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A revision of Phellodendron (Rutaceae)

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  • Beijing Botanical Garden
Article

A revision of Phellodendron (Rutaceae)

Abstract

The genus Phellodendron (Rutaceae) is revised. This work is based on herbarium collections and field observations in eastern Asia. Two species are recognized: Phellodendron amurense Rupr. and Phellodendron chinense C.K.Schneid. All other specific and infraspecific names published in the genus are reduced to synonymy or excluded from the genus. Detailed morphology, character variability, an identification key, a distribution map, line drawings, and taxonomic treatments and lists of specimens examined are provided.
A REVISION OF PHELLODENDRON (RUTACEAE)
JINSHUANG MA
1
,WEI CAO
2
,QUANRU LIU
3
,MING YU
3
&LIJUAN HAN
4
The genus Phellodendron (Rutaceae) is revised. This work is based on herbarium
collections and field observations in eastern Asia. Two species are recognized:
Phellodendron amurense Rupr. and Phellodendron chinense C.K.Schneid. All other specific
and infraspecific names published in the genus are reduced to synonymy or excluded from
the genus. Detailed morphology, character variability, an identification key, a
distribution map, line drawings, and taxonomic treatments and lists of specimens
examined are provided.
Keywords. East Asia, morphology, Phellodendron, revision, taxonomy.
INTRODUCTION
Phellodendron Rupr., a small genus in Rutaceae,Toddalioideae,Toddalineae,
Phellodendrinae (Engler, 1931; Huang, 1958; Chase et al., 1999), is easily recognized
by its thickened corky bark, deciduous leaves and dioecy.
The first species, Phellodendron amurense Rupr., was described by F. J. Ruprecht
(1857) based on R. Maack’s 1855 collections from the Amur Valley of the Russian
Far East. A second taxon, Phellodendron amurense var. sachalinense F.Schmidt, was
described in 1868 based on collections by F. Schmidt from Sakhalin Island, Russia,
and by M. Albrecht near Hokkaido, Japan (Schmidt, 1868). In 1871 Phellodendron
japonicum Maxim. was described based on C. J. Maximowicz’s 1862 collections from
Mt. Fuji, Japan (Maximowicz, 1871).
The genus was first revised by Sargent (1905) based on the collections at A and
GH, as well as on living collections at the Arnold Arboretum of Harvard University.
Sargent recognized and illustrated three species: Phellodendron amurense
(Manchuria, Mongolia, central China), P. sachalinense (Korea; Japan: Hokkaido),
and P. japonicum (Japan; China: Hubei and Sichuan). He also raised Phellodendron
amurense var. sachalinense to species status as P. sachalinense because he perceived
it to differ from P. amurense in the darker colour of the branchlets; the thinner,
not corky, bark; the rufous, rather than silvery-pubescent, winter buds; the
leaflets not lustrous adaxially and glabrous on the margins; and the glabrous
inflorescence.
1
Brooklyn Botanic Garden, 1000 Washington Avenue, Brooklyn, NY 11225-1099, USA. E-mail:
jinshuangma@bbg.org
2
Institute of Applied Ecology, Chinese Academy of Sciences, Shenyang, 110016, China.
3
Department of Botany, College of Biological Science, Beijing Normal University, Beijing, 100875, China.
4
Department of Biology, Changchun Normal College, Changchun, 130032, China.
E D I N B U R G H J O U R N A L O F B O T A N Y 63 (2&3): 131–151 (2006) 131
doi:10.1017/S0960428606000515 ETrustees of the Royal Botanic Garden Edinburgh (2006)
Issued 30 November 2006
Schneider (1907) recognized specimens from central China as a new species,
Phellodendron chinense, based on the shape of the leaf and the form of the panicle.
These had been previously identified by Sargent (1905) as Phellodendron amurense
and P. japonicum. At the same time Schneider also accepted the three species
recognized by Sargent (1905).
Dode (1908) published another four new species: Phellodendron lavallei from
Japan (but based on cultivated materials in Segrez), and P. macrophyllum,P. sinense
and P. fargesii from Sichuan, China. He also accepted the four species described
previously.
Just before 1920, two species were described from Korea and Japan, respectively:
Phellodendron insulare Nakai (1918) and P. molle Nakai (1919). However, these
names have not been used in modern taxonomic work and were only ever applied to
plants cultivated in parks, arboreta, and gardens in Europe and in North America
(Rehder, 1940).
Sprague (1920) completely revised the genus based on the herbarium specimens
and living collections in the Royal Botanic Gardens, Kew. He recognized three
species: Phellodendron amurense (Russian Far East, Chinese Manchuria, northern
China, Korea and Japan), P. japonicum (Japan: Fujiyama) and P. chinense (China:
Hupeh), and two varieties: P. amurense var. lavallei (Japan) and P. amurense var.
sachalinense (Russian Sakhalin and Korea).
New species which were described from the 1920s to the 1960s but which are not
accepted today are: Phellodendron wilsonii Hayata & Kanehira (1920), from Taiwan,
which was reduced to a variety, P. amurense var. wilsonii, by Chang & Hartley
(1993); P. piriforme E.Wolf (1925), from a cultivated plant grown in Berlin with an
unknown origin; P. kodamanum Makino (1929), from Japan, not accepted by Ohba
(1999); P. nikkomontanum Makino (1931), from Japan, not accepted by Ohba (1999);
P. sinii Y.C.Wu (1940), from Guizhou, China, not accepted by Huang (1997); and
P. burkillii Steenis (1960), from Peninsular Malaysia. The last species is excluded
from the genus as a synonym of Tetradium glabrifolium (Champ. ex Benth.)
T.G.Hartley (1981).
In local floras Shishkin & Bobrov (1949) recorded only one species (without
synonyms), Phellodendron amurense (Russian Far East: Amur, Primorye and
Ussuri), but did not mention P. amurense var. sachalinense or P. sachalinense of
Sakhalin Island. In another recent work from the region, two species were recorded
by Kharkevych (1989): Phellodendron amurense (Russian Far East: Amur, Primorye
and Ussuri; China, Japan, Korea) and P. sachalinense (southern Sakhalin, Kurie
Island, Japan and China). The difference between these species was described as bark
thin or bark thick although no detailed measurements were provided.
Huang (1958) revised the Chinese species and recognized six species, including
some from Dode’s earlier work, plus three new varieties: Phellodendron chinense
var. omeiense (from Sichuan), P. chinense var. yunnanense (from Yunnan), and P.
chinense var. falcatum (from Yunnan). Later, in Flora Reipublicae Popularis Sinicae
(Huang, 1997), he recorded two species and one variety: Phellodendron amurense
132 J I N S H U A N G M A ET AL.
Rupr. (eastern, northern and northeastern China, Japan, Korea, Russian Far East),
P. chinense C.K.Schneid. (central China: Hubei, northwest Hunan, and east
Sichuan), and P. chinense var. glabriusculum C.K.Schneid. (central, south and
southwest China).
In the English edition of the Flora of Japan (Ohwi, 1965) only Phellodendron
amurense was accepted at the rank of species, with three varieties: P. amurense var.
sachalinense F.Schmidt (Hokkaido and Honshu), P. amurense var. japonicum
(Maxim.) Ohwi (Honshu, Shikoku and Kyushu), and P. amurense var. lavallei
(Dode) Sprague (Hokkaido and Honshu). This treatment was largely adopted in the
recently updated Flora of Japan (Ohba, 1999), except that Phellodendron amurense
var. sachalinense was synonymized into P. amurense (Hokkaido, Honshu, Shikoku
and Kyushu, the temperate deciduous forests, also in Sakhalin and Amur).
In total, 16 species and 15 infraspecific taxa were described or recombined in the
genus from 1857 to 2005. These are from East Asia, i.e. Russian Far East, Japan, the
Koreas, north and northeast to central and southwestern China. The main goals of
this revision are to determine which taxa should be recognized, what are their
distributions, and which characters used to identify the species are reliable and
useful. An identification key, descriptions, distribution map and line drawings are
also provided.
MATERIALS AND METHODS
More than 1500 specimens, including types, from different herbaria in North
America, Europe and East Asia have been studied by the authors and are cited here
(except for a few notes), plus several trips were undertaken to observe and measure
the plants in their native environment. More than 113 measurements of bark and
trunk characters were recorded from plants growing in the field in East Asia.
RESULTS AND DISCUSSION
Bark
The typical character for the genus is corky bark, hence the name, cork-tree; Phellos
– cork, dendron – tree, in Greek. There are two layers in the bark, a true bark, or
phloem, its inner part usually yellow to dark yellow, and an outer corky bark,
usually grey to dark grey depending upon the time of the collection as well as its
geographical distribution. To test the usefulness of the thickness of bark as a
taxonomic character, both the outer and inner bark of two taxa from different
locations were measured in the field (the taxa from northern and northeastern China,
Japan and central China are used here).
These measurements show that the circumference of the trunk at breast height
(c.1.3 m high) is positively linearly related to the thickness of the bark in both taxa,
A REVISION OF PHELLODENDRON 133
i.e. the larger the trunk circumference, the thicker the bark (see Fig. 1). From these
results, we found that the taxon in cold temperate regions (northeastern and
northern China, Japan, i.e. Phellodendron amurense) has thick corky bark, and the
taxon in warm temperate regions (central China, i.e. P. chinense) has thin corky
bark. In Phellodendron amurense, the outer layer is much thicker than the inner layer
(see Figs 2, 4B: 3–4), and the average ratio of outer to inner is 1.128:0.368 cm (total
1.496 cm thick, average data of 85 examples); but in P. chinense, the outer layer is
much thinner than the inner layer (see Figs 3, 4A: 3–4), and the average ratio of
outer to inner is 0.127:0.336 cm (total 0.463 cm thick, average data of 28 examples).
All 113 voucher specimens were collected by J. S. Ma and are kept in BKL (see the
index of specimens at the end of this paper). The thickened bark in Phellodendron
amurense could serve to protect the trees from damage in deep winter in the cold and
dry conditions of the northern temperate region. No such thick bark is found in
Phellodendron chinense which grows in the warm and humid conditions (even in
winter) of central China.
Previous authors have noted a few exceptions where the bark is thinner than usual
for Phellodendron amurense from southern Sakhalin Island and northern Japan (i.e.
former P. sachalinense,orasP. amurense var. sachalinense). With detailed comments
on the taxon from Sakhalin Island and Hokkaido, Sprague (1920) noted that this
was easily observable in living trees. Hara (1935), in his work from southern
Hokkaido, stated that ‘In Saghalien and North Yezo (Hokkaido), the bark of
Phellodendron sachalinense is often more smooth than that of the southern plant and
such a form seems to be the typical P. amurense var. sachalinense’, without further
FIG. 1. The relationship between circumference and bark thickness of Phellodendron.
134 J I N S H U A N G M A ET AL.
information. In this revision, additional such examples were also observed from
Taiwan (E.H. Wilson 10909, A & K), Japan (E.H. Wilson 6870, A), and Korea (E.H.
Wilson 8507, A). Since there were no detailed records noting the position of the bark
collected for these specimens we are not sure if the bark samples were collected from
the upper part of the trunk or from large branches where the bark is usually thinner
than on the trunk of the same tree.
FIG. 2. The relationship between outer and inner bark of Phellodendron amurense.
FIG. 3. The relationship between outer and inner bark of Phellodendron chinense.
A REVISION OF PHELLODENDRON 135
Hairs on the leaves
The usefulness of this character has been argued for a long time (Sargent, 1905, 1914;
Schneider, 1907, 1912; Sprague, 1920; Hara, 1935; Huang, 1958, 1997; Ohwi, 1965;
Chang & Hartley, 1993; Ohba, 1999). After reviewing the available specimens for the
FIG. 4. The differences between Phellodendron amurense and Phellodendron chinense.A:P.
chinense from J.S. Ma 5055 and 5063 (BKL); B: P. amurense from J.S. Ma 5032 and 5036
(BKL). 1, branches; 2, infructescence; 3, inner bark; 4, outer bark.
136 J I N S H U A N G M A ET AL.
present study and by identifying the species both in their native area (Japan and
China) and in cultivation in northeastern North America, continuous variation from
glabrous to glabrescent (such as Phellodendron amurense), from pubescent when
young to less pubescent when old, and from pubescent along the midribs to
pubescent all over the leaves (represented by P. amurense or P. chinense), was found.
Inflorescence
This is the only character that effectively distinguishes the two accepted species. In
Phellodendron amurense the panicle is 8.5–13.5 66.5–9 cm, loose, the peduncle is
5–8.5 cm long, and the secondary axes have branches at least 1 cm long. In
Phellodendron chinense, the panicle is 6.5–9.5 64–6.5 cm, compact, the peduncle is
4–6 cm long, and the secondary axes are very short or absent, i.e. unbranched (see
Fig. 4).
TAXONOMIC TREATMENT
Phellodendron Rupr., Bull. Cl. Phys.-Math. Acad. Imp. Sci. Saint-Pe´tersbourg, se´r.
2, 15: 353 (1857). – Type: Phellodendron amurense Rupr.
Tree, deciduous, crown spreading and rounded, usually with secretory cavities
containing aromatic ethereal oils scattered throughout the parenchymatous tissues.
Stems upright, when young with many spreading branches, terete; twigs sparsely
pilose or not; nodes prominent when young; pith present, white or light brown to
brown, round, continuous, sometimes spongy; bark corky; buds solitary, small,
always hidden beneath leaf petiole, naked after leaves have fallen; lenticels white,
slightly expanded on young branches; buds pubescent, 2 per node, opposite, obovoid
to obtriangular, pointed, sessile; vascular bundle scars 3, circular, 2 upper and
1 lower. Leaves odd-compound, opposite; exstipulate; petiole terete, pilose or
glabrous, 5.5–7.5 mm long; leaflets (7 or)9(or 11), petiolules 2–3(4) mm long, pilose
or glabrous; leaflet blade base attenuate, sometimes slightly oblique, margin
subentire or with minimal and fine serrulations which are not easily observed, apex
acute or acuminate, sometimes caudate, pilose when young or glabrous, but mostly
becoming glabrous at maturity, adaxial surface bright green, glabrous, abaxial
surface pale green at maturity, sometimes pubescent along main vein; lateral veins
pinnate, 6–11 pairs, mostly not prominent abaxially, curved forward to acute, re-
divided and disappearing before reaching margin; strongly aromatic with pellucid
glands on margin. Inflorescences dioecious, nearly corymbose, terminal or opposite
to young stem, with many flowers in several clusters. Flowers 5-merous; male flowers
with stamens longer than petals, filaments linear, anthers 2-lobed, longitudinally
dehiscent, with very small disc around pistillode, pistillode clavate; female flowers
with staminodes clavate, carpels with 1 ovule per locule, style very short or nearly
A REVISION OF PHELLODENDRON 137
absent, stigma capitate, 5-lobed, much shorter than ovary, persistent. Fruit a drupe,
5-locular, stone-like, glabrous, most with 5 grooves and angles when dry. Seed 1 per
locule, brown, sometimes with black pits, ellipsoid, to 4.5 62.5–3 mm, slightly
compressed, shiny; endosperm oily, cotyledons flattened, embryo straight; germina-
tion epigeal (Tiffney, 1980; Zhou et al., 1999, 2002).
The genus is very similar to Tetradium (a genus from eastern and southern Asia and
the Himalayas with 8–9 species), especially when sterile due to the imparipinnate
compound and opposite leaves. However, in Phellodendron the buds are sunken in
the base of the petiole, protected by the leaf petiole, and are exposed only after the
leaves have fallen, but in Tetradium the buds are always exposed in the leaf axil,
without protection. Additionally in Phellodendron the fruit is syncarpous and
drupaceous whereas in Tetradium the fruit is apocarpous or subapocarpous and
follicular (Hartley, 1981). These differences place them in different subfamilies
according to the system by Engler (1931).
Key to the species
1a. Panicle 8.5–13.5 66.5–9 cm, loose, peduncle 5–8.5 cm long, secondary axes
with branches at least 1 cm long; bark with outer layer 3–4 times thicker
than inner layer, surface striped or fissured at maturity; tree 25–35 m high
(Russian Far East, Japan, Koreas, eastern, northeastern and northern China,
Taiwan) ______________________________________________ 1. P. amurense
1b. Panicle 6.5–9.5 64–6.5 cm, compact, peduncle 4–6 cm long, secondary axes
without branches or nearly so; bark with outer layer 3–4 times thinner than
inner layer, surface smooth; tree 15–20(–25) m high (central and southwestern
China) ________________________________________________ 2. P. chinense
1. Phellodendron amurense Rupr., Bull. Cl. Phys.-Math. Acad. Imp. Sci. Saint-
Pe´tersbourg, se´r. 2, 15: 353 (1857). – Type: Russia, Am rechten Amur-Ufer
unterhalb der Sungari-Mundung beim Flusse Bukatscha, R. Maack 16 (lecto LE,
designated here; iso P; photocopy NA). Figs 2, 4B: 1–4.
Phellodendron amurense var. sachalinense F.Schmidt, Mem. Acad. Imp. Sci. Saint-
Pe´tersbourg, se´r. 7, 12: 120–121 (1868), as sachalinensis.–Phellodendron sachalinense
(F.Schmidt) Sarg., Trees & Shrubs 1: 199, t. 94 (1905). – Type: Russia, Insula
Sachalin, Tunai, 9 vii 1860, fruit, F. Schmidt s.n. (lecto K, designated here; iso LE).
Phellodendron japonicum Maxim., Bull. Acad. Imp. Sci. Saint-Pe´tersbourg, se´r. 3, 16:
212 (1871). – Phellodendron amurense var. japonicum (Maxim.) Ohwi, Fl. Jap. 584
(1965). – Type: Japan, Mt. Fuji, 1862, fruit, C.J. Maximowicz s.n. (lecto GH,
designated here; iso K, L, LE 2 sheets, P, US, W; photocopy NA).
Phellodendron lavallei Dode, Bull. Soc. Bot. France 55: 648 (1908). – Phellodendron
amurense var. lavallei (Dode) Sprague, Bull. Misc. Inform. Kew 1920: 235 (1920).
– Type: Culta (originally from Japan): Segrez, 1 vi 1920 & 12 x 1917, fruit &
flower, Lavallee s.n. (lecto P, designated here; iso K 2 sheets), syn. nov.
138 J I N S H U A N G M A ET AL.
Phellodendron insulare Nakai, Bot. Mag. Tokyo 32: 107 (1918). – Type: Korea, in
insula Oorgongto, in monte Joho, 600 m, 31 v 1917, T. Nakai 4379 (lecto TI,
designated here), syn. nov.
Phellodendron molle Nakai, Bot. Mag. Tokyo 33: 58 (1919). – Phellodendron
amurense var. molle (Nakai) S.H.Li & S.Z.Liou in S.H.Li (ed.), Fl. Liaoning. 1:
1051 (1988). – Phellodendron amurense f. molle (Nakai) Y.C.Zhu in Y.C.Zhu (ed.),
Pl. Medic. Chin. Bor.-Orient. 660 (1989). – Type: Corea sept., in dumosis secus
flumen Horogawa, T. Nakai 7218 (lecto TI, designated here), syn. nov.
Phellodendron wilsonii Hayata & Kanehira in Hayata, Icon. Pl. Formosan. 9: 8
(1920). – Phellodendron amurense var. wilsonii (Hayata & Kanehira) C.E.Chang,
Quart. J. Chin. Forest 7(4): 58 (1974). – Type: Taiwan, Mt. Arison, [27] x 1918,
fruit, R. Kanehira & S. Sasaki s.n. (holo TI), syn. nov.
Phellodendron amurense Rupr. var. angustifolium E.Wolf, Mitt. Deutsch. Dendrol.
Ges. 1925: 215 (1925). – Type: No type information was given in the original
description and no type has been traced.
Phellodendron amurense Rupr. var. latifolium E.Wolf, Mitt. Deutsch. Dendrol. Ges.
1925: 215 (1925). – Type: No type information was given in the original
description and no type has been traced.
Phellodendron piriforme E.Wolf, Mitt. Deutsch. Dendrol. Ges. 1925: 215 (1925).
Type: No type information was given in the original description and no type has
been traced.
Phellodendron kodamanum Makino, J. Jap. Bot. 6: 5 (1929). – Type: Japan, Harima:
Tokura-toge, 1928, T. Makino s.n. (lecto MAK, designated here; iso TI 2 sheets).
Phellodendron nikkomontanum Makino, J. Jap. Bot. 7: 18 (1931). – Type: Japan,
Shimotsuke, Mt. Nikko, T. Makino s.n. (holo TI).
Phellodendron sachalinense (F.Schmidt) Sarg. var. suberosum H.Hara, Bot. Mag.
Tokyo 49: 863 (1935). – Phellodendron amurense Rupr. var. suberosum (H.Hara)
H.Hara, Sci. Res. Ozegahara Moor 446 (1954). – Type: Japan, Kitami, Asajino,
viii 1933, Y. Tomimoto 2799 (no type material traced, including at TI), syn. nov.
Tree, deciduous, to 25–35 m high, and 60–100 cm dbh. Bark corky, total c.1.5 cm
thick, phloem (inner part) yellow, c.0.37 cm thick, and cork (outer part) grey,
c.1.13 cm thick, dark, deeply striped or fissured along the main trunk. Buds 4–4.5 6
3–4 mm. Leaf scars 7–8 mm in diameter. Leaves (17–)21–32 613–16 cm; leaflet
blades green, elliptic to ovate-oblong, (6–)8–11 63.5–4.5(–5) cm, coriaceous to
thickly papyraceous. Inflorescence panicles 8.5–13.5 66.5–9 cm, loose, nearly
corymbose; peduncle 5–8.5 cm long, pubescent or without scales; secondary axes up
to 5 cm long. Male flowers with pedicels c.2 mm long, thin and erect, sepals
triangular, c.1.5 mm long, pubescent outside, petals yellow-green, oblong-ovate, 3.5–
4.5 61.5–2 mm, stamens longer than petals, filaments linear, usually pubescent at
base, anthers yellow, globose, c.1 61 mm, 2-lobed, longitudinally dehiscent, with
very small disc around pistillode, pistillode clavate, white pubescent at apex; female
flowers with pedicels 2–3 mm long, thin and erect, sepals small, triangular, pubescent
A REVISION OF PHELLODENDRON 139
outside, petals pale- or yellow-green, oblong-ovate, 3–4.5 61.5–2 mm, staminodes
clavate, carpels black when dried, 3.5–4 6c.2.5 mm, stone-like, ovules 1 per locule,
style very short or nearly absent, stigma capitate, 5-lobed, much shorter than ovary,
persistent. Fruit a drupe, black, 8–9.4 67.5–8.7 mm, subglobose, stone-like,
glabrous, mostly with 5 grooves and angles when dry, on a pedicel c.0.4 mm long.
Distribution and habitat. Mixed forests, 0–2700 m. China (Beijing, Hebei,
Heilongjiang, Jilin, Liaoning, East Nei Mongol, Shandong) at 300–1850 m, Taiwan
at 2000–2700 m, Japan at 60–1800 m, Korea at 0–1500 m and the Russian Far East
(see Fig. 5); also cultivated in central Asia, Caucasus, parts of Europe and North
America (Kern, 1931; Elias, 1986; Noach, 1990). Flowering from late April to early
June. Pollination is entomophilous and fruit is present from mid-August to late
October but usually remains on the tree until the following spring and is then mainly
dispersed by birds (Zhu & Dong, 1990; Lu et al., 2005) or small animals (Tiffney,
1980). It is a shade-intolerant species (Wang & Tao, 1998; Yoshida & Kamitani, 1999).
A dormancy period is required for the germination of seeds (Starshova, 1979; Zhu &
Dong, 1990; Lin et al., 1994a, 1994b) but a scarification process can promote
germination (Goo et al., 1997). Chromosome number: 2n578 (Guerra, 1984).
Economic uses. The inner part, or true bark, is used in traditional Chinese medicine
under the names of ‘Huang Bai’ or ‘Huang Po’. The tree has frequently been
exploited in its native area, not only for timber (especially by the military in northern
FIG. 5. Native distribution of Phellodendron in East Asia.
140 J I N S H U A N G M A ET AL.
and northeastern China; Zhu & Dong, 1990), but also for medicine (i.e. important
alkaloids) (Gan & Dai, 1990).
Selection of specimens examined
CHINA.BEIJING: Changping, vii 1930, H.F. Chow 40541 (PE 2 sheets); Mentougou
(Miaofengshan), 1100 m, 18 viii 2003 (bark), J.S. Ma 5074 (BKL); Miyun, 9 v 1972, Miyun
Exped. 142 (PE). HEBEI: Chengde, in 1959, Nankai University Exped. 54 (PE); Qinhuangdao,
20 viii 1951, F.T. Wang 169 (PE); Qinglong, 21 xii 1959 (fr), S. Coll. 1086 (PE); Wulingshan
(Xinglong), 1000 m, 29 viii 1953 (fr), X.Y. Liu 1622 (IBSC, KUN, PE); Zunhua (Dongling), 22
viii 1930, H.F. Chow 40940 (IBSC). HEILONGJIANG: Aihui, 3 vii 1954, G.Z. Wang 184 (IFP);
Baoqing, 19 vii 1959 (fr), Y.L. Zhang et al. 1785 (IBSC, IFP, PE); Harbin, 22 ix 1950 (fr), G.Z.
Wang et al. 162 (IFP, PE, cult.); Jixian, 22 vii 1959 (fr), Y.L. Zhang et al. 1718 (IFP, PE);
Hulin, vii 1955, Plants Exped. s.n. (IFP); Mishan, ix 1955, G.Z. Wang 4021 (IFP); Nenjiang,
18 vii 1958, Y.L. Zhang 39 (IFP); Pingshan, 1 ix 1993 (fr), 44.57˚N, 127.23˚E, NACPEC HLJ-
23 (MOAR, MOR, NA); Shangzhi, 16 ix 1950, G.Z. Wang & Q.T. Li 109 (IFP, PE); Shuanghe
Railway Station (Mudanjiang), 9 viii 1951, K.S. Hao 16188 (PE 2 sheets); Tonghe, 20 vii 1986,
Harbin Normal University Exped. 9226 (IBSC); Wuchang, 19 vii 1983, Harbin Normal
University Exped. 8391 (IBSC); Yichun, 20 vii 1956 (fr), China-German Exped. (T.N. Liou)
7608 (IBSC, IFP, KUN, PE). JILIN: Antu, 600 m, 9 ix 1951, T.N. Liou 4227 (IBSC, IFP, PE);
Changbaishan, 900 m, 15 viii 1962 (fr), Temperate Forest Group 451 (PE); Fusong, 780 m, 14
vii 1950 (fr), M. Noda et al. 120 (IBSC, IFP, PE); Helong, 26 viii 1958 (fr), C.S. Wang et al.
2513 (IBSC, IFP 2 sheets); Huadian, 600 m, 8 viii 1950 (fr), Y.C. Ma 96 (PE); Hunchun, 17 vii
1959, P.Y. Fu 795 (IFP); Jiaohe, 31 viii 1950 (fr), Y.L. Zhang et al. 959 (IBSC, IFP 2 sheets,
PE); Jiutai, 300 m, 28 viii 1950, Y.L. Zhou 2076 (IFP); Linjiang (Hunjiang), 1100 m, 8 ix 1963
(fr), S.X. Li et al. 1185 (IFP 2 sheets); Liuhe, 640 m, 14 ix 2003 (bark & photo), J.S. Ma 5034
(BKL); Manjiang (Jingyu), 11 viii 1957, Northeast Normal University Exped. 882 (PE);
Wangqing (Changhuangtsailing), 800 m, 24 vii 1931 (fr), H.W. Kung 1914 (K, NY, PE, SZ).
LIAONING: Benxi, 19 v 1950, M. Noda et al. 641 (IFP); Beizhen, 12 vi 1951, Y.L. Zhou et al.
2921 (PE 3 sheets); Caohekou (Benxi), A. Baranov & D.C. Zhao 641 (PE); Dandong, 3 v 1950,
M. Noda et al. 94 (IFP); Faku, 28 ix 1988 (fr), S.Z. Liu 352 (IFP); Fengcheng, 15 ix 1920, K.
Kondo s.n. (TI); Fushun, 18 vii 1959 (fr), W. Wang et al. 511 (IFP, PE); Fuxian, 23 viii 1958,
IFP Survey Team 26 (IFP); Haicheng, viii 1895, K. Jimbo s.n. (TI); Jianchang, 23 ix 1959 (fr),
C.S. Wang et al. 3265 (IFP 2 sheets); Huanren, 23 viii 1964, S.C. Cui 101 (IFP); Kuandian,
820 m, 11 ix 2003 (bark & photo), J.S. Ma 5001 (BKL); Luda (Dalian), 30 vii 1929 (fr), M.
Kitagawa s.n. (TI); Mukden (Shenyang, Fengtian), xii 1880 (near Mukden, Lao-Yeh Ling &
other Hills), H.E.M. James s.n. (K); Qianshan (Anshan), 16 viii 1909, Y. Yabe s.n. (NAS);
Qingyuan, 740 m, 13 ix 2003 (bark), J.S. Ma 5024 (BKL); Shenyang, vi 1980, S.X. Li 3596
(IFP); Suizhong, 20 vii 1959, S.X. Li et al. 529 (IBSC, IFP); Tieling, viii 1959, H.Z. Cao 1133
(IFP); Xifeng, 500–600 m, 20 vi 1983, S.X. Li 5895 (IFP); Xinbin, 23 ix 1959, X.F. Tian 194
(IFP); Xiongyue (Gaixian), 20 v 1950, T.N. Liou et al. 265 (IFP, PE cult.); Xiuyan, 8 ix 1959,
W. Wang 1330 (IFP); Yingkou, 23 ix 1959 (fr), Yingkou Exped. 146 (IFP); Yixian, 19 vi 1956,
S.X. Li 68 (IFP, PE); Zhuanghe, 14 ix 1959, First Group 162 (IFP). MANCHURIA (NORTHEAST
CHINA): North Manchuria, 12 viii 1922 (fr), B.V. Skvortzow s.n. (A 2 sheets); Xingan
(Chingan), in 1859 (fr), V.L. Komarov s.n. (BM), 3 vi 1895, V.L. Komarov 1013 (BM, K, NY,
P, TI, W). NEI MONGOL: Butha (Zalantun), 1 vii 1959, S.Q. Zhou 1181 (HIMC); Horqin Left
Wing Bear, 25 viii 1980, Y.C. Ma 101 (HIMC); Jalaid, 30 vi 1984 (fr), Medical Herbs Exped.
2290 (IFP 2 sheets); Ningcheng, 11 viii 1973, Y.L. Yang 798 (IFP); Oroqen Bear, 30 vi 1986, R.
Cao 83-2 (HIMC). SHANDONG: Jinan, 12 v 1979, P.C. Tang 790001 (MASS, cult.); Taishan, 13
A REVISION OF PHELLODENDRON 141
ix 1959, T.Y. Zhou et al. 7346 (NAS); Weihai, 6 vi 1959, T.Y. Zhou et al. 2178 (NAS). TAIWAN:
Arishan, 10 iv 1926, S. Saito s.n. (TI), Alishan, 2300 m, 20 viii 1963 (fr), M. Tamura,T. Shimizu
& M.T. Kao 22196 (TI); Morrison, 2666 m, 26 x 1918 (fr), E.H. Wilson 10909 (A, K, US).
JAPAN.AKITA, 10 vii 1954, A. Kimura et al. s.n. (NA). AOMORI, 25 vii 1953 (fr), H. Hara s.n.
(TI). EHIME, 4 viii 1927 (fr), Z. Tashiro s.n. (MAK). FUKUSHIMA, 14 vii 1935, M. Honda 58
(TI). GIFU, 28 vii 1940, K. Kisaut 2850 (TI). GUNMA, 13 x 1953, H. Funakoshi 680 (TI).
HOKKAIDO, 22 vi 1983, K. Deguchi 4541 (A, CM), near Hakodate, M. Albrecht s.n. (iso GH,
NY), Aze-Akaigawa in Mori-Machi, 6 x 1982 (fr), F.G. Meyer, S.G. March, M. Kaware, D.G.
Nielsen & H. Takahashi 19258 (NA), Kirkham, Coode-Adams, 20 ix 1997 (fr), Howick and
McNamara – Expedition to Hokkaido EHOK 14 (K), 31 v 1975, S. Kurosawa & T. Tataishi s.n.
(TI). Hokodate, vii 1887, Y. Tokubuchi s.n. (K), 13 viii 1899 (fr), J. Uatsumura s.n. (TI);
Fukagawa, 15 viii 1961, S. Kobayashi s.n. (MAK); Hidaka, 11 ix 1974 (fr), H. Hara, S. Kurosawa
& Y. Tataishi s.n. (TI); Iburi, 28 vii 1933 (fr), H. Hara s.n. (TI); Ishikari, 20 vii 1933 (fr), H. Hara
s.n. (TI); Kitami, vi 1972, S. Kobayashi s.n. (MAK); Kushiro, D.E. Boufford & E.W. Wood
19759 (A); Obihiro, 29 viii 1931 (fr), S. Saito s.n. (TI); Rishiri, 6 ix 1926, K. Kondo 7623 (TI 2
sheets); Sapporo, in 1884, W.P. Brooks 34 (A); Shiribeshi, 17 vii 1956, H. Kanai s.n. (TI); Teshio,
25 vi 1975, M. Furuse 8919 (K, MOR, NA), Is. Kunashiri, 7 viii 1889 (fr), U. Faurie 5106 (P,
photocopy NA). HONDO, 1800 m, 26 v 1914, E.H. Wilson 6770 (A). HONSHU, 11 vi 1926, T.
Kobayashi 2146 (A, TI). HYOGO, 950–1000 m, 11 viii 1993 (fr), N. Fukuoka et al. 7727 (MAK).
IWATE, 22 vii 1978, J. Murata, H. Ohba & S. Akiyama 5730 (TI). KANAGAWA, 1905 (fr), U.
Faurie 6871 (A, BM, P). KOCHI, viii 1934, T. Makino s.n. (MAK), 10 vii 1988, Y. Koukami 3645
(MAK). KOTSUNE, 16 vii 1909 (fr), K. Sakurai s.n. (A). KYOTO, 680 m, 11 vi 1991, T. Fuji et al.
TWT-14821 (TI). MIE, in 1934, F. Maekawa s.n. (TI). MIYAGI, 13 vi 1960, Y. Hayashi & Y.
Takeuchi s.n. (CM). MUTSU, 22 vi 1952, K. Hosoi 10987 (A, TI). NAGANO, 30 v 1953, H. Kanai
2421 (TI). NAGASHAKI, 100–430 m, 22 viii 1982, K. Ueda, S. Terabayashi & Y. Ueda 1321 (TI 2
sheets). NARA, 900 m, 3 viii 1962 (fr), N. Fukuoka & M. Hotta s.n. (TI). NIIGATA, 28 vi 1961 (fr),
M. Furuse s.n. (A, NA). OKAYAMA, vii 1988, S. Fujii 197 (CM). SAITAMA, 11 vii 1984 (fr), H.
Funakoshi 712 (TI). SHIGA, 4 vi 1997, S. Fujii 5692 (CM). SHIMANE, 6 ix 1905 (fr), J.G. Jack s.n.
(A). SHIZUOKA, 950 m, 9 v 1954, H. Kanai 5998 (TI). TOCHIGI, vii 1928 & vii 1929, T. Nakai s.n.
(TI). TOKUSHIMA, 14 viii 1894 (fr), J. Nakai 1299 (TI). TOKYO, 24 v 1988, H. Funakoshi 1202
(TI). TOTTORI, 25 ix 1955 (fr), H. Muroi 5496 (A). TOYAMA, 11 vi 1991, J. Julila & H. Fujino 408
(A). WAKAYAMA, viii 1921, S. Coll. s.n. (TI). YAMAGATA, 700 m, 24 vii 1980 (fr), D.E. Boufford
et al. 22293 (CM, GH). YAMANASHI, 980 m, 23 ix 1958 (fr), M. Furuse s.n. (A, NA).
KOREA.KANGWON (KANGWON-DO,GANGWON-DO), Chung-wang san, 700 m, 3 vi 1989,
Plant Exploration in the Republic of Korea 170 (MOAR, NA), Mt. Keumkang, 5 viii 1916,
T. Nakai 5597 (TI); Mt. Odae, 25 vii 1946 (bark), I.C. Chung 1172 (F 2 sheets, MICH), Mt.
Sorak, 21 vii 1936 (fr), T. Nakai 17482 (TI 3 sheets), Mt. Whaak, 1458 m, 27 vi 1967,
Smithsonian-Korea Ecological Project (Y.N. Lee & Y.S. Lee) 287 (TI, US). NORTH
KYONGSANG (KYONGSANG BUKTO), 620 m, 15 x 1982 (fr), Beyer, Erskine & Cowley 301
(K). NORTH HAMGYONG (HAMGYONG BUKTO), Daehungli, 18 vii 1914 (fr), T. Nakai 2047
(TI), Taiyudo, 334–1000 m, 15 vi 1917, E.H. Wilson 8599 (A, US 2 sheets), Takkori,100–1000 m,
25 vi 1917, E.H. Wilson 8696 (A 2 sheets, E, K, US 2 sheets). NORTH KAMYONG (HAMGYONG
BUKTO), 15 viii 1917 (fr), E.H. Wilson 8902 (A), Kyonsung, 13 vii 1918, T. Nakai 7219 (TI),
17 vii 1918 (fr), T. Nakai 7220 (TI 2 sheets). NORTH KYONGSANG (GYEONGSANG BUK DO),
Ullung Do (Dagelet Island), 1 x 1982 (fr), Beyer, Erskine & Cowley 101 (K). NORTH PYONGAN
(PYONGAN-BUKTO), 3 viii 1912 (fr), T. Nakai 185 & 186 (TI). SOUTH HWANGHAE (HWANGHAE
NAMDO), Changsan, 4 viii 1929, T. Nakai 13084 (TI), Taechong, 26 vii 1929, T. Nakai 13087
(TI). SOUTH HAMGYONG (HAMGYONG NAMDO), 567–867 m, 2 viii 1917 (fr), E.H. Wilson 8874
(A 2 sheets, E, K, US 2 sheets). SOUTH KYONGSANG (GYEONGSANG NAMDO), Mt. Chiri, 7 vii
1913 (bark), T. Naikai 656 (TI). SOUTH PYONGAN (PYONGAN NAMDO), 15 vii 1917, T. Nakai s.n.
142 J I N S H U A N G M A ET AL.
(TI); In silvis Ohbokdong, 3 vi 1917, T. Nakai 4380 (TI), 7 vi 1917, and in silvis Miroppon,
T. Nakai 4381 (TI 2 sheets).
RUSSIA,FAR EAST.AMUR, 20 vii 1891 (fr), L. Kozshinsky s.n. (A), viii 1900 (fr), G. Littledale
s.n. (K). KHABAROVSK, 23 viii 1903 (fr), C.S. Sargent s.n. (A 2 sheets). KURILES, Shikotan, 11
viii 1927, K. Kondo 7880 (TI 3 sheets), 31 viii 1933 (fr), Y. Tomimoto s.n. (TI). PRIMORSKY,2
vii 1972 (fl & fr), T. Neczaeva & V. Verholat 5584 (#3627) (A, E, K, MICH, NY, P, US, W).
SACHALIN, 31 viii 1936, H. Hara s.n. (TI). USSURISK, 1 ix 1920, A. Kiss s.n. (MICH).
2. Phellodendron chinense C.K.Schneid., Ill. Handb. Laubholzk. 2: 126, fig. 79 c–d
(1907). – Type: China, W Hupeh, fruit, E.H. Wilson (for J. Veitch & Sons) 1972
(lecto K, designated here; iso W). Figs 3, 4A: 1–4.
Phellodendron chinense var. glabriusculum C.K.Schneid., Ill. Handb. Laubholzk. 2:
126 (1907). – Type: No type information was given in the original description and
no type has been traced, syn. nov.
Phellodendron fargesii Dode, Bull. Soc. Bot. France 55: 649 (1908). – Type: China,
Ssu-tchuen oriental, district de Tchen-keou-tin, flower and fruit, R.P. Farges 77
bis. (lecto P, designated here; photocopies NA 2 sheets).
Phellodendron macrophyllum Dode, Bull. Soc. Bot. France 55: 648 (1908). – Type:
China, Sichuan, 1400 m, 25 vi 1893, flower, and 2 xii 1908, fruit, R.P. Farges 1284
(lecto P, designated here; drawn copy L; photocopy NA).
Phellodendron sinense Dode, Bull. Soc. Bot. France 55: 649 (1908). – Type: China,
Sichuan, 1863, fruit, Simon 16 (lecto K, designated here; iso P; photocopy NA).
Phellodendron sinii Y.C.Wu, Bot. Jahrb. Syst. 71: 185 (1940). – Type: China,
Kweichow, Yun-wu-shan, 1700 m, 13 vii 1931, Sin 50118 (holo K, but no material
traced).
Phellodendron sachalinense f. longipes Y.C.Wu, Bot. Jahrb. Syst. 71: 185 (1940). –
Type: China, Kweichow, Yun-wu-shan, 1500 m, 5 vii 1931, Sin 50389 (holo B,
untraced and probably destroyed in WW II).
Phellodendron chinense var. omeiense Huang, Acta Phytotax. Sin. 7(4): 335 (1958). –
Type: China, Szechuan, Mt. Omei, x 1952, W.P. Fang & T.W. Sung 33099 (lecto
PE, designated here; iso SZ).
Phellodendron chinense var. falcatum Huang, Acta Phytotax. Sin. 7: 336 (1958). –
Type: China, Yunnan, in declivibus montis I’cho, Liang-shan, 2100 m, arbor 13 m
alta, fruct, immt. Viridulis punctatis, 12 viii 1932, H.T. Tsai 51252 (lecto A,
designated here).
Phellodendron chinense var. yunnanense Huang, Acta Phytotax. Sin. 7: 336 (1958). –
Type: China, Yunnan, ad collum proper Far-dor, Si-chour-hsien, 1450 m, 17 ix
1947, K.M. Feng 11915 (lecto PE, designated here; as Phellodendron yunnanense
C.Y.Wu ined.).
Tree, deciduous, to 15–20(–25) m high, and 40–60 cm dbh. Bark corky, total
c.0.47 cm thick, phloem (inner part) yellow, c.0.34 cm thick, and cork (outer part)
grey, c.0.13 cm thick, dark, usually smooth along the main trunk. Buds 3–4 63–4 mm.
Leaf scars 7.5–8 mm in diameter. Leaves (19–)21–32 613–16 cm; leaflet blades green,
A REVISION OF PHELLODENDRON 143
elliptic to ovate-oblong, 8–11.5 63.5–5 cm, thinly coriaceous to papyraceous.
Inflorescence panicles 6.5–9.5 64–6.5 cm, compact, nearly corymbose; peduncle
4–6 cm long, pubescent or without scales; secondary axes 0–5 cm long. Male flowers
with pedicels c.2 mm long, thin and erect, sepals triangular, c.1.5 mm long, pubescent
outside, petals yellow-green, oblong-ovate, 3.5–4.5 61.5–2 mm, stamens longer
than petals, filaments linear, usually pubescent at base, anthers yellow, globose, c.1 6
1 mm, 2-lobed, longitudinally dehiscent, with very small disc around pistillode,
pistillode clavate, white pubescent at apex; female flowers with pedicels 2–3 mm long,
thin and erect, sepals small, triangular, pubescent outside, petals light or yellow-green,
oblong-ovate, 3–4.5 61.5–2 mm, staminodes clavate, carpels black when dried,
3.5–4 6c.2.5 mm, stone-like, ovules 1 per locule, style very short or nearly absent,
stigma capitate, 5-lobed, much shorter than ovary, persistent. Fruit a drupe, black,
8.4–9.3 67.5–8.5 mm, subglobose, 5-locular, stone-like, glabrous, mostly with 5
grooves and angles when dry, on a pedicel 0.3–0.4 mm long.
Distribution and habitat. Mixed forests, 400–2300 m. China: ?Anhui, ?Fujian,
?Guangdong, ?Guangxi, ?Guizhou, Hubei, Hunan, ?Jiangsu, ?Jiangxi, Shaanxi,
Sichuan, Yunnan, ?Zhejiang. Since the species has been long cultivated or
naturalized in some places (noted above with ‘?’ before the province name), its
native distribution cannot be determined with certainty. This species is also
cultivated in Europe and North America, but mainly in gardens and arboreta. It
flowers from late April to early June, has entomophilous pollination, and fruits from
mid-August to late October. The fruit usually remains on the tree until the following
spring and is then dispersed mainly by birds or small animals (Tiffney, 1980). It is a
fast-growing tree in full sun. Chromosome number: 2n578 (Guerra, 1984).
Economic uses. The inner part, or true bark, is used in traditional Chinese medicine
under the names of ‘Guan Huang Bai’ or ‘Guan Huang Po’. The tree has frequently
been exploited in its native area, especially for medicinal uses (i.e. important
alkaloids) (Gan & Dai, 1990). This exploitation has increased in the past thirty years
and it is now more difficult to find a living plant in the wild, but rather only in semi-
cultivation. During a visit to the Shenongjia and Metasequoia areas in Hubei, central
China, in 2003, the authors did not find any wild populations during a week-long
field expedition. None of the cultivated trees found were larger than 25 cm dbh. This
species is now very rare in central China.
Selection of specimens examined
CHINA.ANHUI: Jinzhai, 14 vii 1959, S. Coll. 61017 (NAS). FUJIAN: Congan, 15 ix 1980,
Wuyishan Exped. Team 2081 (NAS). GUANGDONG: Heping, 12 x 1958, Heping Exped. 600
(IBSC); Lechang, 11 viii 1968, P.Y. Chen et al. 2852 (IBSC); Liannan, 29 v 1951, Z.S. Zhu 754
(IBSC); Lianshan, 7 v 1958, Nanling Exped. 622 (IBSC); Lianxian, 16 xi 1930, S.P. Ko 50896
(IBSC 2 sheets, PE); North Guangdong, in 1940, T.S. Lau 301 (IBSC); Shaoguan, 16 vi 1960,
Shaoguan Exped. 1111 (IBSC); Wengyuan, 4 ix 1984, Wengyuan Herbs Exped. 64 (PE);
144 J I N S H U A N G M A ET AL.
Yangshan, 1100 m, 5 vi 1956, L. Deng 1299 (IBSC, KUN); Yingde, 12 vii 1972, H.S. Lo 705
(IBSC, cult.). GUANGXI: Jiuxiu (Dayaoshan), 10 xi 1958, Y.C. Chen 1103 (IBK, cult.); Lingle,
13 iv 1960, F.S. Huang 2106 (KUN); Lingyun (Yeo iii Shan), 1600 m, 27 viii 1928 (fr), R.C.
Ching 7168 (A, NY, PE); Longsheng, 20 ix 1984, B.N. Chang 406131 (IBK, cult.); Rongshui,
700 m, 27 vi 1957 (fr), T.C. Chen 860 (IBK, KUN cult.); Tianlin, 18 vi 1958 (fr), Z.D. Li
600845 (KUN); Ziyuan, 8 viii 1958, Z.Z. Chen 51884 (IBK, cult.). GUIZHOU: Anlong, 1300 m,
25 v 1960, Z.S. Zhang & Y.T. Zhang 3013 (KUN); Guiyang (Qianlingshan), 1400 m, 6 viii
1958, Z.Y. Cao 190 (HGAS, PE 2 sheets); Jiangkou, 750–1000 m, 11 ix 1986 (fr), B.
Bartholomew et al. 1152 (A, cult.); Guiding (Pin-fa), 1908 (fr), J. Cavalerie 3437 (E 2 sheets,
K); Leishan, 5 x 1977, Sichuan Forest Institute 926 (SCFI); Shiqian, 750 m, 26 vii 1988 (fr),
Wulingshan Plants Exped. 1654 (KUN 2 sheets); Xingren, 1200 m, 25 viii 1960 (fr), Z.S. Zhang
& Y.T. Zhang 8469 (PE); Wuyi, 28 x 1976 (fr), Sichuan Forest Institute 192 (SCFI); Xishui, 19
xi 1927, P.C. Tsoong 68 (PE). HUBEI: Badong, 1800 m, 6 x 1958, F.H. Chen 5420 (HIB);
Changyang, 1800 m, 25 vii 1959, F.S. Peng 1307 (HIB, cult.?); Enshi, 1500 m, 3 ix 1957 (fr),
G.X. Fu & Z.S. Zhang 1508 (IBSC, NAS, PE 3 sheets); Hefeng, 1050 m, 24 viii 1958, H.J. Li
6346 (HIB, SZ); Ichang, 1887 (fl), A. Henry 4003 (GH, K, P, US, photocopy NA); Jianshi, 6
vii 1951, L.Y. Dai & C.H. Qian 115 (PE); Lichuan, 1180 m, 23 ix 2003 (bark & photo), J.S. Ma
5060 (BKL 2 sheets); Shennongjia, 1260 m, 21 ix 2003 (fr), J.S. Ma 5046 (BKL 5 sheets); West
Hubei (Changyang), 1100–1200 m, 7 vi & 7 ix 1907 (fl & fr), E.H. Wilson 161 (A 2 sheets, BM,
E, GH, K, US); Wufeng, 1500 m, 31 vii 1974 (fr), Y.Z. Ma 457 (HIB); Wuhan, 2 vi 1987 (fr),
Z.E. Zhao 2381 (HIB, cult.); Xingshan, 1300 m, 17 vi 1983, G.H. Chen 98 (HIB); Xuanen,
1800 m, 1 vii 1958, H.J. Li 4395 (HIB 2 sheets); Yichang, 1300 m, 3 vii 1983 (fr), G.H. Chen
398 (HIB). HUNAN: Daoxian, 12 vii 1959, P.C. Tam 62780 (IBK, cult.); Dayong, vi 1978, Z.S.
Shen 1491 (IBSC); Guzhang, 610 m, 17 vii 1985, Q.S. Wang 2494 (HIB), Sangzhi, 1150 m, 12
ix 1990 (fr), T.R. Cao 90249 (KUN); Jiangyong, 9 vii 1959, P.C. Tam 62264 (IBK, cult.?);
Xinning, 17 vii 1979, Z.C. Luo 57 (IBSC, PE 2 sheets); Xuefengshan (Qianyang), in 1954, Z.H.
Li s.n. (PE); Yongxing, 14 vii 1979, W.S. Liao 15132 (IBSC). JIANGSU: Lianyungan, 14 vii
1974, R.X. Mi et al. 74131 (NAS); Xuzhou, 20 vi 1974, P.P. Ling et al. 74018 (NAS). JIANGXI:
Ruichang, 17 viii 1995, C.M. Tan 95589 (IBSC); Suichuan, 28 iv 1959, S.K. Lai 296 (PE);
Wugongshan (Pingxiang), 14 ix 1954, Jiangxi Exped. 1264 (PE), 21 viii 1963, J.S. Yue 3610
(PE). SHAANXI: Mianxian, 28 x 1939, T.N. Liou 11993 (PE); Yangxian, 2 viii 1952, P.C. Kuo
1936 (CDBI, IBSC, NAS, PE). SICHUAN: Baoxing (Mupin), 1600 m, 25 v 1958 (fl), Sichuan
Agriculture University Team 4966 (CDBI, SCFI); Ebian (Washan), 24 x 1938, T.N. Liou 12392
(PE), 17 ix 1908, E.H. Wilson 3228 (A, US); Emei Shan (Omeishan), 25 vii 1957, S.Y. Chen
3783 (NAS), 5 xi 1952, W.P. Fang & T.W. Sung 33486 (IBK, PE); 17 vii 1952 (fr), J.H. Xiong
et al. 31653 (NAS, PE, SCFI, SZ, ut W.P. Fang & S.C. Tsiang 31653); Bao-shing-hsien, 4 vii
1936, K.L. Chu 3073 (E, PE); 10 xi 1933, D.H. Du (T.H. Tu) 4922 (PE); Kuan-hsien, 1000–
1200 m, 8 vii 1928 (fr), W.P. Fang 2091 (A, E 2 sheets, K, NY, PE 2 sheets); 1600 m, 3 viii 1930
(fr), F.T. Wang 22110 (A, KUN, NAS, PE 2 sheets); Changning, 800 m, 18 v 1959, Sichuan
Economic Plants Exped. 514 (KUN, PE); Fengjie, 1200 m, 27 ix 1964 (fl), H.F. Chou & H.Y. Su
110570 (SCFI); Ganluo, 25 viii 1959, Sichuan Economic Plants Exped. 4421 (PE); Guanxian
(Kuan Hsien), 8 x 1938, T.N. Liou 9975 (PE); Gulan, 1300 m, 28 vii 1983 (fr), K.H. Mu & Z.H.
Dai 848 (SCFI 3 sheets); Hejiang, 920 m, 20 vii 1990, G.T. Gong & G.Z. Han 70 (SCFI 2
sheets); Hongqi (Meigu), 1900 m, 19 vii 1959, Sichuan Economic Plants Exped. 1374 (CDBI,
KUN, PE); Hongya (Hungyah), 12 vi 1955 (fr), Sichuan Forest Institute 2200 (CDBI, SCFI);
Junlian, 21 iv 1959, Sichuan Economic Plants Exped. 86 (PE); Lixian, in 1956, D.P. He 47435
(SZ); Miyi, 28 v 1989 (fl), H.Y. Liu 89576 (SCFI); Nanchuan (Jinfoshan, Chin-fu-shan), 14 vi
1935, K.L. Chu 1329 (IBSC, PE); Nanjiang, 1060 m, 5 vi 1959, Sichuan Economic Plants
Exped. (B.L. Chen) 2570 (CDBI, KUN); Pingwu, 920 m, 25 iv 1958, S. Coll. 10094 (SCFI);
Yongchuan, 21 x 1982 (fr), D.H. Du 207 (CM, MICH, NY, PE). YUNNAN: Daguan, 1800 m,
A REVISION OF PHELLODENDRON 145
18 viii 1972 (fr), Northeast Yunnan Exped. 238 (KUN 2 sheets); Kunming, Botanical Garden,
1900 m, 22 i 1988, D.D. Tao 88002 (KUN 2 sheets, cult.); Tengchong, 1740 m, 14 xi 1984 (fr),
C.H. Yang & Q.T. Zhang 84-667 (KUN, cult.); Weixi, 3 xi 1940, K.M. Feng 8716A (KUN 2
sheets, PE); Yongde, 2300 m, 22 iv 2002, E.D. Liu 353 (KUN); Zhenxiong, 1800 m, 29 ix 1957,
Kunming Station Exped. (P.H. Yu) 1065 (KUN 2 sheets, PE). ZHEJIANG: Tianmushan
(Linan), 1 ix 1959, Zhejiang Plant Inventory Team 29493 (HZU, NAS); Tiantai, 18 vii 1959,
Zhejiang Plant Inventory Team 28249 (HZU).
EXCLUDED SPECIES
Phellodendron burkillii Steenis, Gard. Bull. Singapore 17: 357 (1960). – Type: Malay
Peninsula, Kedah: Enggang Forest Reserve, Sik, PEP 78904 (holo L; iso K, KEP,
SING n.v.) 5Tetradium glabrifolium (Champ. ex Benth.) T.G.Hartley.
There are a few fossil species reported from Europe (Huang, 1958; Tiffney, 1980)
and North America (Tiffney, 1980).
ACKNOWLEDGEMENTS
We are grateful to the curators of A, BH, GH, IMC, MASS, MOAR, MOR, NY,
NYS, PE, PH, SCFI, TI, US, W and WU for use of their collections; to the curators of
E, HIB, IFP, K, KUN, KYO, MAK and P for loaning specimens; to Xinfen Gao
(CDBI), Bonnie L. Issac and Cynthia Morton (CM), Xiaodong Li and Jianqiang Li
(HIB), Rui Cao (HIMC), Chengxin Fu (HZU), Fanan Wei (IBK), Dezhi Fu and
Dianxiang Zhang (IBSC), Haiying Ma and Shukun Chen (KUN), Qixin Liu and Rong
Mo (NAS), Yuying Geng (PE) and Jie Bai (SZ) for providing distribution data; and to
Yuri Roskov (LE) for providing photos of types at LE, Chinlong Zheng (Department
of Forestry, the National Taiwan University) for providing literature, and Shuren
Zhang (PE) and Mark Tebbitt (BKL) for their kind help in obtaining photos from LE
and K, respectively. The senior author’s field trip was supported by Brooklyn Botanic
Garden and the Institute of Applied Ecology, Chinese Academy of Sciences in
Shenyang; Xiaodong Li and Jianqiang Li (HIB) provided key help during our field trip
to central China in 2003, and Jin Murata (TI) and Hiroko Murata (Setsunan
University, Osaka) provided key help during our field work in Japan in 2004. The
senior author especially thanks Kerry Barringer (BKL) for his help with loans, Paul
Harwood (BKL) for his drawing, Carsten Glaeser (Glaeser Horticultural Consulting)
for his help around the New York Metropolitan Area, and Michael Nee (NY) for his
primary work on the identification of the cultivated species of the genus from the New
York Botanical Garden and vicinity. We also thank Jacquelyn Kallunki (NY),
Anthony Brach (MO at A/GH), anonymous reviewers and David Middleton, the
editor of EJB, for their valuable comments and suggestions.
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Received 25 January 2006; accepted for publication 15 August 2006
INDEX TO THE SPECIMENS EXAMINED:(1)P. AMURENSE AND
(2) P. CHINENSE
M. Albrecht s.n. (1).
A. Baranov & D.C. Zhao 641 (1); B. Bartholomew et al. 1152 (2); Beyer, Erskine & Cowley
101 (1), 301 (1); E. Bodinier 7689 (2); D.E. Boufford & E.W. Wood 19759 (1); D.E. Boufford
et al. 22293 (1); W.P. Brooks 34 (1), 704 (1).
H.Z. Cao 1133 (1); R. Cao 83-2 (1); T.R. Cao 90249 (2); Z.Y. Cao 190 (2); J. Cavalerie 1776
(2), 3437 (2); B.N. Chang 406131 (2); H.F. Chao 40699 (1); F.H. Chen 398 (1), 5235 (2), 5420
148 J I N S H U A N G M A ET AL.
(2), 5557 (2); G.H. Chen 98 (2), 398 (2); P.Y. Chen et al. 2852 (2); S.Y. Chen 3783 (2); T.C.
Chen 860 (2); Y.C. Chen 1103 (2); Z.Z. Chen 51884 (2); W.C. Cheng 6795 (2); China-German
Exped. (T.N. Liou) 7608 (1); R.C. Ching 7168 (2); H.F. Chou & H.Y. Su 110570 (2); H.F.
Chow 40541 (1), 40940 (1), 41053 (1), 42007 (1); K.L. Chu 1329 (2); S.H. Chun 15546 (2), 15808
(2), 16640 (2); W.Y. Chun 3578 (2), 3662 (2), 3923 (1); I.C. Chung 1172 (1), 3220 (1), 3259 (1);
T.H. Chung 1170 (1), 9949 (1); S.C. Cui 101 (1).
L.Y. Dai & C.H. Qian 115 (2), 1583 (2); R.S. David s.n. (1); Dayaoshan Exped. 10345 (2),
12011 (2); K. Deguchi 4541 (1); L. Deng 1299 (2); D.H. Du (T.H. Tu) 207 (2), 4948 (2).
W.P. Fang 10184 (2), 15616 (2); U. Faurie 179 (1), 767 (1), 3374 (1), 5044 (1), 5106 (1), 6871
(1); K.M. Feng 8716A (2); G. Fenzel 44 (1); First Group 162 (1); G.X. Fu & Z.S. Zhang 1007 (2),
1508 (2), 1667 (2); P.Y. Fu 795 (1); S. Fujii 197 (1), 5692 (1); T. Fujii et al. TWT-14821 (1);
N. Fukuoka & M. Hotta s.n. (1); N. Fukuoka et al. 7727 (1); H. Funakoshi 680 (1), 712 (1), 1202
(1); M. Furuse 6815 (1), 7185 (1), 8854 (1), 8919 (1), 9549 (1), 11055 (1), s.n. (1); N. Furuse
34591 (1).
G.T. Gong & G.Z. Han 70 (2); J.L. Gressitt 2451 (2).
K.S. Hao 16188 (1); H. Hara s.n. (1); H. Hara et al. s.n. (1); Harbin Normal University
Exped. 8391 (1), 9226 (1); Y. Hayashi & Y. Takeuchi s.n. (1); H. Hayata s.n. (1); D.P. He 47435
(2); X.Y. He 5542 (2), 7106 (2); Heping Exped. 600 (2); M. Hiroe 6828 (1); K. Hisauchi 826 (1),
2374 (1); M. Honda 58 (1); K. Hosoi 10987 (1), 11013 (1); Howick and McNamara – Expedition
to Hokkaido EHOK 14 (1); C.M. Hu 7502 (2); D.A. Huang 60341 (2), 60937 (2); F.S. Huang
2106 (2); R.H. Huang 655 (2); Hubei Bot. Exped. 2403 (2); J. Hurusawa s.n. (1); C.T. Hwa 13
(2).
H. Idzumi 680 (1); IFP Survey Team 26 (1); H.T. Im 10501 (1); Institute of Botany Exped.
30316 (1); J. Ishidoya 73 (1); H. Ito s.n. (1).
J.G. Jack s.n. (1); H.E.M. James s.n. (1); Jiangxi Exped. 1264 (2); K. Jimbo s.n. (1); J. Julila
& H. Fujino 408 (1).
Y. Kadota 2442 (1); H. Kanai 1664 (1), 2421 (1), 3502 (1), 4069 (1), 4671 (1), 5998 (1), 7299
(1), 731260 (1), s.n. (1); B. Kasapligil 3570 (1); A. Kimura et al. s.n. (1); K. Kisaut 2850 (1);
A. Kiss s.n. (1); M. Kitagawa s.n. (1); S.P. Ko 50896 (2); S. Kobayashi s.n. (1); T. Kobayashi
2146 (1); G. Koidzumi s.n. (1); G. Koidzumi & Y. Okamoto s.n. (1); V.L. Komarov s.n. (1), 1013
(1); S. Komat s.n. (1); Y. Komori 1964; K. Kondo 7623 (1), 7880 (1), s.n. (1); Y. Koukami 3645
(1); N. Kozlow 14023 (1); L. Kozshinsky s.n. (1); Y. Kumori s.n. (1); Kunming Station Exped.
(P.H. Yu) 1065 (2); H.W. Kung 1914 (1); P.C. Kuo 1936 (2); S. Kurosawa s.n. (1); S. Kurosawa
& T. Tataishi s.n. (1).
S.K. Lau 296 (2), 28764 (2); T.S. Lau 301 (2); T.C. Lee 4588 (2); B.G. Li & S.F. Wan 750105
(2); C.F. Li 10047 (1); G.F. Li 1957 (2), 61492 (2), 61954 (2), 62315 (2), 63312 (2), 63613 (2),
64267 (2); H.J. Li 4395 (2), 6346 (2), 8069 (2); S.X. Li 68 (1), 3596 (1), 5895 (1); S.X. Li et al.
529 (1), 1185 (1); Y.K. Li & D.S. Zhang 8253 (2); Z.D. Li 600845 (2); Z.H. Li s.n. (2); C.F.
Liang 34348 (2); W.S. Liao 15132 (2); P. Licent 8526 (1); W.B. Lin 485 (2); Z.Q. Lin 10018 (2);
P.P. Ling et al. 74018 (2); T.N. Liou 1269 (1), 4227 (1), 9975 (2), 11993 (2), 12392 (2); T.N.
Liou & P.C. Tsoong 3746 (2); T.N. Liou et al. 265 (1), 3468 (1); G. Littledale s.n. (1); D. Litvino
s.n. (1); E.D. Liu 353 (2); H.Y. Liu 89576 (2); L.H. Liu 9071 (2); S.Z. Liu 352 (1), 600 (1); X.Y.
Liu 1622 (1); Z.Y. Liu 469 (2), 1973 (2), 8782 (2), 10641 (2), 14767 (2); H.S. Lo 705 (2); Z.C.
Luo 57 (2).
J.S. Ma 5001 (1), 5002 (1), 5003 (1), 5004 (1), 5005 (1), 5006 (1), 5007 (1), 5008 (1), 5009 (1),
5010 (1), 5011 (1), 5012 (1), 5013 (1), 5014 (1), 5015 (1), 5016 (1), 5017 (1), 5018 (1), 5019 (1),
5020 (1), 5021 (1), 5022 (1), 5023 (1), 5024 (1), 5025 (1), 5026 (1), 5027 (1), 5028 (1), 5029 (1),
5030 (1), 5031 (1), 5032 (1), 5033 (1), 5034 (1), 5035 (1), 5036 (1), 5037 (1), 5038 (1), 5039 (1),
5040 (1), 5041 (1), 5042 (1), 5043 (1), 5044 (1), 5046 (2), 5047 (2), 5048 (2), 5049 (2), 5050 (2),
5051 (2), 5052 (2), 5053 (2), 5054 (2), 5055 (2), 5056 (2), 5057 (2), 5058 (2), 5059 (2), 5060 (2),
A REVISION OF PHELLODENDRON 149
5061 (2), 5062 (2), 5063 (2), 5064 (2), 5065 (2), 5066 (2), 5067 (2), 5068 (2), 5069 (2), 5070 (2),
5071 (2), 5072 (2), 5073 (2), 5074 (1), 5075 (1), 5076 (1), 5077 (1), 5078 (1), 5079 (1), 5080 (1),
5081 (1), 5082 (1), 5083 (1), 5084 (1), 5085 (1), 5086 (1), 5087 (1), 5088 (1), 5089 (1), 5090 (1),
5091 (1), 5092 (1), 5093 (1), 5094 (1), 5095 (1), 5096 (1), 5097 (1), 5098 (1); Y.C. Ma 96 (1), 101
(1); Y.Z. Ma 333 (2), 457 (2); R. Maack s.n. (1); F. Maekawa s.n. (1); T. Makino s.n. (1); Mr.
Maries s.n. (1); J. Matsumura s.n. (1); C.J. Maximowicz s.n. (1); Medical Herbs Exped. 2290
(1); F.G. Meyer et al. 19003 (1), 19258 (1); R.X. Mi et al. 74131 (2); O. Michihito 38533 (1); K.
Miyabe s.n. (1); Miyun Exped. 142 (1); M. Mizushima 241 (1), 2290 (1), 2821 (1), 11621 (1),
13808 (1), s.n. (1); Y. Momiyama s.n. (1); K.H. Mu & Z.H. Dai 848 (2); T. Murakami 177 (1);
H. Muramatu s.n. (1); G. Murata & M. Togashi 167 (1); J. Murata 30395 (1); J. Murata & T.T.
Chen 7666 (1); J. Murata,H. Ohashi & Y. Tateishi 1804 (1); J. Murata,H. Ohba & S. Akiyama
5730 (1); H. Muroi 391 (1), 3389 (1), 3573 (1), 3751 (1), 4336 (1), 4624 (1), 4680 (1), 5383 (1),
5496 (1).
NACPEC HLJ-23 (1); T. Nakai 185 (1), 186 (1), 656 (1), 1299 (1), 2047 (1), 2048 (1), 2049
(1), 2050 (1), 4380 (1), 4381 (1), 5597 (1), 5598 (1), 7219 (1), 7220 (1), 12385 (1), 13084 (1),
13085 (1), 13086 (1), 13087 (1), 17136 (1), 17482 (1), s.n. (1); T. Nakai et al. s.n. (1); Nankai
University Exped. 54 (1); Nanling Exped. 622 (2); T. Neczaeva & V. Verholat 5584 (#3627) (1);
M. Noda et al. 94 (1), 120 (1), 641 (1); Northeast Normal University Exped. 882 (1); Northeast
Yunnan Exped. 238 (2).
H.G. Oga 5748 (1); H. Ohashi & Y. Fukuda s.n. (1); H. Ohashi & Y. Tateishi 686 (1).
N. Palczewsky 3627 (1), s.n. (1); C. Pei 7213 (2), 10239 (2); PE Exped. 1239 (1); F.S. Peng
473 (2), 1307 (2); Plant Exploration in the Republic of Korea 170 (1); Plant Geography Exped.
723 (2); Plants Exped. s.n. (1).
Y. & N. Quadota 3329 (1).
G. Raddle s.n. (1); Regel s.n. (1).
S. Saito s.n. (1), A-189 (1), 3582 (1), 7151 (1); K. Sakurai s.n. (1); C.S. Sargent s.n. (1); J.
Sato s.n. (1); M. Savatier s.n. (1); T. Sawada s.n. (1); L. Schrenk s.n. (1); Shaoguan Exped. 1111
(2); Z.S. Shen 1491 (2); Shennongjia Plants Exped. 11638 (2); Sichuan Agriculture University
Team 4966 (2), 5202 (2), 5254 (2); Sichuan Economic Plants Exped. 86 (2), 305 (2), 514 (2), 1374
(2), 2570 (2), 4421 (2); Sichuan Forest Institute 192 (2), 926 (2), 2200 (2), 6317 (2); Sino-
American Bot. Exped. 4173 (2), 4514 (2), 5241 (2); B.V. Skvortzow s.n. (1); Smithsonian-Korea
Ecological Project (Y.N. Lee & Y.S. Lee) 287 (1); C.S. Song 200 (1), 82-2 (1); South Central
China Forest University Exped. 30914 (2); T. Sugawara 1072905 (1), 2080913 (1), s.n. (1); T.
Sugawara & A. Makmoto s.n. (1); T. Sugawara & Y. Sugawara 1090805 (1); D.Y. Sun 80018
(1); S.C. Sun & K. Chang s.n. (2).
M. Takahasi s.n. (1); P.C. Tam 62264 (2), 62780 (2); M. Tamura et al. 22196 (1), 22202 (1);
C.M. Tan 95589 (2); G.G. Tang et al. 388 (2); P.C. Tang 790001 (1); D.D. Tao 11534 (2), 88002
(2); L. Taquet 4088 (1); Z. Tashiro s.n. (1); Y. Tateishi et al. 15055 (1); Temperate Forest Group
451 (1); X.F. Tian 194 (1); M. Togashi 668 (1), 7630 (1), s.n. (1); Y. Tokubuchi s.n. (1); Y.
Tomimoto s.n. (1), 2797 (1); P.C. Tsoong 68 (2); K. Tsuchiya 1390 (1); M. Tsuchiya 5919 (1);
M.K. Tsugaru 3213 (1); S. Tsugaru & T. Takahashi 6816 (1), 13565 (1), 13636 (1), 13655 (1),
13714 (1), 26198 (1), 27840 (1), 29433 (1).
J. Uatsumura s.n. (1); K. Ueda,S. Terabayashi & Y. Ueda 1321 (1); K. Uno 24139 (1); F.H.
Utech & M. Hoshi 89-155 (1); F.H. Utech et al. 91-360 (1).
C.S. Wang 4375 (1); C.S. Wang et al. 2513 (1), 3265 (1); D. Wang 2290 (1); F.T. Wang 169
(1); G.Z. Wang 184 (1), 4021 (1); G.Z. Wang & Q.T. Li 109 (1); G.Z. Wang et al. 162 (1); Q.L.
Wang & J.L. Zhang 188 (1); Q.S. Wang 1961 (2), 2102 (2), 2494 (2); W. Wang 1330, 2418 (1);
W. Wang et al. 511 (1); Y.M. Wang 457 (2); Z. Wang & Y.X. Liu 1458 (1); Z.T. Wang et al.
870190 (2); Wengyuan Herbs Exped. 64 (2); E.H. Wilson 161 (2), 1286 (2), 2739 (2), 3227 (2),
3228 (2), 3566 (2), 3567 (2), 4217 (2), 6770 (1), 6771 (1), 6870 (1), 6893 (1), 6898 (1), 7058 (1),
150 J I N S H U A N G M A ET AL.
7626 (1), 8507 (1), 8599 (1), 8696 (1), 8874 (1), 8902 (1), 9606 (1), 10909 (1), s.n. (1); Q.R. Wu 9
(1); S.Z. Wu 70092 (2); Wulingshan Plants Exped. 1381 (2), 1654 (2), 2896 (2); Wuyishan
Exped. Team 2081 (2); Xinning Exped. 1028 (2).
J.H. Xiong & Z.L. Zhou 91117 (2), 91464 (2), 91916 (2), 92732 (2); J.H. Xiong et al. 33484
(2).
Y. Yabe s.n. (1); J. Yamazaki 162 (1); T. Yamazaki M75-324 (1), s.n. (1); Yanbian First
Group 322 (1); Yanbian Second Group 60 (1), 761 (1); C.H. Yang & Q.T. Zhang 84-667 (2);
G.H. Yang 56125 (2); W.H. Yang 96 (1); Y.L. Yang 798 (1); Z.C. Ye 193 (2), 504 (2), 671 (2);
Yingkou Exped. 146 (1); I. Yokouchi s.n. (1); K. Yonekura 5959 (1); O. Yongsok 6083 (1); S.W.
Yu 86025 (2); J.S. Yue 3610 (2); J.S. Yue et al. 4233 (2).
G.C. Zhang 351 (2); J.Z. Zhang & X.R. Chen s.n. (2); Y.L. Zhang 39 (1); Y.L. Zhang et al.
294 (1), 959 (1), 1718 (1), 1785 (1); Z.S. Zhang & Y.T. Zhang 3013 (2), 3383 (2), 8469 (2); Z.E.
Zhao 2381 (2); Zhejiang Plant Inventory Team 28249 (2), 29493 (2); S.Q. Zhou 1181 (1); T.Y.
Zhou et al. 2178 (1), 7346 (1); Y.L. Zhou 2076 (1); Y.L. Zhou et al. 2921 (1); H.Q. Zhu 679 (2),
773 (2); Z.S. Zhu 754 (2).
A REVISION OF PHELLODENDRON 151
... Prior to a recent revision of the genus Phellodendron [9], the species may have been overlooked as an introduced member of the local flora due to confusion in the nomenclature. Greller [10] and Bertin et al. [11] both reported P. japonicum, a species now included within the variable P. amurense, as a part of their floristic works in the northeastern region. ...
... Their work reports that cultivated collections at the New York Botanical Garden contained P. amurense, P. chinense, P. japonicum, P. lavallei and P. sachalinense. With the exception of P. chinense, the additional four species have all now been designated as P. amurense [9]. At the site of a large invasion within the hemlock forest of the New York Botanical Garden, the P. amurense population has shown wide diversity in its morphology in both the leaflet base shape and the leaflet tomentum, [12] possible character differences which may continue to lead to confusion in correctly identifying this species. ...
... and P. amurense Rupr. (Ma et al. 2006). Phellodendron chinense is endemic to Anhui, Hubei, Hunan, Sichuan and Yunnan provinces, southern China, scattering in subtropical broad-leaved forests or mixed forests (Zhang et al. 2019). ...
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... 생물종 내 유전다양성 과 지역적인 분포 및 유전구조의 특성 평가는 환경변화에 대한 잠재적인 적응성을 평가하는 척도가 될 수 있으며, 생물다양성 보존을 위한 효율적인 전략을 제시 할 수 있 다 (Hughes et al., 2008). (Ma et al., 2006). ...
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... Although a " primitive " position of this group within Rutaceae has been suggested by Waterman and Grundon (1983) and by Da , it is not supported by the genera included in the current study. Phellodendron (Toddalioideae, two species in Japan, China, and eastern Russia; Ma et al., 2006) is close to Zanthoxylum in the trnL-trnF study of Scott et al. (2000) only in the neighbor-joining tree and in the study of Chase et al. (1999) only in the atpB , not the rbcL , analysis. In a more recent study of ITS and trnL-trnF regions ( Poon et al., 2007 ), Phellodendron , Tetradium , Toddalia , and Zanthoxylum ( Fagaropsis was not sampled) indeed formed a clade, and, thus, the proximity of these genera suggested by the presence of 1-BTIQ alkaloids was supported. ...
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