A revision of Elaeagnus L. (Elaeagnaceae) in mainland China

Abstract

A revision of Elaeagnus L. for mainland China is provided based on field observations and herbarium studies. Forty-two morphological characters are selected and coded, then the matrix is prepared following cluster analysis. Morphological characters and species delimitation are re-evaluated resulting in the recognition of 36 species, one subspecies and five varieties in mainland China. Thirteen species names and three variety names are regarded as new synonyms. Species information covers full synonyms, type information, description, taxonomic remarks, distribution range, occupied habitats, examined specimens, relevant illustrations, and references to selected published illustrations and reports. A key for whole species determination is provided. The lectotypification of Elaeagnus magna (Serv.) Rehd. has been designated.

Full text

Journal of Systematics and Evolution 48 (5): 356–390 (2010) doi: 10.1111/j.1759-6831.2010.00085.x
A revision of Elaeagnus L. (Elaeagnaceae) in mainland China
1,2Miao SUN§1Qi LIN∗
1(State Key Laboratory of Systematic and Evolutionary Botany, Institute of Botany, Chinese Academy of Sciences, Beijing 100093, China)
2(Graduate University of Chinese Academy of Sciences, Beijing 100049, China)
Abstract A revision of Elaeagnus L. for mainland China is provided based on field observations and herbarium
studies. Forty-two morphological characters are selected and coded, then the matrix is prepared following cluster
analysis. Morphological characters and species delimitation are re-evaluated resulting in the recognition of 36
species, one subspecies and five varieties in mainland China. Thirteen species names and three variety names are
regarded as new synonyms. Species information covers full synonyms, type information, description, taxonomic
remarks, distribution range, occupied habitats, examined specimens, relevant illustrations, and references to selected
published illustrations and reports. A key for whole species determination is provided. The lectotypification of
Elaeagnus magna (Serv.) Rehd. has been designated.
Key words cluster analysis, Elaeagnaceae, Elaeagnus, relationships, synonymy, taxonomic revision.
Elaeagnaceae is a small family with three genera:
Elaeagnus L., Hippophae L. and Shepherdia Nutt. Tem-
perate Eurasian Hippophae as well as North Ameri-
can Shepherdia only consists of three species, whereas
Elaeagnus L. is the largest with a wide distribution in
eastern Asia, extending to Southeast Asia and Queens-
land in northeastern Australia. A few species occur in
southern Europe and North America (Heywood et al.,
2007). The diversity of the genus Elaeagnus is centered
in China. It is mainly distributed from the Yangtze River
valley to its southern region, but is also found in north-
west China, growing in diverse habitats at elevation 50–
3100 m, such as thicket near seashore, streamside in
valleys, forests, roadside, shrub land, and rocky slope.
This genus was established by Linnaeus (1753).
Although Britton & Brown (1913) designated Elaeag-
nus angustifolia as the type species of Elaeagnus,it
turned out to be ineffective (Tropicos, available at
http://www.tropicos.org). The lectotypification desig-
nated by Hitchcock & Green (1929) is acceptable. On
the worldwide scale, taxonomic studies were conducted
by Schlechtendal (1857, 1860) and Servettaz (1908,
1909). Schlechtendal (1860) recognized 28 species in
this genus, but only four of them occur in China. Knowl-
edge about Elaeagnus was greatly updated in Servettaz’s
monograph (Servettaz, 1909), where 44 species were re-
ported with more than 20 species occurring in China.
Received: 30 December 2009 Revised: 5 May 2010 Accepted: 25 May
2010
∗Author for correspondence. E-mail: linqi@ibcas.ac.cn; Tel.: 86-10-
62836476; Fax: 86-10-62590296.
§E-mail: cactus@ibcas.ac.cn.
Additionally, for the first time he divided this genus
into two sections: sect. Sempervirentes Serv. and sect.
Deciduae Serv. Both sections’ names were accepted by
Zoku (1965) and Chang (1983).
Chang (1983) systematically studied Elaeagnus
from China and prepared a taxonomic treatment of
the genus Elaeagnus for Flora Reipublicae Popularis
Sinicae with 55 species recognized. Authors of most
local Chinese floras (Yao, 1985; Jin, 1993; Ding &
Wang, 1997; Zhu, 1997; Ruan, 2000; Du, 2006) fol-
lowed Chang’s treatment. Since Chang’s work (1983),
several new species of Elaeagnus have been described
(Chang, 1984, 1985, 1986; Xu, 1985; Fang & Liang,
2000; Qi & Lin, 2000). The number of published species
has increased from the 55 recognized by Chang in 1983
to 67 discerned in the recent taxonomic treatment made
by Qin & Gilbert (2007), in whose study, even some
newly described species were not recorded, e.g., E.
hunanensis C. J. Qi & Q. Z. Lin (Qi & Lin, 2000).
To a certain extent, this genus still remains poorly
known. Qin & Gilbert (2007) noticed that: (i) quantita-
tive characters are frequently used as distinguished char-
acters among species of Elaeagnus; (ii) there is a lack
of information available regarding the infraspecific mor-
phological variation; (iii) the key was prepared mostly
based on vegetative and floral characters; (iv) fruit char-
acters of many species are not recorded; and (v) more
comprehensive study of Elaeagnus should lead to a sig-
nificant reduction in the number of recognized species.
Du (2006) made an effort to combine five previously
described species. His work was based on the exami-
nation and revision of a limited number of herbarium
specimens without the advantage of field work. In
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SUN & LIN: A revision of Elaeagnus in China 357
summary, the studies undertaken hitherto may not com-
pletely reflect the diversity of the entire genus of Elaeag-
nus in China.
Our work presented here summarizes the results of
taxonomic research on the genus Elaeagnus in mainland
China. Species of Elaeagnus for the most part are well
defined, some specimens appear to be both morpholog-
ically and geographically intermediate among several
species. Some names represent as synonyms and need
to be combined. Our knowledge about some species of
Elaeagnus is still inadequate (e.g., Elaeagnus bambuse-
torum Hand.-Mazz.). Additional collections are needed
to determine their appropriate status. An effort should
be made to find a more precise range of variation of mor-
phological characters in recognized species. The rele-
vant studies should be undertaken which have power to
prove or reject hybrid hypotheses. Our study will in-
troduce several nomenclatural and taxonomic changes,
and will provide a taxonomic framework for further
floristic studies prior to a complete revision of this
genus.
1 Material and methods
Extensive field and herbarium studies were car-
ried out in Chongqing, Guangdong, Guangxi, Guizhou,
Hubei, Jiangxi, Sichuan, Xizang, Yunnan, and Zhejiang.
Comprehensive bibliographic research on the taxonomy
of Elaeagnus was carried out, with a special focus on
species in Guizhou, Sichuan, Xizang, and Yunnan. The
protologues, local floras, and monographs were con-
sulted to document all reports of Elaeagnus from this
area. More than 7000 specimens (including 161 type
specimens) were examined from the following herbaria:
CDBI, GXMI, HGAS, HHBG, IBK, IBSC, K, KUN,
LBG, N, NAS, NAU, P, PE, SYS, SWFC, SZ, YAF,
YCP, and YUKU (abbreviations according to Holmgren
& Holmgren, 1998– and Fu, 1993). The type specimens
of all recognized species and nearly all types of their
synonyms were examined.
The taxonomic analysis was based on extensive
morphological and anatomical comparisons. Then 42
morphological characters (Table 1) were selected for
the descriptions and differentiation among the species.
Some anatomical sections such as details on flowers
were prepared from dry herbarium specimens after hy-
dration in a soap solution and observed under a stere-
omicroscope (SMZ1000; Nikon, Japan).
Each studied species was described using charac-
ters mentioned in Table 1. In this way matrix data were
obtained (Table 2), which were processed with statisti-
cal analysis in order to find possible relationships among
studied species of Elaeagnus. The matrix data were pre-
pared (Table 2), following the principles as outlined
below: (i) each character with three states was assigned
with equal weight, and constantly presents on most spec-
imens studied; (ii) the coding data in matrix did not have
priority; (iii) code numbers represented no taxonomic
meanings (e.g., compared with “1”, “0” does not rep-
resent the character in a plesiomorphic state, and vice
versa); and (iv) the missing characters were represented
by “?”. Both cluster analyses were carried out using
STATISTICA version 8.0 software (StatSoft, 2007).
2 Results and discussion
Results of cluster analysis and observation of spec-
imens concerning 42 characters indicated that the good
characters for distinguishing infrageneric taxa in genus
Elaeagnus are habit, anthesis, calyx shape, presence
of different types of trichomes (Zhang et al., 1992)
on branches as young, leaves and calyxes and fruits,
fruit texture, and presence of fruit wing (Table 1). Most
species (22/36) of this genus are evergreen distribut-
ing in the Yangtze River valley and its southern region,
flowering at autumn and winter. Combined with leath-
ery leaves, these species can be immediately separated
from the rest of the species in the genus. Species with
evergreen habit are generally uniform (leathery leaves,
covered with scales, and juicy fruits), but they can be
easily divided into several small groups by the calyx
shape (campanulate, tubular, and urceolate), then iden-
tified by other secondary characters (shape of leaf blade,
color below leaf surface, or length of pedicel). The re-
mainder of this genus are in defoliation during autumn
and winter with papery leaves. Although some of them
seem semi-evergreen (e.g., E. stellipila), in fact they
are usually deciduous at spring and easily recognized
by stellate hairs or long-stalked stellate hairs on young
branches, leaves, and flowers. Elaeagnus mollis is eas-
ily distinguished by its fruit wing and covering stellate
hairs. Likewise, fruit texture and silvery scales are use-
ful characters for E. oxycarpa and E. angustifolia. The
rest can be recognized by secondary characters (e.g.,
length of pedicel, fruit shape). In summary, these are
key characters, constant, reliable, and easily recognized
on live plants as well as herbarium specimens. They
have significantly contributed to our identification key
and determination of species.
Results of cluster analysis indicated potential rela-
tionships among species of Elaeagnus and resulted in
two overall clusters: I, Sempervirentes; and II, Elaeag-
nus (Deciduae). Each cluster comprises two groups: IA
and IB; and IIAand IIB(Fig. 1). These two main clusters
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358 Journal of Systematics and Evolution Vol. 48 No. 5 2010
Tab l e 1 Morphological characters and states and code numbers used in the data matrix of cluster analysis of Elaeagnus L. (Elaeagnaceae) in
mainland China
1. Habit: evergreen (0); semievergreen or deciduous (1);
2. Type of life form: climbing shrub (0); erect shrub (1); tree (2);
3. Spines on branches: absent (0); present (1);
4. Shape of young branches: columned (0); woodiness up-ridge (1);
5. Color of hairs on branches: yellowish white or silvery (0); rust-colored or ferruginous (1);
6. Type of hairs on young branches: stellate (0); stellate-peltate (1); scales (2);
7. Texture of leaves: leathery or sub-leathery (0); papery or membranous (1);
8. Speckles on leaves: absent (0); present (1);
9. Color below leaf surface: yellowish white or silvery (0); rust-colored or ferruginous (1);
10. Shape of leaf blade: round or ovate (0); oblong or elliptic (1); lanceolate (2);
11. Shape of leaf apex: round or obtuse (0); acute or acuminate (1);
12. Shape of leaf margin: revolute (0); entire(1);
13. Shape of leave base: round or obtuse (0); attenuate or cuneate (1);
14. Hairs on adaxial leaf surface: stellate (0); stellate-peltate or mixture (1); scales (2);
15. Hairs on abaxial leaf surface: stellate (0); stellate-peltate or mixture (1); scales (2);
16. Type of inflorescence: solitary or 1–3 (including 3) (0); around 5 in axils (1); racemose (2);
17. Length of flower pedicel: short or nearly absent (0); ≤5mm(1);>5mm(2);
18. Flower color: yellowish white or white (0); yellow or yellowish green (1);
19. Shape of calyx: urceolate (0); campanulate (1); tubular or tubular-funnelform (2);
20. Hairs outside of calyx: stellate (0); stellate-peltate or mixture (1); scales (2);
21. Texture of calyx: membranous (0); papery (1); sub-leathery (2);
22. Calyx tube length: ≤5mm(0);>5mm(1);
23. Calyx tube width: ≤3mm(0);>3mm(1);
24. Shape of calyx lobe: triangular or ovate-triangular (0); ovate or round (1);
25. Presence of hairs inside calyx: stellate (0); stellate-peltate or mixture (1); scales (2);
26. Calyx lobe length: ≤3mm(0);>3mm(1);
27. Calyx lobe width: ≤3mm(0);>3mm(1);
28. Anther shape: oblong (0); ellipsoid (1);
29. Filament length: nearly absent or ≤1mm(0);>1mm(1);
30. Type of style: erect (0); curve or S shape (1); curve in the upper part (2);
31. Presence of hair on style: absent (0); present (1);
32. Stigma position: under or equal height to stamen (0); beyond stamen (1);
33. Shape of fruit: nearly globose or obovoid (0); ellipsoid (1);
34. Fruit size: ≤8 mm (0); 8–15 mm (1); 15–20 mm (2); >20 mm (3);
35. Presence of wings on fruit surface: absent (0); present (1);
36. Fruit surface type: hairy (0); scaly (1);
37. Ripe fruit color: red (0); yellowish brown or yellow to orange (1); yellowish gray (2);
38. Texture of fruit flesh: dryish mealy (0); fleshy and juicy (1);
39. Presence of ribs on seed surface: inconspicuous (0); conspicuous (1);
40. Type of fruit pedicel: erect, robust (0); slender, nodding (1);
41. Length of fruit pedicel: short or nearly absent (0); ≤5mm(1);>5mm(2);
42. Anthesis: spring, summer (0); autumn, winter (1);
responded with traditional circumscriptions of sections:
Section I. Sempervirentes Serv. and Section II. Decid-
uae Serv. (Elaeagnus L. is correct, and see Section II.
Elaeagnus L. in text for explanation), which were dis-
tinguished by Servettaz (1909) based on leaves ever-
green or deciduous, floral position, and anthesis (Fig. 1).
Cluster I included 22 evergreen species characterized by
leaves evergreen and leathery, and flowering mainly in
winter. Cluster II contained 14 species characterized by
leaves mostly deciduous, papery, and flowering mainly
in summer.
Species in Cluster I are all evergreen shrubs, some-
times climbing, and flowering in autumn or winter,
rarely in spring. Its component groups IAand IBap-
parently seem to be divergent from their calyx shapes,
otherwise most other characters are more or less uni-
form in this section. Group IAis well characterized by
the tubular calyx. However, E. oldhamii seems to be
an exception for its different characters, obovate leaf
blade, and cup-shaped calyx tube. Because its mor-
phological differences would be weakened after taking
into consideration E. tucheri,E. pungens, and other re-
lated species from Taiwan (e.g., E. thunbergii Serv.),
further study is needed in coastland areas, especially
Taiwan. With regard to E. viridis, we have expanded
its circumscription and access in section 3 “Taxonomic
treatment”. Elaeagnus tubiflora seems to be the clos-
est relative to E. delavayi, although its calyx tube is
up to 10 mm long. Eleven species in group IBwere
characterized by campanulate calyx and color below
leaf surface (scales), although their calyx sizes varied
from species to species. Elaeagnus luxiensis seemed to
be morphologically intermediate between E. macran-
tha and E. loureiroi, and the circumscription between
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SUN & LIN: A revision of Elaeagnus in China 359
Tab l e 2 Data matrix of 36 species of Elaeagnus L. (Elaeagnaceae) from mainland China and 42 morphological characters used for construction
of cluster analysis
Tax a Characters†
000000000111111111122222222223333333333444
123456789012345678901234567890123456789012
E. angustata 110112100111012020220101000000111101010120
E. angustifolia 121002100201112110121000201000011101100110
E. argyi 111102100001002211222101100101010101011120
E. bockii 001012010210122210221100100001111101010111
E. calcarea 111010100111000210211000000100111201011010
E. cinnamomiflia 00010200001002222022211011100000??01? 1???1
E. conferta 000112100111022200011010200000111201011011
E. courtoisii 110100100011000020210100010000011100011120
E. delavayi 010112010101122221221110110000101101110121
E. difficilis 001012001111022211121011100001011101011111
E. glabra 000012001110022211221001011000011201011111
E. gonyanthes 000012011101022221111011211000011201010021
E. grijsii 101010001011000210101100000000101101011021
E. guizhouensis 11011100101100020022110100000001110101?010
E. henryi 011112000010022210222101000000011201010111
E. jingdongensis 110010101111000220201001000110011100010020
E. lanceolata 011002000210022210211100100001101101011111
E. liuzhouensis 01001210100010121022100000000010??01?1?111
E. loureiroi 000012001111022211121111211011111201011121
E. luxiensis 010112101111022211121111011000011201011121
E. macrantha 010012101110021210111110011010011301011021
E. macrophylla 010112100010022110121011000001000201011111
E. magna 111102110011122100221101000001111101011010
E. mollis 110100100001000111001011000000110110201010
E. multiflora 111112101011011020211101011000011101011120
E. oldhamii 011012101000122210121010000000100001011111
E. oxycarpa 111102100201111110111000000000100101100110
E. pingnanensis 001102000011022211122110211010110201010111
E. pungens 011012001100022210211001000000101101010111
E. sarmentosa 001112101111022120221110010000011301011121
E. stellipila 111010101011000201201000010000011101011110
E. tonkinensis 011112111101022210221001000001101101010111
E. tubiflora 01001211121102222022110101000110??01?1?121
E. tutcheri 010?12001100011210121011000000011101010111
E. umbellata 111102100011012210220100000000100001011010
E. viridis 01111200100002221022100100000011110101?121
Cluster analysis was based on information from herbarium studies and field observations. †Descriptions and codings of characters and states are detailed
in Table 1. ?, missing information.
E. cinnamomifolia and E. pingnanensis is not clear
enough. In summary, additional collections and field
work relevant to these taxa are needed.
However, in Cluster II, groups IIAand IIBare char-
acterized by the presence of different types of trichomes,
anthesis, and fruit texture. Group IIAincluded E. mol-
lis,E. oxycarpa, and E. angustifolia. On the basis of
its fruit wing and stellate hairs (Table 2), E. mollis
was distinctively separated from the above two species
as a sister group. It is possible to distinguish some
groups within 11 members of group IIB. The five species
E. guizhouensis,E. calcarea,E. stellipila,E. grijsii,
and E. jingdongensis seem to form a natural group re-
stricted only to limestone areas, sharing characters such
as semi-evergreen shrubs, seasonal dimorphiced leaves,
and of stellate hairs on abaxial leaf surfaces and young
branches. Among the rest, E. umbellata,E. magna,
E. multiflora and their infraspecific taxa seem to form
a natural group characterized by deciduous shrubs, pa-
pery leaf blades, leaves densely covered with silvery
scales, tubular calyx, and flowering in spring or sum-
mer. Problematically, E. courtoisii seems to be an in-
termediate between the two groups mentioned above,
with several combinations of characters, such as stellate
hairs and long pedicels. Elaeagnus wushanensis and
E. nanchuanensis were considered as regional variation
of E. magna with few distinct characters. They may al-
ternatively be treated as varieties of a single variable
and widespread species, E. magna.
Elaeagnus multiflora Thunb. var. longipedunculata
(N. Li & T. N. Wu) M. P. Deng & K. Yao was described
as a variety of E. multiflora, distinctive due to its long
pedicels. Given the protologue of this variety is not
available to us at present and also that the International
Plant Names Index (available at http://www.ipni.org)
indicates that this name is not valid, we suggest treating
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360 Journal of Systematics and Evolution Vol. 48 No. 5 2010
Fig. 1. Tree diagram for 36 species of Elaeagnus L. (Elaeagnaceae) from mainland China. Ward’s method using the Euclidean distance shows the
interrelationships between species. The whole cluster of Elaeagnus is contained in two overall clusters: I, Sempervirentes; and II, Elaeagnus (Deciduae).
Each cluster comprises two groups, IAand IB,andII
Aand IIB.
this variety within the circumscription of E. multiflora
and will verify the name in the future.
In conclusion, based on examination of specimens,
field observations, and statistical analyses, 36 species,
one subspecies and five varieties in mainland China
are accepted in this study. Elaeagnus angustata (Rehd.)
C. Y. Chang var. songmingensis W. K. Hu & H. F.
Chow is regarded as a new synonym of E. angustata
(Rehd.) C. Y. Chang; E. angustifolia L. var. orientalis
(L.) Kuntze as a new synonym of E. angustifolia L;
E. bockii Diels var. muliensis C. Y. Chang as a new
synonym of E. bockii Diels; E. difficilis Serv. var. bre-
vistyle W. K. Hu & H. F. Chow as a new synonym of
E. difficilis Serv.; E. geniculata D. Fang as a new syn-
onym of E. gonyanthes Benth.; E. heterophylla D. Fang
& D. R. Liang and E. luoxiangensis C. Y. Chang as new
synonyms of E. glabra Thunb.; E. hunanensis C. J. Qi
& Q. Z. Lin as a new synonym of E. lanceolata Serv.
subsp. grandifolia Serv.; E. jiangxiensis C. Y. Chang as
a new synonym of E. grijsii Hance; E. longiloba C. Y.
Chang, E. retrostyle C. Y. Chang, and E. xingwenensis
C. Y. Chang as new synonyms of E. delavayi Lecomte;
E. micrantha C. Y. Chang as a new synonym of E. ar-
gyi L´
evl.; E. obovatifolia D. Fang as a new synonym of
E. pingnanensis C. Y. Chang; E. obtusa C. Y. Chang as
a new synonym of E. pungens Thunb.; E. xichouensis
C. Y. Chang and E. xizangensis C. Y. Chang as new
synonyms of E. viridis Serv. Two species, E. nanchua-
nensis C. Y. Chang and E. wushensis C. Y. Chang, are
reinstated as E. magna (Serv.) Rehd. var. nanchuanensis
(C. Y. Chang) M. Sun & Q. Lin and E. magna (Serv.)
Rehd. var. wushanensis (C. Y. Chang) M. Sun & Q. Lin,
respectively. Species that have long been recognized
and are also accepted by us have not been included in
the taxonomic treatment below. They are E. calcarea
Z. R. Xu, E. cinnamomifolia W. K. Hu & H. F. Chow,
E. conferta Roxb., E. courtoisii Belval, E. guizhouensis
C. Y. Chang, E. henryi Warb. , E. jingdongensis C. Y.
Chang, E. lanceolata Warb. subsp. lanceolata Warb.,
E. liuzhouensis C. Y. Chang, E. loureiroi Champ., E. lux-
iensis C. Y. Chang, E. macrantha Rehd., E. macrophylla
Thunb., E. mollis Diels, E. multiflora Thunb., E. old-
hami Maxim., E. oxycarpa Schlechtend., E. sarmentosa
Rehd., E. stellipila Rehd., E. tonkinensis Serv., E. tubi-
flora C. Y. Chang, E. tutcheri Dunn and E. umbellata
Thunb.).
3 Taxonomic treatment
Elaeagnus L., Sp. Pl. 1: 121. 1753.—Oleaster
Heist. ex Fabr. in Enumeratio Methodica Plantarum 214.
1759.—Octarillum Lour., Fl. Cochinch. 90. 1794.—
Aeleagnus Cav., Descr. Pl. Lecc. Publ. 350. 1802. Lec-
totype: E.angustifolia L. (Hitchcock & Green, 1929).
Deciduous or evergreen; shrubs, erect or some-
times climbing, or small trees, sometimes spiny, con-
spicuously covered by silvery white or rust-colored
scales or stellate hairs. Leaves alternate, papery, mem-
branous, or leathery, petiolate, blade margin usually
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SUN & LIN: A revision of Elaeagnus in China 361
entire. Flowers bisexual, 3 or more clustered on short ax-
illary shoots, sometimes solitary. Calyx tubular, 4-lobed,
constricted above ovary and breaking at constriction at
fruiting stage; lobes usually spreading, deciduous, white
or yellow inside. Stamens 4, inserted in mouth of calyx
tube, alternate with lobes. Style linear. Fruit globose
or ellipsoid, rarely longitudinally winged (e.g., E. mol-
lis Diels); seed usually 8-ribbed, with a large straight
embryo.
Key to sections and species
This key has been prepared for all 36 species we
recognize, and the order of species that appears in the
detailed description below is concordant with the order
arranged in the key.
1a. Evergreen shrubs, erect or climbing; leaf blades
leathery or nearly so, rarely papery; flowering
in autumn or winter, rarely in spring; fruiting in
spring and summer: Sect. I. Section Semper-
virentes ..................................2
1b. Deciduous or semi-evergreen trees or shrubs,
rarely climbing; leaf blades papery or membra-
nous, rarely leathery; flowering in spring or sum-
mer; fruiting in summer or autumn: Sect. II. Sec-
tion Elaeagnus .......................... 23
2a. Calyx tube campanulate or broadly tubular . . . 3
2b. Calyx tube tubular or funnelform-tubular. . . .13
3a. Flower bigger, calyx tube 7–10 mm. . . . . . . . . . 4
3b. Flower smaller, calyx tube 2–6 mm . . . . . . . . . . 8
4a. Leaves abaxially with dense silvery or yellow-
ish white scales; flowers with dense silvery and
yellowish white scales . . . . . . . . . . . . . . . . . . . . . . 5
4b. Leaves abaxially with dense or sparse rust-
colored scales; flowers with dense or sparse rust-
colored scales . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
5a. Leaf blade papery, elliptic; flowers densely with
white scales, calyx papery, 7 mm long, lobes
5mmwide................. 1. E. macrantha
5b. Leaf blade leathery, ovate or obovate; flowers
densely with yellowish white scales, calyx thick,
4–5.5 mm long, lobes 4–4.5 mm wide . . . . . . . . .
..........................2. E. pingnanensis
6a. Calyx tube copular, conspicuously 4-angled, 4–
6 mm, lobes inside densely with peltate scales
only; fruiting pedicels 1.2–2.5 cm long, robust,
drupe broadly ellipsoid or ovoid-ellipsoid, 1.5–
2.2cm ..................... 3. E. gonyanthes
6b. Calyx tube broadly campanulate, faintly 4-
angled, 6–11 mm, lobes inside densely with stel-
late hairs and peltate scales; fruiting pedicels
7–11 cm long, slimly bending, drupe ellipsoid,
1.5–2.2 cm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
7a. Leaves abaxially with dense rust-colored peltate
scales, lateral veins abaxially inconspicuous and
not raised; flowers densely with rust-colored
peltate scales, lobes 5–7 mm . . . . 4. E. loureiroi
7b. Leaves abaxially with dense silvery white scales
and sparsely rust-colored peltate scales, lateral
veins abaxially conspicuous and raised; flow-
ers densely with silvery white peltate scales
and sparsely with ferruginous scales, lobes
4–4.5 mm . . . . . . . . . . . . . . . . . . . . . 5. E. luxiensis
8a. Spines present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
8b. Spines absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
9a. Leaves abaxially densely with rust-colored
peltate scales; flowers densely with ferruginous
scales; drupe 1.3–1.8 cm . . . . . . . . 6. E. difficilis
9b. Leaves abaxially densely with silvery white
peltate scales; flowers densely with silvery white
scales; drupe 0.7–1.2 cm . . . . . . . . . . . . . . . . . . 10
10a. Leaf blade papery, elliptic, base obtuse, apex
obtuse; calyx tube 3.5–4 mm; drupe narrowly
ellipsoid, 1.2 cm . . . . . . . . . . . . 7. E. tonkinensis
10b. Leaf blade nearly leathery, obovate or ovate-
lanceolate, base cuneate, apex rounded; calyx
tube ca. 2 mm; drupe globose or nearly so,
0.7–0.9 cm . . . . . . . . . . . . . . . . . . . 8. E. oldhamii
11a. Young branches prismatic; leaf blade ovate or
nearly orbicular, base rounded or cordate, petiole
1.1–1.2 cm, robust . . . . . . . . . 9. E. macrophylla
11b. Young branches cylindrical; leaf blade elliptic,
base rounded or cuneate, petiole 0.6–1 cm, not
robust...................................12
12a. Leaf blade leathery, surface often bullate, apex
obtuse; calyx tube camupanulate, lobes erect;
drupe 1–1.2 cm . . . . . . . . . . . . . . . 11. E. tutcheri
12b. Leaf blade papery, surface flat, apex cuspidate;
calyx tube urceolate, lobes not erect; drupe 1.3–
4cm.........................10. E. conferta
13a. Scandent shrub, sometimes erect . . . . . . . . . . . 14
13b.Erectshrub .............................. 17
14a. Calyx tube cylindrical . . . . . . . . . . . . . . . . . . . . . 15
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362 Journal of Systematics and Evolution Vol. 48 No. 5 2010
14b. Calyx tube funnelform-tubular . . . . . . . . . . . . . 16
15a. Spines present. Leaf blade leathery, lanceolate,
abaxially densely with yellowish white peltate
scales; flowers densely with yellowish white
scales, calyx tube 5–6 mm . . . 12. E. lanceolata
15b. Spines absent. Leaf blade papery, oblong or
elliptic, abaxially densely or sparsely with
rust-colored peltate scales; flowers densely or
sparsely with rust-colored scales, calyx tube 8–
9mm.....................13. E. sarmentosa
16a. Branches yellowish white, no spines; lateral
veins abaxially conspicuous and raised, peti-
ole 0.8–1.6 cm, robust, densely with yellow-
ish white and light brown scales; calyx tube
8–9 mm, densely with yellowish white peltate
scales. . . . . . . . . . . . . . . . . 15. E. cinnamomifolia
16b. Branches brown, rarely spiny; lateral veins abax-
ially inconspicuous, petiole 0.5–1 cm, slim,
densely with rust-colored or brown scales; ca-
lyx tube 4.5–6 mm, densely with brown peltate
scales . . . . . . . . . . . . . . . . . . . . . . . . . 14. E. glabra
17a. Young branches rust-colored; calyx tube
funnelform-tubular . . . . . . . . . . . . . . . . . . . . . . . 18
17b. Young branches yellowish brown, rarely brown;
calyx tube cylindrical . . . . . . . . . . . . . . . . . . . . . 20
18a. Spines present; leaf blade leathery, abaxially
with peltate scales; calyx tube ca. 7 mm, style
glabrous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
18b. Spines absent; leaf blade thick papery, abax-
ially with stellate-peltate scales; calyx tube
4–4.5 mm, style with sparse white stellate
hairs ......... ...........18. E. liuzhouensis
19a. Leaves broad lanceolate to ovate, speckled, base
cuneate to rounded, apex cuspidate, petiole ro-
bust, with brown scales; pedicel up to 6 mm long;
drupe oblong, up to 2 cm . . . . . . . . 16. E. henryi
19b. Leaves elliptic or oblong-lanceolate, not speck-
led, base cuneate, apex obtuse or acumi-
nate, petiole slim, with rust-colored scales;
pedicel up to 12 mm long; drupe ellipsoid,
ca.1.2cm....................17. E. delavayi
20a. Spines present; leaf blade abaxially with silvery
white or sparsely ferruginous peltate scales; ca-
lyx tube 4–7 mm . . . . . . . . . . . . . . . . . . . . . . . . . 21
20b. Spines absent; leaf blade abaxially with
rust-colored peltate scales; calyx tube
9–10 mm . . . . . . . . . . . . . . . . . . . . 19. E. tubiflora
21a. Leaf blade oblong or oblong-lanceolate, abax-
ially with silvery white peltate scales; base
cuneate; calyx tube sometimes up to 10 mm long,
lobe triangular, inside densely stellate-peltate
scales . . . . . . . . . . . . . . . . . . . . . . . . . .20. E. bockii
21b. Leaf blade elliptic, base rounded, abaxially with
silvery white or sparsely ferruginous peltate
scales; calyx tube up to 8 mm long, lobe ovate,
inside densely stellate hairs . . . . . . . . . . . . . . . . 22
22a. Leaf blade leathery, abaxially densely or
sparsely with brown peltate scales; calyx tube
densely with brown or ferruginous scales;
pedicel 3–5 mm, erect; fruiting pedicel 4–
6mm........................21. E. pungens
22b. Leaf blade nearly leathery or papery, abaxi-
ally densely with grayish white peltate scales,
sparsely rust-colored scales; calyx tube densely
with silvery white scales, sparsely ferruginous
scales; pedicel 6–18 mm, nodding; fruiting pedi-
celca.10 mm..................22. E. viridis
23a. Leaf blade nearly leathery or papery; fruit
juicy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26
23b. Leaf blade papery; fruit not juicy . . . . . . . . . . . 24
24a. Spines absent; leaf blade elliptic or oblong;
young branches, leaves, flowers, fruit densely
with stellate hairs; fruit 8-winged, flesh mealy
soft............................23. E. mollis
24b. Spines present; leaf blade lanceolate or ellip-
tic; young branches, leaves, flowers densely
with peltate scales or stellate-peltate; fruit no
8-winged, flesh juicy . . . . . . . . . . . . . . . . . . . . . . 25
25a. Floral disk usually glabrous, rarely hairy;
drupe yellowish brown, oblong or narrowly
cylindric, both ends obtuse, 0.9–2.5 ×0.5–
1.3cm....................24. E. angustifolia
25b. Floral disk pubescent; drupe chestnut yellow,
globose-ovoid or subglobose, both ends pointed,
0.7–1.2 ×0.6–0.8 cm . . . . . . . . 25. E. oxycarpa
26a. Leaf blade abaxially densely with stellate
hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27
26b. Leaf blade abaxially densely with peltate
scales . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33
27a. Young branches densely with peltate scales,
sparsely with stellate hairs or inconspicuous; ca-
lyx tubes densely with scales or stellate-peltate
scales . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28
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SUN & LIN: A revision of Elaeagnus in China 363
27b. Young branches densely with stellate hairs or
short-stalked stellate hairs; calyx tubes densely
with stellate hairs . . . . . . . . . . . . . . . . . . . . . . . . . 30
28a. Leaf blade dimorphic by season, spring leaves
small, elliptic to oblong, autumn leaves larger,
oblong-ovate or broadly elliptic; calyx tube
funnelform-tubular, 5.5–6 mm, thick; fruiting
pedicel 0.5–1 cm, slim, drupe obovoid ... ... ..
................................26. E. argyi
28b. Leaf blade not dimorphic by season or incon-
spicuous; calyx tube tubular, 4–4.5 mm, pa-
pery; fruiting pedicel 0.3–0.4 cm, dumpy, drupe
globose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29
29a. Leaf blade leathery, obovate, rarely elliptic-
lanceolate; drupe 1.5 cm . . . . . . . . . . . . . . . . . . . . .
.........................27. E. guizhouensis
29b. Leaf blade papery, elliptic; drupe 2.5–
3cm.........................28. E. calcarea
30a. Lobes lanceolate or triangular; fruiting pedicel
up to0.3cm..................29. E. stellipila
30b. Lobes ovate or ovate-triangular; fruiting pedicel
up to0.5–2cm ........................... 31
31a. Leaf blade leathery, ovate or broadly
ovate...........................30. E. grijsii
31b. Leaf blade papery, elliptic or obovate . . . . . . . 32
32a. Old leaf blade abaxially densely with stellate
hairs; fruiting pedicel 0.8–1.5 cm, drupe oblong
to broadly fusiform . . . . . . 31. E. jingdongensis
32b. Old leaf blade only veins abaxially with stel-
late hairs; fruiting pedicel up to 5 cm, drupe
ellipsoid . . . . . . . . . . . . . . . . . . . . 32. E. courtoisii
33a. Fruiting pedicel slim bending, 1.5–5 cm ...
...... ...................................34
33b. Fruiting pedicel fleshy, erect or nearly absent,
0.4–1cm................................ 35
34a. Leaf blade lanceolate; drupe ovoid, densely with
silvery white peltate scales . . . 33. E. angustata
34b. Leaf blade ovate or obovate; drupe ellip-
soid, densely with rust-colored peltate scales ...
...... .....................34. E. multiflora
35a. Calyx tube tubular; drupe ellipsoid, 1.2–2 cm
...............................35. E. magna
35b. Calyx tube funnelform-tubular; drupe globose or
subdlobose, 0.5–0.7 cm . . . . . . 36. E. umbellata
Section I. Elaeagnus L. section Sempervirentes
Serv. in Beih. Bot. Centralbl. 25(1): 25, 1909. Type:
Elaeagnus latifolia L.
Evergreen; shrubs, erect or climbing; leaf blade
leathery or nearly so, rarely papery; flowering in autumn
or winter, rarely in spring; fruiting in spring and summer.
Twenty-two species in this section chiefly occuring in
Yangtze River valley and its southern region.
Characters in this section are relatively uniform. In
wild, species of this section can be easily recognized by
leaves and flowers.
2. Elaeagnus pingnanensis C. Y. Chang in J. Sichuan
Univ., Nat. Sci. Ed. 4: 91. 1984. Type: China. Guangxi:
Pingnan, Penghua, C. Wang 40325 (holotype, IBSC!),
Fig. 2: A.
Elaeagnus obovatifolia D. Fang in Acta Phyto-
tax. Sin. 38: 289. 2000. Type: China. Guangxi: Napo,
Longhua, D. Fang & Z. G. Wang 78477 (holotype,
GMXI!), Fig. 2: B.
Elaeagnus obovatifolia F. Du, Fl. Yunnan. 12. 758.
2006, nom. invalid. Type: China. Yunnan: Funing, Z. H.
Hu 78250 (holotype, YUKU!), Fig. 2: C.
Evergreen; shrubs, climbing, ca. 4 m. Spines
present; young branches up-ridge, densely with silvery
white or yellowish white scales. Petiole reddish brown,
8–1.5 cm; leaves thickly leathery; blade green above,
yellowish white below, obovate or elliptic, apex bluntly
acute with triangular mucro or rarely obtuse, margin
slightly revolute or undulate, base rounded or broadly
cuneate; 5–12.7 ×3.5–7.3 cm, adaxially glossy, abax-
ially with overlapping yellowish silvery scales; lateral
veins 6–8 per side of midrib, robust, conspicuous on
both surfaces. Flowers ca. 7 in axils or subumbellate on
axillary shoots, 2–5 mm; pedicels 4–5 mm. Flowers yel-
low; tube broadly campanulate, slightly 4-angled when
dried, outside with dense yellowish white and sparse red-
dish brown scales; sub-leathery; 4–7.5 mm, 5–6 mm in
diam.; lobes broadly triangular, apex bluntly acuminate,
abruptly constricted above ovary, inside with deeply di-
vided scales, 4–5.5 ×4 mm; anthers oblong, ca. 1.5 mm;
filaments ca. 1.5 mm; style erect, glabrous or with sparse
white stellate hairs; stigma slightly exserted beyond sta-
mens. Fruit narrowly ellipsoid or oblong, 1.6–2 cm,
densely yellowish white scales, red when ripe, juicy;
ribs on seed inconspicuous; fruit pedicels slender, ca.
5 mm. Fl. Oct.–Dec., fr. Feb.–Mar.
Limestone hills or by streamside; alt. 1000–
1400 m. Endemic to China. Guangxi, Guizhou, and
Yunnan.
Additional specimens examined:
China. Guizhou: Ceheng, Z. Y. Cao 1121 (PE). Guangxi: Precise
locality unknown, s. n. (PE Herb. Code No. 01015281). Napo, H. N. Qin
et al. 2453 (PE). Pingnan, C. Wang 40325 (IBK, IBSC). Yunnan: Funing,
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364 Journal of Systematics and Evolution Vol. 48 No. 5 2010
Fig. 2. A, Holotype of Elaeagnus pingnanensis C. Y. Chang (C. Wang 40325, IBSC). B, Holotype of Elaeagnus obovatifolia D. Fang ( D. Fang & Z. G.
Wang 78477, GMXI). C, Holotype of Elaeagnus obovatifolia F. Du, nom. invalid, (Z. H. Hu 78250, YUKU). D, Holotype of Elaeagnus geniculata D.
Fan g (J. J. Wang 5449, GXMI).
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SUN & LIN: A revision of Elaeagnus in China 365
Z. H. Hu 78250 (YUKU), Z. Z. Ding & J. X. Wang 1262 (NAS). Malipo,
K. M. Feng 12906 (PE).
Elaeagnus pingnanensis C. Y. Chang was de-
scribed by Chang (1984) based on one collection made
in 1936 from Pingnan, Guangxi (Fig. 2: A) without
fruits. Likewise, 16 years later, a new species recog-
nized by Fang & Cactus Liang (2000) from Guangxi
under the name E. obovatifolia was also based on one
flowering specimen without fruits (Fig. 2: B). Fang and
Liang (2000) stated that E. obovatifolia was similar to
E. pingnanensis, but characters like leaf blade shape,
color, anthers, filaments, and stigma could be used to
distinguish them. After examining the type specimens
of the two, we found that the shape of leaf blade was
not a good distinguishing character, because ovate, el-
liptic, and obovate leaves were all present on the type
specimens under both names. Furthermore, the obser-
vation under microscope also suggested that the length
of anthers and filaments from type specimen of E. ping-
nanensis was 1.5–2.5 mm long and not all the stigmas
of E. obovatifolia were coiled into a circle. Therefore,
these diagnostic characters used by Chang are not reli-
able. In addition, the holotype of E. obovatifolia F. D u
(Du, 2006) (Fig. 2: C) turns out to be an intermediate
plant between E. pingnanensis and E. obovatifolia, al-
though E. obovatifolia F. Du is an invalid name. It seems
clear that all the type specimens under these three names
represent only one taxon (Fig. 2: A–C), which are recog-
nized here as E. pingnanensis. Collections from Yunnan
and Guizhou strongly support this treatment.
3. Elaeagnus gonyanthes Benth. in Hooker’s J. Bot.
Kew Gard. Misc. 5: 196. 1853. Type: China. Hong
Kong: Lindley s. n. (holotype, BM).
Elaeagnus geniculata D. Fang in Acta Phytotax.
Sin. 38: 291. 2000. Type: China. Guangxi: Longzhou,
Xiangshui, J. J. Wang 5449 (holotype, GXMI!),
Fig. 2: D.
Evergreen; shrubs, climbing, ca. 4 m or more tall.
Spines usually absent; young branches columned, slen-
der, densely with brownish red fringed scales. Petiole
brown, 4–8 cm; leaves leathery; speckled; blade dark
green above, rust-colored below, elliptic or oblong-
elliptic, apex obtuse, margin narrowly revolute, base
rounded; 3–13 ×1.2–5.5 cm, both surfaces scaly, later
adaxially glabrescent; lateral veins 7–10 per side of
midrib, raised on both surfaces. Flowers several in a
shortened raceme, rarely solitary; pedicels 3–6 mm.
Flowers yellowish green; calyx broadly campanulate,
slightly 4-angled, outside with dense silvery and sparse
reddish brown stellate-scales; papery; ca. 5 mm, 5–
6 mm in diam.; lobes ovate-triangular, apex acute, con-
spicuously constricted above ovary, inside with white
scales, 3.5–4.5 ×4 mm; anthers oblong, ca. 1.2 mm;
filaments ca. 1 mm; style erect, glabrous; stigma slightly
exserted beyond stamens. Fruit broadly ellipsoid, 1.5–
2.2 cm, densely yellowish white scales, red when ripe,
juicy; ribs on seed inconspicuous; fruit pedicels slender,
1.2–2.5 cm. Fl. Oct.–Nov., fr. Feb.–Mar.
Wet valleys, streamside, montane forests or road
sides; alt. 150 to 1 900 m. Guangdong, Guangxi, Hunan,
Macao and Yunnan.
Additional specimens examined:
China. Precise locality unknown, Sino-Vietnam Exped. s. n. (KUN
Herb. Bar Code No. 0629703), Y. Tsiang 1863 (N). Guangdong: Pre-
cise locality unknown, anonymous 147 (IBSC), S. P. Ko 6226 (IBSC,
PE), Sino-Vietnam Exped. 1433 (KUN), SYS Coast Exped. 3069 (SYS).
Conghua, L. Deng 8324 (IBSC, KUN, NAS, PE). Dinghushan, G. L.
Shi 12467 (IBSC), 3000 (IBSC), 240 (IBSC), K. C. Ting & G. L. Shi
1070 (IBSC, NAS, YUKU), 1935 (IBSC). Doumen, W. C. Ko et al. 832
(IBSC). Fengkai, K. C. Ting et al. 6288 (IBSC, CDBI), 6370 (IBSC,
CDBI). Gaoyao, C. Wang 162302 (IBSC), Y. G. Liu 01950 (IBSC, PE).
Guangzhou, C. M. Hu 7259 (IBSC), H. G. Ye et al. 194 (IBSC), L. Deng
10397 (CDBI, IBSC), 10442 (IBSC), S. H. Chun 8180 (IBK, IBSC,
KUN, PE), 8182 (IBK, IBSC, KUN, PE), S. K. Lau 25692 (IBK, KUN),
H. S. Lo 515 (IBSC), 1036 (IBSC). Longmen, X. G. Li 200225 (IBSC).
Luoding, H. G. Ye et al. 2478 (IBSC), N. Liu et al. 2105 (IBSC). Tais-
han, YUE-73 02702 (IBK, IBSC), 02749 (IBK, IBSC), P. Y. Chen et al.
0024 (IBSC). Xinyi, C. Wang 38159 (IBSC). Xuwen, Xuwen Exped.145
(KUN, NAS, PE), 14 (PE), N. Liu 145 (IBSC). Yangchun, anonymous
3025 (SYS), 1043 (SYS), H. G. Ye et al. 6567 (IBSC), N. Liu et al. 977
(IBSC), C. Wang 38557 (IBSC). Yangjiang, C. Wang 38822 (IBK, IBSC).
Yunan, N. Liu 2813 (IBSC). Yunfu, R. K. Huang 1863 (IBSC), C. Wang
37581 (IBK, IBSC, NAS, PE). Zhuhai, B. H. Chen 839 (IBSC), YUE-73
2946(IBSC), W. C. Ko et al. 291 (IBSC, HGAS), Z. X. Li et al. 682
(IBSC). Guangxi: Precise locality unknown, X. R. Liang 68193 (IBK).
Beiliu, M. J. Lu 4032 (GXMI). Bobai, anonymous 16045 (IBSC), D. Fang
& X. P. Liao 22034 (GXMI), F. S. Huang 16427 (GXMI, IBSC), G. F.
Wu 16045 (GXMI), G. M. Huang 22025 (GXMI). Baise, R. C. Ching
7455 (IBSC, NAS, PE). Cenxi, Cengxi Exped. 7–657 (GXMI). Debao,
C. C. Chang 14201 (IBSC). Dongxing, Naqin Exped. 358 (GXMI), S. H.
Chun 4024 (IBSC). Jingxi, C. C. Chang 14886 (IBK, IBSC). Longzhou,
anonymous 5965 (IBSC), J. J. Wang 5449 (GXMI), P. C. Tam 57063
(IBSC). Luchuan, anonymous 15503 (GXMI), K. X. Chen 13503
(GXMI). Ningming, H. H. Su 68193 (IBSC). Qinzhou, S. H. Chun
3764 (IBSC). Rongxian, W. Chen 82160 (IBSC, PE). Shangsi, W. T.
Tsang 24513 (IBSC, SYS). Shiwandashan, X. R. Liang 69972 (IBK,
IBSC). Tianyang, anonymous 218 (IBK). Xinyi, S. P. Ko 51335 (NAS).
Xing’an, X. R. Liang 67467, 67401 (IBK, IBSC). Yulin, Y. G. Yang68292
(GXMI). Hainan: Precise locality unknown, anonymous s. n. (SYS Herb.
Bar Code No. 001401617), 6226 (SYS), 4883 (SYS), Hainan Veg. Ex-
ped. 584 (KUN), J. Linsley Gressitt 1171 (SYS), P. Zeng 11659 (SYS),
W. Y. Chun 5716 (N), 5743 (N), 5812 (N), 6400 (N), Y. H. Huang 667
(SYS). Baisha, Hainan Veg. Exped. 732 (IBSC, PE), S. K. Lau 25692
(IBSC), 25578 (IBSC, KUN, PE). Baoting, E. Hainan Exped. 953 (IBK,
IBSC, NAS, PE), 954 (CDBI), Diaoluoshan Exped. 2357 (IBK, IBSC,
PE), 2942 (IBK, IBSC, PE), 3201 (IBK, IBSC, PE), F. C. How 73426
(IBK, IBSC, NAS, PE), 72736 (IBK, IBSC, NAS, PE), S. H. Chun 7741
(IBSC). Between Baoting & Sanya, Ganshiling, H. Ohashi et al. 7008
(IBSC, PE). Changjiang, S. K. Lau 3300 (SYS), 1374 (SYS), X. R. Liang
66567 (IBK, IBSC, KUN, PE); Bawangling,G. A. Fu 3505 (IBSC), 2837
(IBSC), Z. X. Li et al. 3699 (IBSC). Chengmai, C. I. Lei 283 (IBSC,
NAS, SYS, PE), 452 (IBSC, NAS, SYS, PE). Ding’an, C. Wang 35750
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366 Journal of Systematics and Evolution Vol. 48 No. 5 2010
(IBK, IBSC), X. R. Liang 68060 (IBSC, NAS, PE), 64673 (IBSC, NAS,
PE), 64245 (IBK, IBSC, KUN, PE), 64573 (IBK, IBSC, KUN, PE), S. P.
Ko 52290 (IBK, IBSC). Dongfang, Hainan Veg. Exped. 00355 (IBSC),
H. T. Zhang 3653 (IBSC), S. K. Lau 5330 (SYS), 3473 (SYS), X. R.
Liang 65219 (IBK, IBSC, PE). Lingshui, C. Wang 36206 (IBK, IBSC,
PE), 36693 (IBK, IBSC, PE), Hainan Exped. 1892 (PE), L. Deng 2969
(KUN, PE), X. R. Liang 61676 (IBK, IBSC, KUN, PE), YUE-76 5363
(IBSC). Qionghai, Y. Zhong 4692 (IBSC, PE). Qiongzhong, 236–6 Ex-
ped. 2002 (PE), L. Deng 3404 (IBSC, KUN, PE). Sanya, 236–6 Exped.
1631 (PE, YUKU), C. L. Tso et al. 44709 (IBSC, NAS, PE), C. Wang
32966 (IBK, IBSC, PE), N. Q. Chen 70096 (IBSC), 44709 (IBK, KUN),
S. K. Lau 6004 (SYS), X. R. Liang 62195 (IBK, IBSC, KUN, PE), Z. X.
Li et al. 2718 (IBSC), 1348 (IBSC). Wanning, G. A. Fu 5489 (IBSC),
F. W. Xing 5786 (IBSC), X. R. Liang 68421 (IBSC). Wenchang, G. A.
Fu 2691 (IBSC). Zhanzhou, Hainan Exped. 00584 (IBSC, CDBI, PE).
Macao: H. G. Ye et al. 92430 (IBSC), 9056 (IBSC). Yunnan: Hekou,
W. X. Liu 606 (PE), Y. M. Shui 973 (PE). Malipo, C. W. Wang & Y. Liu
86227 (IBSC, KUN, PE), K. M. Feng 13313 (KUN, PE), 13830 (KUN),
Q. A. Wu 62–328 (KUN). Maguan, H. T. Tsai 58–8273(KUN). Mengla,
S. S. Zhou 632 (PE). Pingbian, K. M. Feng 05187 (KUN, PE), Y. Tsiang
13540 (IBSC, NAS). Vietnam. Precise locality unknown, C. Y. Wu et al.
774 (KUN), 54 (KUN), 904 (KUN), 884 (KUN, NAS), Sino-Vietnam
Exped. 54 (IBSC), 884 (IBSC), 904 (IBSC), 1433 (IBSC, NAS), 2674
(IBSC, KUN, NAS), 2065 (IBSC, KUN).
Elaeagnus gonyanthes Benth. extends from Guang-
dong to Yunnan and can be easily distinguished from
other members of this genus by its leaf blade abaxially
with dense brownish red scales and calyx tube broadly
campanulate. Although Fang & Cactus Liang (2000)
pointed out that E. geniculata D. Fang was similar to
E. gonyanthes, it can be recognized by its ovate and
smaller leaf blade, flat and inconspicuous lateral veins,
and axillary flowers. However, E. geniculata is known
to us only from the protologue and the type specimens.
After further investigation, Fang’s viewpoint has been
refuted since we find these two species are, in fact, so
close that they cannot be satisfactorily distinguished
at the specific level, as detailed below. Young leaf of
E. gonyanthes is firstly ovate, then becoming elliptic
afterwards. Likewise, lateral veins are inconspicuous
on both surfaces at first then become conspicuous, so
are lateral flowering shoots, which may be later up to
7.5 cm long (Fig. 3: A). Moreover, other specimens Y. G.
Yang 68292 (GXMI), Z. X. Li & F. W. Xing 682 (IBSC),
and S. K. Lau 25692 (IBSC) also indicate that these
two species shared a few characters and should not be
treated as two distinctive species.
6. Elaeagnus difficilis Serv. in Bull. Herb. Boissier, s´
er.
2. 8(6): 386. 1908. Type: China. Hubei: Badong, Henry
1451 (holotype, P; isotype, K!), Fig. 3: B.
Elaeagnus cuprea Rehd., Pl. Wilson. 2(2): 414.
1915. Type: China. Hubei: Changyang, E. H. Wilson
3565 (holotype, A).
Elaeagnus difficilis var. brevistyla W. K. Hu &
H. F. Chow, Fl. Sichuan. 1: 464. 1981. Type: China.
Chongqing: Nanchuan, Jinfoshan, K. F. Lee 61039
(holotype, SZ!; isotypes, KUN!; PE!), Fig. 3: C.
Evergreen; shrubs, spreading or climbing, 2–3 m
tall. Spines present; young branches columned, densely
with reddish brown and off-white scales. Petiole pale
brown, 8–20 mm; leaves sub-leathery to leathery; blade
green above, rust-colored below, ovate-elliptic, rarely to
lanceolate, apex slightly acuminate, margin entire, base
rounded; 7–13.5 ×2.2–6 cm, adaxially glabrous except
for brown scales on midrib, abaxially with scales over-
lapping, silvery or ferruginous; lateral veins 5–9 per
side of midrib, inconspicuous on both surfaces. Flow-
ers several in a shortened raceme; pedicels 1–5 mm,
rarely longer. Flowers yellowish green; calyx short cam-
panulate, slightly 4-angled, outside with dense red-
dish brown scales; sub-leathery; 4.5–5.5 ×2.7–4 mm;
lobes ovate, apex acute or apiculate, constricted above
ovary, abruptly widening below lobes, inside white
stellate-pilose, 2.5–3 ×ca. 2.5 mm; anthers oblong,
1.2–1.7 mm; filaments ca. 0.6 mm; style flexuous,
glabrous; stigma slightly beyond stamens or not so.
Fruit ellipsoid or nearly cylindric, 1.3–1.8 ×0.6–
0.9 cm, densely yellowish white scales, salmon or or-
ange red when ripe, juicy; 8-ribbed seed conspicuous;
fruit pedicels slender, less than 0.5 cm. Fl. Dec.–Mar., fr.
May.–Jun.
Streamsides, scrublands, open slopes, and sparse
forests; alt. 300–1 900 m. Endemic to China.
Chongqing, Fujian, Guangdong, Guangxi, Guizhou,
Hubei, Hunan, Jiangxi, Sichuan, and Yunnan.
Additional specimens examined:
China. Precise locality unknown, anonymous 5690 (IBSC), 3870
(KUN), 027 (SWFC). Chongqing: Fengjie, H. F. Zhou et al. 111595
(IBSC, NAS, PE), 111413 (SZ), 107970 (IBSC, PE, SZ), 108302 (IBSC,
PE, SZ), 107964 (IBSC, PE, SZ), M. Y. Fang 24089 (IBSC, NAS, PE,
SZ), 24051 (PE, SZ). Jingyunshan, anonymous 10318 (NAS). Kaixian,
T. L. Dai 1000389 (PE, SZ). Nanchuan, G. F. Li 62166 (KUN, SZ),
61990 (KUN, PE, SZ), 61039 (KUN, PE, SZ), 64626 (KUN, NAS, PE,
SZ) 62327 (IBSC, KUN, NAS), J. H. Xiong et al. 93475 (KUN), S. X. Tan
092 (PE), W. P. Fang 841 (NF), 41 (IBSC, PE), 5690 (PE), Y. C. Yang 3100
(PE, SZ), 3106 (PE), X. Y. He 09 (NAS), K. L. Chu 896 (PE). Shizhu,
Y. Chen 968 (SZ), 0191 (SZ), 0021 (SZ), 631 (CDBI). Wulong, S. R.
Yi 00005 (PE). Wushan, G. H. Yang 57607 (IBSC, PE, SZ). Youyang,
Dept. Biol. S.W. Norm. Univ. 02648 (PE). Fujian: Shunchang, G. S.
He 0350 (PE). Wuyishan, M. K. Wang 3373 (PE). Guangdong: Precise
locality unknown, anonymous 200529 (IBK), S. P. Ko 57627 (SZ), S. Q.
Tang et al. 187 (SYS). Ruyuan, L. Deng 5800 (IBSC, KUN, PE, SZ),
5720 (IBSC, KUN, PE, SZ, NAS), 5625 (IBSC, KUN, PE, SZ, NAS), C.
Wang 44297 (IBSC, KUN, PE). Xinyi, C. Wang 38003 (IBK). Yangshan,
P. C. Tam 60329 (IBSC, KUN, NAS, SZ), 60391 (IBSC, KUN, NAS),
S. P. Ko 51027 (IBK, IBSC). Yingde, X. G. Li 200529 (IBSC). Guangxi:
Precise locality unknown, C. H. Tsoong 86313 (IBK). Baise, IBSC Ex-
ped. 2638 (IBSC). Damiaoshan, T. C. Chen 41 (IBK, IBSC, KUN), S. H.
Chun 16792 (IBK, IBSC, KUN, NAS, PE). Debao, C. C. Chang 13683
(IBK). Guanyang, Z. Z. Chen 52493 (IBK, IBSC). Huanjiang, Beijing
Exped. 892945 (PE). Jinxiu, Dayaoshan Pl. Exped. 12860 (IBSC), 10550
(IBSC), 11485 (IBSC). Lingui, L. Q. Chen 94620 (IBK, IBSC, PE).
Lingyun, C. C. Chang 11139 (IBK, IBSC, KUN). Longsheng, Guangfu
Pl. Exped. 242 (IBSC, PE, SYS), S. F. Yuan et al. 5093 (IBSC, SYS),
X. W. Tian et al. 182 (PE), Z. T. Li & Y. C. Chen 600492 (IBK, IBSC,
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SUN & LIN: A revision of Elaeagnus in China 367
Fig. 3. A, Flowering branch of Elaeagnus gonyanthes Benth. is longer than 7.5 cm (Cengxi Exped. 7–657, GXMI). B, Isotype of Elaeagnus difficilis
Serv. (Henry 1451, K); the white arrow shows the spine. Ba, Style of E. difficilis is longer than stamens, marked by white arrows. Bb, Style of E. difficilis
is as long as stamens, marked by white arrows. C, Isotype of E. difficilis var. brevistyla W. K. Hu & H. F. Chow (K. F. Lee 61039,PE).
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368 Journal of Systematics and Evolution Vol. 48 No. 5 2010
KUN). Luocheng, C. H. Tsoong 83731 (IBK), L. Q. Chen 91648 (IBK).
Xiuren, C. Wang 40573 (IBK, IBSC), 40516 (IBK, IBSC). Tianlin, C. L.
Gao & Z. R. Liu 26537 (GXMI), Hongshuihe Pl. Exped. 432 (KUN,
PE). Guizhou: Precise locality unknown, anonymous F0061 (PE), S.
Guizhou Exped. 870 (HGAS), Y. C. Qin et al. 58 (NAS), Y. Tsiang 7747
(NAS), 7547 (IBSC, NAS), 7539 (IBSC, NAS), 7749 (IBSC, NAS).
Dejiang, N. Guizhou Exped. 2891 (HGAS). Duyun, Y. K. Li 10451
(HGAS). Guiding, M. Z. Yang 1210 (HGAS). Jiangkou, Z. S. Zhang
402173 (IBSC), 401892 (IBSC), 400512 (IBSC). Kaili, S. Guizhou Ex-
ped. 2005 (HGAS). Leishan, C. P. Tsien et al. 50136 (KUN), 50638
(KUN, HGAS), S. Guizhou Exped. 870 (KUN), 879 (HGAS), Y. K. Li
10366 (HGAS), 10919 (HGAS). Pingtang, anonymous 0156 (HGAS);
Xingren, C. Z. Dang 1475 (HGAS). Wugongshan, Jiangxi Exped. 1234
(PE), Yinjiang, Z. S. Zhang et al. 401079 (HGAS), 400749 (IBSC).
Zhenfeng, C. Z. Dang 2525 (HGAS). Hubei: Precise locality unknown,
H. J. Li 1653 (IBSC). Badong, M. Z. Qian 1715 (NF), R. H. Xuan
1392 (NF), Servettaz 1451 (IBSC), Z. W. Liu s. n. (NAS). Enshi, H. J.
Li 8721 (IBSC, NAS, PE), 8757 (IBSC, NAS, PE), 8845 (IBSC, NAS,
PE). Hefeng, F. S. Peng 274 (PE), H. J. Li 6539 (IBSC, PE, SZ), 6645
(IBSC, PE, SZ), 8496 (IBSC, NAS, PE), 8526 (IBSC, NAS, PE), 8524
(IBSC, NAS, PE), 8180 (IBSC, PE, SZ), 6647 (PE), 6004 (SZ). Huang-
gang, X. Q. Huang 243 (NF), 245 (NF), 365 (NF). Laohushan, H. J. Li
1653 (SZ). Lichuan, G. G. Tang 982 (NF), 25 (NF), 071 (NF), G. X. Fu
et al. 1863 (IBSC, NAS, PE), 1898 (KUN, NAS, PE), Sino-American
Exped. 2077 (KUN, NAS). Jiang’en, H. J. Li 5422 (SZ). Jianshi, W. B.
Lin 64 (PE). Xianfeng, H. J. Li 9084 (IBSC, NAS, PE), 8524 (NF).
Xingshan, E. H. Wilson 59 (IBSC). Hunan: Precise locality unknown,
anonymous 0078 (KUN), M. X. Huang 111568 (IBSC), X. G. Li 200529
(PE). Guidong, Hunan Exped. B299 (PE). Nanyue, F. C. How 74192
(KUN, PE), G. X. Zhu 107 (IBK, IBSC), H. T. Chang 3232 (IBK,
IBSC), J. S. Yue 1742 (NAS), 1682 (NAS), M. H. Li & Y. Q. Kuang
646 (PE), Z. D. Chen 38 (IBSC), Z. H. Hu 605 (PE). Shaoyang, L. D.
Duan 2523, 2085 (PE). Wugang, Y. F. Deng 10004 (NF, PE), 10069
(NF, PE). W. Hunan, L. H. Liu 1830 (IBSC, KUN). Xinning, L. B. Luo
1154 (PE), 1215 (PE), Ziyunshan Exped. 348 (PE). Yangmingshan, S. H.
Chun & Y. Tsiang 708 (IBK, IBSC). Yizhang, L. H. Liu 490 (NAS,
YUKU), 1830 (NAS, PE), P. H. Liang 83885 (IBK, IBSC), 85160 (IBK,
IBSC), S. H. Chun 2774 (IBK, IBSC), 2332 (IBSC). Zhijiang, Wuling
Pl. Exped. 1743 (IBSC). Jiangxi: Precise locality unknown, anonymous
6135 (IBSC, PE), R. C. Ching 12133 (NAS). Dexing, anonymous 11732
(NAS). Sichuan: Precise locality unknown, anonymous 3100 (NAS),
C. Y. Chang 2470 (SZ), T. L. Dai 100389 (CDBI). Emei, Emei Mountain,
W. P. Fang 15364 (IBSC, KUN, PE, SZ), 15313 (KUN, SZ). Hongya,
H. L. Tsiang 11204 (SZ). Jinyang, Dept. Biol. S. W. Norm. Univ. 02470
(PE). Xingwen, Pan 800277 (YUKU). Yunnan: Luquan, P. I. Mao 719
(KUN, NAS). Zhejiang: Longquan, anonymous 3444 (SZ). Yunhe, X. Y.
He 3535 (NAS).
Elaeagnus difficilis var. brevistyla was thought to
be easily separated from E. difficilis by the presence of
spines, leaf leathery, and style slightly longer than sta-
mens (position) (Chang, 1981). Our study suggests that
these two taxa cannot be distinguished by the following
characters. (1) Spines are present on isotype specimen
of E. difficilis (Fig. 3: B, white arrow) and other common
specimens under this name such as Ziyunshan Exped.
348 (PE) and M. H. Li & Q. Y. Kuang 646 (PE). (2)
Young leaves of both taxa are papery, but the older ones
tend to be subleathery and leathery (G. H. Yang 57607
(IBSC, PE, SZ), H. F. Zhou & H. Y. Su 107970 (IBSC,
PE. SZ), H. F. Zhou & H. Y. Su 111595 (IBSC, NAS,
PE), K. L. Chu 896 (PE), S. R. Yi 00005 (PE), and W. P.
Fang 5690 (PE)). (3) Under the microscope, style of
E. difficilis is sometimes shorter than stamens (Fig. 3:
Ba, Bb, also see L. D. Duan 2085 (PE) and Y. F. Deng
10069 (NF, PE)). Hence it seems more appropriate to
expand the circumscription of E. difficilis by including
E. difficilis var. brevistyla.
12. Elaeagnus lanceolata Warb. subsp. grandifolia
Serv. in Bull. Herb. Boissier, s´
er. 2, 8(6): 388. 1908
et in Beih. Bot. Centralbl. 25(1): 88. 1909, ampl. de-
scr. Type: Chongqing: Nanchuan, Rosthorn 1103,1104
(syntypes, B).
Elaeagnus hunanensis C. J. Qi & Q. Z. Lin in J.
Centr. S. Forest. Univ. 20(2): 89–90. 2000. Type: China.
Hunan: Tongdao, J. G. Xiao & C. J. Qi 3376 (holotype,
CSFI!; isotypes, CSFI!; IBSC!; PE!), Fig. 4: A.
Evergreen; shrubs, erect or slightly scandent, to
4 m tall. Spines absent or present on old branches;
young branches densely with reddish brown scales.
Petiole red-brown, 10–15 mm; leaves leathery; blade
green above, rust-colored below, elliptic or oblong to
oblong-lanceolate, apex acuminate, margin entire or
revolute, base obtuse; 5–18 ×3–8 cm, adaxially scaly
when young, glabrescent later, abaxially with dense
rust-colored or brown scales; lateral veins 8–12 per
side of midrib, conspicuous on both surfaces. Flowers
often 3–5 in axils; pedicels to 10 mm long, slender.
Flowers yellowish white; calyx tubular or obconic-
tubular, outside with reddish brown scales; 6–8 mm; an-
thers oblong, ca. 1.5 mm; filaments very short or absent;
style erect, glabrous, or sparsely stellate-hairy; stigmas
2–3 mm. Fruit ellipsoid, 1.2–1.5 ×0.5–0.6 cm, densely
red-brown scales, red when ripe, juicy; Fl. Aug.–Oct.,
fr. Apr.–May.
Valley forests, limestone hills, and scrublands; alt.
300–2600 m. Endemic to China. Chongqing, Guangxi,
Hubei, Hunan, Sichuan, and Yunnan.
Additional specimens examined:
China. Chongqing: Nanchuan, G. F. Li 63558 (PE), K. C. Kuan
et al. 2243 (PE). Guangxi: Damiaoshan, S. H. Chun 16790 (IBSC, PE).
Longsheng, F. H. Xie 3068 (IBK). Tian’e, Beijing Exped. 89326 (PE).
Hubei: Enshi, H. J. Li 6073 (PE). Hefeng, H. J. Li 6679 (IBSC, KUN,
PE), 8073 (PE), 8276 (PE), 8424 (PE). Hunan: W. Hunan, L. H. Liu
490 (IBSC). Xinning, L. B. Luo 1216 (PE). Xinshao, B. Y. Li 6274 (PE).
Sichuan: Precise locality unknown, anonymous 705 (NAS), 3878 (PE),
J. H. Xiong & Z. L. Zhou 90043 (IBSC), T. Tang 23310 (PE), X. Y. He
6596 (IBSC, NAS), Z. W. Yao 4707 (NAS, PE), 2986 (NAS, PE). Emei,
anonymous 527 (NAS), C. W. Yao 3163 (PE), G. H. Yang 57481 (IBSC,
KUN, NAS, PE, SYS), 57366 (IBSC, KUN, NAS, PE, SYS), 53560
(IBSC, SYS), 57422 (KUN, NAS, PE), 52697 (SYS), 52733 (SYS),
52642 (SYS), J. H. Xiong et al. 30496 (IBK, IBSC), 30456 (IBK, IBSC,
NAS, PE), 32100 (IBK, IBSC, NAS, PE), 32373 (IBK, IBSC, NAS, PE),
3606 (IBK, IBSC), S. X. Wang426 (PE), 398 (PE), T. C. Loe 3693 (KUN),
3606 (KUN), 5378 (KUN), 4430 (KUN), W. P. Fang 17562 (IBSC, PE),
13842 (KUN, NAS, PE), 15207 (KUN, NAS, PE), Y. L. Qiao 00591
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SUN & LIN: A revision of Elaeagnus in China 369
Fig. 4. A, Holotype of Elaeagnus hunanensis C. J. Qi & Q. Z. Lin (J. G. Xiao & C. J. Qi 3376, CSFI). B, Holotype of Elaeagnus heterophylla D. Fa ng
&D.R.Liang(T. Ji 8553, GXMI). C, Isotype of Elaeagnus luoxiangensis C. Y. Chang (S. S. Sin 3984, IBSC). D, Holotype of Elaeagnus pilostyla C. Y.
Chang (H. T. Tsai 58–8144, KUN).
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370 Journal of Systematics and Evolution Vol. 48 No. 5 2010
(YUKU). Leibo, anonymous 0289 (PE).Tianquan, K. C. Kuan & W. T.
Wang 2216 (PE), H. L. Tsiang 35245 (IBSC, PE, SWFC), 37556 (IBSC,
PE, SWFC), Y. B. Yang 21725 (IBSC). Yunnan: Daguan, NE. Yunnan
Exped. 117 (KUN). Lanping, H. T. Tsai 56264 (IBSC, PE). Maguan, X.
Wang et al. 100457 (IBSC). Precise locality unknown, Z. X. Zhao 17
(KUN).
Qi & Lin (2000) suggested that E. hunanensis was
closely related to E. pungens, but could be distinguished
by leaf shape, size, pedicel length, and calyx shape. Nev-
ertheless, it seems that Qi & Lin ignored the affinity
between E. lanceolata subsp. grandifolia and E. huna-
nensis. Indeed, in E. lanceolata subsp. grandifolia the
leaves are elliptic or oblong to oblong-lanceolate, 5–
18 ×1.5–8 cm, pedicels 10 mm, calyx tube 6–8 mm and
style erect, glabrous or sparsely stellate-hairy. All these
characters strongly suggest these two species could not
be successfully separated, morphologically supported
by specimens such as B. Y. Li 6274 (PE), L. B. Luo 1216
(PE), and H. J. Li 8424, 8276, 6073 (PE), 6679 (IBSC,
KUN, PE).
14. Elaeagnus glabra Thunb. in Murray, Fl. Jap. 67.
1784. Type: unknown.
Elaeagnus heterophylla D. Fang & D. R. Liang
in Acta Phytotax. Sin. 38: 292. 2000. Type: China.
Guangxi: Jinxiu, T. Ji 8553 (holotype, GXMI!),
Fig. 4: B.
Elaeagnus luoxiangensis C. Y. Chang in Bull. Bot.
Lab. N. E. Forest. Inst., Harbin 1980(6): 108. 1980.
Type: China. Guangxi: Jinxiu, Luoxiang, S. S. Sin 3984
(holotype, IBK; isotypes, IBSC!), Fig. 4: C.
Evergreen; shrubs, climbing. Spines normally ab-
sent; young branches columned, with reddish brown
scales. Petiole brown, 7–10 mm; leaves thinly leathery;
blade green above, partially rust-colored below, elliptic
to obovate, rarely lanceolate, apex acute, rarely obtuse
or rounded, margin revolute, base obtuse or rounded;
1.2–11.5 ×1.2–3.5 cm, adaxially glabrous or sparsely
brown scales, abaxially covered with brown and rarely
silvery scales. Flowers several in a shortened raceme;
pedicels 3–4 mm. Flowers yellow or sometimes reddish
brown; calyx tubular-funnelform to slightly tubular-
campanulate, outside with dense reddish brown scales;
papery; 4.5–5.5 ×2.7 mm; lobes triangular-ovate, apex
acute, inside sparsely with white stellate, 2–4 ×2 mm;
anthers oblong, 1.5–1.8 mm; filaments ca. 0.8 mm; style
erect, glabrous, scarcely stellate-hairy; stigma beyond
stamens. Fruit broadly elliptic to elliptic, 1.4–2 ×0.6–
0.9 cm, densely ferruginous scales, orange-red when
ripe, juicy; ribs inconspicuous; fruit pedicels slender,
up to 8 mm long. Fl. Sep.–Nov., fr. Apr.–May.
Streamside, scrublands, and thickets; alt. ca.
2200 m. Anhui, Chongqing, Fujian, Guangdong,
Guangxi, Guizhou, Hainan, Hubei, Hunan, Jiangsu,
Jiangxi, Sichuan, Taiwan, Xizang, Yunnan, and
Zhejiang.
Additional specimens examined:
China. Precise locality unknown, anonymous s. n. (KUN Herb. Bar
Code No. 0629769), B78 (IBSC), D51002 (KUN), 44 (SYS), 2079 (SYS),
Q. Feng 10777 (SYS), Sw. Univ. Exped. 6075 (KUN), S. J. Liang 66
(SYS), Sw. Univ. Exped. 6291 (KUN). Anhui: Precise locality unknown,
anonymous 1009 (NAS), M. B. Deng 0046 (NAS). Guangde, D. X. Zuo
et al. 3083 (NAS). Huangshan, Z. Deng 183 (NAS). Huoshan, M. B.
Deng & H. T. Wei 80125 (NAS). Jinzhai, M. B. Deng 81143 (HHBG).
Qimen, M. B. Deng et al. 5078 (HHBG, IBSC, NAS, PE), 5318 (NAS,
PE), 4934 (NAS). Shexian, anonymous 2096 (NAS), 1675 (NAS), 2418
(NAS). Taihu, S. Chen 2641 (N). Xiuning, R. H. Shan 2537 (NAS).
Xuancheng, anonymous 402 (NAS). Chongqing: Fengjie, H. F. Zhou
et al. 111413 (IBSC, PE). Nanchuan, G. F. Li 64855 (KUN, NAS, PE,
SZ), 62166 (NAS), 60735 (PE), M. Z. She et al. 6447 (NAS). Xiushan,
D. H. Du 3966 (IBSC). Fujian: Anonymous 5714 (N), L. K. Ling 1534
(KUN), Y. Lin 4449 (PE). Chun’an, M. K. Wang et al. 3476 (NAS, PE).
Dehua, S. Y. Huang 05333 (IBSC). Guangze, H. Y. Zou et al. 20381
(SWFC). Liancheng, D. S. Wang 1053 (KUN, PE), Y. Lin 4036 (PE).
Minhou, H. C. Zhou 5417 (IBSC). Minnan, G. S. He 1542 (PE). Nanjing,
S. Y. Huang 04704 (IBSC). Shaowu, H. C. Zhou 5313 (IBSC). Shunchang,
G. S. He 0341 (PE). Yanping, G. S. He 3746 (PE). Yong’an Y. Lin 540
(PE). Yongtai, H. H. Chung 3210 (IBSC). Zhangping, Y. Lin 5051 (PE).
Guangdong: Precise locality unknown, anonymous s. n. (IBSC Herb.
Bar Code No. 0386545), s. n. (IBSC Herb. Bar Code No. 0386555),
3084 (SYS), 3094 (IBK), 46 (SYS), 40455 (SYS), 8039 (SYS), H. Tse
92 (SYS), L. Deng 4035 (PE), SYS Exped. 005 (SYS), S. P. Ko 51580
(KUN), 50939 (KUN), W. Y. Chun 5626 (IBSC, NAS, PE), 5804 (PE).
Baiyunshan, S. H. Chun 7098 (IBK, IBSC, PE). Boluo, L. Deng 356
(IBK, IBSC), N. Q. Chen 40409 (IBSC, SYS), Q. Feng A-513 (SYS),
S. P. Ko 52438 (IBK, IBSC), 57258 (IBK), T. M. Tsui s. n. (KUN), 23
(SYS), Y. F. Deng 15701 (PE), Y. Tsiang 1650 (SYS, IBSC, N, NAS,
PE). Chao’an, IBSC Geobotanical Exped. 20 (IBSC). Conghua, C. Wang
44451 (KUN), L. Deng 8570 (IBSC, NAS, PE), 8719 (NAS, PE), Sino-
Germany Shaanxi Exped. 890 (IBSC), 1141 (IBSC). Deqing, Y. G. Liu
01337 (HHBG, IBSC, NAS). Guangzhou, F. W. Xing 048 (IBSC). Huaiji,
W. T. Tsang 23305 (IBSC), Y. G. Liu 2744 (HHBG, IBSC, PE). Huidong,
P. Y. Chen et al. 94 (IBSC). Jiaoling, Nanling Exped. 2337 (IBSC).
Lechang, N. Q. Chen 42378 (IBSC), 42926 (IBSC, KUN, NAS, PE),
Nanling Exped. 3186 (IBSC), S. P. Ko 51850 (IBK, IBSC, NAS, PE),
W. T. Tsang 20679 (IBSC, NAS, PE, SYS), X. R. Liang et al. 31529
(SYS, IBK, IBSC, NAS), Y. Tsiang 11130 (IBK, IBSC). Liannan, Z. S.
Zhang 618 (IBSC). Lianping, YUE-73 2004 (IBSC). Lianshan, Nanling
Exped. 556 (IBSC). Lianxian, P. C. Tam 60022 (IBSC, KUN, NAS, PE),
S. P. Ko 50939 (IBK, IBSC, NAS, PE, SZ). Longmen, S. Q. Zhong 132
(IBK), X. W. Wang et al. 8172, 8236 (HGAS, IBSC). Pingyuan, L. Deng
4053 (IBSC, KUN). Raoping, N. Q. Chen 42926 (IBK). Ruyuan, H. S.
Lo 1793 (IBSC), Y. Li 2177 (IBK, IBSC, KUN, NAS, PE, SZ), YUE 71
172 (IBSC). Wengyuan, S. K. Lau 25237 (IBSC, KUN, PE), 774 (SYS).
Xinfeng, Y. M. Tamn 330 (SYS). Xinyi, Chang s. n. (SYS00101579), C.
Wang 32149 (IBSC, SYS), 31948 (IBSC), S. P. Ko 51335 (IBSC, KUN,
PE). Xingning, IBSC Geobotanical Exped. 7682 (IBSC). Yangshan, P. C.
Tam 60329 (KUN, NAS, PE). Yingde, C. Wang 30397 (IBK, PE), L.
Deng 952 (IBSC, KUN, PE), R. K. Huang 30397 (IBSC, SYS), IBSC
Geobotanical Exped. 6136 (IBSC). Yongshun, C. J. Qi 3640 (IBSC).
Guangxi: Anonymous 120 (IBSC), C. C. Chang 10488 (IBSC), C. H.
Tsoong 85056 (IBK, IBSC), 85048 (IBK, IBSC), 808883 (IBK, IBSC),
85045 (IBK), 86531 (IBK), H. H. Fang 2–333 (GXMI), W. T. Tsang
28549 (IBSC). Binyang, Binyang Exped. 2–230 (GXMI). Chacheng,
Y. K. Li 403010 (IBK), 402920 (IBK). Debao, C. C. Chang 13686 (IBSC).
Guilin, Sino-Germany Shaanxi Exped. 1283 (IBK), IBSC Geobotanical
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SUN & LIN: A revision of Elaeagnus in China 371
Exped. 4978 (IBK, IBSC). Gongcheng, Y. K. Li 403010 (IBSC), 402920
(IBSC). Huanjiang, C. C. Wang 17197 (GXMI), F. N. Wei & Y. Liu
M0419 (IBK), Guangxi Med. Pl. Exped. 000377 (GXMI), YGG Exped.
70132 (IBK, KUN), 70170 (IBK). Jinxiou, anonymous 1085 (IBSC),
C. F. Liang 34340 (IBK), Dayaoshan Exped. 811925 (GXMI), 65193
(GXMI), 12650 (IBSC), 10431 (IBSC), 11498 (IBSC). Jingxi, C. C.
Chang 14533 (IBK, IBSC), Y. C. Chen 1085 (IBK), Z. T. Li 602622 (IBK,
IBSC). Leye, H. N. Qin et al. 3157 (PE), W. Y. Yao & K. J. Yan 73893
(GXMI), 73904 (PE). Liangfeng, C. H. Tsoong 808780 (IBK). Lingui,
C. F. Liang 30942 (IBK, KUN, PE), Huaping Nature Reserve Pl. Exped.
H0130 (PE), L. Q. Chen 94403 (IBK), 93237 (IBK, IBSC, PE), S. K. Lee
200337 (IBK), Y. B. Xu 10149 (IBSC, IBK), 10164 (IBK, IBSC, KUN).
Lingyun, C. C. Chang 10488 (IBK), C. Y. Chiao et al. 91 (N). Liuzhou,
C. H. Tsoong 84577 (IBK), G. Z. Li 12875 (IBK), L. Q. Chen 91768
(IBK). Longlin, T. H. Wei 3–34136 (GXMI). Longsheng, C. C. Chang
14145 (IBSC), F. S. Tan & D. Fang 43184 (GXMI), Guangfu Exped.
00250 (IBK, IBSC), 242 (IBSC), Longsheng Exped. 50169 (IBK, IBSC,
KUN, NAS, PE), 50123 (IBK, IBSC, NAS, PE, SYS, YUKU). Luocheng,
C. H. Tsoong 83829 (IBK), L. Q. Chen 91592 (IBK). Luoxiang, S. S. Sin
9884 (IBSC). Napo, C. C. Chang 14145 (IBK), D. Fang & D. B. Liu 22130
(GXMI), 22231 (GXMI). Rongxian, L. Q. Chen 91694 (IBK), Z. Y. Wei
et al. 40068 (IBK). Rongshui, Beijing Exped.892308 (PE), 891773 (PE),
892218 (PE). Sanjiang, C. H. Tsoong 84013 (IBK), L. Q. Chen 92878
(IBK). Shangsi, S. H. Chun 4690 (IBSC). Tian’e, Beijing Exped. 890411
(PE), 890195 (PE). Wuming, D. Fang 22591 (GXMI). Xiuren, C. Wang
40662 (IBK, IBSC). Zhaoping, Y. K. Li 402552 (IBK, IBSC). Guizhou:
Guizhou Exped. 4689 (IBSC), 9232 (PE), H. T. Chang 7362 (SYS), 7368
(SYS), 7378 (SYS), 7340 (SYS), 7350 (SYS), 40455 (SYS). Anlong,
Z. S. Zhang & Y. T. Zhang 3640 (PE). Anshun, S. W. Teng 0079 (IBSC).
Ceheng, F. C. Wang 0575 (PE), Y. Tsiang 9232 (IBSC, NAS). Chishui,
P. C. Tsoong 238 (PE). Guiding, anonymous 1675 (HGAS), P. Zhao et al.
679 (HGAS). Guiyang, anonymous 031 (HGAS), S. Guizhou Exped. 58
(KUN, HGAS, PE), Z. J.Ren 3699 (PE). Jiangkou, Fanjingshan, Y. Tsiang
7533 (IBSC, NAS). Leishan, C. P. Tsien 50136 (PE), 50638 (PE), Y. S.
Chen 4167 (PE). Libo, D. F. Huang 1615 (HGAS), Y. K. Li 10009 (IBSC,
HGAS), 10040 (IBSC, HGAS), 9372 (IBSC), X. M. Wang 234 (HGAS).
Pingtang, anonymous 0399 (HGAS). Pu’an, C. Z. Dang 1219 (HGAS).
Puding, S. W. Teng 342 (IBSC). Tiantaishan, P. C. Tsoong 238 (KUN).
Xingren, Y. G. G. Exped. 50986 (KUN). Wangmo, G. F. Wang 1–1003
(PE). Zhenfeng, Y. Tsiang 4665 (NAS, SYS), 4687 (NAS). Zunyi, N.
Guizhou Exped. 0137 (IBK). Hainan: Changjiang, X. R. Liang 66567
(NAS). Hubei: Badong, H. C. Zhou 94 (N), Q. L. Chen et al. 1835
(PE). Enshi, H. J. Li 8757 (NAS, PE), 8845 (NAS, PE). Hefeng, H. J.
Li 8356 (NAS), 8362 (PE), 8364 (PE), Sangzhi Forestry Institute 0526
(KUN). Laifeng, H. J. Li 7291, 7457 (PE). Luotian, Q. G. He 85–17 (PE).
Yongshun, anonymous 00998 (KUN). Hunan: Precise locality unknown,
anonymous 6291 (SWFC). Chengbu, F. J. Huang 0476 (PE). Dongkou,
L. H. Liu et al. 16608 (IBSC, KUN, PE). Hongjiang, C. T. Lee 1653
(IBSC, PE). Nanyue, anonymous 6025 (SWFC), F. C. How 74192 (IBK,
NAS), X. X. Wang 255 (PE), Z. H. Hu 605 (PE). Sangzhi, Sangzhi
Forestry Institute 0687 (KUN), T. R. Cao 90531 (KUN). Shaodong, J. H.
Liu 7488 (PE). Shaoyang, L. D. Duan 20030610 (PE). Suining, D. Z.
Wang 868 (IBSC). Tongdao, T. C. Chen 1185 (IBSC). W. Hunan, Hunan
Exped. 0291 (PE). Xinning, L. B. Luo 1107 (PE), 419 (PE), Z. C. Luo
1736 (PE). Xinshao, B. Y. Li 6340 (PE). Yizhang, B. Z. Xiao 3928
(PE), S. H. Chun 154 (IBK, IBSC), P. C. Tsoong 667 (PE). Yongshun,
anonymous 3640 (PE). Jiangsu: Yixing, F. X. Liu & Z. Y. Huang 2844
(NAS),M.B.Dengs.n.(NAS),S.H.Mao012(IBK,KUN,NAS,PE,SZ).
Jiangxi: Jiangxi Univ. Exped. 5667 (KUN), 6135 (KUN), 12603 (KUN),
12711 (KUN), X. S. Yang et al. 830082 (IBSC), 10248 (IBSC). Anfu,
J. S. Yue 2772 (NAS, PE). Dayu, J. S. Yue 1415 (NAS, PE), 1523 (NAS,
PE). Guangchang, J. S. Yue 2340 (IBSC, NAS, PE). Guangfeng, M. X.
Nei & S. S. Lai 5667 (PE), 5973 (PE). Guixi, anonymous 900456 (NAS).
Huichang, W. H. Wan et al. 1994 (IBSC, NAS, PE), 1923 (PE). Jing’an,
Lushan Botanical Garden Exped. DJD2004–0464 (PE), S. S. Lai 001394
(PE). Jinggangshan, J. S. Yue et al. 4897 (NAS, PE). Jiujiang, C. M.
Tan 941626 (HHBG, PE), 84190 (IBSC), 98454 (IBSC), 951344 (IBSC),
92363 (IBSC, SYS), 941361 (PE), 941273 (PE), H. Migo s. n. (NAS), H.
Zou 1200 (NAS), Y. G. Xiong 1007 (NAS), 7070 (NAS), 9794 (NAS, PE),
9722 (PE). Lichuan, M. K. Wanget al. 2362 (IBSC, NAS, PE). Nanchang,
anonymous s. n. (PE). Ningdu, C. M. Hu 5465 (IBSC, KUN). Poyang,
J. S. Yue 2021 (NAS), M. K. Wang 4002 (NAS, PE). Shicheng, W. H. Wan
et al. 1923 (IBSC, NAS). Suichuan, 236 Exped. 597 (PE), J. S. Yue et al.
4202 (PE), S. S. Lai et al. 5533 (IBSC), 5439 (IBSC). Wuning, C. M. Tan
941257 (HHBG, PE), 9611177 (KUN). Wuyuan, J. S. Yue 4202 (NAS),
M. K. Wang 3954(IBSC, NAS, PE), Q. H. Li & C. Chen 97 (PE). Xiushui,
C. M. Tanet al. 951073 (IBSC), S. L. Liou 9080 (PE). Xunwu, anonymous
1192 (IBSC, PE), J. S. Yue 1917 (NAS, PE), X. S. Yang et al. 12603,
12711 (IBSC). Sichuan: Precise locality unknown, W. P. Fang 5690 (N,
NAS), Z. W. Yao 5282 (NAS), 2986 (PE). Dˆ
erong, H. Sun et al. 2498
(KUN). Xingwen, anonymous 0376 (PE). Xiushan, anonymous 3996
(PE). Youyang, Dept. Biol. Univ. Sichuan 02218 (PE). Xizang: Precise
place unknown, H. Green 1130 (IBSC). Mˆ
edog,B.S.Li&S.Z.Cheng
04323 (PE), H. Sun et al. 5313 (KUN). Yunnan: Gongshan, Nujiang
Exped. 790277 (KUN). Menglian, Z. H. Hu et al. 7900623 (YUKU).
Wenshan, Y. M. Shui 001407 (PE). Yingjiang, G. D. Tao 013062 (SYS).
Zhejiang: Precise place unknown, M. Chen 928 (IBSC), S. Y. Chang
3995 (IBSC), 9950 (NAS). Chun’an, L. Hong 1456 (HHBG), M. Z. She
26981 (NAS). Hangzhou, anonymous 1982 (HHBG, KUN,PE), 702 (PE),
13 (SZ), H. Migo s. n. (NAS), H. X. Zhou 808 (HHBG), S. Y. Chang 1662
(HHBG, NAS, PE). Jingning, S. Y. Chang 3995 (HHBG, PE), 22951
(NAS). Lin’an, anonymous 30799 (HHBG, PE), 28739 (HHBG, PE),
X. Y. He 26933 (HHBG, IBSC, NAS, PE), 22762 (HHBG, IBSC, NAS,
PE), 30799 (IBSC), 30842 (IBSC), Zhejiang Pl. Exped. 29830 (NAS).
Longquan, anonymous 21713 (HHBG), 23319 (HHBG), 22951 (HHBG),
22882 (HHBG), D. X. Zuo et al. 22273 (HHBG, NAS), 25549 (HHBG,
NAS), 21799 (HHBG, NAS), 23319 (NAS), S. Y. Chang 7173 (PE), 4530
(HHBG, PE), 4541 (HHBG, PE), 4315 (HHBG, IBSC, KUN, PE), 22882
(NAS), X. Y. He 03450 (NAS), 3417 (NAS, SYS). Ningbo, H. Migo
s. n. (NAS), L. Hong 3734 (HHBG). Pan’an, L. Hong 2714 (HHBG).
Pingyang, D. X. Zuo et al. 24602 (HHBG, NAS), D. X. Zuo 24818 (NAS).
Suichang, M. Z. She 26361 (NAS, PE), 26646 (NAS, PE), Zhejiang
Pl. Exped. 26361 (HHBG), 26645 (HHBG), 25689 (HHBG, NAS, PE).
Taishun, D. X. Zuo 25413 (HHBG, NAS), 23835 (HHBG, NAS), 23981
(NAS), 23559 (NAS). Tianmushan, anonymous 30842 (HHBG), 189
(PE), X. Y. He 271 (HHBG, IBSC). Tiantai, anonymous 1241 (NAS).
Wenling, HHBG 4039 (HHBG, PE).
Elaeagnus heterophylla D. Fang & D. R. Liang
was recognized as a new species from Guangxi, with
remarks that this species appeared to be closely related
to E. liuzhouensis, but differed in shape of leaf blade,
lateral veins, and flowers (Fang & Liang, 2000). Nev-
ertheless, Qin & Gilbert (2007) argued that E. hetero-
phylla is known only from the protologue. Our investiga-
tions suggest that Fang & Liang used a narrow concept
and failed to compare E. heterophylla with E. glabra.
This explains why collections of E. heterophylla are
only restricted in type specimens. In fact, E. glabra is
widespread in southern China, as well as Guangxi. The
shape of leaf blade ranges from orbicular-ovate or ellip-
tic to lanceolate with an ecologically varied distribution
which indicates a geographical and morphological in-
clusion of E. heterophylla (S. H. Mao 012 (IBK, KUN,
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372 Journal of Systematics and Evolution Vol. 48 No. 5 2010
NAS, PE, SZ), S. Y. Chang 1662 (HHBG, NAS, PE),
Y. G. Xiong 9794 (NAS, PE), N. Q. Chen 42926 (IBSC,
KUN, NAS, PE), and L. Q. Chen 93237 (IBK, IBSC,
PE)). Hence it seems improper to treat E. heterophylla
as a new species.
Because of the limited knowledge and a relatively
narrow concept of species in Elaeagnus, Chang (1980)
also considered E. luoxiangensis as new to Guangxi,
stating that E. luoxiangensis had an affinity with E. dif-
ficilis. After our comparison with E. difficilis,wehave
found E. luoxiangensis and E. difficilis can be easily dis-
tinguished, but type specimens of E. luoxiangensis are
hardly distinguishable from specimens of E. glabra, es-
pecially the collections from Guangxi and Guangdong.
These following specimens with overlapping vegetative
and floral characters appear to combine E. luoxiangen-
sis and E. glabra as one taxon: C. O. Levine 243 (PE),
S. P. Ko 50939 (IBK, IBSC, NAS, PE, SZ), Y. Tsiang
1650 (SYS, IBSC, N, NAS, PE), L. Q. Chen 91649
(IBK), 93237 (IBK, IBSC, PE), and S. H. Mao 012
(IBK, KUN, NAS, PE, SZ).
17. Elaeagnus delavayi Lecomte in Notul. Syst. (Paris)
3: 156. 1915. Type: China: Yunnan, Tch´
eou-tchoui (=
Heqing), Ta-pin-tz´
e(=Dapingzi), Delavay s. n. (holo-
type, P).
Elaeagnus pilostyla C. Y. Chang, Fl. Sichuan. 1:
463. 1981. Type: China. Yunnan: Wenshan, H. T. Tsai
58–8144 (holotype, KUN!; isotype, KUN!), Fig. 4: D.
Elaeagnus longiloba C. Y. Chang in Bull. Bot. Lab.
N. E. Forest. Inst., Harbin 1980(6): 113. 1980. Type:
China. Guizhou: Bijie, Y. Tsiang 9049 (lectotype, des-
ignated by Q. Lin in Bull. Bot. Res., Harbin 26(6): 657.
2006: IBSC!), Fig. 5: A.
Elaeagnus retrostyla C. Y. Chang in Bull. Bot. Lab.
N. E. Forest. Inst., Harbin 1980(6): 112. 1980. Type:
China. Guizhou: Bijie, P. H. Yu 283 (holotype, KUN!;
isotype, PE!), Fig. 5: B.
Elaeagnus xingwenensis C. Y. Chang, Fl. Sichuan.
1: 464. 1981. Type: China. Sichuan: Xingwen, Xian-
feng, C. Y. Chang 5 (holotype, SZ!; isotypes, SZ!),
Fig. 5: C.
Evergreen; shrubs, erect. Spines absent; young
branches up-ridge, densely with rust-colored or brown
scales. Petiole red-brown, 0.8–1.5 cm; leaves thinly
leathery; speckled; blade yellowish green above, yellow-
ish white below, narrowly elliptic or oblong-lanceolate,
apex rounded or obtuse, sometimes acuminate, mar-
gin entire, base cuneate; 5–12 ×1.6–4 cm, adaxially
glabrous or sparsely yellowish brown scales, abaxi-
ally silvery or rust-colored scales; lateral veins 5–8 per
side of midrib, faintly conspicuous on both surfaces.
Flowers often 5–7 forming a shortened raceme; pedicels
up to 12 mm. Flowers white; calyx tubular or obconic-
tubular, faintly 4-ribbed, outside with reddish brown
scales; sub-leathery; 6–7 ×2–3 mm; lobes triangular,
apex acuminate, conspicuously constricted above ovary,
inside densely with scales and stellate, 2.5–5 ×ca.
2 mm; anthers oblong, ca. 1 mm; filaments ca. 0.5 mm;
style erect, densely white stellate-villous; stigma not be-
yond stamens. Fruit elliptic, 1. 2 ×0.6 cm, densely with
scales, yellow when ripe, juicy; ribs inconspicuous; fruit
pedicels slender, 6–10 mm. Fl. Sep.–Dec., fr. Feb.–May.
Open forests or thickets on eastern slopes; alt.
1300–3100 m. Endemic to China. Chongqing, Guangxi,
Guizhou, Sichuan and Yunnan.
Additional specimens examined:
China. Precise locality unknown, s. n. (SWFC Herb. Bar Code
No. 0036228), anonymous 981 (IBSC), 1294 (KUN), 845 (KUN), 075
(KUN), 123 (HGAS), 43 (SWFC), H. T. Tsai 5472 (KUN), 54742 (KUN),
H. T. Tsai & J. S. Xin 845 (KUN). Chongqing: Nanchuan, F. T. Wang
10293 (PE), 10082 (PE), G. F. Li 65051 (IBSC, KUN, NAS, PE), 65046
(KUN, PE), J. H. Xiong & Z. L. Zhou 93475 (PE), K. C. Kuan et al. 2290
(PE), 2243 (PE), 2241 (PE). Wulong, S. R. Yi 00007 (PE). Guizhou:
Precise locality unknown, Y. Tsiang 7635 (IBSC), 7741 (IBSC). Anlong,
S. K. Wu 100993 (KUN), D. J. Liu 87–53 (HGAS), 282 (HGAS). Bi-
jie, P. H. Yu 283 (IBSC, KUN, PE), S. Guizhou Exped. 00210 (HGAS).
Daozhen, G. A. Zhang & W. L. Liu 028 (HGAS). Leishan, Q. H. Chen
3124 (HGAS). Luodian, S. Guizhou Exped. 210 (PE). Qingzhen, S. W.
Teng 90134 (IBSC). Xishui, K. M. Lan 92–0234 (PE). Xingren, C. Z.
Dang 1579 (HGAS), 80558 (HGAS). Yinjiang, C. P. Tsien et al. 31053
(HGAS), Z. S. Zhang et al. 401121 (IBSC). Zhenfeng, D. J. Liu 831
(HGAS). Sichuan: Precise locality unknown, anonymous 3758 (IBSC),
20348 (SYS), G. H. Yang 59742 (IBSC), Sichuan Exped. 3377 (IBSC),
Z. X. Zhao 1732 (KUN). Emei, D. H. Du 705 (IBSC), G. H. Yang 57422
(IBSC), H. Z. You 249 (KUN), Y. X. Xiao 48172 (GXMI). Xizang:
Zay¨
u, Qinghai-Xizang Exped. 10864 (KUN). Mˆ
edog, H. Sun et al. 3562
(KUN), 4127 (KUN). Yunnan: Precise locality unknown, S. K. Wu
et al. 804 (KUN), T. N. Liou 17891 (IBSC), Z. X. Zhao 1732 (KUN).
Binchuan, H. Sun et al. 451 (KUN), Yunnan Univ. Exped. 0265 (YUKU),
0235 (YUKU). Cangyuan, Yunnan Univ. Exped. 844 (YUKU). Daguan,
N. E. Yunnan Exped. 117 (YUKU). Dali, H. C. Wang 2058 (IBSC).
Dˆ
eqˆ
en, T. T. Yu 9634 (KUN). Fengqing, J. Chen 72 (KUN). Gongshan,
Drungjiang, Drungjiang Exped. 2191 (KUN), 1038 (KUN), 5442 (KUN),
1550 (KUN), 4648 (KUN), 2240 (KUN), 5513 (KUN), 5717 (KUN),
6208 (KUN), 5446 (KUN), H. Li et al. 8125 (KUN), Qinghai-Xizang Ex-
ped. 9874 (KUN). Guangnan, anonymous 9082 (KUN). Jingdong, B. Y.
Qiu 53468 (KUN), Z. H. Yang et al. 101399 (KUN). Kunming, D. S.
Feng s. n. (SWFC), H. T. Jiang s. n. (SWFC), K. M. Feng 10405 (KUN),
Q. Jiang s. n. (SWFC), Z. Q. Yao & W. Y. Zhang 13388 (YUKU), 13836
(YUKU). Luquan, G. H. Yang s. n. (YUKU), P. I. Mao 01905 (KUN).
Lushui, H. Sun 1596 (KUN). Shiping, S. K. Wu 914 (KUN). Tengchong,
780 Exped. 521 (KUN, PE), H. Li et al. 21611 (KUN). Tonghai, Yunnan
Univ. Exped. 19376 (YUKU). Wenshan, H. Sun et al. 411 (KUN), H. T.
Tsai 58–8144 (KUN), 51620 (IBSC), K. M. Feng 1905 (IBSC), 22237
(IBSC), Y. M. Shui et al. 591 (KUN). Wuding, H. Li et al. 1466 (KUN),
0941 (KUN), J. M. Hui 1092 (YUKU), S. S. Sin 2 (KUN). Yimen, H. K.
Liu 23 (YCF). Yingjiang, G. D. Tao 013062 (KUN). Yunlong, Westfor-
est Exped. 162 (SWFC). Zhenxiong, Spicy Pl. Exped. 870471 (KUN),
870470 (KUN), N. E. Yunnan Exped. 1083 (KUN), P. H. Yu 23 (KUN),
S. S. Sin 923 (IBSC).
Du (2006) considered E. pilostyla is a synonym
of E. delavayi. After carefully studying specimens and
protologue, we agree with his treatment.
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SUN & LIN: A revision of Elaeagnus in China 373
Fig. 5. A, Lectotype of Elaeagnus longiloba C. Y. Chang (Y. Tsiang 9049, IBSC). B, Holotype of Elaeagnus retrostyla C. Y. Chang (P. H . Y u 2 8 3 ,
KUN). C, Isotype of Elaeagnus xingwenensis C. Y. Chang (C. Y. Chang 5,SZ).D, Lectotype of Elaeagnus bockii var. muliensis C. Y. Chang (S. K. Wu
3510,PE).
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374 Journal of Systematics and Evolution Vol. 48 No. 5 2010
Elaeagnus retrostyla and E. longiloba were pub-
lished by Chang (1980) who proposed that both species
had an affinity with E. lanceolata, but they could be
distinguished from E. lanceolata in leaves, veins, and
lobes. However, these two species are known to us only
from protologues and their collections are restricted in
type specimens from the same locality (Bijie, Guizhou).
The holotypes of these two species represent com-
pletely overlapping morphological characters with each
other as well as E. delavayi, since E. delavayi is nar-
rowly elliptic or oblong-lanceolate, lateral veins 5–8
per side of midrib, and lobes triangular, 2.5–4 mm, apex
acuminate. Moreover, our observation on the holotype
of E. retrostyla under the microscope shows that not
all of its style was coiled into a spiral up to base of
lobes, but only a few, which seems to be caused by
their anomalous lobes. Therefore it is probably true
that its style was pressed by lobes forcing it coiled.
This case is not alone, and can be frequently observed
among other species, such as E. difficilis. Some overlap-
ping features (oblong-lanceolate leaf blade, long pedi-
cel, and obconic-tubular calyx) suggest that E. delavayi
includes E. longiloba and E. retrostyla, which is sup-
ported by specimens including G. F. Li 65051 (IBSC,
KUN, NAS, PE), C. Z. Dang 1579 (HGAS), S. S. Sin 2
(KUN), K. M. Feng 10405 (KUN), and B. Y. Qiu 53468
(KUN).
Elaeagnus xingwenensis was a different case, be-
cause Chang (1981) already realized that E. xingwenen-
sis had an affinity with E. delavayi, and he discussed
that E. xingwenensis could be easily recognized by its
petiole 8–1.2 cm, leaf blade bigger, lateral veins 5–6 per
side of midrib, impressed or flat abaxially, apex obtuse
or acuminate. But Qin & Gilbert (2007) suggested that
E. xingwenensis was described only from fruiting mate-
rials and could not be included in the identification key.
In addition, they also pointed out that E. xingwenensis
was known to them only from the protologue. We had
the opportunity to access the type materials of E. xing-
wenensis consisting of 12 sheets of fruiting specimens.
Therefore, one single collection with poor diagnostic
characters is insufficient to separate E. xingwenensis
from E. delavayi as a new species. Specimens: G. F.
Li 65051 (IBSC, KUN, NAS, PE), C. Z. Dang 1579
(HGAS), S. S. Sin 2 (KUN), K. M. Feng 10405 (KUN),
and B. Y. Qiu 53468 (KUN), are good examples on this
point. We thus suggest that E. retrostyla and E. longiloba
as well as E. xingwenensis be included in E. delavayi,as
they are all uniformly well characterized and also have
overlapping distribution areas.
20. Elaeagnus bockii Diels in Bot. Jahrb. Syst. 29:
482. 1900. Type: China. Sichuan: Nanchuan, Rosthorn
3144 (lectotype, designated by A. Rehder in Sargent,
Pl. Wilson. 2: 416. 1915: B).
Elaeagnus bockii var. muliensis C. Y. Chang, Fl.
Sichuan. 1: 463. 1981. Type: China. Sichuan: Muli, S. K.
Wu 3510 (lectotype, designated by Q. Lin in Bull. Bot.
Res., Harbin 26(6): 656. 2006: PE!), Fig. 5: D.
Evergreen; shrubs, erect, 1–3 m tall. Spines
present, thick; young branches columned, densely with
rust-colored or brown scales. Petiole brown, 0.4–0.8 cm;
leaves leathery, rarely sub-leathery; speckled; blade pale
green above when dried, yellowish white below, lanceo-
late, sometimes oblong-elliptic, apex acuminate, rarely
rounded, margin narrowly revolute, base cuneate; 3–
9×0.8–3.5 cm, adaxially glabrous or sparsely yellowish
brown scales, abaxially with overlapping silvery scales,
scales on older leaves deeply fimbriate-lacerate; lat-
eral veins 5–7 per side of midrib, slightly raised on
both surfaces. Flowers often 5–7 forming a shortened
raceme; pedicels 3–8 mm. Flowers yellowish white; ca-
lyx tubular or funnelform-tubular, outside with silvery
and reddish brown scales; thick papery; 4–7 ×2–3 mm;
lobes triangular or ovate-triangular, apex acute, inside
sparsely with white scales and stellate, 2.5–3 ×1.5–
2 mm; anthers oblong, ca. 1.2 mm; filaments ca. 0.5 mm;
style curved, densely whitish stellate-hairy, sometimes
subglabrous; stigma beyond stamens. Fruit elliptic, 1–
1.2 ×0.6 cm, densely with scales, red when ripe, juicy;
ribs inconspicuous; fruit pedicels slender, 4–5 mm,
sometimes longer. Fl. Oct.–Nov., fr. Mar.–Apr.
Mountain slopes and roadsides; alt. 350–2900 m.
Endemic to China. Chongqing, Gansu, Guangdong,
Guangxi, Guizhou, Hubei, Hunan, Shaanxi, Sichuan,
and Yunnan.
Additional specimens examined:
China. Precise locality unknown, Q. Y. Liu 4136 (IBSC), W. K.
Hu 1124 (SZ), 55 (SZ), H. S. Kung 3626 (SZ), S. Y. Hu 217, 215 (SZ),
Y. Han 1979 (SZ), Z. X. Zhao 225 (KUN). Chongqing: Precise locality
unknown, T. N. Liou 9724 (PE), T. Y. Cheo & K. C. Tsu 80 (N). Beibei,
C. Pei 7777 (NAS, PE), 7709 (NAS), 10278 (NAS), K. L. Chu 4152
(IBSC), 6650 (NAS, PE), R. H. Shan 1241 (NAS), T. T. Yu 7 (IBSC),
2803 (PE), Jinyunshan, S. C. Wang 1424 (NAS), 1481 (NAS), Y. Z. Sun
1461 (NAS), 1395 (IBSC, NAS, PE), 1447 (PE). Chengkou, P. Y. Li 6529
(PE), T. L. Dai & Z. He 103543 (IBSC, NAS, PE, SZ). Fengjie, H. F.
Zhou et al. 110523 (IBSC), 110646 (IBSC), 111636 (IBSC), 0111038
(IBSC), 111252 (IBSC), 107562 (IBSC, PE), 107595 (IBSC, PE), 11711
(PE), 107715 (PE). Jiangbei, T. P. Wang 12132 (PE), Z. He 14357 (SZ).
Nanchuan, F. T. Wang10439 (PE), G. F. Li 64622 (IBSC, KUN, SZ, NAS,
PE), 60156 (IBSC, KUN, SZ, NAS, PE), 60062 (IBSC, KUN, SZ, NAS,
PE), 60226 (IBSC, KUN, SZ, NAS, PE), 64947 (IBSC, KUN, PE, SZ),
65046 (IBSC), 60008 (PE, SZ), J. H. Xiong & Z. L. Zhou 90070 (IBSC,
SZ, PE), 9307 (IBSC, PE), 90371 (SZ), K. C. Kuan et al. 193 (CDBI),
Y. C. Yang 3036 (PE, SZ). Qijiang, Z. He 12091 (SZ). Wulong, S. R.
Yi et al. 0018 (PE). Wushan, G. H. Yang 65565 (KUN). Wuxi, K. L.
Chu 2015 (IBSC), G. H. Yang 65114 (NAS). Gansu: Bailongjiang, C. K.
Chow 90 (NAS), 63 (NAS). Lixian, Z. P. Huang et al. 2030 (KUN, NAS,
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SUN & LIN: A revision of Elaeagnus in China 375
PE), S. S. Chien s. n. (KUN). Kangxian, Z. Y. Zhang 17265 (PE), 17289
(PE). Wenxian, Baishuijiang Exped. 3613 (PE), 3693 (PE), 4845 (PE),
4789 (PE), 4902 (PE), J. X. Yang et al. 3453, 3519 (IBSC, PE), X. Wang
60 (PE), X. Y. Zhu et al. 20228 (PE), Y. C. Hou 00976, 01245 (PE), Z. Y.
Zhang 14019 (PE), 14219 (PE), 14799 (PE), 15366 (PE), 15567 (PE).
Guangdong: Lechang, X. R. Liang & R. K. Huang 31568 (IBSC, SYS).
Guangzhou, C. M. Hu 7280 (IBSC). Huaiji, G. L. Shi 15039 (IBSC).
Ruyuan, 75-Improved Group 20205 (SYS). Guangxi: Guilin, anonymous
92838, 4631 (IBK), G. Z. Li 13419 (PE), Sino-Germany Shaanxi Exped.
096 (IBSC). Lingui, C. H. Tsoong 808511 (IBK), G. Z. Li 16273 (PE).
Longlin, anonymous s. n. (KUN Herb. Bar Code No. 0622191), C. C.
Chang 10707 (KUN). Rongxian, C. H. Tsoong 83790 (IBK). Tian’e,
G. D. Tao & L. H. Tan 3–1627 (GXMI). Xing’an, G. Z. Li 16725 (PE).
Guizhou: Precise locality unknown, S. J. Wang 1424 (IBSC), S. W.
Teng 1578 (IBSC). Fanjingshan, Y. Tsiang 7989 (NAS, IBSC). Jiangkou,
C. Y. Chiao et al. 946 (PE, SYS). Renhuai, F. Gao 2047 (IBSC, PE),
P. C. Tsoong 128 (KUN, PE). Xishui, C. W. Wu et al. 1067 (IBSC, PE).
Xingyi, Z. Y. Deng 2413 (KUN). Yanhe, J. W. Xiao 5530 (PE). Zhenfeng,
Y. Tsiang 4565 (IBSC). Hubei: Badong, Z. S. Lu 17 (NF). Fangxian, G. R.
Liu 403 (IBSC, NF). Lichuan, G. G. Tang 471 (NF), W. C. Cheng & C. T.
Hwa 800 (PE, NF), Tieshan, Y. Chen 918 (NF). Xianfeng, H. J. Li 7596
(IBSC, PE). Zhuxi, Z. Zheng 91–316 (KUN). Zigui, H. J. Li 1908 (IBSC).
Hunan: Sangzhi, anonymous 1767 (KUN), Sangzhi Veg. Exped. 948
(IBSC). Taojiang, L. D. Duan 3629 (PE), 4651 (PE). Yongshun, C. J. Qi
3662 (IBSC, PE), H. Li et al. 1594 (KUN). Shaanxi: Baxian,K.T.Fu
5605 (KUN, PE), 6132 (PE), T. N. Liou 11961 (PE), T. P. Wang 10191,
9061 (PE). Chenggu, K. T. Fu 5453 (IBK, IBSC, PE), T. N. Liou 11257
(PE), Z. Y. Zhang 18120 (IBSC). Foping, X. M. Zhang 609 (KUN).
Lueyang, C. L. Tang 291 (KUN), 534 (NAS, KUN, PE), K. T. Fu 5753
(IBK, IBSC, KUN, PE), T. P.Wang 18639 (KUN). Meixian, K. T.Fu 3930
(KUN, PE), T. P. Wang 9061 (KUN), 10191 (KUN). Mianxian, K. T. Fu
5605 (IBSC), T. P. Wang 9061 (CDBI). Nanzheng, X. X. Hou 241 (IBSC),
Z. X. Hu et al. 900 (IBSC). Ningqiang, T. N. Liou 11912 (PE). Ningshan,
J. Q. Xing 4210 (IBSC). Shiquan, J. Q. Xing 3341 (IBSC). Xixiang, P. C.
Tsoong 4043 (PE). Yangxian, P. C. Kuo 1968 (IBK, IBSC, PE), T. P.
Wang 16335 (KUN, PE), J. X. Yang 1561 (PE). Sichuan: Precise locality
unknown, anonymous 5574 (IBSC), E. H. Wilson 4420 (IBSC), H. Smith
13478 (PE), 13568 (PE), 13603 (PE), H. F. Zhou et al. s. n. (IBSC), H. Yu
279 (PE), S. J. Wang 1424 (PE), S. N. Xu s. n. (NAS), T. P. Wang 9061
(KUN), T. T. Yu 1255 (IBSC), 4127 (IBSC), W. C. Cheng 3039 (IBSC),
3429 (IBSC), W. K. Hu 9 (SZ), 12 (SZ), 14 (SZ), 15 (SZ), 16 (SZ), 21
(SZ), 31 (SZ), 37 (SZ), 61 (SZ), 68 (SZ), 102 (SZ), W. P. Fang 121022
(KUN), 12194 (IBSC), 9726 (NAS), 9468 (SZ), 5071 (NAS), Y. C. Yang
4179 (NAS, PE), 3016 (PE). Baoxing, K. C. Kuan & W. T. Wang 2549,
2786 (PE), K. L. Chu 6276 (NAS, PE), 3878 (PE), T. H. Tu 4866 (PE),
T. P. Soong 38207 (IBSC, NAS, PE), 1954 (KUN), T. P. Wang 10101 (PE),
Sichuan Econ. Pl. Exped. 00054 (PE), Sw. Univ. Exped. 1498 (KUN),
T. T. Yu 14380 (KUN), S. K. Wu 3510 (KUN, PE), 2565 (KUN, PE),
X. S. Chang & Y. X. Ren 7522 (PE), 7443 (PE), 7299 (PE). Chendu,
C. S. Fan 1459 (N), H. T. Chang 5446 (SYS), X. Y. He et al. 4 (SZ), 9
(SZ), 3 (SZ), T. P. Wang 10101 (KUN), 13246 (IBSC), W. K. Hu 18 (SZ),
W. P. Fang 13246 (KUN), 13164 (KUN), 12022 (SZ), Y. B. Yang 8249
(CDBI), Y. Chen 7850 (NF). Dajin, X. Li et al. 76904 (IBSC, SZ, PE,
NAS), 76956 (NAS, PE, SZ). Dujiangyan, K. C. Kuan 0006 (PE), X. Li
47356 (IBSC, PE, SZ), 47306 (IBSC, PE, SZ), D. H. Zhu et al. 1262 (PE).
Emei, anonymous 4893 (PE), C. H. Li 96–41 (PE), 97–345 (PE), G. F. Li
53239 (IBSC, SZ), G. H. Yang 53974 (IBSC), 57544 (NAS, SZ), G. S.
Zhou 81230 (PE), H. L. Tsiang 33254 (IBK), H. L. Tsiang & S. S. Chang
33512 (IBK, IBSC, NAS, PE), J. H. Xiong et al. 33254 (IBSC, NAS, PE),
33325 (IBK, IBSC, NAS, PE), 33394 (IBK, IBSC, SZ, NAS, PE), 32972
(IBK, IBSC, PE, SZ), K. C. Kuan et al. 2785 (PE), P. J. Dai 0049 (PE),
S. L. Sun 1378 (KUN), Dept. Biol. Univ. Sichuan s. n. (IBSC), 5472
(SYS), 53102 (SYS), 53572 (SYS), 48280 (SYS), 53529 (SYS), 53974
(SYS), 57544 (PE, SYS), 53005 (SZ, SYS), 52949 (SZ, SYS), 53090 (SZ,
SYS), 52356 (SZ, SYS), 53174 (SZ, SYS), 53383 (SZ, SYS), T. N. Liou
12744 (PE), T. T. Yu 262 (PE), W. K. Hu 6 (SZ), 9200 (SZ), W. P. Fang
19362 (PE), 15761 (PE), 13777 (KUN), 14934 (KUN), 13899 (KUN),
15633 (KUN, PE), 14959 (SZ), 13724 (KUN, SZ), 13899 (KUN, SZ),
10591 (KUN, SZ), 15475 (KUN, SZ), 15966 (IBSC, PE), 15891 (IBSC),
15832 (IBSC), 17561 (IBSC), 18085 (IBSC), 15381 (KUN, IBSC, PE),
15455 (NAS), s. n. (KUN Herb. Bar Code No. 0622199), W. R. He 569
(KUN), Y. B. Yang 53573 (SZ), Y. Chen 7021 (IBSC, NF), 5018 (NF),
Y. S. Shiao 48071, 48337 (IBSC, SZ), Z. S. Zheng 301 (KUN), Z. W. Yao
3290 (NAS, PE). Guangyuan, P. Y. Li 991 (SZ), T. N. Liou & C. Wang 164
(PE), Y. C. Hou 1535 (PE), Z. P. Wei 3441 (SZ). Hanyuan, C. Pei 8302
(PE), T. P. Wang 9620 (PE). Hechuan, C. J. Tsiang 5311 (PE). Jiange,
T. N. Liou 11748 (PE). Jiangjin, C. Pei 7308 (PE). Jiangyang, Y. P. Xu
466 (SZ). Kangding, Q. S. Cao et al. 110840 (CDBI, PE, SZ), W. K. Hu
& Z. He 11389 (PE, SZ), 11349 (PE, SZ), Z. J. Zhao et al. 115392 (SZ),
113310 (SZ). Leshan, Z. X. Li et al. 8790 (CDBI). Leibo, Z. J. Ren 3941
(PE). Lixian, X. Li 46694 (IBSC, PE, SZ). Longchang, Y. P. Xu 285(SZ),
Z. He et al. 5793 (NAS). Luding, anonymous 25033 (CDBI). Mianning,
T. T. Yu 1755 (PE). Maoxian, anonymous 8913 (CDBI), W. P. Fang 5574
(PE). Muli, G. Yao 127, 448 (NAS), Q. S. Cao et al. 4320 (SZ), S. K. Wu
2565 (SZ), T. T. Yu 14380 (PE). Neijiang, Y. P. Xu 367 (SZ). Pengzhou,
D. H. Zhu et al. 20070480 (PE). Pingwu, Z. L. Wang 0009 (SZ), P. Y.
Li 991 (SZ). Shimian, C. J. Xie 39999 (IBSC, PE). Songpan, K. T. Fu
2249 (PE). Tianquan, D. Y. Peng 46774 (IBSC), 46731 (IBSC), K. L.
Chu 2288 (PE), 2295 (IBSC, PE, SZ), K. Y. Lang et al. 241 (PE), Q. S.
Cao et al. 116232 (SZ), F. J. Tai 5315 (KUN, SZ), X. Y. He 6907 (NAS,
PE), Y. Chen 5792 (NF). Wanyuan, K. L. Chu 2216 (PE). Wenchuan,
H. Z. You et al. 532 (KUN), 465 (KUN), 532 (KUN), C. Y. Wu et al.
86–1184 (KUN). Xichang, Sichuan Agri-Bio Institute 95 (SZ). Ya’an,
X. Y. He 6907 (NAS). Yingjing, T. P. Wang 9915 (PE), W. K. Hu & Z.
He 12091 (PE). Ziyang, Y. P. Xu 409 (SZ). Yunnan: Dali, T. N. Liou
15238 (KUN). Kunming, H. K. Teng 335 (KUN). Lijiang, T. T. Yu 11093
(KUN). Songming, Y. P. Chang 80 (KUN). Suijiang, X. X. Li 12 (IBSC).
Wuding, H. Li et al. 0848 (KUN).
Elaeagnus bockii is a long recognized species
widespread in Sichuan Province and nearby areas.
Chang (1983) remarked that E. bockii represented a
wide variation range. For example, the length and shape
of calyx tube of this species is variable from place to
place, even individuals from Chengdu city to Mt. Emei.
The differences were even extraordinary, when compar-
ing collections from Baxian, Jiangjin, and Guangyuan
to other adjoining provinces like Gansu and Shaanxi.
However, Chang considered that E. bockii var. muliensis
can be easily separated from E. bockii by being barely
spiny on young branches, smaller leaves, scaly style,
and 1.3 cm long fruiting pedicels. However, examina-
tion of type specimens of E. bockii var. muliensis and
specimens of E. bockii (K. T. Fu 2249 (PE), 3930 (KUN,
PE), X. Y. He 6907 (NAS, PE), X. Li 76956 (NAS, PE,
SZ), C. L. Tang 534 (NAS, KUN, PE), and Y. C. Hou
00976 (PE)) suggest that this variety should be included
in E. bockii.
21. Elaeagnus pungens Thunb. in Murray, Syst. Veg.,
ed. 14. 164. 1784. Type: unknown.
Elaeagnus obtusa C. Y. Chang in J. Sichuan
Univ., Nat. Sci. ed., 4: 92. 1984. Type: China. Hunan:
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2010 Institute of Botany, Chinese Academy of Sciences

376 Journal of Systematics and Evolution Vol. 48 No. 5 2010
Yizhang, S. H. Chen 5314 (holotype, IBSC!),
Fig. 6: A.
Evergreen; shrubs, erect, 3–4 m tall. Spines
present; young branches columned, densely with brown
scales. Petiole brown, 0.5–1.8 cm; leaves leathery; blade
green above, grayish brown below, elliptical to oblong,
apex rounded or obtuse, margin revolute, base rounded;
5–18.5 ×1.8–6.5 cm, adaxially glabrous or sparsely yel-
lowish brown scales, abaxially with dense grayish white
and brown scales; lateral veins 7–9 per side of midrib.
Flowers several in a shortened raceme; pedicels brown,
ca. 5 mm. Flowers yellowish white; calyx funnelform-
tubular, outside with silvery and reddish brown scales;
thick papery; 3–5 ×2–3 mm; lobes ovate, apex rounded,
inside sparsely with white stellate, 1.5–3 ×1.5–2 mm;
anthers oblong, ca. 1.2 mm; filaments ca. 0.5 mm; style
erect, densely whitish stellate-hairy, sometimes sub-
glabrous; stigma beneath stamens. Fruit elliptic, 1.2–
1.5 ×0.6 cm, densely with scales, red when ripe, juicy;
ribs inconspicuous; fruit pedicels slender, 4–5 mm. Fl.
Sep.–Dec., fr. Apr.–Jun.
Open slopes, roadsides, thickets, and seashores;
below 1000 m in altitude. Anhui, Fujian, Guangdong,
Guangxi, Guizhou, Hubei, Hunan, Jiangsu, Jiangxi, and
Zhejiang.
Additional specimens examined:
China. Precise locality unknown, anonymous s. n. (HHBG), 187
(IBSC), A215 (KUN), R. C. Ching 2610 (N), T. Y. Cheo 2231 (IBSC),
Z. X. Zhao 994 (KUN). Anhui: Precise locality unknown, anonymous
2610 (IBSC), Y. F. Xiao & W. Z. Xie 23 (IBSC), 29 (IBSC). Chuxian, A. N.
Steward 1084 (N), M. Chen 1047 (N, PE), 1442 (PE), M. K. Wang 4054
(NAS), W. C. Cheng 5221 (PE). Huangshan, H. Zou 01549 (PE), 01745
(PE), J. S. Yue 1886 (NAS), M. Chen 1090 (IBSC, PE), 1047 (IBSC, PE),
Statio Orientali-Sinensis 6167 (NAS, PE), W. C. Cheng 4042 (NAS, PE),
4604 (NAS, PE), W. Z. Fang 205 (NAS), Z. Zheng 0173 (NAS), 0207
(NAS), 0219 (NAS). Huoshan, M. B. Deng 81273 (NAS), 81474 (NAS).
Jinzhai, M. B. Deng 81178 (HHBG, NAS), 81780 (NAS), 81697 (NAS),
Y. F. Xiao & W. Z. Xie 31 (IBSC). Jingxian, anonymous 375 (NAS), 618
(NAS). Jiuhuashan, anonymous 5120 (NAS), R. C. Ching 2610 (N, NAS),
Statio Orientali-Sinensis 5984 (NAS, PE), 4737 (NAS, PE). Langyashan,
J. S. Yue 0095 (NAS), 274 (NAS), M. B. Deng 3012 (NAS). Qimen,
M. B. Deng 4850 (IBSC, NAS, PE), 5223 (IBSC, NAS, PE). Qianshan,
B. A. Shen 0309 (PE), 0152 (PE), Statio Orientali-Sinensis 7169 (N,
NAS). Shexian, anonymous 0025 (NAS), 1130 (NAS). Shucheng, M. B.
Deng 90130 (PE), Statio Orientali-Sinensis 4460 (NAS, PE), 4086 (NAS,
PE), 4218 (NAS, PE). Tongcheng, An-Tong Exped. 032 (NAS). Xiun-
ing, M. B. Deng A0594 (NAS). Xuancheng, anonymous99 (NAS). Yuexi,
Statio Orientali-Sinensis 6991 (IBSC, NAS, PE), 6920 (IBSC, NAS, PE).
Chongqing: Chengkou, T. L. Dai 104226 (IBSC). Fengjie, H. F. Zhou
& H. Y. Su 111381 (IBSC). Wushan, G. H. Yang 65565 (IBSC), 65114
(IBSC). Fujiang: Precise locality unknown, P. C. Tsoong 323 (IBSC).
Dehua, P. C. Tsoong 231 (IBSC, PE). Dongshan, G. S. He 525 (PE).
Fuzhou, H. T. Chang 7222 (SYS), 7223 (SYS). Wuyishan, M. K. Wang
et al. 1857 (NAS). Guangdong: Precise locality unknown, F. A. Mcclure
1901 (SYS), T. S. Liu 47 (IBSC), 88 (IBSC), 67 (IBSC). Fengkai, G. Q.
Ding et al. 6796 (IBSC). Huaiji, G. L. Shi 15004 (IBSC). Lechang, Nan-
ling Exped. 3053 (IBSC), N. Q. Chen 42019 (IBSC). Lianxian, C. L. Tso
22628 (IBSC). Qingyuan, F. C. How 74166 (IBK). Ruyuan, 2366 Exped.
3042 (IBSC, PE), C. Wang 42309 (IBSC, SYS), 44401 (IBSC), H. Y.
Mark 122 (IBSC), Improved Exped. 20205 (SYS), Nanling Exped. 1980
(IBSC), S. B. Guo 80019 (IBK, IBSC), S. K. Lau 28921 (IBK, IBSC),
S. H. Chun 239 (IBK, IBSC), S. P. Ko 53752 (IBK, IBSC, KUN, NAS,
PE), 53349 (IBK, IBSC), 53520 (IBSC), 54643 (PE). Yangshan, Nanling
Exped. 1261 (IBSC), IBSC Geobotanical Exped. 454 (IBSC). Yingde, Y.
Tsiang 491 (IBSC). Guangxi: Precise locality unknown, C. H. Tsoong
94258 (IBK). Fuzhong, anonymous 13986 (IBSC). Guilin, C. H. Tsoong
808511 (IBSC), L. Q. Chen & Z. N. Deng 13270 (IBSC). Lingui, C. H.
Tsoong 808511 (PE), 808780 (PE), S. Q. Zhong A61250 (KUN, PE). Li-
uzhou, C. H. Tsoong 84515 (IBK). Longsheng, B. Liu 0506 (PE). Luzhai,
M. Y. Wang 12 (PE). Napo, Z. J. Li et al. 1166 (IBK). Quanzhou, Q. Li
0007 (PE), Z. Z. Chen 52722 (IBSC, IBK, KUN). Xing’an, F. Liu 0003
(PE), S. L. Yu 900006 (IBSC). Yangshuo, R. H. Shan 603, 789 (PE).
Guizhou: Libo, Q. H. Chen 2094 (HGAS). Qingzhen, S. W. Teng 90134
(IBK, NAS). Weng’an, Libo Exped. 1993 (HGAS), 1966 (HGAS), 1992
(PE). Zunyi, Sichuan-Guizhou Exped. 1322 (PE). Henan: Luanchuan,
K. C. Kuan 2477 (IBSC). Shangcheng, anonymous 43 (PE). Songxian,
anonymous 924 (PE). Zhenping, Dept. Forest. Henan 1501 (PE). Hunan:
Precise locality unknown, H. F. Zhou 45160 (PE), Y. F. Deng et al. 11810
(PE), 11993 (PE), 11829 (PE). Changsha, J. S. Yue 1442 (NAS). Chengbu,
P. C. Tsoong 1446 (PE). Daweishan, C. J. Qi 6753(IBSC). Dong’an, J. K.
Liu 382 (PE), Y. Liou 886 (NAS). Dongkou, Z. Y. Yang 710 (IBSC).
Fenghuang, anonymous 2 (NAS). Guangjisi, B. G. Li et al. 565 (IBSC).
Hengshan, C. X. Ye, 4931 (SYS), F.C . How 74166 (IBSC), Hunan Exped.
792 (IBSC, PE), Lee 22 (IBSC), Z. H. Hu 489 (PE). Jianghua, B. G. Li
et al. 05411 (IBSC, PE). Nanyue, M. H. Li & Y. Q. Kuang 1130 (PE), 644
(PE), 654 (PE), Z. H. Hu 613 (PE). Qidong, J. D. Li 092 (PE), 445 (PE).
Qianyang, Z. T. Li 1679 (IBSC, PE). Shaodong, J. H. Liu 7023 (PE),
7473 (PE). Shaoyang, L. D. Duan 2074 (PE), 0637 (PE). Taoyuan, G. F.
Liao 420 (PE). Tongdao, C. J. Qi 3376 (IBSC, PE). Wugang, H. T. Chang
4659 (IBSC, SYS), L. H. Liu et al. 15950 (IBSC, PE). W. Hunan, L. H.
Liu 10065 (IBSC). Xinning, anonymous 32 (PE), L. H. Liu 15501 (IBSC,
KUN, PE), Z. C Luo 1717 (PE). Xinshao, B. Y. Li 6441 (PE). Yizhang,
P. H. Liang 83011 (IBK, IBSC), 546 (IBSC), M. X. Huang 113115
(IBSC), S. H. Chun 36 (IBK, IBSC), 5314 (IBSC). Yuanling, F. J. Zhou
015 (PE). Jiangsu: Precise locality unknown, anonymous 1287, 1119,
7287, 380 (IBSC), A. N. Steward 893 (N), C. C. Kung 10 (IBSC), C. L.
Tso 225 (IBSC), H. C. Chang s. n. (SYS Herb. Bar Code No. 00101584),
J. S. Yue 9 (PE), T. Y. Cheo 38 (N), W. C. Cheng 4693 (N), W. X. Wu
2426 (NAS). Changshu, T. Y. Cheo 2231 (NAS). Jiangning, M. B. Deng
3080 (NAS), Nanlin Exped. 6739 (IBSC). Jiangpu, A. N. Steward 2270
(N), A. N. Steward & H. C. Cheo 3504 (N), M. B. Deng 3791 (KUN,
NAS, PE). Jiangyin, T. Y. Cheo et al. 934 (NAS). Jurong, Chen & Teng
134 (NAS), C. H. Chun 2243 (NAU, PE), K. C. Kuan 172 (PE), M. B.
Deng 3448 (KUN, NAS, PE), 2982 (NAS), J. J. Gong 549 (NAS), 275
(IBSC, NAS), J. S. Yue 825 (NAS), W. Z. Fang 058791 (NAS), Y. Tsiang
10831 (NAS). Nanjing, anonymous 153 (IBK, NAS), A. N. Steward 2770
(SYS), C. C. Chen 1208 (IBSC), Chen & Teng 3903 (IBSC, NAS), C. L.
Tso 43 (NAS, PE), 969 (NAS, PE), C. N. Chen 8694 (NAS), C. Y. Luh
315 (NAS), 335 (NAS), 356 (NAS), 380 (NAS), F. X. Liu 891 (IBSC,
NAS, PE), G. Yao 8468 (NAS), K. Ling 1388 (N), J. J. Gong 273 (IBSC),
275 (PE), 10 (NAS), J. S. Yue 0009 (NAS), S. J. Yang et al. 178 (NAS),
T. Y. Cheo et al. 508 (IBSC, PE), W. C. Cheng 4222 (NAS), Y. N. Xiong
149 (NAS). Suzhou, P. Z. Ling & S. X. Sun 67 (IBSC, KUN, NAS, PE),
W. Z. Fang et al. 88 (IBSC, NAS, PE). Wuxi, G. Yao 10895 (NAS),
W. X. Wu 9643 (NAS). Wuxian, Wuxian Exped. 111 (PE). Yixing, T.
Shen 987 (NAS), 1061 (NAS, SYS), Z. Z. Ding 35, 846 (NAS). Jiangxi:
Anonymous 356 (IBSC), y0218 (PE), C. Liu 127 (PE), H. H. Hu 2082
(PE), Jiangxi Univ. 5428 (IBSC, KUN), 5603 (KUN), 5645 (KUN), 6291
(KUN), 5124 (KUN), 4360 (KUN), M. K. Wang 3999 (IBSC), M. X. Nie
6715 (IBSC), S. T. Shen 00058 (IBSC), Y. Tsiang 10831 (IBSC). Anfu,
J. S. Yue 3745 (NAS, PE). Boyang, M. K. Wang 3999 (NAS). De’an,
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SUN & LIN: A revision of Elaeagnus in China 377
Fig. 6. A, Holotype of Elaeagnus obtusa C. Y. Chang (S. H. Chen 5314, IBSC). Aa, Floral buds. B, Isotype of Elaeagnus viridis Serv. (Henry 1105,
K). C, Holotype of Elaeagnus xichouensis C. Y. Chang (H. T. Tsai 58–8484, KUN). D, Holotype of Elaeagnus xizangensis C. Y. Chang (S. Z. Cheng &
B. S. Li 03399,PE).
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378 Journal of Systematics and Evolution Vol. 48 No. 5 2010
C. M. Tan 9611031 (IBSC, NAS). Dexing, M. X. Nie & S. S. Lai 5603
(IBSC, PE). Fengxin, G. Yao 1337 (NAS), S. S. Lai 000970 (PE). Fengyi,
X. P. Ding 133 (PE), S. S. Lai 006 (PE). Guixi, anonymous 900456
(NAS). Jing’an, J. H. Zhang 97117 (IBSC, PE). Jingdezhen, X. X. Yang
et al. 17515 (IBSC). Jinggangshan, Jiangxi Exped. 0904 (PE), J. Xiong
2849, 2847 (PE), S. S. Lai 5006 (IBSC). Jiujiang, anonymous 10271
(PE), C. M. Tan 92139 (HHBG, NAS), 98453 (IBSC), 95012 (HHBG,
IBSC, NAS), 92110 (HHBG, IBSC, NAS), 941685 (PE), 951374 (PE),
G. Yao 8599 (NAS), A. N. Steward & H. C. Cheo 385 (N, NAS), H.
Migo s. n. (NAS), H. Zou 1019 (NAS), J. Xiong 7076 (PE), M. K. Wang
01166 (NAS), 199 (NAS), 0180 (NAS), 0002 (NAS), M. X. Nie et al.
7934 (KUN, PE), 92371 (PE), R. C. Ching 12111 (NAS), S. T. Shen
825 (IBSC), Y. G. Xiong 857 (NAS), 4997 (NAS). Guling, H. Zou 1019
(IBSC). Guangfeng, M. X. Nie 5645 (PE). Lichuan, M. K. Wang et al.
2310 (HHBG, IBSC, NAS, PE), X. X. Yang et al. 560023 (IBSC, PE).
Pengze, S. X. Yu & Q. G. Shen 3772 (PE). Pingxiang, Jiangxi Exped. 2478
(PE), 2838 (PE). Qianshan, C. P. Tsien et al. 400798 (PE), M. X. Nie &
S. S. Lai 4360 (PE). Shangrao, anonymous 5030 (IBSC, PE), M. X. Nie &
S. S. Lai 5124 (IBSC, PE), 5428 (PE). Suichuan, 236 Exped. 0043 (PE),
J. S. Yue 4565 (IBSC, NAS). Taihe, X. M. Mo 20497 (IBSC), 20413
(IBSC). Tonggu, S. S. Lai 03708 (PE). Wugongshan, Jiangxi Exped.
00435 (PE), 1641 (PE), 1205 (PE), Wuning, C. M. Tan 951261 (IBSC,
PE), 9604097 (NAS, PE), 941201A (NAS, PE). Wuyuan, C. Chen &
Q. H. Li 007 (PE), M. B. Deng 87179 (NAS). Xiushui, C. S. Ye 907
(NAS), S. S. Lai 03387 (PE). Yushan, M. X. Nie et al. 6075 (IBSC, PE),
6291 (IBSC, PE). Shanghai: Precise locality unknown, C. N. Yen 811
(HHBG), F. X. Liu 508 (NAS), H. Migo s. n. (NAS). Sichuan: Precise
locality unknown, W. P. Fang 9882 (NAS). Dujiangyan, K. C. Kuan et al.
0006 (CDBI, PE). Yongzhou, D. H. Du 161 (PE). Zhejiang: Precise local-
ity unknown, C. Pei 2461 (NAS). Anji, HHBG 003234, 3281 (HHBG).
Changhua, anonymous 69 (HHBG), 782358 (IBSC), X. Y. He 30327
(HHBG, NAS), 29429 (HHBG, NAS), 29813 (HHBG, NAS), 30985
(HHBG, IBSC, PE), 30847 (HHBG, IBSC, PE), 30565 (HHBG, IBSC,
PE), 0125 (HHBG, IBSC, PE), 2069 (HHBG, IBSC, PE), 1767 (HHBG,
IBSC, PE), 1096 (HHBG, IBSC, PE), 25992 (HHBG, IBSC, NAS, PE),
26658 (HHBG, IBSC, NAS, PE), 2006 (HHBG, IBSC, NAS, PE), 23109
(NAS), H. Q. Zhu 722 (IBSC), J80 J8026–183 (IBSC). Chun’an, L.
Hong 1478 (HHBG), M. L. She et al. 37539 (NAS). Hangzhou, anony-
mous 000722 (HHBG, PE), 1957 (HHBG, PE), 1390 (HHBG, PE), 2017
(HHBG, PE), 276 (HHBG, PE), 1430 (HHBG, PE), 1865 (HHBG, PE),
1916 (HHBG, KUN, PE), 0514 (NAS), C. Y. Chiao 30 (N), S. Y. Chang
1512 (HHBG, NAS, PE), 043 (HHBG, NAS, PE), 277 (NAS), 276 (PE),
T. N. Liou L401 (PE), X. Y. He 20499 (HHBG, IBSC, NAS). Jingn-
ing, anonymous 24632 (HHBG), 24169 (HHBG). Lishui, anonymous
3789 (NAS). Lin’an, Z. M. Ye J8111–031 (IBSC, PE). Moganshan, C. P.
Tsien 61180 (PE). Ningbo, anonymous 26 (HHBG), H. X. Zhou 512
(HHBG), K. M. Wang et al. 1093 (NAS), 1146 (NAS), L. Hong 3305
(HHBG), X. Y. Liou 1701 (PE). Pan’an, L. Hong 2850, 2597 (HHBG).
Putuo, C. Pei 2520 (NAS), HHBG 2345 (HHBG). Qingtian, Zhejiang Pl.
Exped. 8553 (NAS). Taishun, anonymous 25435 (HHBG). Tiantaishan,
A. N. Steward 1178 (N), C. Y. Chiao 1180 (N). Tianmushan, anonymous
001767 (NAS), G. R. Chen 2358 (PE), H. Q. Zhu 000722 (NAS), K. K.
Tsoong 425 (PE), L. Q. Qiu & R. L. Lu 1752 (IBK), T. N. Liou L6582
(PE), T. Shen 411 (NAS), 100 (N, NAS, SYS), 41 (SYS), 71 (SYS),
W. B. Wei 35210 (N), W. C. Cheng 2312 (PE), X. Y. He 0125 (NAS),
1096 (NAS), 20969 (NAS), 20965 (PE), Y. H. Liu 1119 (N), Y. W. Law
1119 (NAS, PE), 1287 (NAS, PE), Zhejiang Pl. Exped. 29429, 29006
(NAS). Tiantai, Zhejiang Med. Pl. Exped. 1439 (NAS). Wenling, HHBG
4086 (HHBG, PE), 4019 (HHBG, PE). Zhoushan, anonymous 4374, 4354
(HHBG).
Elaeagnus pungens has long been recognized as
a distinct species. Specimens of this species indicate a
wide range of distribution and morphological variation.
However, it can be easily identified based on its elliptical
leathery leaves and short funnelform calyx tube. Chang
(1984) considered E. obtusa as a new taxon, without
comparing it with E. pungens but with E. gonyanthes.
He stated that E. obtusa and E. gonyanthes are fairly
similar in flower, but with the following differences: (i)
young branches are grayish brown; (ii) adaxial leaves
surfaces are light green or green when dried, abaxially
white and petioles 5–18 mm long; and (iii) flowers 1–3,
clustered in axils of short branches, lobes inside sparsely
with white stellate and scales and style sparsely stellate-
hairy or glabrous. After examining the type specimens,
we found that Chang’s description was not accurate. He
did not realize that the “flowers” on the type specimen
of E. obtusa were actually buds closed in a 4-angled
campanulate shape, not yet open (Fig. 6: A, Aa). In
fact, all flowers of this genus are in a 4-angled cam-
panulate shape before anthesis. Conversely, leaves of
E. gonyanthes are densely covered with rust-colored
scales, pedicel 3–6 mm long, which can be easily sep-
arated from E. obtusa. But morphological characters
and distribution areas suggest that the circumscriptions
between E. obtusa and E. pungens are not clear. Our
reasonings are: (i) leaves of both species are elliptical to
narrowly oblong, 7–14 mm long, 3.8–6 cm wide, base
rounded, apex obtuse to bluntly acute, petiole robust; (ii)
both flower in December; and (iii) E. obtusa is known
to us only from type specimens and its type locality is
included in distribution areas of E. pungens. The spec-
imens of J. S. Yue et al. 1442 (NAS), Y. F. Deng et al.
11810 (PE), 11993 (PE), L. H. Liu et al. 15950 (IBSC,
PE), J. K. Liu 382 (PE), J. H. Liu 7023 (PE), and B. Y. Li
6441 (PE) also support expanding the circumscription
of E. pungens by treating E. obtusa as its synonym.
22. Elaeagnus viridis Serv. in Bull. Herb. Boissier, s´
er.
2. 8(6): 388. 1908. Type: China. Hubei: Yichang, Henry
1105 (holotype, P; isotype, K!), Fig. 6: B.
Elaeagnus viridis var. delavayi Lecomte in Bull.
Mus. Hist. Nat. Paris 21: 166. 1915. Type: China. Yun-
nan: Heqing, Delavay 247 (holotype, P).
Elaeagnus pallidiflora C. Y. Chang in Bull. Bot.
Lab. N. E. Forest. Inst., Harbin 1980(6): 107. 1980.
Type: China. Yunnan: Shuangbai, W. C. Yin 411 (holo-
type, YUKU!; isotype, KUN!).
Elaeagnus taliensis C. Y. Chang in Acta Phytotax.
Sin. 23: 376. 1985. Type: China. Yunnan: Dali, H. C.
Wang 4568 (holotype, IBSC!; isotypes, KUN!; PE!) (as
“4368” on p. 376, err. typogr.).
Elaeagnus wenshanensis C. Y. Chang, Fl. Sichuan.
1: 463. 1981. Type: China. Yunnan: Wenshan, H. T. Tsai
51496 (holotype, SZ!; isotypes, IBSC!; PE!).
Elaeagnus xichouensis C. Y. Chang in Acta Phyto-
tax. Sin. 23: 377. 1985. Type: China. Yunnan: Xichou,
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SUN & LIN: A revision of Elaeagnus in China 379
H. T. Tsai, 58–8484 (holotype, KUN!; isotype, KUN!),
Fig. 6: C.
Elaeagnus xizangensis C. Y. Chang in Bull. Bot.
Res., Harbin 6(2): 74. 1986. Type: China. Xizang:
Mˆ
edog, S. Z. Cheng & B. S. Li 03399 (holotype, PE!;
isotype, PE!), Fig. 6: D.
Evergreen; shrubs, erect, to 2 m tall. Spines present,
reflexed, ca. 1 cm; young branches up-ridged, densely
with rust-colored or yellowish white scales. Petiole
brown, 0.5–0.7 cm; leaves leathery or nearly papery;
blade light green above, yellowish white below, ellip-
tical to oblong, apex round or obtuse, margin revolute,
base rounded or obtuse; 2.5–11 ×1.2–3.5 cm, adaxially
glabrous or sparsely yellowish brown scales, abaxially
with dense silvery scales; lateral veins 6 or 7 per side
of midrib, slightly conspicuous on both surfaces. Flow-
ers several in a shortened umbellate-raceme; pedicels
brown, 2–3 mm. Flowers white; calyx broadly tubu-
lar, outside with silvery and few reddish brown scales;
papery; 3–8 ×2–3 mm; lobes broadly ovate or ovate-
triangular, apex acuminate, inside sparsely with white
stellate, 2.5 ×2 mm; anthers oblong, ca. 1.2 mm;
filaments ca. 0.5 mm; style erect, densely whitish
stellate-hairy; stigma beyond stamens. Fruit elliptic, ca.
1.3 ×0.7 cm, densely with silvery scales, red when ripe,
juicy; fruit pedicels slender, ca. 10 mm. Fl. Oct.–Nov., fr.
Mar.–May.
Shrubland, streamside, and thicket; alt. 500–
3150 m. Endemic to China. Chongqing, Guizhou,
Hubei, Hunan, Shaanxi, Shanxi, Sichuan, Xizang, and
Yunnan.
Additional specimens examined:
China. Chongqing: Fengjie, H. F. Zhou & H. Y. Su 11309 (PE),
111116 (IBSC, NAS, PE), 11440 (IBSC, NAS, PE), 110646 (IBSC, NAS,
PE), 110685 (IBSC, NAS, PE), 110309 (IBSC, NAS, PE), 11637 (IBSC,
NAS, PE). Nanchuan, G. F. Li 64357 (KUN, PE), J. H. Xiong et al. 92886
(KUN), 93072 (KUN). Wushan, G. H. Yang 65565 (PE), H. F. Zhou &
H. Y. Su 110285 (NAS). Guizhou: Precise locality unknown, S. W. Teng
90134 (KUN). Dafang, Bijie Exped. 946 (PE). Hezhang, W. Guizhou Ex-
ped. 2080 (HGAS), 1770 (HGAS), 1568 (HGAS). Jiangkou, C. Y. Chiao
et al. 946 (N), H. Peng 1086 (KUN). Panxian, Anshun Exped. 1002
(HGAS). Weng’an, Libo Exped. 1992 (KUN). Yinjiang, B. Bartholomew
et al. 1415 (HGAS). Hubei: H. J. Li 4845 (PE), W. Y. Chun 3790 (N),
3897 (N). Badong, H. C. Zhou 1644 (N). Lichuan, G. X. Fu & Z. S. Zhang
1556 (IBSC, KUN, NAS, PE). Yichang, Henry 1105 (IBSC, PE). Zigui,
H. J. Li 1908 (PE). Zhuxi, Z. Zheng 91–564 (KUN). Hunan: Yongshun,
H. Li et al. 1826 (KUN). Shaanxi: Zhouzhi, Zhu 1873 (KUN). Sichuan:
H. F. Zhou 111381 (IBSC), K. L. Chu 2216 (IBSC, PE). D ˆ
erong, H. Sun
et al. 2440(KUN). Mianning, T. T. Yu 18093 (IBSC, KUN, PE). Pingwu,
C. Y. Wu et al. 86–217 (KUN), 095 (KUN). Yanbian, X. F. Gao 3582
(PE), 3529 (PE). Xizang: Zay¨
u, Qinghai-Xizang Exped. 10745 (KUN).
Mˆ
edog, B. S. Li & S. Z. Cheng 01396 (PE), H. Sun et al. 5602 (KUN).
Yunnan: C. W. Wang 70427 (IBSC), 84530 (KUN), 84071 (KUN), E. E.
Maire s. n. (SYS), 3835 (IBSC, NAS), 68 (IBSC), 30 (NAS), H. T.
Tsai 51496 (IBSC, SZ), K. M. Feng 2341 (PE), M. Chen 2141, 3297
(SYS), S. T. Yue 1934 (KUN), T. N. Liou s. n., 15238 (IBSC), 17925
(IBSC), 17393 (IBSC), 15482 (IBSC), 23104 (IBSC), 14561 (IBSC),
15309 (IBSC), T. T. Yu 16949 (PE), Y. Tsiang 13038 (KUN). Anning,
H. S. Lo 229 (IBSC). Binchuan, H. Li et al. 451 (KUN). Dali, H. C. Wang
2058 (PE), 4104 (IBSC), 4568 (IBSC, KUN, PE), T. N. Liou 15468
(KUN), 15309 (IBSC, KUN, PE), 14353 (IBSC, KUN, PE), 017393
(PE), 017925 (PE). Dayao, Yunnan Tropical Pl. Exped. 60–085 (KUN).
Dˆ
eqˆ
en, K. M. Feng 23942 (KUN, PE), 23791 (KUN, PE), T. T. Yu 9634
(KUN, PE), 10521 (KUN, PE). Dongchuan, R. C. Ching 24893 (KUN,
PE). Drungjiang, anonymous 9874 (KUN), Qinghai-Xizang Exped. 9874
(KUN). Fugong, H. Li et al. 19835 (KUN). Fumin, B. Y. Qiu 596138
(PE, KUN). Funing, Wenshan Exped. 077 (KUN). Gongshan, H. Li et al.
14602 (KUN), 14388 (KUN), 14649 (KUN), K. M. Feng 8554 (KUN,
PE), Qinghai-Xizang Exped. 7611(PE). Jingdong, B. Y. Qiu 53404 (PE),
52921 (PE), Z. H. Yang et al. 101501 (KUN). Kunming, anonymous
s. n. (SWFC Herb. Bar Code No. 0036303), B. Y. Qiu 70194 (CDBI),
70085 (CDBI), 77617 (CDBI, KUN), 55291 (PE), 55718 (PE), G. R.
Wei et al. 86 (SWFC), H. B. Wang 91039 (SWFC), H. T. Chang 5876
(SYS), H. W. Li 9612 (YUKU), Hu s. n. (YUKU Herb. Bar Code No.
02034908), J. S. Yang 91–996 (IBSC, KUN), Q. Fan 8317 (SYS), Q. Xu
02122930 (SWFC), T. N. Liou 20196 (PE), 017177 (PE), 15468 (PE),
15309 (PE), 15238 (PE), 15615 (PE), 14156 (PE), H. S. Lo 172 (IBSC),
Z. Q. Yao et al. 13802 (YUKU). Lijiang, F. C. How 74412 (IBSC), 74263
(IBSC), H. T. Chang 1015 (IBSC), T. T. Yu 11093 (PE), X. L. Huang 5472
(SYS), Y. Tsiang 13038 (NAS). Longling, C. Y. Wu et al. 1232 (PE), J.
Chen 669 (KUN). Luquan, B. S. Sun 64215 (KUN), H. Li et al. 1100
(KUN), P. I. Mao 01465 (IBSC, PE), 719 (NAS), Y. B. Zhang 429 (IBSC).
Lushui, H. Li et al. 24150 (KUN), 24192 (KUN), 10244 (KUN), H. T.
Tsai 54536 (IBSC). Luxi, H. T. Tsai 54536 (IBSC, PE). Mengzi, C. W.
Wang 83525 (KUN). Shuangbai, anonymous 411 (KUN), W. Q. Yin 411
(IBSC). Songming, Yunnan Univ. Exped. 03025 (KUN), Y. B. Zhang 80
(IBSC). Weixi, C. W. Wang 70427 (KUN, PE), K. M. Feng 3859 (KUN,
PE), 8715 (PE), 3741 (PE). Wenshan, H. C. Wang 3676 (PE), H. T. Tsai
51496 (KUN, PE), 51620 (PE). Wuding, H. Li et al. 0925 (KUN), 0430
(KUN), 1146 (KUN). Xichou, C. W. Wang 85391 (KUN, PE). Yanshan,
C. W. Wang 84071 (KUN). Yangbi, R. C. Ching 25183 (KUN, PE). Yil-
iang, N. E. Yunnan Exped. 829 (KUN, YUKU). Yimen, anonymous s. n.
(YUKU Herb. Bar Code No. 02034910), Z. Z. Ding & J. X. Wang 1121
(NAS). Yingjiang, anonymous s. n. (KUN Herb. Bar Code No. 0675894).
Yongning, S. K. Wu 2936 (KUN). Zhaotong, K. M. Feng 73–368 (KUN),
73–369 (KUN). Zhenkang, Yunnan Univ. Exped. A1010 (KUN). Zhenx-
iong, Spicy Pl. Exped. 870470 (KUN). Zhongdian, W. M. Zhu et al. 17367
(YUKU).
Elaeagnus viridis and E. viridis var. delavayi were
problematic and their circumscriptions have been con-
fusing. In 1908, E. viridis was first collected by Henry
from Yichang, Hubei. Later Lecomte (1915) recognized
a new variety as E. viridis var. delavayi with descrip-
tion “Foliis ellipticis basi Paulo rotundatis vel attenu-
atis”. In Flora Reipublicae Popularis Sinicae, Chang
(1983) recorded both taxa and stated E. viridis was dis-
persed in south Shaanxi and west Hubei and E. viridis
var. delavayi only distributed in Yunnan, differing in
leaves and pedicel length. Du (2006) broadened the
circumscription of E. viridis var. delavayi by includ-
ing E. pallidiflora C. Y. Chang, E. wenshanensis C. Y.
Chang, and E. taliensis C. Y. Chang. However, Qin &
Gilbert (2007) treated E. viridis var. delavayi as a syn-
onym of E. viridis and explained that “Fruiting material
has been described as E. viridis var. delavayi but it is
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380 Journal of Systematics and Evolution Vol. 48 No. 5 2010
not possible to assess the status of such plants without
knowledge of comparable material of both var. viridis
and var. delavayi.”
After studying specimens from relevant regions
and investigation on Yunnan and Xizang, we agree with
the treatment of combining E. viridis var. delavayi and
E. viridis, as suggested by Qin & Gilbert (2007). We
were surprised to find that E. pallidiflora C. Y. Chang,
E. wenshanensis C. Y. Chang, E. taliensis C. Y. Chang,
E. xichouensis C. Y. Chang, and E. xizangensi C. Y.
Chang appear to form a natural group recognized here as
E. viridis. All of these species were described by Chang
(1980, 1981, 1985) from Yunnan except E. xizangensi
from Xizang. Their shared characters include oblong-
elliptic leaves with silvery white scales, tubular ca-
lyx, and long, slim pedicels. In summary, we defined
E. viridis more broadly. (1) This species is variable with
a vast geographical distribution range and its leaf blade
ranges from elliptic to oblong-elliptic and size from
small to bigger. Especially in limestone areas, its leaves
become leathery compared to papery on the holotype.
Its silvery white scales, broadly tubular calyx tubes,
and slim pedicels are stable and distinctive characters.
(2) We also update the distribution ranges of E. viridis
based on collections from Chongqing, Guizhou, Hubei,
Hunan, Shaanxi, Shanxi, Sichuan, Xizang, and Yunnan.
Previous authors may have overlooked its variation and
the morphological and geographical intermediates be-
tween E. viridis and E. viridis var. delavayi. We suggest
only one species E. viridis is recognized here.
Section II. Elaeagnus L. section Elaeagnus.(Serv.
in Beih. Bot. Centralbl. 25(1): 25. 1909; C. Y. Chang,
Fl. Reip. Pop. Sin. 52(2): 5. 1983, “Deciduae”).
According to Art. 22.1 of the International Code of
Botanical Nomenclature (Vienna Code) (McNeill et al.,
2006), the name of this section is an autonym. Hence
“Deciduae” described by Servettaz (1909) is actually
nomenclaturally superfluous, and should be rejected.
Deciduous or semi-evergreen tree or shrub, rarely
climbing. Leaf blade papery or membranous, rarely
leathery. Flowering in spring or summer. Fruiting in
summer or autumn. Fourteen species in this section
mostly distributed in Yangtze River valley and its south-
ern region.
Several species massively dominate in north
and northwest China and Elaeagnus umbellata
Thunb. widely spread in China excluding Helongjiang
Province. Generally this section is heterogeneity in
habit, habitat, life form, and morphological characters.
24. Elaeagnus angustifolia L., Sp. Pl. 1: 121. 1753.
Type: Clifford 38 Elaeagnus no. 1 (lectotype, designated
by McKean in Jarvis et al., Regnum Veg. 127: 44. 1993:
BM).
Elaeagnus angustifolia var. orientalis (L.) Kuntze
in Trudy Imp. S.-Peterburgsk. Bot. Sada 10: 235.
1887.——E. orientalis L. in Syst. Nat., ed. 12, 2: 127.
1753. Type: D. D. Royen s.n. (lectotype, designated
by Murray in Rechinger (ed.), Fl. Iranica 55: 3. 1968:
LINN).
Deciduous; trees, sometime shrubs, erect, to 10 m
tall in cultivation. Spines present; young branches
columned, densely with silvery scales, old branches
with reddish brown, somewhat pruinose bark. Petiole
5–20 mm; leaves papery; blade grey-green above, sil-
very grey below, lanceolate, sometimes elliptic, ovate,
or oblong-ovate, apex obtuse, margin entire, base usu-
ally broadly cuneate; 1–8 ×0.4–3.2 cm; adaxially with
white scales when young, then gradually glabrescent or
not, abaxially with overlapping silvery scales; lateral
veins inconspicuous on both surfaces. Flowers several
in a shortened raceme, rarely solitary, nodding; pedicels
2–4 mm. Flowers yellowish white; calyx campanulate
or broadly campanulate, with silvery scales outside; pa-
pery; 4–5 ×2–5 mm; lobes triangular, inside glabrous
or with sparse reddish brown scales, 3 ×2–4 mm; an-
thers oblong, ca. 0.5 mm; filaments short; style erect,
enclosed by tubular disks, glabrous; stigma exserted
beyond stamens. Fruit ellipsoid or subglobose, 0.7–
2.5 ×0.5–1.3 cm, densely scaly, yellowish brown when
ripe, mealy; ribs inconspicuous; fruit pedicels erect and
short, ca. 2 mm. Fl. May.–Jun., fr. Aug.–Oct.
Sea coasts, riversides, lakeshores, dry river beds,
and mountains; alt. 55–1800 m. Beijing, Gansu, Hebei,
Henan, Inner Mongolia, Liaoning, Ningxia, Qinghai,
Shaanxi, Shanxi, and Xinjiang.
Additional specimens examined:
China. Precise locality unknown, anonymous s. n. (KUN Herb.
Bar Code No. 0629691), P. C. Tsoong 8790 (KUN, PE). Beijing: Precise
locality unknown, 236–6 Exped. 310 (PE). Haidian, Botanical Garden,
Z. Y. Cao s. n. (PE), Yintaogou, Forestry Coll. 106 (PE). Shunyi, Bei-
jing Med. Pl. Exped. 300 (PE). Gansu: Precise locality unknown, L. S.
Chow 255, 294 (NAS), Lanzhou Univ. Bio. Dep. 1 (SZ), R. C. Ching
179 (IBSC), Y. C. Wu 153 (NAS), Yellow River Exped. 4969 (KUN),
6738 (NAS), 2910 (PE), 2955 (PE), 8778 (KUN, PE),Y. Y. Pai 243
(PE, SYS). Dunhuang, anonymous 0327 (PE). Ejin, T. N. Liou 2200
(PE), 2227 (PE). Gaolan, T. P. Wang 6338 (PE), Yellow River Exped.
2657 (PE). Huining, Yellow River Exped. 5166 (PE). Jingtai, Y. C. Hou
6225 (PE), 6297 (PE). Jiuquan, anonymous 1868 (PE), 1915 (PE), 1916
(PE). Lanzhou, P. C. Tsoong 8030 (PE), S. X. Zhen 0195 (PE). Lihe,
C. K. Chow 160 (NAS), 166 (NAS). Minle, Y. C. Hou 3232 (PE). Min-
qin, C. W. Yao 7 (N), Qinghai-Gansu Exped. 2945 (PE), 1450 (PE),
Y. C. Hou 3333 (PE). Yongjing, Yellow River Exped. 3735 (KUN, NAS,
PE). Hebei: Beidaihe, C. S. Wang 121 (PE), F. T. Wang 0342 (PE).
Cheng’an, Y. Y. Pai 231 (IBSC, PE). Xiaowutai, anonymous 1840154
(NF, SWFC). Henan: Kaifeng, T. Kanasiro 2902 (PE). Inner Mon-
golia: Precise locality unknown, anonymous 0001 (PE, YUKU). Alxa,
C. W. Yao 1015 (NAS), Y. C. Hou 2724 (PE). Bayanhot, Y. C. Hou
2568 (PE). Dengkou, anonymous Syz115 (SWFC). Huhhot, Z. H. Guo
GIB-0097 (SWFC). Linhe, Z. Shi 000121 (KUN). Wengping, W. Wang
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SUN & LIN: A revision of Elaeagnus in China 381
3364 (PE). Ulanhot, Y. C. Hou 7878 (PE). Ihju Meng, Neimenggu-
Ningxia Exped. 543 (PE). Liaoning: Precise locality unknown, E. H.
Wilson 8790 (IBSC). Dalian, C. Wang 860 (IBSC, PE), Y. F. Cheng
227 (NF). Gaizhou, T. N. Liou et al. 379 (PE). L¨
ushun, Y. Yabe s. n.
(NAS). Ningxia: Precise locality unknown, W. Y. Hsia 3876 (PE). Helan-
shan, Y. C. Hou 02329 (PE). Lingwu, Yellow River Exped. 7881 (PE),
7901 (PE). Pingluo, anonymous 253 (PE). Tongxin, Y. C. Hou 6576
(PE). Yinchuan, Y. C. Hou 2122 (PE), 7377 (PE). Zhongwei, Y. C. Hou
6449 (PE). Shaanxi: Dengkou, anonymous 30 (PE). Dingbian, J. M.
Liu 11595 (PE). Yulin, Yellow River Exped. 6738 (PE), 6792 (PE).
Xinjiang: Precise locality unknown, anonymous 31 (PE), D. Y. Hong
et al. 0003 (PE), 0002 (PE); on the way from Wen’aili to Jiashi, A. J.
Li 7551 (KUN, PE); Xinjiang Exped. 7881 (NAS), 9623 (KUN), 5904
(IBSC), 9369 (KUN); Poa 24 (NAS, PE), 47 (NAS), R. Z. Ma 12 (NAS),
S. Z. Qu s. n. (NF), Xinjiang Exped. 9293 (KUN, NAS), 9223 (KUN,
NAS), 9255 (KUN, NAS), 9604 (PE), 9623 (PE). Aksu, Xinjiang Exped.
8958 (KUN, PE). Anjihai, K. C. Kuan 1161 (IBSC, PE). Artux, A. J. Li
7511 (KUN, PE). Baicheng, A. J. Li et al. 8202 (KUN, PE). Baogdo Ula,
T. N. Liou 3668 (PE). Bole, Xinjiang Exped. 3950 (PE). Dihua, T. N. Liou
2817 (PE). Fuhai, Q. X. Liu & X. Y. Chen 87113 (N), X. Y. Chen 86189
(N). Hami, Luotuojuanzi, J. H. Zhang 87137 (PE), K. C. Kuan431 (IBSC,
PE), R. C. Ching 0109 (PE), Y. G. Man 002 (PE). Hetian, K. C. Kuan
033 (PE), 00105 (PE), 00130 (PE). Kashi, Taxi 0111 (PE), X. J. Ge 086
(IBSC), Xizang Exped. 3060 (PE), 3067 (PE). Korla, anonymous 0008
(PE), 0021 (PE), 0047 (PE), 0174 (PE), A. J. Li 5904 (KUN), Xinjiang
Exped. 430 (PE). Luntai, anonymous 0022 (PE), A. J. Li & J. R. Zhu 7550
(KUN), 8676 (PE). Manas, K. C. Kuan 808 (IBSC), R. S. Zhang & Z. X.
Hu 3126 (SZ). Minfeng, G. L. Chu & Y. W. Xu 7372 (IBSC, NAS, PE),
Xinjiang Exped. 9584 (PE), W. K. Han s. n. (KUN Herb. Bar Code No.
0629834). Minyu, Y. R. Lin 1357 (SYS). Qiemo, Xinjiang Exped. 9535
(KUN, PE). Sandaohe, K. C. Kuan 720 (IBSC). Shache, anonymous 011a
(PE), 031 (PE), 067 (PE), 068 (PE), 069 (PE), 070 (PE), 334 (PE), 337
(PE), 022 (SZ), 018 (SZ), D. Y. Li 83 (PE). Shawan, R. C. Ching 3808
(IBSC). Shanshan, A. J. Li 5421 (KUN, PE), 06721 (KUN, PE). Shihezi,
K. C. Kuan 654 (IBSC, PE), R. Y. Hu s. n. (SZ). Taikesishan, Xinjiang
Exped. 9212 (KUN, NAS), 9213 (KUN, NAS). Tieganli, Xizang Exped.
3018 (PE). Tuksun, anonymous 0049 (PE). Turpan, A. J. Li 5489 (KUN,
PE), 06629 (KUN, PE), T. Y. Cheo et al. 65142 (IBSC, KUN, NAS, SZ,
PE). Urumqi, Xinjiang Exped. 263 (PE), Y. G. Man 088 (PE), Y. R. Lin
74012 (IBSC, PE). Weili, A. J. Li & J. R. Zhu 8521 (NAS, KUN, PE),
8628 (KUN, PE), 8505 (KUN, PE), 8590 (PE), Q. R. Wang et al. 4184
(PE), Y. W. Tian 20478 (PE), 20480 (PE), 20481 (PE), 20482 (PE), 20483
(PE). Wenquan, X. Z. Zheng & M. X. Lai 80747 (GXMI). Xayar, A. J. Li
8772 (KUN, PE), 8773 (KUN, PE), 8774 (PE). Yanqi, Xinjiang Exped.
6883 (PE). Yengisar, anonymous 0092 (PE). Yili, C. Y. Yang & Younusi
741156 (PE), S. H. Yang 0107 (PE). Yining, T. Y. Cheo et al. 650350
(NAS). Yiwu, anonymous 2257 (KUN), K. C. Kuan 4982 (PE). Yumin,
Y. R. Lin 74–1356 (IBSC, PE), 74–1357 (PE).
Chang (1983) recorded that E. angustifolia var. ori-
entalis is distributed in northwest China. Characters he
used to separate E. angustifolia var. orientalis from var.
angustifolia are the leaves on flowering branches, flo-
ral disks, and fruit. Based on the study of Elaeagnus
in Xinjiang, Huang & Maimaitijiang (2005) suggested
leaves (shape and size) on flowering branches are good
characters to distinguish E. angustifolia var. orientalis
and var. angustifolia, supporting the status of E. an-
gustifolia var. orientalis as a variety. Our comprehen-
sive study on these two taxa from Xinjiang, Gansu,
Ningxia, and Inner Mongolia shows that leaves (shape
and size) on flowering branches successively range
from oblong-lanceolate to linear-lanceolate, sometimes
elliptic-lanceolate, ovate, or oblong-ovate (Fig. 7: A).
Moreover, it is a uniform character in this genus that
old leaves tend to be smaller, usually ovate or ellip-
tic (e.g., E. argyi). Thus characters from leaves only
may not be reliable. The dissection of fresh flowers of
E. angustifolia indicates that its floral disks are some-
times glabrous, but sometimes pubescent. We examined
the specimens Yellow River Exped. 6738 (NAS), Y. R.
Lin 74–1356 (IBSC, PE), W. Y. Hsia 3876 (PE), and
anonymous 253 (PE), which were identified by C. Y.
Chang as E. angustifolia var. orientalis. Our study sug-
gests that leaves under fruits (flowering branches) of
these specimens are broadly ovate, but their fruits are
smaller, generally not more than 1 cm (diameter). Al-
though there are some collections of E. angustifolia var.
orientalis with fruits 2.5 cm long (A.J.Li&J.R.Zhu
8958 (KUN, PE), 6721 (KUN, PE), 7551 (KUN, PE),
8628 (KUN, PE), and 7511 (KUN, PE)), notes on these
specimens show that they were all collected from river-
side and edges of fertile farmland. It seems that habitat
with sufficient water and fertile soil contribute greatly to
bigger fruits. Using the data from measurement of fruits
of both taxa, statistical analysis suggests that E. angus-
tifolia var. orientalis can not be separated from E. an-
gustifolia in fruit size (Fig. 7: B). Based on the observa-
tions above, we do not think it is still reasonable to treat
E. angustifolia var. orientalis as a variety. Therefore, we
treat E. angustifolia var. orientalis as a synonym of E.
angustifolia.
26. Elaeagnus argyi L´
eveill´
e in Fedde, Repert. Spec.
Nov. Regni Veg. 12: 101. 1913. Type: China. Jiangsu:
Zuo-Se. Pou-Si. Montagnes, d’Argy (holotype, B).
Elaeagnus micrantha C. Y. Chang in Bull. Bot.
Lab. N. E. Forest. Inst., Harbin 1980(6): 116. 1980.
Type: China. Yunnan: Songming, P. Y. Chu & W. X. Liu
51182 (holotype, KUN!; isotypes, KUN!), Fig. 8: A.
Semi-evergreen; shrubs, erect, 2–3 m tall. Spines
usually present; young branches columned, with dense
yellowish white scales. Petioles yellowish brown, 5–
7 mm; leaves papery; blade grey-green above, silvery
grey below; leaf blades dimorphic by season, spring
leaves small, elliptic to oblong, apex rounded or obtuse,
margin entire, base obtuse, 1–4 ×0.8–2 cm; autumn
leaves larger, obovate, 6–10 ×3–5 cm; adaxially with
yellowish brown stellate when young, abaxially with
white and brown overlapping scales; lateral veins 8–
10 per side of midrib, deeply impressed on above sur-
face of old leaves. Flowers often 5–7 in a fascicle at
base of young branches with new growth, rarely soli-
tary; pedicels ca. 3 mm, slender. Flowers yellow; ca-
lyx funnelform-tubular, with silvery and rust-colored
scales outside; sub-leathery; 5.5–6 ×2–5 mm; lobes
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382 Journal of Systematics and Evolution Vol. 48 No. 5 2010
Fig. 7. Results of statistical analysis showing the variation range of Elaeagnus angustifolia and Elaeagnus angustifolia var. orientalis.A, Leaf size.
B, Fruit size.
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SUN & LIN: A revision of Elaeagnus in China 383
Fig. 8. A, Holotype of Elaeagnus micrantha C. Y. Chang (P. Y. Chu & W. X. Liu 51182, KUN). Aa, Flora buds. B, Holotype of Elaeagnus grijsii Hance
(Grijs s. n. =Herb. Hance no. 6686, K). C, Holotype of Elaeagnus jiangxiensis C. Y. Chang (236 Exped. 491,PE).Ba–Cb, Long-stalked stellate hairs
present on branches of the holotypes of E. grijsii and E. jiangxiensis.Ba, Bb, Long-stalked stellate hairs of branches of E. grijsii (Y. Ling 4472,PE).
Ca, Cb, Long-stalked stellate hairs of branches of E. jiangxiensis (S. S. Lai 0180,PE).Ba, Ca, Young branches. Bb, Cb, Old branches. D, Isolectotype
of Elaeagnus angustata var. songmingensis W. K. Hu & H. F. Chow (P. Y. Chu 51495,PE).E, Lectotype of Elaeagnus umbellata subsp. magna Serv.
(Henry A. 1637, designated here, K).
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384 Journal of Systematics and Evolution Vol. 48 No. 5 2010
ovate-triangular, apex acute, inside with sparse short
villi and scales, 2–3 ×1–2 mm; anthers ellipsoid, ca.
1.2 mm; filaments very short; style erect, glabrous;
stigma beyond stamens. Fruit obovoid-oblong, 1.3–
1.6 ×ca. 0.6 cm, densely silvery scales, and red when
ripe, juicy; ribs conspicuous; fruiting pedicels, slender,
5–10 mm. Fl. Jan.–Mar., fr. Apr.–May.
Streamsides, roadsides, open slopes, and forests,
also cultivated; alt. 100–2250 m. Endemic to China. An-
hui, Guizhou, Henan, Hubei, Hunan, Jiangsu, Jiangxi,
Yunnan, and Zhejiang.
Additional specimens examined:
China. Anhui: Precise locality unknown, anonymous 10896
(NAS), 17164 (NAS), 7688 (NAS), R. C. Ching 2714 (IBSC, PE). An-
qing, H. Migo s. n. (NAS). Chuzhou, W. C. Cheng 5298 (N, PE). Guangde,
R. H. Shan 3377 (NAS). Langyashan, M. B. Deng 3038 (NAS). Xiaoxian,
C. C. Yuan & J. S. Yue 2091 (NAS). Xiuzhou, W. C. Cheng 5298 (SZ).
Guizhou: Weining, anonymous 74–1009 (HGAS). Henan: Shangcheng,
Henan Pl. Exped. 0051 (PE). Hubei: Precise locality unknown, Y. H.
Zhang 4, 111 (PE). Huangpo, Hubei Pl. Exped. D01502 (PE). Jinmen,
H. Migo s. n. (NAS). Macheng, Y. J. Ma 4300 (SZ). Wuchang, C. H.
Tsoong 81879 (IBSC), H. Migo s. n. (NAS), S. W. Teng 257 (NAS),
Wuhan Univ. Campus, S. Y. He 150214 (IBSC). Hunan: Precise local-
ity unknown, anonymous 24 (PE). Jiangsu: Precise locality unknown,
anonymous 1050 (NAS), 1828 (NAS), 2063 (NAS), T. Y. Cheo et al. 115
(N), W. X. Wu 3187 (N), X. Y. He 3375 (IBSC), Z. K. Wei 22820 (NAS),
32018 (NAS). Jiangning, Nanjing Forestry Exped. 6188 (SZ). Jiangpu,
M. Chen 90 (IBSC, PE), 1470 (PE). Jurong, M. B. Deng 3451 (KUN,
NAS, PE). Nanjing, C. L. Tso 944 (NAS, PE), C. N. Chen 8788 (NAS),
K. H. Cheo s. n. (NF Herb. Bar Code No.10900011), K. Ling 1387 (N),
Y. N. Xiong 150 (NAS), Z. Y. Huang & X. Cheng 310 (NAS). Sheshan,
H. Migo s. n. (NAS). Yixing, C. Y. Luh 439 (IBSC, NAS), K. Ling 2604
(NF), Q. W. Yao s. n. (NF Herb. Bar Code No.10900016). Jiangxi: Gul-
ing, C. D. Reeves s. n. (NAS). Jiujiang, C. M. Tan 951334 (IBSC, KUN,
NAS, PE), H. Migo s. n. (NAS), Y. G. Xiong 10092 (NAS, PE), 10075
(NAS, PE), 980(NAS), 10007 (PE). Luxi, W. X. Wu 9515 (NAS). Yun-
nan: Dˆ
eqˆ
en, C. C. Lu 63214 (KUN, PE). Songming, B. Y. Qiu 51637
(IBSC, PE), B. Y. Qiu & W. X. Liu 51182 (IBSC, KUN, PE). Weixi,
K. M. Feng 3595 (KUN). Zhejiang: R. C. Ching 3375 (IBSC). Anji,
Z. X. Zhang s. n. (NF). Hangzhou, anonymous 1921 (HHBG, PE), 1882
(HHBG), 2022 (HHBG), 2023 (HHBG), 1899 (HHBG), 1863 (HHBG),
2022 (KUN, PE), 880 (NAS), 450 (PE), 12 (PE, SZ), 1882 (SZ), 119 (SZ),
84 (SZ), 005 (SZ), 64 (SZ), 13 (SZ), 2 (SZ), A. D. Merrill 30 (NAS), C. Y.
Chang 7 (SZ), C. Y. Chiao 1913 (N), S. Y. Chang 089 (HHBG, NAS,
PE), 441 (HHBG, NAS, PE), W. K. Hu 2 (SZ). Jiangning, anonymous
s. n. (HHBG). Tianmushan, anonymous 190 (PE), H. Migo s. n. (NAS),
T. Shen 117 (NAS, N), 488 (NAS, N), P. C. Tsoong 5 (PE), X. Y. He
1183 (HHBG), 21447 (HHBG, NAS, PE), 506 (HHBG, IBSC, NAS),
West Lake Museum 61 (NAS). Wenling, HHBG 4187 (PE).
Types of Elaeagnus micrantha bear young flow-
ers. Chang remarked that E. micrantha was close to
E. guizhouensis differing in leaf shape and size, quality
of calyx, and length of pedicel. After carefully examin-
ing the holotype of E. micrantha, we found that the only
key character which Chang used to separate E. micran-
tha from E. argyi, is length of calyx tube. Calyx tube of
E. micrantha is shorter than that of E. argyi. The flowers
of E. micrantha are only from holotype specimens, and
they were actually buds (Fig. 8: A, Aa). The specimens
R.C. Ching 2741 (KUN), C. M. Tan 951334 (IBSC,
KUN, NAS, PE), and anonymous 76–1009 (HGAS) also
show that the calyx tube of E. argyi tends to be shorter
in east China compared to southwest China (Yunnan,
type locality of E. micrantha). Likewise, leaves are be-
coming membranous. It seems that the species status
of E. micrantha is poorly supported. In addition, one
specimen (B. Y. Qiu 51637 (IBSC, PE)) determined by
Chang to be E. micrantha is actually E. argyi. Hence
it would be more appropriate to combine E. micrantha
and E. argyi.
30. Elaeagnus grijsii Hance in Ann. Sci. Nat., Bot., s´
er.
4. 15: 227. 1861. Type: China. Fujian: 1861, Grijs s. n.
(=Herb. Hance no. 6686) (holotype, K!), Fig. 8: B.
Elaeagnus jiangxiensis C. Y. Chang in Bull. Bot.
Lab. N. E. Forest. Inst., Harbin 1980(6): 118. 1980.
Type: China. Jiangxi: Suichuan, 236 Exped. 491 (holo-
type, PE!; isotype, PE!), Fig. 8: C.
Semi-evergreen; shrubs, climbing, sometimes
erect, 1–3 m tall. Spines present; young branches
columned, with dense yellowish brown long-stalked
stellate hairs. Petioles yellowish brown, 4–7 mm; leaves
sub-leathery; blade green above, rust-colored below;
blade ovate or broadly ovate, apex bluntly acuminate,
margin entire, base obtuse; 2.7–7 ×1.8–3.8 cm; adax-
ially with yellowish brown long-stalked stellate hairs
when young, then gradually glabrescent or not, abaxially
with dense long-stalked stellate hairs; lateral veins 3–5
per side of midrib, deeply impressed on above surface of
old leaves. Flowers 7–10 in axils, forming a shortened
subumbellate raceme; pedicels 2–3 mm, slender. Flow-
ers yellow; calyx campanulate, with yellowish brown
and rust-colored stellate outside; papery; 6–6.5–6 ×2–
5 mm; lobes ovate-triangular, apex acute, inside with
sparse stellate or glabrous, 2–3 ×1–2 mm; anthers ob-
long, ca. 1.5 mm; filaments very short; style erect, stel-
late or glabrous; stigma not longer than stamens. Fruit
narrowly ellipsoid, ca. 1.8 ×0.6 cm, densely yellowish
brown scales, and red when ripe, juicy; ribs conspicu-
ous; fruiting pedicels, thick, 1–2.5 cm. Fl. Jan.–Feb., fr.
Apr.–May.
Shrublands on south-facing mountain slopes; alt.
600–800 m. Endemic to China. Fujian, Guizhou, and
Jiangxi.
Additional specimens examined:
China. Fujian: Precise locality unknown, anonymous 6686
(IBSC). Anxi, Fujian Exped. 2208 (NAS). Hua’an, W. D. Han 20993
(NF). Zhangping, Y. Ling 4472 (PE). Guizhou: Yanhe, N. Guizhou Ex-
ped. 2319 (HGAS). Jiangxi: Suichuan, 236 Exped. 491 (PE), S. S. Lai
0180 (PE).
Elaeagnus jiangxiensis was recognized by Chang
(1980) based on two sheets of young fruiting specimens
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SUN & LIN: A revision of Elaeagnus in China 385
(236 Exped. 491, PE) without information of flowers,
but he pointed out that this new species was similar to
E. guizhouensis differing in flower and pedicel morpho-
logy. Although we can separate Elaeagnus jiangxiensis
from E. guizhouensis by pedicels and hairs on branches,
the circumscriptions between E. jiangxiensis and E. gri-
jsii seem unclear. Lacking of flowering type specimens
indicate E. jiangxiensis was poorly established and it
seems difficult to separate E. jiangxiensis from E. grijsii.
In contrast, the combination of E. grijsii and E. jiangx-
iensis provides a distinctive circumscription well char-
acterized by ovate leaves, long pedicels and branches,
leaves, and flowers with dense long-stalked stellate hairs
(Fig. 8: Ba–Cb). The species is widely distributed in
Jiangxi (type locality of E. jiangxiensis) and Fujian
provinces.
33. Elaeagnus angustata (Rehd.) C. Y. Chang in Bull.
Bot. Lab. N. E. Forest. Inst., Harbin 1980(6): 121.
1980.——E. multiflora Thunb. f. angustata Rehd. in
Sarg., Pl. Wils. 2: 413. 1915. Type: China. Sichuan:
Kangding, Tachien-Lu (=Dajianlu), E. H. Wilson 1076
(holotype, GH).
Elaeagnus angustata (Rehd.) C. Y. Chang var.
songmingensis W. K. Hu & H. F. Chow in Bull. Bot.
Lab. N. E. Forest. Inst., Harbin 1980(6): 121. 1980.
Type: China. Yunnan: Songming, B. Y. Qiu 51495 (lec-
totype, designated by Q. Lin in Bull. Bot. Res., Harbin
26(6): 656. 2006: SZ!), Fig. 8: D.
Deciduous, shrubs, erect, 1.5–4 m tall. Spines ab-
sent; young branches up-ridge, densely with ferruginous
scales. Petiole rust-colored, 3–5 mm; leaves papery or
membranous; blade light green above, silvery below,
elliptic as young, later becoming oblong or elliptic,
apex acuminate, margin entire, slightly undulate, base
rounded or slightly obtuse; 3–10 ×1.2–2.2 cm; adaxi-
ally with white stellate hairs or deeply lacerate scales or
overlapping shallowly bowl-shaped scales when young,
then gradually glabrescent or not, abaxially with over-
lapping silvery scales, scales mostly white, a few pale
brown; lateral veins 7–10 per side of midrib, slightly
raised on both surfaces. Flowers 1–3 in a shortened
raceme, nodding; pedicels brown, 5–8 mm. Flowers
light white; calyx tubular, with yellowish silvery scales
outside; papery; 4–5 ×3–4 mm; lobes broadly ovate, sil-
very scales inside, 1–3 mm; anthers oblong, ca. 1.5 mm;
filaments ca. 0.6 mm; style erect, with dense stellate
hairs; stigma exserted beyond stamens. Fruit ellipsoid,
ca. 1.4 ×0.5–0.8 cm, densely scaly, red when ripe, juicy;
ribs on seed inconspicuous; fruit pedicels nodding, 1.5–
3 cm. Fl. Apr.–May., fr. Jul.–Aug.
Forest streams and montane scrublands; alt. 580–
3400 m. Endemic to China. Hubei, Sichuan, and
Yunnan.
Additional specimens examined:
China. Precise locality unknown, S. K. Wu 2035 (KUN), T. N.
Liou 7398 (SZ). Hunan: Hunan Exped. 291 (IBSC). Sichuan: Pre-
cise locality unknown, K. L. Chu 7023 (NAS). Baoxing, T. P. Soong
38953 (IBSC, KUN, SZ), 38831(IBSC, PE, SZ), 35670 (IBSC), T. T.
Yu 1980 (PE), 1986 (PE). Hongya, W. K. Bao et al. 2637 (CDBI), 2345
(CDBI), 1882 (CDBI). Huidong, S. K. Wu 1094 (KUN, SZ), s. n. (KUN
0622228). Jinyang, Sichuan Econ. Pl. Exped. 3026 (PE, SZ). Kangding,
anonymous 6228 (CDBI, PE), C. S. Cao et al. 111043 (CDBI, PE, SZ),
119156 (CDBI, SZ), G. Hu et al. 111998 (SZ), H. L. Tsiang 35670
(IBK, IBSC, PE, SWFC, SZ), 36441 (IBK, IBSC, PE, SZ), Z. J. Zhao
et al. 113934 (SZ), Z. P. Huang et al. 1771 (IBSC, PE), Z. Y. Chen
& Z. X. Xiong 112140 (SZ). Leibo, Dept. Biol. Univ. Sichuan 110570
(SZ). Luding, G. R. Xu 23263 (IBSC), 35670 (IBSC), Vegetation Group
31564 (CDBI). Mianning, S. K. Wu 2035 (PE, SZ). Muli, S. Jiang 5689
(PE, SZ). Shimian, C. J. Xie s. n. (HHBG), 40121 (IBSC, PE, SZ),
426190 (IBSC, PE), 41090 (IBSC, PE, SZ), 41276 (IBSC, PE, SZ).
Yunnan: Dongchuan, N. E. Yunnan Exped. 583 (KUN). Guodong, G. R.
Yao et al. 24 (SWFC). Lanping, Beijing Exped. 01187 (PE). Luquan,
W. M. Zhu et al. 00292 (YUKU). Songming, B. Y. Qiu 51495 (IBSC,
PE), 35670 (IBSC), B. Y. Qiu & P. H. Yu 98 (IBSC, PE, KUN), 51637
(KUN), 51495 (SZ, PE), 54135 (PE, KUN), J. G. Wang et al. 55 (YUKU),
W. M. Zhu et al. 03038 (YUKU).
Elaeagnus angustata (Rehd.) C. Y. Chang var.
songmingensis W. K. Hu & H. F. Chow was pub-
lished by Chang (1980) with the description “A typo
recedit ramulis dense argeno et luteo-albo-lepidotis;
foliis supra argentec-lepidotis et squamis luteis con-
speris; pedicellis tubis calycis ovarium includente dense
albo-lepidotis”, and he also stated that young branches
of E. angustata var. songmingensis are white or yellow-
ish white, its leaves adaxially with white and few yellow
scales, and calyx tube and pedicel densely with silvery
scales. These characters can be used to distinguish E. an-
gustata var. songmingensis from E. angustata. However,
these specimens, including anonymous 6228 (CDBI,
PE), Z. P. Huang et al. 1771 (IBSC, PE), C. J. Xie 40121
(IBSC, PE, SZ), 426190 (IBSC, PE), 41090 (IBSC, PE,
SZ), 41276 (IBSC, PE, SZ), S. K. Wu 2035 (PE, SZ),
and T. P. Soong 38831 (IBSC, PE, SZ), identified by
Chang as E. angustata, suggest that E. angustata also
shares the characters such as yellowish white and rust-
colored scales on young branches, leaves, calyx tubes,
and pedicels. Furthermore, field investigation suggests
that white and rust-colored scales or stellate hairs are
not exclusively present, but coexist. Another concern
is that all the specimens under the name E. angus-
tata var. songmingensis are flowering specimens, and
its fruits are unknown. It seems that the characters the
authors used appear insufficient to distinguish E. an-
gustata var. songmingensis from E. angustata, and we
thus include the former in a broader circumscription of
E. angustata.
35. Elaeagnus magna (Serv.) Rehd. in Sargent, Pl. Wil-
son. 2(2): 411. 1915.——E. umbellata Thunb. subsp.
magna Serv. in Bull. Herb. Boissier, s´
er. 2, 8: 383. 1908
et in Beih. Bot. Centralbl. 25(1): 58. 1909. Type: China.
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386 Journal of Systematics and Evolution Vol. 48 No. 5 2010
Hubei: Yichang, Henry A. 1637 (lectotype, designated
here, K!; isolectotype, P) (as “1437”, on p. 58, err. ty-
pogr.), Fig. 8: E.
Servettaz (1908) described E. umbellata Thunb.
subsp. magna as a new subspecies without type ci-
tation. Later he made this name validly published by
designating Henry A. 1437 as types housed at K and
P, respectively (Servettaz, 1909). Rehder (1915) recog-
nized it as E. magna and pointed out that the type of
E. umbellata Thunb. subsp. magna,Henry A. 1637 was
mistakenly quoted by Servettaz as Henry A. 1437.How-
ever, Rehder did not realize that the type designated by
Servettaz contained at least two gatherings separately
deposited in K and P. The lectotypification of this name
is therefore needed. According to Art. 8, Art. 9, and Art.
37 of the International Code of Botanical Nomencla-
ture (Vienna Code) (McNeill et al., 2006), we desig-
nate the specimen Henry A. 1637 deposited in K as the
lectotype.
Key to infraspecific taxa of E. magna
1a. Young branches, leaves and fruit densely silvery
scales . . . . . . . . . . . . . . . . . . . . . . . . . . . var. magna
1b. Young branches, leaves and fruit densely or
sparsely rust-colored scales . . . . . . . . . . . . . . . . . . 2
2a. Spines thick and short. Young branches and leaves
densely rust-colored scales. Fruiting pedicel 3–
9mm.....................var.nanchuanensis
2b. Spines sparse and small. Young branches and
leaves densely yellowish white scales. Fruiting
pedicel 8–16 mm . . . . . . . . . . . . .var. wushanensis
35a. var. magna
Deciduous; shrubs, erect, to 5 m tall. Spines
present; young branches up-ridged, densely with silvery
scales. Petiole 4–10 mm; leaves papery; blade green
above, silvery gray below, ovate or oblong-ovate, apex
bluntly acute to acuminate, margin entire, base usu-
ally cuneate; 4–11 ×1.5–4.2 cm; adaxially with dense
silvery scales when young, then gradually glabrescent
or not, abaxially with overlapping silvery and some-
times scattered yellowish scales; lateral veins 7–10 per
side of midrib, inconspicuous on both surfaces. Flow-
ers 1–3 at base of young branches with new growth
or more than 3 in axils, forming a shortened subum-
bellate raceme; pedicels very short or absent. Flow-
ers yellowish white; calyx tubular, with silvery scales
outside; papery; 5–13 ×2–3 mm; lobes ovate, inside
with sparse stellates, 3–4 ×2 mm; anthers oblong,
ca. 2 mm; filaments very short; style curved, stellate
or glabrous; stigma beyond stamens. Fruit ellipsoid to
fusiform, 1.2–2 ×0.5–1.3 cm, densely scaly, pink-red
when ripe, juicy; ribs conspicuous; fruiting pedicels
erect, ca. 5 mm, or nearly absent, thick. Fl. Apr.–May., fr.
Jun.
Sandy soil in mountains, roadsides, forest margins,
and river banks; alt. 100–2300 m. Endemic to China.
Chongqing, Guangdong, Guangxi, Guizhou, Hubei, Hu-
nan, Jiangxi, Shaanxi, and Sichuan.
Additional specimens examined:
China. Precise locality unknown, anonymous s. n. (KUN Herb.
Bar Code No. 0629821), 3200 (KUN), 50066 (IBK), 50070 (IBK).
Chongqing: Chengkou, T. L. Dai 100517 (SZ), 101373 (CDBI, SZ),
100823 (CDBI, SZ). Fengdu, J. A. Wang et al. 078 (CDBI). Fengjie, C. Y.
Chang 25229 (IBSC, NAS, PE, SZ), 25220 (KUN), 25258 (SZ), H. F.
Zhou et al. 107995 (IBSC, NAS, PE, SZ), 107782 (IBSC, NAS, PE, SZ),
107828 (IBSC, NAS, PE, SZ), 26531 (IBSC, NAS, PE, SZ), 108082,
(IBSC, SZ). Kaixing, T. L. Dai 101763 (SZ). Nanchuan, K. C. Kuan
1046 (CDBI), 1238 (CDBI). Shizhu, Y. Chen 021 (CDBI), 2633 (CDBI,
SZ), 2562 (CDBI, SZ). Wushan, anonymous 58975 (CDBI). Wuxi, G. H.
Yang 65389 (CDBI), 59471 (CDBI, IBSC). Guangdong: Lechang, Nan-
ling Exped. s. n. (IBSC Herb. Bar Code No. 0386932). Ruyuan, Y. R.
Lin et al. 8 (IBSC), 331 (IBSC), YUE-73 1269 (HGAS, IBSC, PE), S. P.
Ko 52676 (IBK, IBSC, KUN, NAS, PE, SZ), 53939 (IBK, IBSC). Yang-
shan, L. Deng 1559 (IBSC). Guangxi: Donglan, Donglan Exped. 179
(GXMI). Leye, Z. F. Wei & D. Fang 3–5738a (GXMI). Luzhai, Y. M.
Wang 290 (PE). Quanzhou, W. T. Tsang 27681 (IBSC). Guizhou: Precise
locality unknown, anonymous 82–84 (IBSC). Songtao, anonymous 75–
1442 (HGAS), Z. F. Liu & T. P. Zhu 1556 (KUN, HGAS), 1545 (KUN,
NAS, HGAS). Tongren, Wuling Pl. Exped. 1416 (KUN). Weining, W.
Guizhou Exped. 23 (HGAS). Hubei: Precise locality unknown, anony-
mous 1143 (NAS), T. P. Wang 11545 (IBSC, PE), W. Y. Chun 3902 (PE),
Y. Tsiang 5916 (NAS). Badong, T. P. Wang 10959 (IBSC, PE), 10951
(KUN). Hefeng, F. S. Peng 78 (PE), H. J. Li 408 (IBSC, PE, SZ), 395
(IBSC, PE, SZ), 6230 (IBSC, PE, SZ), 8242 (NAS, PE), 8243 (NAS,
PE), 8264 (NAS, PE), 8267 (NAS, PE), 6670 (PE, SZ), 3200 (PE, SZ).
Lichuan, W. B. Lin 450 (PE). Jianshi, C. R. Wang 437 (PE). Wuhan,
anonymous 155 (PE). Xingshan, H. J. Li 395 (PE), 408 (PE). Xuan’en,
H. J. Li 3200 (PE). Yichang, anonymous 4059 (PE), Z. Zheng 16 (PE),
122 (PE). Hunan: Precise locality unknown, P. C. Tam 63112 (IBK),
Q. Z. Lin 84092 (SWFC). Chengbu, F. J. Huang 0499(PE), Z. T. Li 1813
(IBSC, PE). Cili, G. X. Zhu 030 (IBK, IBSC). Dong’an, J. K. Liu 040
(PE), 399 (PE). Hengshan, Bio. Dept. Hunan Univ. 232 (NAS), Z. H. Hu
007 (PE). Hongjiang, Z. T. Li 2376 (IBSC, PE). Huaihua, B. Liu 0038
(PE). Jianghua, Y. K. Li 401314 (IBK). Longshan, L. H. Liu 1938 (KUN,
NAS, PE). Nanyue, L. H. Liu 015742 (PE), Y. Q. Kuang 719 (PE), 1154
(PE), Z. H. Hu 152 (PE). Ningyuan, P. C. Tsoong 921 (PE). Qidong, J. D.
Li 268 (PE). Sangzhi, anonymous 420 (PE), 84079 (KUN), Q. Lin 647
(IBSC), Sangzhi Forestry Institute 1041 (KUN). Shaodong, J.H. Liu 7100
(PE), 7246 (PE). Shaoyang, L. D. Duan 5029 (PE), 2318 (PE), 0668 (PE).
Shimen, Hupingshan Exped. 1177 (PE), 1167 (PE), 0993 (PE), 0634 (PE),
0034 (PE). W. Hunan, L. H. Liu 1938 (IBSC). Xinning, anonymous 164
(PE), L. B. Luo 517 (PE), Q. Z. Lin 10101 (IBSC), Z. C. Luo 101 (PE),
Ziyunshan Exped. 1098 (PE). Yangmingshan, Y. Tsiang et al. 386 (IBK,
IBSC). Yizhang, S. H. Chun 1188 (IBK, IBSC), P. H. Liang 83099 (IBK,
IBSC). Yongshun, Beijing Exped. 01104 (KUN). Yuanling, anonymous
059 (IBSC), F. J. Zhou 119 (PE), Wulingshan Pl. Exped. 471 (IBSC).
Zhijiang, P. C. Tam 62498 (IBK), Wulingshan Pl. Exped. 2200 (IBSC).
Jiangxi: Jiangxi Exped. 0576 (PE). Anfu, Jiangxi Exped. 00189, 195
(PE). Fengyi, X. P. Ding 0063 (PE). Wugongshan, Jiangxi Exped. (PE).
Yifeng, S. S. Lai & D. F. Huang 000410 (PE). Shaanxi: T. L. Dai 100517
(SZ). Guang’an, Q. S. Cao et al. 161120 (SZ). Nanzheng, X. X. Hou 294,
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SUN & LIN: A revision of Elaeagnus in China 387
304 (IBSC). Sichuan: Chengdu, X. Q. Li 002 (CDBI). Hongya, W. K.
Bao et al. 3069 (CDBI), Z. W. Wang 00405 (CDBI). Leibo, M. Y. He &
Z. L. Tan 116985 (CDBI, SZ), Q. S. Cao et al. 117264 (CDBI). Pengshui,
Y. L. Cao et al. 0159 (CDBI). Wulong, Y. L. Cao et al. 0463 (CDBI).
Xingwen, C. Y. Chang s. n. (SZ).
35b. Elaeagnus magna (Serv.) Rehd. var. nanchuanen-
sis (C. Y. Chang) M. Sun & Q. Lin, comb. nov. & stat.
nov.——E. nanchuanensis C. Y. Chang, Fl. Sichuan. 1:
464. 1981. Type: China. Chongqing: Nanchuan, C. H.
Hsiung & C. L. Chow 90645 (holotype, SZ!; isotypes,
IBSC!; KUN!; PE!), Fig. 9: A.
Spines thick and short; young branches with rust-
colored scales. Leaf blade elliptic to lanceolate; apex
obtuse to acuminate, base obtuse to cuneate, abaxially
with rust-colored scales. Flowers yellowish green, 5–
6 mm, with dense rust-colored scales. Fruit with dense
rust-colored scales.
Streamside, roadsides and forests; alt. 750–1800 m.
Endemic to China. Chongqing, Gansu, Guangxi,
Guizhou, Hubei, Sichuan, and Yunnan.
Additional specimens examined:
China. Precise locality unknown, anonymous 1960 (SZ), C. W. Yao
3897 (NAS, PE), Z. X. Zhao 836 (KUN). Chongqing: Fengdu,J. A. Wang
& Y. X. Wang 078 (SZ). Fengjie, M. Y. Fang 24257 (KUN). Hechuan,
X. L. Sun 5452 (NAS). Nanchuan, G. F. Li 61091 (IBSC, KUN, NAS, PE,
SZ), 61414 (IBSC, KUN, NAS, PE, SZ), Jinfoshan Exped. 0625 (PE),
0684 (PE), 1249 (PE), 1733 (PE), J. H. Xiong & Z. L. Zhou 90645 (IBSC,
KUN, PE, SZ), 90085 (PE), K. C. Kuan et al. 1046 (PE), 1238 (PE).
Wulong, S. R. Yi etal. 0137 (PE). Wuxi, P. Y. Li 3365 (KUN), G. H. Yang
65389 (KUN), 59471 (KUN), 59461 (KUN, PE). Youyang, Fuling Exped.
02337 (PE, SZ). Zhongxian, D. S. Xiang 1363 (KUN, PE, SZ). Gansu:
Wenxian, Baishuijiang Exped. 4323 (PE), 4336 (PE). Guangxi: Leye,
Hongshuihe Pl. Exped. 1112 (KUN), 1129 (KUN). Guizhou: Y. Tsiang
5102 (IBSC, NAS, PE), 5107 (IBSC), S. S. Sin 50066 (IBSC), 50070
(IBSC). Zheng’an, anonymous 3332 (PE). Zunyi, N. Guizhou Exped.
0137 (PE). Hubei: Precise locality unknown, T. P. Wang 11543 (KUN).
Sichuan: Precise locality unknown, anonymous 3332 (IBSC), 1563 (SZ),
T. T. Yu 5452 (IBSC), W. P. Fang 1803 (NAS), Z. C. Zhong 0092 (SZ).
Chengdu, K. C. Kuan et al. 330 (PE). Ebian, Z. X. Zhao 1072 (KUN).
Emei, S. L. Sun 764 (KUN). Dujiangyan, F. T. Wang 20782 (KUN,
NAS), Z. He 12220 (IBSC, PE, SZ). Fuling, Sichuan Econ. Pl. Exped.
2685 (SZ). Hongya, X. S. Chang 03059 (SZ). Jiangjin, Sichuan Econ. Pl.
Exped. 508 (KUN, PE). Leibo, Dept. Biol. Univ. Sichuan 110303 (SZ),
110413 (SZ), 110455 (SZ), 110256 (SZ), 110299 (SZ), C. S. Cao 117264
(SZ), Taxonomic Improment Class 76–089 (SZ). Longquan, M. Y. He &
H. F. Zhou 1 (SZ), 2 (SZ), 10 (SZ), W. K. Hu 30 (SZ), 51 (SZ), 66 (SZ),
69 (SZ), 81 (SZ), 85 (SZ). Nanjiang, Z. X. Li 2794 (KUN). Nanping, J. H.
Xiong et al. 90085 (IBSC). Pingshan, F. T. Wang 22435 (KUN, NAS).
Tianquan, D. Y. Peng 45608 (IBSC), Sichuan Agr. Coll. 00400 (KUN.
PE). Wanyuan, B. L. Li 2046 (KUN), 2008 (KUN). Yunnan: Suijiang,
B. S. Sun et al. 0118 (IBSC, KUN).
Chang (1981) considered E. nanchuanensis as a
new species closely related to E. magna, but differing
in the following characters: (i) brown or rust-colored
young branches; (ii) brown or light yellow flowers; (iii)
calyx tube 5–6 mm long; and (iv) fruit brown scaly.
Our field observation and herbarium study suggest
that the characters Chang used do not appear constant
enough to justify the species rank. (1) Young branches
of E. magna are also brown or rust-colored (e.g., Z. T.
Li 2376 (IBSC, PE), W. Y. Chun 3902 (PE), S. S. Lai
& D. F. Huang 000410 (PE)). (2) Leaves of E. magna
abaxially with silvery scales, but E. nanchuanensis with
rust-colored ones. (3) Flower color of E. magna is firstly
white, but gradually ranges from white to light yellow
to yellow until withered (e.g., H. F. Chow & H. Y. Su
108082 (IBSC, SZ), L. D. Duan 0668 (PE)). (4) Length
of calyx tube of E. magna is 5–13 mm, include the
length magnitude of E. nanchuanensis (5–6 mm). (5)
Fruit of E. magna is covered by silvery scales, dotted
with ferruginous scales, sometimes even more. Under
this circumstance, there does not appear to be a large
difference between the two (e.g., L. H. Liu 015742 (PE),
H. J. Li 408 (IBSC, PE, SZ), Wulingshan Exped. 1419
(KUN)). (6) Distribution area of E. magna is relatively
wide, which overlaps the distribution of E. nanchuanen-
sis (e.g., H. F. Chow 26531 (IBSC, NAS, PE, SZ), K. F.
Li 61091 (IBSC, KUN, NAS, PE, SZ)). In conclusion,
it would be more appropriate to treat E. nanchuanensis
as a variety of E. magna.
35c. Elaeagnus magna (Serv.) Rehd. var. wushanensis
(C. Y. Chang) M. Sun & Q. Lin, comb. nov. & stat.
nov.——E. wushanensis C. Y. Chang, Fl. Sichuan. 1:
465. 1981. Type: China. Chongqing: Wushan, K. H.
Yang 57882 (holotype SZ!; isotype, PE!), Fig. 9: B.
Leaf blade densely or sparsely with rust-colored
scales. Fruit narrowly ellipsoid, 1.2–1.8 cm, with dense
rust-colored and silvery scales, red when ripe; fruiting
pedicels 8–16 mm, thick.
Forests, slopes, scrublands and roadsides; alt.
1400–2300 m. Endemic to China. Chongqing, Guizhou,
Hubei, and Sichuan.
Additional specimens examined:
China.Chongqing: Chengkou, Bashan Exped. 1630 (PE), T. L.
Dai 101373 (PE), 100517 (PE), 101345 (PE), 100823 (PE). Fengjie, C. Y.
Chang 25258 (PE), 5528 (NAS), H. F. Zhou et al. 108178 (IBSC), 108151
(IBSC), 108176 (PE, SZ), 108361 (IBSC, PE, SZ), M. Y. Fang 24257
(PE, SZ). Kaixian, Bashan Exped. 2254, 2502 (PE), 2134 (PE), T. L. Dai
101763 (PE). Wanyuan, B. L. Li 2046 (SZ), 2008 (SZ), L. N. Zhao 2635
(SZ), 2631 (SZ). Wushan, G. H. Yang 58034 (IBSC, PE, SZ), 57882 (PE,
SZ), 58054 (NAS, PE), 58975 (NAS, PE), H. F. Zhou et al. 109875 (IBSC,
PE, SZ), 109003 (IBSC, PE, SZ). Wuxi, G. H. Yang 65389 (IBSC, NAS,
PE), 59471 (NAS, PE, SZ), 6538 (NAS, PE, SZ), 59461 (NAS, PE, SZ),
M. L. She et al. 64167 (NAS), P. Y. Li 3365 (SZ), B. Z. Ni 00311 (SZ).
Guizhou: Duyun, Y. Tsiang 5916 (IBSC, NAS, PE). Hubei: Badong,
M. X. Nie & Q. H. Li 939 (IBSC, PE), Z. S. Zhang et al. 00939 (IBSC,
NAS, PE). Hefeng, H. J. Li 8243 (IBSC). Sichuan: Nanjiang, Z. X. Li
2794 (SZ).
Compared with E. magna, Chang (1981) consid-
ered E. wushanensis as a new species and distinctive
characters are brown or rust-colored young branches,
rounded or obtuse base and apex, 8–16 mm fruiting
pedicel, 5–6 mm calyx tube, and rust-colored scaly
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388 Journal of Systematics and Evolution Vol. 48 No. 5 2010
Fig. 9. A, Isotype of Elaeagnus nanchuanensis C. Y. Chang (C. H. Hsiung & C. L. Chow 90645,PE).B, Isotype of Elaeagnus wushanensis C. Y.
Chang (K. H. Yang 57882,PE).C, Comparison of fruits from Elaeagnus magna and E. magna var. wushanensis.Ca, Fruits of Elaeagnus magna var.
wushanensis.Cb, Fruits of E. magna.
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SUN & LIN: A revision of Elaeagnus in China 389
fruits. He also added that E. wushanensis was ex-
clusively distributed at an elevation of 1400–2300 m,
whereas E. magna was distributed at 100–1200 m. But
our studies suggest the following differences. (1) Young
branches of E. magna with rusted color scales (Z. T. Li
2376 (IBSC, PE), W. Y. Chun 3902 (PE), S. S. Lai &
D. F. Huang 000410 (PE)). (2) Shapes of these two taxa
are not constant, and leaf base and apex of E. magna
is sometimes rounded, sometimes obtuse, and some-
times coexisting (Z. T. Li 2376 (IBSC, PE), H. J. Li
408 (IBSC, PE, SZ), H. J. Li 8243 (NAS, PE)). (3)
Length of calyx tube of E. magna is 5–13 mm, con-
taining the length magnitude of E. wushanensis (H. F.
Zhou&H.Y.Su108082 (IBSC, SZ), L. D. Duan 0668
(PE)). (4) Fruit of E. magna is densely with silvery
scales, scattered with rust-colored scales (L. H. Liu
15742 (PE), H. J. Li 408 (IBSC, PE, SZ), Wulingshan
Exped. 1419 (KUN, PE)), but fruit of E. wushanensis
is only densely with rust-colored scales. (5) Fruiting
pedicel of E. magna is shorter than 8 mm, but fruit-
ing pedicel of E. wushanensis is 8–16 mm (Fig. 9: C).
(6) Distribution range of E. magna and E. wushanen-
sis are partly overlapping (H. F. Zhou 26531 (IBSC,
NAS, PE, SZ)). In summary, the main differences be-
tween E. wushanensis and E. magna are young branches
and fruiting pedicels. Therefore, it seems more appro-
priate to reinstate E. wushanensis as E. magna var.
wushanensis.
Acknowledgements This work was supported by
the Plant Specimen Digitization and Chinese Virtual
Herbarium Establishment (Grant No. 2005DKA21401).
We are profoundly grateful to the curators and relevant
staff of the following herbaria: CDBI, GXMI, HGAS,
HHBG, IBK, IBSC, K, KUN, N, NAS, NAU, P, PE,
SWFC, SYS, SZ, YAF, YCP, and YUKU, who granted
us access to their collections. We also thank Dr. Zhi-
Duan CHEN for his generous help and constructive
suggestions, Dr. Jun WEN and Dr. Beata PASZKO for
comments on the manuscript, and Dr. Xiang-Yun ZHU
for remarks on issues related to nomenclature code.
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