 116 (11); 1351 (12); 1566 (12); 2791 (11);<br />
5534 (12); 5909 (12); 23499 (12)<br />
Fernald, M. L. s.n. (15)<br />
Fernald M. L. & B. Long 9112 (2)<br />
Fernald, M.L. & K. M. Wieg<strong>and</strong> 5948 (15)<br />
Ferreyra 2751 (19)<br />
Filho, Martins & Stubblebine 13280 (20)<br />
Fink 250 (2); s.n. (15)<br />
Fishbein et al. 704 (3); 725 (3); 1617 (16)<br />
Fitzpatrick s.n. (15)<br />
Flanagan & Kirkham ESUS 130 (7)<br />
Fleck s.n. (3)<br />
Fleming 331 (9)<br />
Foley s.n. (2)<br />
Folsom 3436 (22)<br />
Forbes 1702 (3)<br />
Foreman s.n. (2)<br />
Fosberg 48647 (1); S5286 (1)<br />
Fowler s.n. (2) 1868; s.n. (2), Aug 1893; s.n. (2),<br />
25 Jul 1893; s.n. (15)<br />
Fox, H. E. s.n. (4)<br />
Fox, W. B. & R. K. Godfrey 3388 (2); 5378 (2);<br />
5415 (2)<br />
Freer, R. S. 2442 (2); 2782 (15)
Freer & Hooks 3856 (15)<br />
French s.n. (2)<br />
Fu, G. X. 403 (9)<br />
Fu, G. X. & K. S. Zhang 1397 (7)<br />
Fu, K. J. 5140 (14); 17097 (7)<br />
Fu, L. K. 0776 (7)<br />
Fukuyama & Suzuki-Tokio ST 15202 (6)<br />
Furuse 21403 (13); 26619 (4); 45807 (4)<br />
Gage 2512 (5); 2632 (5)<br />
Galen 1047 (2)<br />
Gales 325 (1)<br />
Gamble 25054 (8); 25231 (8); 27311 (8)<br />
Gammie 712 (9)<br />
Garton 24121 (2); 9786 (11)<br />
Gates 864 (2); 130 (2); 344 (15)<br />
Gay 486 (19); s.n. Jun 1817 (16)<br />
Gayle 714 (15)<br />
Gentry, A. & C. Diaz 58212 (19)<br />
Gentry, A. et al. 44106 (19)<br />
Gentry, H. S. 1516 (16); 4262 (18)<br />
Gentry, H. S. & W. B. Fox 11806 (18)<br />
Giddiqi 4052 (8)<br />
Gierisch 3789 (3)<br />
Gilbert et al. 876 (2)<br />
Giles 544 (8)<br />
Gillett 4985 (15)<br />
Gillis 2964 (15)<br />
Gilman 39 (16)<br />
Glassman 8380 (15)<br />
Glaziou 1895 (20); 6559 (20); 10895 (20);<br />
109416a (20); 19416a (20); s.n. (20)<br />
Glaziou, A. 6559 (20)<br />
Glowenke 5192 (15)<br />
Godfrey, C. C. s.n. (2)<br />
Godfrey, R. K. & J. E. O’Connell 51881 (2);<br />
52001 (2)<br />
Godfrey R. K. et al. 51656 (15)<br />
Gonzalez M. 8408 (17)<br />
González-Medrano 7029 (17)<br />
Gonzalez & Acevedo 2163 (16)<br />
Goodding 324 (3); s.n. (3); 58-61 (16); 230-45<br />
(16)<br />
Goodman & Lawson 8672 (15)<br />
Gorman 1668 (1)<br />
Gorovoy s.n. (7)<br />
Gould et al. 2681 (16)<br />
Grant, G. B. 4450 (1)<br />
Grant, M. L. 2769 (15)<br />
Graves s.n. (2)<br />
Greenman 2554 (2)<br />
Greenman et al. 32 (15)<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
161<br />
Griffin s.n. (1)<br />
Griffiths 7122 (3)<br />
Grimaldo N. 511 (16)<br />
Grisebach s.n. (19)<br />
Grisebach & Lorentz 11/77 (19)<br />
Guan 75598 (7)<br />
Guang-Fu Forestry District Team 00928 (7);<br />
01076 (7)<br />
Guzman, Castro & Montiel 398 (22)<br />
Haber & Shchepanek 1613 (15)<br />
Haines 2245 (8)<br />
Hajra 525 (9)<br />
Hallberg 984 (16)<br />
Hall, H. M. s.n (1); 9383 (1)<br />
Hall, J. s.n. (2)<br />
Hall, W. S. s.n. (15)<br />
Halley 49 (3)<br />
Halse 5634 (1)<br />
Hammond 168 (1)<br />
Hammel 1604 (22)<br />
H<strong>and</strong>el-Mazzetti 7025 (11); 8979 (12)<br />
Hansen, G. 325 (1)<br />
Hansen, G. M. s.n. (1)<br />
Hardin 712a (15); 13220 (15)<br />
Harger s.n. (2)<br />
Harms, L. J. 2673 (2)<br />
Harms, V. L. 17640 (15)<br />
Harshberger s.n. (2)<br />
Harris, G. W. s.n. (2)<br />
Harris, S. K. & C. C. et al. 6976 (1)<br />
Harrison, G. J. 4849 (3)<br />
Harrison, H. H. s.n. (15)<br />
Hartman, C. V. 143 (16)<br />
Hartman, R. L. 17576 (15)<br />
Harvill 17768 (2)<br />
Hatschbach & Silva 50381 (19)<br />
Haught 737 (2)<br />
Hayes 734 (22)<br />
Hayne 958 (15)<br />
Hayward 240 (15); 290 (15); 840 (15); 868 (15);<br />
1131 (15); 1203 (15); 1348 (15); 1439 (15);<br />
1716 (15); 1922 (15)<br />
Hazlett 712 (15)<br />
He, D. P. 45641 (12)<br />
He, Z. & Z. L. Zhou 12948 (10)<br />
Heidenreich 437 (2)<br />
Heller 5717 (1)<br />
Henrickson 11442 (3); 13172 (17)<br />
Henry 6088 (14); 6655 (14); 6655A (14); 6785<br />
(12)<br />
Hermann 7879 (15)
162<br />
Hernández M. 6058 (17)<br />
Herrerá s.n. (16)<br />
Hess, W. 2367 (3)<br />
Hess, W. et al. 7817 (1); 8260 (15); 8684 (15);<br />
8770 (2); 8809 (15)<br />
Higgins 1827 (2)<br />
Hill 9576 (2)<br />
Hickey II 276 (15)<br />
Hinckley s.n. (3); 405 (3)<br />
Hinton, G. B. 1789 (16); 4430 (16); 5141 (16)<br />
Hinto, G. B. n et al. 9605 (16); 11295 (16);<br />
11440 (16)<br />
Hitchcock, A. S. 12017 (15)<br />
Hitchcock, C. L. 17716 (15); 24019 (15)<br />
Hitchcock, C. L. & J. S. Martin 5191 (1)<br />
Ho 561 (6)<br />
Hobson s.n. (9)<br />
Hodgson 3230 (3); 6433 (16); 8121 (18); 8148<br />
(18)<br />
Hogan 3835 (15)<br />
Hong et al. 33.318 (7); 34.433 (7)<br />
Hooker s.n. (8)<br />
Hoolpson s.n. (4)<br />
Horr 4243 (2)<br />
Hotchkiss & Jones 4068 (2)<br />
House 27588 (2); 28182 (2); 29957 (2)<br />
Howard 13 (1)<br />
Howe, C. D. 1338 (15)<br />
Howe, C. D. & W. F. Lang 1307 (15)<br />
Howell 786 (1); s.n. (1)<br />
Hsu 3190 (13)<br />
Hu, W.-H. 1435 (6); 2821 (6)<br />
Hu, S. Y. 1242 (see discussion in 9); 1933 (10);<br />
2075 (10); 2591 (10)<br />
Huang, C.-I. 2343 (6)<br />
Huang, D.-A. 61322 (7)<br />
Huang, S. F. & S. Y. Yang 3765 (6)<br />
Huang, Y.-Y. 99 (6)<br />
Huang, Z. P. et al. 1743 (9); 1854 (9)<br />
Huang, Z. Q. 164 (11)<br />
Huang Tu Team 5939 (10)<br />
Hubbard s.n. (3)<br />
Hui 75-512 (9); 75-896 (9); 75-1051 (9)<br />
Hutchins 3492 (3); 4597 (3)<br />
Hutchinson 7321 (16)<br />
Hyypio 596 (15)<br />
Ignotov s.n. (5)<br />
Imdorf 988 (3)<br />
Iseley 1262 (15)<br />
Ismael C. 23891 (16)<br />
Izaguirre 131 (16)<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
Jacquemont 717 (8); 718 (8); 913 (8); s.n. (8)<br />
James, F. C. 10196 (15)<br />
James, H. E. M. s.n. (7)<br />
Janke 3815 (15)<br />
Janssen 257 (2); 498 (2)<br />
Jenkins 3227 (2)<br />
Jiangxi Team 537 (7)<br />
Jiménez et al. 1085 (22); 1358 (22)<br />
Jing Fu Shan Team 1741 (14)<br />
Johansen 15 (22); 43 (22)<br />
Johnson, R. T. 419 (2)<br />
Johnston, M. C. et al. 11149 (17); 11466 (17);<br />
11484 (17); 11819 (3)<br />
Jones, G. N. 7842 (1); 32369 (15)<br />
Jones, M. E. 25 (16); 139 (15); 2375 (1); 13414<br />
(1); 24945 (3); 27094 (18); s.n. (2); s.n. (16),<br />
25 Aug 1903; 4 Sep 1903; s.n. (3), 21 Sep<br />
1903; s.n. (16); s.n. (16), 24 Sep 1903; s.n.<br />
(3), 26 Sep 1903; s.n. (3), 18 Jul 1922; s.n.<br />
(16), 19 Sep 1931<br />
Jórgensen 3254 (20)<br />
Kane 2481 (15)<br />
Kao 4153 (6); 9477 (6)<br />
Karges 2002-08095-6 (3)<br />
Karwinsky 777 (17); 1248 (17)<br />
Kearney, T. H. Jr. 148 (2)<br />
Kearney, T. H. & R. H. Peebles 9901 (3)<br />
Keckard et al. 5683 (1)<br />
Keil 17760 (15)<br />
Kellerman 5828 (see excluded names)<br />
Kerber 266 (16)<br />
King, A. 22 (15)<br />
King, G. 4436 (9)<br />
Kitz 6 (15)<br />
Kleeberger s.n. (2)<br />
Kluge 50 (22)<br />
Knobloch 5802 (16)<br />
Knowlton s.n. (2)<br />
Ko et al. 1382 (6)<br />
Kologiski & Perino 190 (15)<br />
Komarov 1152 (7)<br />
Kong 41889 (7); 42009 (7)<br />
Kral 10274 (15); 50610 (2); 51004 (2); 51097<br />
(2)<br />
Krautter s.n. (1)<br />
Krishna 2238 (9)<br />
Krotkov 9263 (2)<br />
Krysht<strong>of</strong>ovich s.n. (5); 171 (5)<br />
Kuhlmann 54822 (20)<br />
Kumar & Singh 9562 (9)<br />
Kunming Institute <strong>of</strong> Botany Team 130 (12)
Kunming Work Station 23918 (11)<br />
Kunming Vegetation Team s.n. (12), 14 Jun<br />
1981; s.n. (12), 6 Jul 1981<br />
Kusche s.n. (3)<br />
Lace 1476 (8)<br />
Lai, S. S. et al. 01450 (7)<br />
Lai, S. S. & M. X. Nie (7)<br />
Laing 102 (15)<br />
Lakela 1407 (15)<br />
Lammers 10297 (2)<br />
Lammers & Vincint 9018 (15); 9927 (15)<br />
Lanping Herbal Medicine Co. s.n. (12)<br />
Lansing Jr. 891 (15)<br />
Lathrop 7656 (1)<br />
Leavenworth, W. & M. 1898 (16)<br />
Leavenworth, W. C. & H. Hoogstraal 1684 (16)<br />
Lee 284 (15)<br />
Lehto 2154 (16)<br />
Leiberg 5883 (3)<br />
Lemieux 9837 (15)<br />
Lemmon 564 (1)<br />
Lemmon Herb. 2616 (16); 2716 (16)<br />
León 3560 (18)<br />
Leonard 1699 (2); 2101 (2); 3973 (15)<br />
LeSueur 940 (3); 1153 (16); 1400 (3)<br />
Levy s.n. (2)<br />
Lewis 49158 (1)<br />
Li, B. G. & S. B. Wan 750219 (14)<br />
Li, G. F. 62967 (14)<br />
Li, G. Z. 15425 (7)<br />
Li, H. J. 3446 (14); 3577 (7); 5264 (7); 5744 (7);<br />
5754 (14)<br />
Li, M. K. 0720 (11)<br />
Li, Q. 73940 (10); 73998 (10); 78132 (10)<br />
Li, Q. X. 422 (10)<br />
Lian et al. 93-339 (10)<br />
Liebmann 33 (16); 12135 (16)<br />
Liesner 17208 (2)<br />
Lillo 163 (19)<br />
Lin, C.-H. 617 (6)<br />
Lin, C. Q. 961 (7); 1048 (7)<br />
Lin, M.-J. 154 (6); 285 (6)<br />
Lingriff 788 (4)<br />
Liou 3112 (7); 3607 (7); 3850 (7); 5019 (7)<br />
Liou & P. C. Tsoong 2175 (10); 2616 (7)<br />
Liu 11808 (7); 11836 (7)<br />
Lorentz & Hieronymus 214 (19); 1173 (19)<br />
Lott 964 (22); 1055 (22); 3802 (22)<br />
Louis-Marie s.n. (2)<br />
Lowry II et al. 4969 (6)<br />
Lu, F. Y. & C. H. Ou 1507 (6); 1836 (6)<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
163<br />
Lu, S. Y. 5971 (6); 12676 (6); 14919 (6)<br />
Lucien & Louis-Marie 1202 (2)<br />
Lucy 8779 (2)<br />
Ludlow, F. et al. 21302 (9)<br />
Ludlow, F. & G. Sherriff 535 (9); 7831 (8); 9366<br />
(8)<br />
Lund et al. 263 (3)<br />
Lundell s.n. (15), 1 Jul 1905; s.n. (15), Ramsey<br />
Co.<br />
Luo, Y. B. 2487 (7); 2769 (7)<br />
Luo, Z. C. 07 (14); 1661 (7)<br />
Macbride et al. 882 (1)<br />
MacDougal 465 (3)<br />
MacFadden s.n. (1)<br />
Mackenzie 429 (2); 1607 (2); 3895 (2); s.n. (15)<br />
Macon s.n. (2)<br />
Magallanes 1110 (23); 777 (22)<br />
Maire, E. E. s.n. (11), Jul 1913; s.n. (11), Oct<br />
1913; s.n. (11), Ma-li-ouan; 294 (11)<br />
Maire, R. P. s.n. (11), 10 Aug 1921; s.n. (11),<br />
near Ma-li-ouan<br />
Makino s.n. (4), 14 Aug 1900; s.n. (4), 4 Oct<br />
1926; s.n. (4), 2 Nov 1931; s.n. (4), 19 Sep<br />
1937<br />
Malte 700/22 (2)<br />
M<strong>and</strong>on 570 (19)<br />
Mao 207 (11)<br />
Marul<strong>and</strong>a & Betancur 2128 (22)<br />
Marcelline 2269 ½ (3); 1905 (3)<br />
Marie-Victorin, et al. 48990 (2); 56252 (2)<br />
Marie-Victorin & Roll<strong>and</strong>-Germain 18507 (15);<br />
55285 (15); 56602 (15)<br />
Marie-Victorin, Roll<strong>and</strong>-Germain & Meilleur<br />
44562 (2)<br />
Marin 88 (16)<br />
Marley 912 (3)<br />
Marsh Jr. 828 (3)<br />
Martin et al. 78 (16)<br />
Martineau 359 (1)<br />
Mason 5759 (1)<br />
Matsushita 196 (4)<br />
Matsumura s.n. (13)<br />
Matuda-Eizi s.n. (6)<br />
Maysilles 10 (15)<br />
Maximowicz s.n. (4); s.n. (7), 1860<br />
Maximowicz, C. J. 10960 (4)<br />
Maxon 6181 (15)<br />
McCallum s.n. (1)<br />
McClatchie s.n. (1)<br />
McCleary s.n. (3)<br />
McCoy 585 (2)
164<br />
McDonald Jr. 18 (15); s.n. (15)<br />
McDougall 1413 (2)<br />
McGill & Lehto L20373 (16)<br />
McVaugh 8794 (2); 10348 (17)<br />
McMurphy 4 (1)<br />
Mead 2628 (15)<br />
Mearns 23 (15)<br />
Mell 130 (2)<br />
Melvin 3393 (2)<br />
Merrill 583 (2)<br />
Metcalfe 303 (3)<br />
Meyer, C. V. s.n. (1), 30 Jul 1968<br />
Meyer, F. s.n. (19), 15 Sep 1967<br />
Michener & Bioletti s.n. (1)<br />
Miller, G. S. 2101 (2)<br />
Miller, J. S. et al. 8889 (2)<br />
Miller, J. S. & N. Snow 8345 (2)<br />
Milligan s.n. (2)<br />
Mills s.n. (7), 25 Jul 1911; s.n. (7), 3 Aug 1911<br />
Millspaugh s.n. (2); 3552 (15)<br />
Minott s.n. (15)<br />
Minshall 106 (15)<br />
Mir<strong>and</strong>a 8820 (22)<br />
Mir<strong>and</strong>a 9068 (16)<br />
Mittleman 550 (16)<br />
Miyamoto et al. 10271 (9); 20169 (9)<br />
Mohr s.n. (2)<br />
Moir 4221 (15)<br />
Moldenke, A. L. & H. N. 30238 (1); 31095 (2)<br />
Moldenke, H. N. 2395 (15); 8741 (2); 9558 (15);<br />
13532 (2); 19455 (15); 20270 (2)<br />
Molina R., A. 4077 (16); 5414 (16); 7039 (22);<br />
12849 (16); 15228 (16); 22478 (16); 31748<br />
(16)<br />
Molina R., A. & A. R. Molina 27963 (16);<br />
30843 (16)<br />
Monetti 1748 (19)<br />
Mooar 8003 (15)<br />
Moodie 2 (15); 36 (15); 930 (15)<br />
Moore, J. K. 2488 (2)<br />
Moore, J. W. & M. F. 11032 (15)<br />
Moran 11545 (18); 11554 (18); 11586 (18);<br />
20405 (18); 23846 (18)<br />
Moreno 25344 (22)<br />
Morton 1903 (15); 11608 (15)<br />
Motorina s.n. (5)<br />
Mt. Jolmo Lungma Exped. Team 758 (9)<br />
W. C. Muenscher & Bechtel 477 (2)<br />
Mühlbauer 01 (19)<br />
Mullen II 590 (2); s.n. (2)<br />
Murata, G. 16727 (13)<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
Murata, J. 10106 (4)<br />
Murat, J. et al. 5993 (4)<br />
Murdoch, J. 1367 (2)<br />
Murdoch Jr., J. 4143 (15)<br />
Mustard 319 (15)<br />
Nan Shui Bei Diao Team 9245 (12)<br />
Nash, G. V. s.n. (15)<br />
Nash, L. & E. Lehto L19400 (16)<br />
Nee, M. 10784 (22); 33832 (19); 40304 (19)<br />
Nee, M. & J. Wen 53807 (19); 53845 (19);<br />
53878 (19)<br />
Nee, M. et al. 53714 (19)<br />
Neill 1082 (22); 3458 (22)<br />
Nelson, A. 2140 (15); 9456 (15)<br />
Nelson, D. G. 633 (1)<br />
Nelson, E. H. s.n. (1)<br />
Nelson, E. W. 1435 (16); 1952 (16); 3237 (16);<br />
3451 (16); 4463 (17); 6141 (3)<br />
Nelson, E. W. & E. A. Goldman 7189 (18)<br />
Nicely 3126 (2)<br />
Nichols s.n. (15)<br />
Nie et al. 788 (9); 938 (9); 1023 (9)<br />
Norris 72 (15)<br />
Northeast Yunnan Team 677 (7)<br />
Northwest Yunnan Jingshajiang Team 6546 (11)<br />
Nourse 34 (1)<br />
Ohashi s.n. (4)<br />
Ohba 680914 (4); 69828 (4)<br />
Ohlendorf 4154 (2)<br />
Ohwi s.n. (4)<br />
Oldham 310 (4); 837 (4); s.n. (4), 1862<br />
Olds s.n. (2)<br />
Omer & Qaiser 2748 (8)<br />
Oosting 34303 (15)<br />
Ortega 5051 (16); 5388 (16)<br />
Ou, C. H. 2909 (6)<br />
Ou, C. H. et al. s.n. (6), 20 Jun 1990; s.n. (6), 24<br />
Aug 1990<br />
Over 1790 (15); 1951 (2); 14415 (2)<br />
Ownbey & Meyer 2221 (1)<br />
José Padilla 16 (16)<br />
Palczewsky s.n. (7)<br />
Palmer, E. 110 (16); 2408 (1)<br />
Palmer, E. J. 34334 (3)<br />
Palmer, W. & W. H. King 126 (2); 142 (2)<br />
Pammel & Ball 740 (15)<br />
Parfitt, B. D. 4387 (16)<br />
Parfitt, B. D. & C. M. Christy 4696 (16); 4698<br />
(16)<br />
Parish, S. B. s.n. (1)<br />
Parish, S. B. & W. F. 437 (1)
Parks, H. E. 710 (1)<br />
Parks, H. E. & S. E. 24030 (1)<br />
Parmelee 687 (2); 1769 (2); 1911 (2)<br />
Parmelee & Hyypio 155 (15)<br />
Parodi 4/83 (20)<br />
Pase, C. P. 1413 (3)<br />
Pase, C. & D. Keil 3461 (3)<br />
Patterson 37 (15)<br />
Peattie 1212A (2); 1244 (2)<br />
Peck 8914 (1)<br />
Peebles et al. 4486 (3)<br />
Peirce s.n. (2)<br />
Peng 7263 (6); 14460 (6)<br />
Pérez 111 (22); 958 (22); 1770 (22)<br />
Pérez & Pérez 834 (22)<br />
Petrov s.n. (7)<br />
Pfeifer 1531 (16); 1753 (16)<br />
Phelps 721 (15)<br />
Phillips 2467 (3)<br />
Pinfold 252 (8)<br />
Pinkava et al. 16143 (3)<br />
Pinto & Bautista 91/83 (23)<br />
Pittillo, D. 346 (2); 3161 (2)<br />
Pittillo, J. D. 2877 (2)<br />
Plant Chemistry Voucher 683 (7)<br />
Plowman 13862 (2); 10536 (15)<br />
Pollard s.n. (1)<br />
Polunin 56/606 (8); 1639 (9)<br />
Polunin et al. 5045 (8)<br />
Popov s.n. (5)<br />
Poretzky 279 (7)<br />
Powell & Powell 3921 (3)<br />
Preble & Cary 127 (15)<br />
Presl s.n (16)<br />
Price s.n. (2)<br />
Prince & Atwood 764 (2)<br />
Pringle 4366 (16); 5144 (16); 6173 (16); 6237<br />
(16); s.n., 28 Sep 1891 (16)<br />
Pucha Team 20325 (7)<br />
Purpus 5010 (17); 5011 (17); 9160 (16); 9232<br />
(16); s.n. (18);<br />
Qi 612694 (11)<br />
Qinghai-Xizang Team 1071A (12); 1850 (12);<br />
2663 (12); 6340 (9); 6856 (12); 7597 (12);<br />
8574 (12); 10555 (12); 74-2149 (9); 74-2440<br />
(9); 75-34 (9)<br />
Qinghai-Xizang Additional Collections Team<br />
751563 (9); 751842 (9)<br />
Qiu, B.-Y. 50212 (11); 50428 (11); 54837 (11);<br />
54965 (11); 57877 (11); 596355 (11)<br />
Qiu, P. X. 1873 (7)<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
Radford & Stewart 1769 (2)<br />
Radford, A. E. 6258 (2); 17228 (2)<br />
Raizada 7263 (8)<br />
Raju 4128 (9)<br />
Ramseur 4655 (15)<br />
R<strong>and</strong> s.n. (2)<br />
R<strong>and</strong>all 6 (1)<br />
Rattan s. n. (1)<br />
Ratter et al. 5047 (20)<br />
Raup, H. M. 6142 (15)<br />
Raup, H. M. & E. C. Abbe 4561 (15)<br />
Rea et al. 74 (19)<br />
Rebman et al. 1420 (18)<br />
Rebman 2372 (18)<br />
Redden s.n. (1)<br />
Redfearn Jr & Weber 27516 (2)<br />
Redfield s.n. (2); 2036 (2)<br />
Reed s.n. (1)<br />
Reeves 3889 (3)<br />
Reko 4977 (16)<br />
Rich 1239 (8)<br />
Ricksecker s.n. (2)<br />
Rixford s.n. (1)<br />
Robbins 2066 (1)<br />
Robinson 221 (15)<br />
Rodin 311 (1)<br />
Valerio Rodriguez 480 (16); 1531 (16)<br />
Roll<strong>and</strong>-Germain 3200 (2)<br />
Roller 348 (2)<br />
Romírez 367 (22)<br />
Rosales 787 (22); 1279 (22)<br />
Rose 3049 (16)<br />
Rose & Painter 8145 (15)<br />
Rose et al. 13097 (16)<br />
Rose 38304 (1); 49158 (1); 66045 (1)<br />
Ross, T. S. & A. H. Ross 6025 (1)<br />
Rothrock s.n. (2); s.n. (15)<br />
Rousseau & Rouleau 5 (15)<br />
Roy 1203 (2)<br />
Royle s.n. (8)<br />
Rudd 685 (15); 802 (15); 807 (15)<br />
Rutter s.n. (15)<br />
Rydberg 732 (15); 733 (15)<br />
Safford 94 (15)<br />
Salinas T., A. et al. F-3144 (16)<br />
Salinas, N. 3471 (19)<br />
S<strong>and</strong>berg 772 (15)<br />
S<strong>and</strong>ers 11212 (22)<br />
S<strong>and</strong>ers et al. 5195 (16)<br />
S<strong>and</strong>ino 3492 (22)<br />
Saravia et al. 35A (19)<br />
165
166<br />
Sasaki s.n. (6)<br />
Sather & Caddell 2479 (4); 4914 (4); 5544 (4);<br />
6605 (4)<br />
Savatier 523 (4); 3835 (4)<br />
Savernin 107 (7)<br />
Scamman 3082 (15)<br />
Schaeffer Jr. 34359 (2); 37866 (2)<br />
Scheruk s.n. (2)<br />
Schlagintweit 3796 (8)<br />
Schmidt, F. s.n. (5), 1860; s.n. (5), Aug 1860;<br />
18202 (5)<br />
Schmidt, H. H. & M. Merello 1082 (2)<br />
Schneider 758 (15); 1648 (15)<br />
Schott s.n. (2)<br />
Schreiber 685 (1)<br />
Schreiter 11380 (19)<br />
Schuel 189 (19)<br />
Schuette s.n. (2), 3 Aug 1886; s.n. (2), 31 Jul<br />
1887; s.n. (2), 2 Aug 1887<br />
Schwerdtfeger 3644 (8); 14783 (8)<br />
Scoggan,H. J. 12795 (2); 13255 (2)<br />
Scoggan, H. J. & D. Erskine 12953 (2)<br />
Seibert s.n. (1)<br />
Seler 3030 (16)<br />
Senn, H. A. 1385 (15); 1786 (15)<br />
Senn, H. A. et al. 1073 (2)<br />
Seymour, F. C. 52 (2); 26295 (2)<br />
Seymour, F. C. & W. D. Countryman 22569 (2)<br />
Seymour, F. C. & H. K. Svenson 25868 (2)<br />
Shanxi Team 02243 (7)<br />
Sharsmith, C. W. 4038 (15)<br />
Sharsmith, H. K. 3721 (1)<br />
Sheldon s.n. (15)<br />
Sherff s.n. (2)<br />
Shetler & Stone 3061 (15)<br />
Shilom T. 2692 (16); 2889 (16)<br />
Shimek s.n. (2), 1 Jul 1918; s.n. (15), 21 Aug<br />
1920; s.n. (2), 8 Aug 1921; s.n. (2), 18 Jul<br />
1927;<br />
Short s.n. (15)<br />
Shrieve 5087 (3)<br />
Shui et al. 43014 (14)<br />
Shvedtchikova s.n. (5), 19 Jul 1985; s.n. (5), 20<br />
Aug 1986; s.n. (5), 26 Aug 1986<br />
Shyu 635 (6)<br />
Sichuan Economic Botany Liangshan Team<br />
0801 (see 9 for discussion); 1510 (see 9 for<br />
discussion); 1674 (7)<br />
Sichuan Economic Plants 4304 (10)<br />
Sichuan Economic Plants Liangshan 1526 (14)<br />
Siebold 107 (4)<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
Singh & Long 17851 (9)<br />
Small s.n. (2), 9 Aug 1888; s.n. (15), 21 May<br />
1890; s.n. (2), 19-23 Aug 1890; s.n. (15), 25<br />
May 1892; s.n. (2), 23 Jul 1892; s.n. (2), 5<br />
Aug 1892; s.n. (2), 7 Sep 1892<br />
Smith, A. J. 80 (15)<br />
Smith, C. L. 898 (16)<br />
Smith, G. L. & C. R. Wheeler 7288 (1)<br />
Smith, H. H. 1595 (22); 3942 (1); 5357 (1)<br />
Smith, J. F. s.n. (2)<br />
Smith, L. 45 (2)<br />
Smith, L. B. & A. R. Hodgdon 3935 (2)<br />
Smith, L. B. & R. M. Klein 14120 (20)<br />
Smith, W. R. 22148 (15)<br />
Soejarto 5446 (2); 5133 (2); 5034 (2)<br />
Solomon & M. Nee 18122 (19)<br />
Soong 38957 (11); 38937 (7); 39067 (7); s.n.<br />
(12)<br />
Sorrie 9845 (2)<br />
Soukup 5417 (19)<br />
Soulié 1143 (12)<br />
Soza, Grijalva & Ar<strong>and</strong>a 81 (22)<br />
Sperry T198 (3)<br />
Stainton, J. D. A 5147 (9)<br />
Stainton et al. 3474 (9); 7904 (9)<br />
St<strong>and</strong>ley, P. C. 6136 (3); 10160 (16); 12067 (16);<br />
12252 (16); 12621 (16); 14698 (3); 16765<br />
(16); 18277 (15); 20829 (22); 20932 (22);<br />
22409 (16); 25481 (22); 25558 (22); 25892<br />
(22); 27023 (22); 27858 (22); 28218 (22);<br />
28941 (22); 29415 (22); 30706 (22); 31813<br />
(22); 32104 (22); 40062 (22); 76513 (16);<br />
76837 (16); 76887 (16); 77119 (16); 81503<br />
(16); 82862 (16)<br />
St<strong>and</strong>ley, P. C. & H. C. Bollman 9982 (2); 10700<br />
(3); 11875 (15)<br />
St<strong>and</strong>ley, P. C. & L. O. Williams 435 (16)<br />
Steele 25 (2)<br />
Stevens, A. F. s.n. (15)<br />
Stevens, O. A. & D. R. Moir 1690 (2)<br />
Stevens, W. D. 6192 (22); 10139 (16); 14925<br />
(16); 22963 (22)<br />
Stewart s.n. (8); s.n. (15)<br />
Steyermark, J. A. 1017 (2); 4115 (2); 5946 (2);<br />
22371 (15); 25258 (2); 26090 (2); 28689<br />
(15); 29555 (16); 30322 (16); 30647 (16);<br />
32024 (16); 50543 (16); 51650 (16); 63558<br />
(15); 72377 (2); 72525 (2); 74610 (15);<br />
89765 (22)<br />
Steyermark, J. A. , R. Smith, S. Nehlin & M.<br />
Lobo 109907 (22)
Stiles 409 (2)<br />
St. John 312 (1)<br />
Stolarskaya & Maschkova s.n. (5)<br />
Storms s.n. (15)<br />
Stutz O. 2280 (20)<br />
Suksdorf 484 (1)<br />
Swift 40 (1)<br />
Swink 1465 (2); 1587 (2); 2083 (15); 2268 (2)<br />
Tagawa 224 (4)<br />
Takahashi, H. 2853 (4)<br />
Takahashi, H. et al. 6599 (4)<br />
Takahashi, T. et al. 1393 (4)<br />
Tan 63652 (7)<br />
Tang 23346a (10)<br />
Taquet 138 (7); 891 (7)<br />
Tateishi et al. 4489 (4); 16089 (4); 17783 (6);<br />
40570 (4)<br />
Taylor, J. & C. 18659 (15)<br />
Taylor , M. S. & R. Wessel 5028 (1)<br />
Taylor, N. & R. C. Schneider 0619 (2)<br />
Ten s.n. (11)<br />
Tenorio L. 1881 (16); 14778 (16)<br />
Thieret 1578 (2); 1590 (2); 2427 (2);<br />
Thomas, J. H. 2439 (1); 3467 (1)<br />
Thomas, R. D. et al. 141164 (2)<br />
Thomas, R. D. & B. R. Thomas 149978 (2)<br />
Thompson, J. W. 9207 (15); 12422 (1)<br />
Thompson, J. W. & E. M. 661 (15)<br />
Thomson s.n. (8), 28 Aug 1849; s.n. (8), 28 Sep<br />
1848; s.n. (8), Garhwal; s.n. (8), 10 Aug 1851<br />
Thornber 182 (16)<br />
Thone 97 (2)<br />
Thorne, R. F. 13220 (2); 31188 (2); 35425 (1)<br />
Thorne, R. F. et al. 35137 (1); 42376 (1)<br />
Thunberg 7577 (4)<br />
Tierney & Foxx 17 (3)<br />
Togashi et al. s.n. (4)<br />
Tonduz 13823 (22)<br />
Torres 45 (16)<br />
Townsend & Barber 183 (3)<br />
Tracy 3320 (1); 416 (1)<br />
True, G. H. 8455 (1)<br />
True, G. H. & J. T. Howell 2453 (1)<br />
Tryon Jr. 1935 (2)<br />
Tsai 50227 (7)<br />
Tsoong, P. G. 3538 (14)<br />
Tsugaru & Takahashi 16775 (4)<br />
Tucker, G. C. 36 (2)<br />
Tucker, H. H. s.n. (2)<br />
Tuhy 2326 (3)<br />
Ule 7073 (20)<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
167<br />
Umbach 776 (15); s.n (2), 27 Jul 1899; s.n. (2),<br />
21 Aug 1897; s.n. (15)<br />
Valasek 889 (2)<br />
Vargas C. 14342 (19)<br />
Vasey 233 (1); s.n. (1), 1875<br />
Vega et al. 2645 (16)<br />
Ventura & Lopez 7998 (17); 8241 (17); 9217<br />
(see discussion in 17)<br />
Venturi 1081 (19); 1081b (19)<br />
Vervoorst & Cuezze 7632C (19)<br />
Villarreal 4377 (17); 4388 (17); 7807 (17)<br />
Visher s.n. (15)<br />
Wagner 264 (3)<br />
Waldo s.n. (2)<br />
Walker, E. H. 2301 (15); 2677 (2); 2746 (15);<br />
3212 (2)<br />
Walker, H. A. 401 (1); s.n. (1)<br />
Walker, J. B. et al. 872 (1)<br />
Wallich 4934 (9)<br />
Walpole 57 (1)<br />
Walther s.n. (1)<br />
Wan, B. B. & K. S. Chow 81106 (7)<br />
Wan¸ X. B. 283 (7)<br />
Wang & Wen 694 (see discussion in 9)<br />
Wang, C. M. 01714 (6); 08185 (6)<br />
Wan, C. S. et al. 3858 (7); 3859 (7)<br />
Wang, C. W. 60812 (7); 63731 (12); 63870 (12);<br />
64935 (12); 71583 (11)<br />
Wang, J. C. et al. 10093 (6)<br />
Wang, J. C. & K. C. Yang 4895 (6)<br />
Wang, K.-Y. s.n. (6)<br />
Wang, T. P. 9674 (7)<br />
Wang, W. C. 2473 (7)<br />
Wang, W. S. 652 (6)<br />
Wang, Z. B. 14444 (10); 14625 (10)<br />
Warming s.n. (20)<br />
Warnock, B. H. 7511 (3); 21518 (3)<br />
Warnock, M. J. 926 (15)<br />
Warrington 373 (2)<br />
Weberbauer 5668 (19); 5840 (19)<br />
Webster, G. L. 26161 (18); 26312 (18)<br />
Webster, G. L. et al. 12906 (16); 12725 (16)<br />
Wei 390 (10)<br />
Welsh 23829 (3)<br />
Wells s.n. (2)<br />
Wen, J. 83 (15); 84 (2); 85 (15); 87 (2); 95 (2);<br />
100 (15); 109 (2); 148 (2); 149 (2); 518 (11);<br />
522 (11); 548 (11); 610 (7); 625 (2); 626<br />
(15); 747 (1); 748 (1); 751 (1); 752 (1); 753<br />
(1); 755 (1); 756 (1); 757(1); 758 (1); 759<br />
(1); 762 (1); 763 (1); 764 (1); 765 (1); 766
168<br />
(1); 774 (1); 775 (1); 776 (1); 771 (1); 772<br />
(1); 773 (1); 783 (2); 796 (2); 817 (2); 818<br />
(15); 833 (15); 849 (15); 855 (2); 880 (2);<br />
883 (2); 895 (2); 906 (2); 1071 (2); 1216<br />
(11); 1362 (14); 1442 (10); 1514 (2); 1526<br />
(15); 1535 (15); 1800 (2); 1810 (15); 1824<br />
(2); 2456 (4); 2476 (4); 2487 (15); 3048<br />
(11); 3057 (11); 3079 (10); 3129A (7); 4704<br />
(15); 4714 (2); 4717 (15); 4735 (15); 4739<br />
(15); 4743 (15); 4885 (7); 4972 (3); 4974<br />
(16); 4977 (3); 4979 (3); 4980 (3); 4981 (3);<br />
4982 (3); 4983 (3); 5426 (7); 5446 (10); 5455<br />
(10); 5545 (7); 6234 (15); 6250 (2); 6269<br />
(15); 6270 (2); 7104 (1); 7250 (3); 8138 (7);<br />
8196 (14); 8542 (4); 9300 (7); 9330 (7); 9424<br />
(6); 9697 (2); 9840 (15); 9841 (2); 9931 (2);<br />
9940 (15); 9949 (2); 10440 (15); 10466 (15);<br />
10482 (2); 10483 (2); 10489 (2); 10507 (2);<br />
10640 (14); 11550 (7)<br />
Wen, J. et al. 130 (2); 131 (15); 132 (2); 133<br />
(15); 134 (2); 136 (15); 138 (2); 139 (2); 179<br />
(15); 182 (2); 186 (15); 787 (15); 982 (2)<br />
Wen, J. et al. (Tibet-MacArthur Expedition) 926<br />
(11); 1241 (12); 1446 (12); 1487 (11); 1917<br />
(11); 2283 (12); 2382 (12); 2680 (12)<br />
Wen, J. & R. Aguilar 6779 (22)<br />
Wen, J. & T. Lammers 88 (2)<br />
Wen, J. & X. P. Li 599 (11)<br />
Wen, J. & E. Martínez 384 (17); 8679 (16), 8707<br />
(16)<br />
Wendt, T. & A. Adamcewicz 462 (3)<br />
Wendt, T. & F. Chiang 185 (17)<br />
Wharton 5177a (2); 4992 (2)<br />
Wheeler 7626 (1)<br />
White 2728 (16)<br />
Whitford 210 (15)<br />
Wiegend et al. 5841 (19)<br />
Wiggins 5489 (18); 7415 (16); 13455 (1)<br />
Wilbur 4592 (2)<br />
Wilcox 309 (16)<br />
C. Wilford s.n. (4), 1859; s.n. (4), Jul 1859<br />
Williams, L. H. J. 903 (9)<br />
Williams, L. O. & A. Molina R. 10659 (16);<br />
10712 (16); 10889 (16); 13236 (16); 13291<br />
(16)<br />
Williams, L. O. et al. 22369 (16)<br />
Williams, T. A. s.n. (2), Aug 1889; s.n. (2), 23<br />
Aug 1892<br />
Wilson, E. H. 982 (12); 3698 (10); 3710 (see<br />
discussion in 9); 4285 (see discussion in 9)<br />
Windham 95-268 (3)<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
Winton, A. L. s.n. (2)<br />
Winton, Jr., A. L. s.n. (15)<br />
Wolf 3060 (1); 3071 (3); 5831 (1); 9047 (1)<br />
Wood 10637 (19); 11444 (19)<br />
Woodbury 27 (3)<br />
Woody Oil Plants Team 65-0329 (11); 65-0429<br />
(11)<br />
Woolston 1260 (20)<br />
Wooton s.n. (3)<br />
Worthington 12276 (3)<br />
Wright , C. 8 (22)<br />
Wright, T. 92-332 (3)<br />
Wright, W. F. 163 (2)<br />
Wu, C. Y. et al. 75-531 (9); 75-1043 (9)<br />
Wu, S. G. 151 (11); 771 (11); 2061 (7)<br />
Wu, Z. H. 90-808 (14)<br />
Wuyi Expedition Team 00210 (7); 00482 (7)<br />
Wuyishan Team 80-0118 (7)<br />
Xiong, J. 2873 (7)<br />
Xiong, J. H. & Z. L. Zhou 92161 (14)<br />
Xiong, J. H. et al. 42236 (14)<br />
Xu 960329 (10)<br />
Kana & Yamashita 8963 (13)<br />
Yamazaki, T. 3740 (13); 9779 (4)<br />
Yamazaki, T. & F. 5069 (4)<br />
Yamazaki, T. & K. Asano 7518 (4)<br />
Yanbian Group I 416 (7)<br />
Yanbian Group II 962 (7)<br />
Yang, B.-P. 74 (6)<br />
Yang, G. H. 59114 (14)<br />
Yang, J.-S. 5042 (11)<br />
Yang, J. X. 681 (10); 01603 (7);<br />
Yang, T. Y. 89 (6)<br />
Yang, T. Y. A. 11188 (6)<br />
Yang, T. Y. A. & Y. B. Cheng 06356 (6)<br />
Yang, T. Y. A. et al. 13875 (6)<br />
Ye 3310 (7); 2002 (7)<br />
Yi et al. 0234 (14)<br />
Ying et al. 0135 (7)<br />
Yoshitake & Kaneda s.n. (4)<br />
Young s.n. (8)<br />
Yu 222 (7)<br />
Yü, T. T. 2368 (7); 9913 (12); 14050 (11); 22458<br />
(12)<br />
Yue et al. 4253 (7)<br />
Yuncker, E. C. & T. G. 5440 (15)<br />
Yuncker, T. G. et al. 5800 (16)<br />
Yuncker, T. G. & E. C. 6709 (15)<br />
Yunnan Tropical Biological Resources Team 60-<br />
160 (11); s.n. (12)<br />
Zamora 194 (19)
Zerny s.n. (8)<br />
Zhang 1056 (7)<br />
Zhi Yun Shan Team 360 (14); 388 (14); 920 (14);<br />
929 (14); 931 (14); 931A (14); 933 (14); 962<br />
(14)<br />
Zhmylev s.n. (7)<br />
Zhong 5929 (9)<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
169<br />
Zhongdian Team 1018 (11); 2289 (11); 63-3193<br />
(11); 632539 (12)<br />
Zhou, G. S. et al. 469 (7); 106 (7)<br />
Zhou, T. Y. & G. J. Xu 466 (10)<br />
Zhu et al. 2435 (10)<br />
1984 Sino-Amer. Bot. Exped. 928 (11)
170<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
INDEX TO SCIENTIFIC NAMES<br />
(Accepted names are in bold; synonyms are in italics; main entry for each is in bold;<br />
<strong>and</strong> figures <strong>and</strong> plates are in parentheses)<br />
Abies 70, 71, 72, 75, 79, 92,94, 102, 103, 113<br />
Acacia berl<strong>and</strong>ieri 129<br />
Acacia crassifolia 129<br />
Acanthophora Merr. 34<br />
Acer 38, 40, 42, 44, 47, 48, 66, 70, 71, 72, 75, 85, 86, 113,<br />
114, 115, 116, 117<br />
Acer rubrum 48, 117<br />
Acer saccharum 44, 66, 115<br />
Agave lechuguilla 129<br />
Agave macroculmis 129<br />
Agave parryi 126<br />
Anadenanthera macrocarpa 136, 138<br />
Andropogon 126<br />
Apiaceae 5<br />
<strong>Aralia</strong> L. 5, 22, 34<br />
<strong>Aralia</strong> apioides H<strong>and</strong>.-Mazz. 13, 28, 36, 100, 102, 103, (15,<br />
58, 101, 102)<br />
<strong>Aralia</strong> arizonica Eastw. 68<br />
<strong>Aralia</strong> armata (Wall. ex Don) Seem. 22, 30, 93<br />
<strong>Aralia</strong> atropurpurea Franch. 10, 13, 22, 28, 36, 96, 99,<br />
100, 103, (16, 57, 95, 97)<br />
<strong>Aralia</strong> bahiana J. Wen 14, 30, 33, 34, 134, 149, (137, 150)<br />
<strong>Aralia</strong> bicrenata Wooton & St<strong>and</strong>l. 8, 9, 28, 32, 33, 35, 68,<br />
73, (24, 51, 69, 71)<br />
<strong>Aralia</strong> bipinnata Blanco var. bipinnata 30<br />
<strong>Aralia</strong> bipinnata Blanco var. apoensis (Elmer) J. Wen 30<br />
<strong>Aralia</strong> brevifolia Marchal 9, 118<br />
<strong>Aralia</strong> cachemirica Decne. 6, 10, 28, 33, 36, 87, 90, 93,<br />
(88, 89)<br />
<strong>Aralia</strong> caesia H<strong>and</strong>.-Mazz. 29<br />
<strong>Aralia</strong> californica S. Watson 6, 8, 9, 22, 28, 33, 35, 36, (23,<br />
37, 39, 49)<br />
<strong>Aralia</strong> californica S. Watson var. acuminata S. Watson ex<br />
Howell 36<br />
<strong>Aralia</strong> castanopsisicola (Hayata) J. Wen 29<br />
<strong>Aralia</strong> chilapensis Sessé & Moç. 9, 118, 127<br />
<strong>Aralia</strong> chinensis L. 10, 22, 30<br />
<strong>Aralia</strong> continentalis Kitag. 10, 28, 33, 35, 83, 87, (54, 79,<br />
84)<br />
<strong>Aralia</strong> cordata Thunb. 6, 8, 10, 13, 28, 33, 36, 73, 80, 83,<br />
151, (52, 74, 76)<br />
<strong>Aralia</strong> cordata Thunb. var. continentalis (Kitag.) Y. C. Zhu<br />
83<br />
<strong>Aralia</strong> cordata Thunb. var. sachalinensis (Regel) Nakai 77<br />
<strong>Aralia</strong> dasyphylla Miq. 7, 31<br />
<strong>Aralia</strong> dasyphylloides (H<strong>and</strong>.-Mazz.) J. Wen 7, 13, 22, 30<br />
<strong>Aralia</strong> debilis J. Wen13, 30<br />
<strong>Aralia</strong> decaisneana Hance 13, 30<br />
<strong>Aralia</strong> delavayi J. Wen 29<br />
<strong>Aralia</strong> dumetorum H<strong>and</strong>.-Mazz. 96, 99, 100<br />
<strong>Aralia</strong> echinocaulis H<strong>and</strong>.-Mazz. 30<br />
<strong>Aralia</strong> edulis Siebold & Zucc. 10, 73, 77<br />
<strong>Aralia</strong> elata (Miq.) Seem. var. elata 26, 30, 75, (14)<br />
<strong>Aralia</strong> elata (Miq.) Seem. var. m<strong>and</strong>shurica (Rupr. &<br />
Maxim.) J. Wen 26, 30<br />
<strong>Aralia</strong> elata (Miq.) Seem. var. ryukyuensis J. Wen 30<br />
<strong>Aralia</strong> elata (Miq.) Seem. var. inermis (Yanagita) J. Wen 30<br />
<strong>Aralia</strong> excelsa (Griseb.) J. Wen 14, 22, 29, 30, 33, 34, 134,<br />
142, 148, 149, (63, 145, 147)<br />
<strong>Aralia</strong> fargesii Franch. 4, 8, 10, 15, 28, 33, 36, 94, 96, 99,<br />
105, (56, 95)<br />
<strong>Aralia</strong> ferox Miq. 30<br />
<strong>Aralia</strong> finlaysoniana (Wall. ex Don) Seem. 30<br />
<strong>Aralia</strong> fluminensis Glaziou 140, 142<br />
<strong>Aralia</strong> foliolosa Seem. ex C. B. Clarke 22, 30<br />
<strong>Aralia</strong> franchetii J. Wen 27, 29<br />
<strong>Aralia</strong> frodiniana J. Wen 30<br />
<strong>Aralia</strong> gigantea J. Wen 17<br />
<strong>Aralia</strong> gintungensis C. Y. Wu 14, 30<br />
<strong>Aralia</strong> glabra Matsum. 4, 6, 10, 28, 33, 36, 103, 105, (104,<br />
105)<br />
<strong>Aralia</strong> glabrifoliolata (C. B. Shang) J. Wen 29<br />
<strong>Aralia</strong> henryi Harms 10, 22, 28, 33, 35, 106, 108, 109, (64,<br />
107, 108)<br />
<strong>Aralia</strong> hiepiana J. Wen & Lowry 30<br />
<strong>Aralia</strong> hispida Vent. 9, 13, 22, 26, 30, 32<br />
<strong>Aralia</strong> humilis Cav. 9, 13, 22, 28, 32, 33, 118, 127, 131, (14,<br />
19, 21, 25, 60, 119, 120, 121, 123)<br />
<strong>Aralia</strong> hypoglauca (C. J. Qi & T. R. Cao) J. Wen & Y.-F.<br />
Deng 29<br />
<strong>Aralia</strong> kansuensis G. Hoo 94, 96<br />
<strong>Aralia</strong> kingdon-wardii J. Wen, Lowry & Esser 29<br />
<strong>Aralia</strong> laevis J. Wen 29<br />
<strong>Aralia</strong> leschenaultii (DC.) J. Wen 22, 27, 29<br />
<strong>Aralia</strong> lihengiana J. Wen, L. Deng & X. Shi 29<br />
<strong>Aralia</strong> macrophylla Lindl. 87, 90<br />
<strong>Aralia</strong> malabarica Bedd. 22, 30<br />
<strong>Aralia</strong> melanocarpa (H. Lév.) Lauener 96, 99, 100<br />
<strong>Aralia</strong> merrillii C. B. Shang 30<br />
<strong>Aralia</strong> montana Blume 30<br />
<strong>Aralia</strong> nudicaulis L. 8, 9, 11, 13, 22, 29, 31, 47, 73, 109,<br />
117, 118, (14, 21, 23, 59, 110, 112)<br />
<strong>Aralia</strong> nudicaulis f. prolifera (Apgar) 109<br />
<strong>Aralia</strong> nudicaulis var. prolifera Apgar 109<br />
<strong>Aralia</strong> nudicaulis var. elongata Nash 109<br />
<strong>Aralia</strong> nudicaulis f. depauperata Marie-Vict.109<br />
<strong>Aralia</strong> nutans Franch. & Sav. 10, 73<br />
<strong>Aralia</strong> <strong>of</strong>ficinalis Z. Z. Wang 14, 30<br />
<strong>Aralia</strong> parasitica (D. Don) J. Wen27, 29<br />
<strong>Aralia</strong> pilosa Franch. 10, 106<br />
<strong>Aralia</strong> pinnata Sessé & Moç. 9, 118<br />
<strong>Aralia</strong> plumosa H. L. Li 29<br />
<strong>Aralia</strong> pubescens DC. 9, 32, 118, 127<br />
<strong>Aralia</strong> racemosa L. 6, 7, 8, 9, 28, 31, 32, 33, 34, 35, 42, 67,<br />
68, 73, 90, 114, (14, 18, 20, 43, 45, 50)<br />
<strong>Aralia</strong> racemosa var. foliosa Vict. & J. Rousseau42, 67<br />
<strong>Aralia</strong> racemosa L. f. foliosa (Vict. & J. Rousseau) Scoggan<br />
42, 67<br />
<strong>Aralia</strong> racemosa subsp. bicrenata (Wooton & St<strong>and</strong>l.) S. L.<br />
Welsh & J. T. Atwood 68
<strong>Aralia</strong> racemosa L. var. sachalinensis Regel 77, 80<br />
<strong>Aralia</strong> regeliana Marchal 9, 28, 33, 118, 127, 131, (26, 61,<br />
128, 130)<br />
<strong>Aralia</strong> rex (Ekman ex Harms) J. Wen 30, 33, 134, 142,<br />
(143, 144)<br />
<strong>Aralia</strong> sachalinensis Hort. ex Sieb. et Voss 77<br />
<strong>Aralia</strong> scabra C. Presl ex DC. 9, 118<br />
<strong>Aralia</strong> scaberula G. Hoo 30<br />
<strong>Aralia</strong> schmidtii Pojark. 6, 10, 28, 33, 35, 77, 80, 83, (52,<br />
78, 79)<br />
<strong>Aralia</strong> scopulorum Br<strong>and</strong>egee 9, 11, 22, 28, 33, 118, 127,<br />
131, (27, 61, 130, 132)<br />
<strong>Aralia</strong> searelliana Dunn 22, 31<br />
<strong>Aralia</strong> shangiana J. Wen 29<br />
<strong>Aralia</strong> sololensis Donnell-Smith 15<br />
<strong>Aralia</strong> soratensis Marchal 19, 30, 33, 34, 134, (20, 24, 62,<br />
135, 137)<br />
<strong>Aralia</strong> spinifolia Merr. 7, 22, 30<br />
<strong>Aralia</strong> spinosa L. 6, 8, 9, 13, 14, 22, 27, 30, 34<br />
<strong>Aralia</strong> stellata (King) J. Wen 29<br />
<strong>Aralia</strong> stipulata Franch. 6, 13, 14, 30<br />
<strong>Aralia</strong> subcordata (Don) J. Wen 29<br />
<strong>Aralia</strong> taiwaniana Y. C. Liu & F. Y. Lu 6, 10, 28, 33, 35,<br />
80, 82, 83, (17, 53, 81)<br />
<strong>Aralia</strong> thomsonii Seem. ex C. B. Clarke 13, 30<br />
<strong>Aralia</strong> tibetana G. Hoo 10, 28, 33, 36, 90, 93, 94, (55, 91,<br />
93)<br />
<strong>Aralia</strong> tomentella Franch. 29<br />
<strong>Aralia</strong> tripinnata Blanco 90<br />
<strong>Aralia</strong> undulata H<strong>and</strong>.-Mazz. 8, 14, 30<br />
<strong>Aralia</strong> urticifolia Miq. 7, 31<br />
<strong>Aralia</strong> verticillata (Dunn) J. Wen 29<br />
<strong>Aralia</strong> vietnamensis T.-D. Ha 31<br />
<strong>Aralia</strong> warmingiana (Marchal) J. Wen 8, 9, 30, 33, 34, 134,<br />
138, 142, 149, (25, 139, 141)<br />
<strong>Aralia</strong>? weberbaueri Harms 8, 19<br />
<strong>Aralia</strong> wilsonii Harms 29<br />
<strong>Aralia</strong> yunnanensis Franch. 96, 99<br />
<strong>Aralia</strong> sect. Anomalae Harms 8, 15, 28, 34<br />
<strong>Aralia</strong> sect. <strong>Aralia</strong> 5, 6, 10, 13, 26, 28, 31, 33, 34, 35, 109<br />
<strong>Aralia</strong> sect. Arborescentes Harms 30<br />
<strong>Aralia</strong> sect. Capituligerae Harms 30<br />
<strong>Aralia</strong> sect. Digitatipanicula Hoo 30<br />
<strong>Aralia</strong> sect. Dimorphanthus (Miq.) Miq. 6, 13, 22, 30, 33,<br />
34<br />
<strong>Aralia</strong> sect. Echinatae C. B. Shang & X. P. Li 7<br />
<strong>Aralia</strong> sect. Genuinae Harms 28, 34<br />
<strong>Aralia</strong> sect. Glabrae Z. Z. Wang 7<br />
<strong>Aralia</strong> sect. Humiles Harms 13, 28, 33, 118, 127<br />
<strong>Aralia</strong> sect. Nanae Harms 28, 119<br />
<strong>Aralia</strong> sect. Pentapanax (Seem.) J. Wen 6, 13, 29, 99<br />
<strong>Aralia</strong> sect. Sciadodendron (Griseb.) J. Wen 13, 22, 29, 33,<br />
133, 134<br />
<strong>Aralia</strong> sect. Undulatae C. B. Shang & X. P. Li 7<br />
<strong>Aralia</strong>ceae 5, 7, 8, 9, 10, 14, 22, 26, 31, 100, 148, 151<br />
Arctostaphylos 114, 126, 129, 130<br />
Azorella 5<br />
Cercocarpus mojadensis 129<br />
Cereus comarapanus 138<br />
Cnidoscolus cnicodendron 138<br />
Coemansia Marchal 34, 133<br />
Coemansia warmingiana Marchal 9, 138<br />
Cornus 40, 47, 70, 71, 113, 114<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
171<br />
Corylus cornuta 47, 114<br />
Coudenbergia Marchal 8, 29, 34, 133<br />
Coudenbergia angelicifolium (Griseb.) C. B. Shang 8, 134<br />
Coudenbergia ulei (Harms) C. B. Shang 138<br />
Coudenbergia warmingiana (Marchal) Marchal 8, 29, 34,<br />
133, 138<br />
Cwangayana Rauschert 34<br />
Cwangayana sc<strong>and</strong>ens (Merrill) Rauschert 34<br />
Dasylirion 126, 130<br />
Dendropanax Decne. & Planch. 5, 9<br />
Dimorphanthus Miq. 34<br />
Dimorphanthus edulis (Siebold & Zucc.) Miq. 73<br />
Dimorphanthus elatus Miq. 34<br />
Eleutherococcus melanocarpus H. Lév. 96, 99<br />
Eragrostis 126<br />
Flourensia 129<br />
Fraxinus 47, 114, 116<br />
Fraxinus americana 117<br />
Garrya 125, 126, 129<br />
Hechitia 129<br />
Hydrophyllum virginicum 117<br />
Impatiens 47, 66, 117<br />
Kalopanax 79, 86<br />
Lindleya mespiloides 129<br />
Maianthemum 115<br />
Megalopanax Ekman ex Harms 9, 29, 34, 133<br />
Megalopanax rex Ekman ex Harms 9, 29, 34, 133, 142<br />
Neoacanthophora Bennet 34<br />
Neoacanthophora sc<strong>and</strong>ens (Merr.) Bennet 34<br />
Onoclea sensibilis 117<br />
Oreopanax Decne. & Planch. 5, 9<br />
Osmorhiza claytonii 47, 114<br />
Osmorhiza longistylis 47, 114<br />
Panax L. 5<br />
Panax tripinnatum Wall. ex Don 90<br />
Parapentapanax Hutch. 29, 34<br />
Parapentapanax racemosus (Seem.) Hutch. 29, 34<br />
Parthenocissus quinquefolia 47, 114<br />
Pentapanax Seem. 29, 34<br />
Pentapanax angelicifolius Griseb. 9, 134<br />
Pentapanax granatensis Rusby 9, 144<br />
Pentapanax leschenaultii (DC.) Seem. 29, 34<br />
Pentapanax mexicanus C. B. Shang & X. P. Li 118, 127<br />
Pentapanax scopulorus (Br<strong>and</strong>egee) C. B. Shang 131<br />
Pentapanax ulei Harms 9, 138<br />
Pentapanax warmingianus (Marchal) Harms 138<br />
Picea 48, 68, 72, 85, 100, 102, 103, 113<br />
Pinus 40, 70, 71, 72, 85, 92, 98, 113, 123, 124, 125, 126,<br />
130<br />
Polyscias J. R. Forst. & G. Forst. 5, 10<br />
Porlieria microphylla 138<br />
Prunus 47, 70, 79, 114<br />
Prunus serotina 47, 73, 114<br />
Pseudopanax K. Koch 5<br />
Pteridium 42, 115<br />
Quercus emoryi 126<br />
Quercus rubra 115, 116<br />
Reynoldsia americana Donn.Sm. 9, 144<br />
Rhus virens 129<br />
Rubus 38, 44, 70, 72, 117<br />
Sambucus 42, 66<br />
Sambucus simpsonii Rehder 151<br />
Schefflera J. R. Forst. & G. Forst. 5
172<br />
Schinopsis haenkena 138<br />
Schizeilema 5<br />
Sciadodendron Griseb. 8, 9, 29, 34, 133<br />
Sciadodendron excelsum Griseb. 9, 29, 34, 133, 144, 148<br />
Spiraea 129<br />
Stilbocarpa (Hook. f.) Decne. & Planch. 5<br />
Tilia 47, 70, 86, 114<br />
<strong>Systematics</strong> <strong>of</strong> <strong>Aralia</strong><br />
Tilia americana 44, 47, 114, 117<br />
Trillium 47, 114<br />
Ulmus americana 65, 117<br />
Vaccinium 114, 115<br />
Vauquelinia californica 129<br />
Viburnum 42, 48, 70