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FOC | Family List | FOC Vol. 12 | Rhamnaceae | Rhamnus

2. Rhamnus subg. Rhamnus

鼠李亚属 shu li ya shu

Shrubs or trees, usually spinescent, rarely unarmed, with terminal scaly buds and axillary buds. Leaves alternate, subopposite to opposite, or fascicled at short shoots. Flowers unisexual and plants dioecious, rarely polygamous, usually 4(or 5)-merous. Sepals adaxially with conspicuous or inconspicuous midvein. Petals spatulate, base shortly clawed, apex entire or 2-fid. Male flower with developed stamens; female flowers with rudimentary stamens. Disk thin, lining calyx tube. Ovary superior, globose, not immersed in disk, 2-4-loculed; style 2-4-cleft. Berrylike drupe subglobose or obovoid-globose, base with persistent calyx tube, with 2-4 one-seeded stones. Seeds with a ± long, narrow to gaping, often distinctly margined abaxial or lateral furrow.

About 150 species: mainly in E Asia, a few species in tropical Africa, Europe, and North America; 52 species (34 endemic) in China.

The Chinese species of Rhamnus subg. Rhamnus belong to R. sect. Tetrarhamnus Grubov, R. sect. Pseudoceanothus Grubov, and R. sect. Rhamnus.

1 Plants only with long shoots, without short shoots, unarmed; leaves alternate; flowers 4- or 5-merous; petals present or absent   (2)
+ Plants with both long and short shoots, apices of short shoots often terminating in a woody spine; leaves opposite or alternate on long shoots, clustered or fascicled on short shoots; flowers unisexual and plants dioecious, 4-merous; petals present (R. sect. Rhamnus)   (16)
2 (1) Flowers polygamous, 4-merous; petals absent; ovary 4-loculed, rarely 3-loculed; flowers usually fascicled in axils of leaves; inflorescences never with leaflike bracts (R. sect. Tetrarhamnus)   (3)
+ Flowers unisexual and plants dioecious, rarely polygamous, 5-merous; petals usually present; ovary 3-loculed; flowers solitary or few fascicled in axillary cymose racemes or cymose panicles; inflorescences with small, leaflike bracts (R. sect. Pseudoceanothus)   (5)
3 (2) Leaf blade 2-4.5 cm, apex obtuse or rounded, margin entire, both surfaces yellow tomentulose, velutinous, lateral veins 5 or 6(-8) pairs.   8 R. ninglangensis
+ Leaf blade 5-20 cm, apex acuminate or long acuminate, margin serrate, abaxially puberulent on veins or barbate in axils of veins, lateral veins 9-17 pairs   (4)
4 (3) Deciduous trees or shrubs; young branches puberulent; leaves papery, abaxially sparsely pubescent on veins.   7 R. sargentiana
+ Evergreen trees or shrubs; young branches glabrous; leaves leathery, abaxially glabrous or barbellate in axils of leaves.   6 R. hemsleyana
5 (2) Flowers few, solitary or 2-6-fascicled in axils of leaves   (6)
+ Flowers numerous, in cymose racemes or cymose panicles   (9)
6 (5) Leaf blade to 4 cm, lateral veins 2-4 pairs   (7)
+ Leaf blade usually longer than 4 cm, lateral veins 4-6 pairs   (8)
7 (6) Leaves papery, petioles 2-7 mm; flowers solitary, unisexual and plants dioecious; petals present; pedicels 1-2 mm.   9 R. heterophylla
+ Leaves thinly leathery, petioles shorter than 2 mm; flowers polygamous; petals absent; pedicels 5-6(-10) mm.   10 R. procumbens
8 (6) Leaves leathery, lateral and reticulate veins adaxially conspicuously impressed; seeds broad at both ends, abaxially narrowly furrowed at middle part.   11 R. bodinieri
+ Leaves papery to thickly papery, lateral veins prominent on both surfaces; seeds with base broader than middle, abaxially longitudinally furrowed in upper part.   12 R. xizangensis
9 (5) Flowers polygamous or unisexual; inflorescences often with many persistent leaflike bracts; seeds shortly furrowed for 1/2 length   (10)
+ Flowers unisexual and plants dioecious; inflorescences without leaflike bracts or with few caducous leaflike bractlets; seeds abaxially longitudinally furrowed for 3/5 length   (11)
10 (9) Drupe 3-4 mm in diam., base with persistent sepals and calyx tube.   13 R. formosana
+ Drupe 5-6 mm in diam., sepals caducous, base with persistent calyx tube only.   14 R. hainanensis
11 (9) Leaves abaxially hairy or only hairy on veins and petioles; petals present   (12)
+ Leaves and petioles glabrous or leaves abaxially clustered hairy in vein axils; petals present or absent   (14)
12 (11) Leaves thickly leathery, abaxially densely tomentulose; seeds with equally broad, longitudinal furrow for 4/5 length.   15 R. kwangsiensis
+ Leaves papery or thickly papery, abaxially puberulent or only on veins; seeds abaxially furrowed for 3/4 length or for nearly entire length   (13)
13 (12) Leaf blade narrowly elliptic or oblanceolate-elliptic; petiole shorter than 1 cm; flowers often in short cymose racemes; pedicel and calyx puberulent; seeds abaxially with furrow extending over nearly entire length.   16 R. esquirolii
+ Leaf blade elliptic to broadly elliptic; petiole 1.2-2 cm; flowers usually in large panicles; pedicel and calyx glabrous; seeds abaxially furrowed for 3/4 length.   17 R. nigricans
14 (11) Leaf blade usually broadly elliptic or broadly oblong; young branches and inflorescence rachises puberulent; inflorescences to 12 cm; petals present.   18 R. napalensis
+ Leaf blade elliptic or oblong-elliptic; young branches and inflorescence rachises glabrous; inflorescences shorter than 5 cm; petals absent   (15)
15 (14) Leaf blade papery or thinly leathery, apex caudate-acuminate, abaxially pale purple when dry, petiole to 18 mm.   19 R. subapetala
+ Leaf blade leathery, apex shortly acuminate or acuminate, abaxially not drying purple, petiole to 7 mm.   20 R. coriophylla
16 (1) Leaves opposite or subopposite, rarely alternate   (17)
+ Leaves and branches alternate, rarely subopposite   (34)
17 (16) Leaves not longer than 3 cm, narrow, usually less than 1 cm wide, lateral veins in 2 or 3, rarely 4, pairs   (18)
+ Leaves longer than 3 cm, ca. 1.5 cm wide, lateral veins (3 or)4-7 pairs   (22)
18 (17) Leaves subleathery, both surfaces glabrous, petioles 5-20 mm, glabrous; seeds abaxially shortly furrowed for 1/2 length, obcordate.   21 R. maximovicziana
+ Leaves papery or thickly papery, both surfaces and petioles sparsely puberulent; seeds abaxially or laterally furrowed for ca. 4/5 of length   (19)
19 (18) Leaf blade oblong or elliptic, petiole 1-3 mm   (20)
+ Leaf blade ovate, rhombic-obovate, or rhombic-elliptic, petiole 4-15 mm   (21)
20 (19) Branchlets dark brown; bark scabrous, longitudinally fissured; leaf blade usually elliptic, abaxially black when dry, clustered hairy at vein axils or glabrous, reticulate veins inconspicuous; seeds abaxially deeply furrowed for 4/5 length.   22 R. gilgiana
+ Branchlets silvery-gray or gray-brown, slightly glabrous; leaf blade usually oblong or ovate-elliptic, abaxially golden-yellow or yellowish when dry, reticulate veins conspicuous, both surfaces puberulent, rarely subglabrous; seeds abaxially shallowly furrowed for nearly entire length.   23 R. flavescens
21 (19) Leaf blade papery, ovate or ovate-lanceolate, abaxially yellow when dry, veins or vein axils white puberulent; seeds abaxially with broad furrow.   24 R. bungeana
+ Leaf blade thickly papery, rhombic-obovate or rhombic-elliptic, abaxially gray-white when dry, areole of vein axils puberulent; seeds abaxially with narrow furrow.   25 R. parvifolia
22 (17) Leaf blade ovate-cordate or ovate-orbicular, base cordate or rounded, margin densely sharply serrate; fruiting pedicel 1.3-2.3 cm; seeds abaxially furrowed for 4/5 length.   26 R. arguta
+ Leaf blade not ovate-cordate, base cuneate or subrounded, margin obtusely serrate or crenate-serrate; fruiting pedicel not longer than 1.2 cm   (23)
23 (22) Petioles usually longer than 1-1.5 cm; seeds with short abaxial furrow restricted to 1/3 of length   (24)
+ Petioles shorter than 1 cm; seeds abaxially or laterally longitudinally furrowed for more than 1/2 length (except in R. dumetorum)   (28)
24 (23) Branchlets hairy or glabrous; leaves abaxially often turning yellow or golden-yellow when dry, veins and vein axils golden-yellow pubescent, petioles 5-15 mm.   37 R. utilis
+ Branchlets glabrous; leaves abaxially greenish when dry, glabrous, or abaxially midvein sparsely white hairy, petioles 1.5-3 cm   (25)
25 (24) Leaf blade glabrous, margin densely crenate, elliptic or broadly elliptic, lateral veins 3 or 4 pairs.   33 R. cathartica
+ Leaf blade and petiole subglabrous, margin sparsely obtusely serrate, or crenate, orbicular, narrowly to broadly elliptic, or oblong, lateral veins 4-6 pairs   (26)
26 (25) Leaf blade suborbicular, ovate-rhombic, or elliptic; axillary buds small; seeds easily separated from endocarp, exposing a nearly circular, distinctly margined furrow.   34 R. diamantiaca
+ Leaf blade narrowly to broadly elliptic or oblong; axillary buds 4-8 mm; seeds exposing a short, never circular furrow   (27)
27 (26) Branches terminating in a spine or large bud; leaf blade narrowly elliptic or narrowly oblong.   35 R. ussuriensis
+ Branches terminating in a large bud, rarely spinescent; leaf blade broadly elliptic or ovate-oblong.   36 R. davurica
28 (23) Young branches, annual branches, both surfaces of leaves, or veins and petioles puberulent; flowers and pedicels sparsely puberulent; leaf blade obovate-orbicular, ovate-orbicular, or suborbicular.   27 R. globosa
+ Young branches, annual branches, and petioles glabrous or nearly so; flowers and pedicels glabrous; leaf blade not obovate-orbicular or suborbicular   (29)
29 (28) Leaves adaxially glabrous, abaxially vein axils clustered hairy   (30)
+ Leaves adaxially sparsely pubescent at least on veins, abaxially areoles of veins clustered hairy or rarely glabrous   (31)
30 (29) Leaf blade narrowly elliptic or oblanceolate-elliptic, 1-2.2 cm wide, apex caudate-acuminate or long acuminate, lateral veins (4 or)5 or 6(or 7) pairs, petiole 2-6 mm.   28 R. iteinophylla
+ Leaf blade obovate or obovate-elliptic, 2-5 cm wide, apex shortly acuminate or acute, lateral veins 3-5 pairs, petiole 7-20 mm.   29 R. leptophylla
31 (29) Branchlets pale gray or gray-brown; bark scabrous, not nitid; seeds black, abaxially at base with short furrow.   30 R. dumetorum
+ Branchlets red-brown, purple-red, or dark brown; bark laevigate, shiny; seeds red-brown or brown, abaxially with longitudinal furrow extending over 2/3 of length   (32)
32 (31) Leaf blade rhombic-obovate or rhombic-elliptic, both surfaces glabrous or abaxially areoles of vein axils sparsely puberulent, lateral veins 2-4 pairs.   25 R. parvifolia
+ Leaf blade elliptic, obovate-elliptic, or obovate-lanceolate, adaxially sparsely hirsutulous, abaxially glabrous or areoles of vein axils sparsely pilose, lateral veins 4 or 5 pairs   (33)
33 (32) Leaf blade usually elliptic or obovate-elliptic, abaxially yellowish or gray when dry, reticulate veins inconspicuous, vein axils often with areole, papery to thickly papery; young branches glabrous.   31 R. tangutica
+ Leaf blade usually obovate-lanceolate, abaxially often reddish when dry, reticulate veins conspicuous, vein axils without areoles, rarely with areoles, thinly papery or papery; young branches slightly pubescent or glabrous.   32 R. virgata
34 (16) Leaves usually shorter than 1.2 cm, narrow; seeds abaxially or laterally with over 2/3 entire length longitudinal furrow   (35)
+ Leaves usually more than 3 × 2 cm; seeds abaxially or laterally with long or short furrow   (42)
35 (34) Leaf blade linear, oblanceolate, or narrowly oblanceolate, both surfaces glabrous; seeds abaxially with narrow furrow for 4/5 entire length   (36)
+ Leaf blade elliptic, obovate-elliptic, or spatulate, both surfaces puberulent, or abaxially puberulent on veins, rarely glabrous; seeds with broad or narrow furrow   (37)
36 (35) Leaf blade narrowly lanceolate to narrowly obovate, 3-5 cm, apex acute or obtuse, petiole 3-15 mm; branches terminating in a spine; erect shrubs or small trees.   38 R. erythroxylum
+ Leaf blade oblanceolate or narrowly oblanceolate, 1-2.5 cm, apex obtuse or emarginate, petiole 2-3 mm; branches terminating in a small bud, not spinescent; procumbent small shrubs.   39 R. tzekweiensis
37 (35) Leaves at both surfaces, or at least major veins, yellowish verrucose-pilosulose; flowers and pedicels sparsely pilose.   40 R. aurea
+ Leaves glabrous or puberulent or abaxially vein axils clustered hairy; flowers and pedicels glabrous   (38)
38 (37) Leaves leathery or nearly so; seeds abaxially or laterally with deep furrow   (39)
+ Leaves papery; seeds abaxially with broad, shallow furrow   (40)
39 (38) Branchlets spreading, often virgate, purple-red or dark purple, smooth, shiny; leaf blade subleathery, oblanceolate or obovate-elliptic, apex obtuse or emarginate, glabrous or subglabrous; seeds abaxially with broad furrow.   41 R. leptacantha
+ Branchlets not virgate, gray-brown or dark brown, scabrous, with longitudinally arranged slits, not shiny; leaf blade leathery, spatulate or rhombic-elliptic, apex truncate or acute, adaxially glabrous or midvein sparsely puberulent, abaxially with ± distinct bearded domatia in axils of primary and lateral veins; seeds abaxially with narrower furrow.   42 R. rosthornii
40 (38) Leaf blade elliptic or oblong, margin entire or abaxially inconspicuously serrulate above middle, glabrous, lateral veins 3 or 4 pairs; flowers 10-20-clustered at leaf axils.   43 R. songorica
+ Leaf blade obovate or obovate-elliptic, rarely elliptic, both surfaces sparsely puberulent, margin sparsely serrulate, lateral veins 2 or 3 pairs; flowers 2-5-clustered at leaf axils or terminal on short shoots   (41)
41 (40) Leaves 5-10 mm, obovate; profusely shortly branched, pulvinate compact shrubs, 10-25 cm tall.   44 R. minuta
+ Leaves ca. 2.2 mm, elliptic or ovate-elliptic; prostrate, small shrubs, to 200 cm tall.   45 R. prostrata
42 (34) Young branches, leaves, petioles, flowers, and pedicels glabrous   (43)
+ Young branches, both surfaces of leaves, leaf veins, or vein axils hairy; flowers and pedicels puberulent or glabrous   (46)
43 (42) Leaf blade membranous, oblong-ovate or ovate-orbicular, margin crenate to serrate; terminal bud not spinescent.   46 R. nakaharae
+ Leaf blade papery, elliptic, oblong-elliptic, or broadly elliptic, margin hooklike crenate; terminal bud spinescent   (44)
44 (43) Leaf blade apically shortly acuminate, petiole 1-1.5 cm; seeds abaxially with margined furrow for 5/7 of length.   49 R. hupehensis
+ Leaf blade apically caudate-acuminate or long acuminate, petiole not longer than 1 cm; seeds with margined furrow for 1/2 of length   (45)
45 (44) Petioles 2-4 mm; seeds abaxially with margined furrow for 2/5-1/2 of length, apex without fissured line.   47 R. wilsonii
+ Petioles 5-10 mm; seeds abaxially with margined furrow for 1/4-1/3 of length, apex with fissured line.   48 R. lamprophylla
46 (42) Calyx and pedicel sparsely puberulent; annual branches, both leaf surfaces, or at least leaf veins pubescent   (47)
+ Calyx and pedicel glabrous; leaves subglabrous or ± hairy   (49)
47 (46) Leaf blade thickly papery, obovate-elliptic or obovate-orbicular, adaxially veins conspicuously impressed, distinctly rugose when dry; seeds abaxially with margined furrow nearly as long as entire length, above furrow without fissured line.   50 R. rugulosa
+ Leaf blade papery or thinly papery, veins adaxially not rugose when dry; seeds abaxially with fissured furrow for 1/4-2/5 of length, above furrow with fissured line   (48)
48 (47) Leaf blade broadly elliptic, obovate-elliptic, or ovate-orbicular, apex shortly acuminate or acuminate, petiole 7-25 mm; fruiting pedicel 7-14 mm.   51 R. koraiensis
+ Leaf blade oblanceolate-oblong or elliptic, apex long acuminate or caudate-acuminate, petiole 3-8 mm; fruiting pedicel 4-7 mm.   52 R. grandiflora
49 (46) Leaf blade membranous, ovate or elliptic-ovate, glabrous, or midvein adaxially and veins abaxially sparsely pilose.   53 R. liukiuensis
+ Leaf blade papery; seeds abaxially with, long or short furrow   (50)
50 (49) Seeds abaxially with margined furrow for nearly entire length; leaves abaxially sparsely clustered hairy in vein axils   (51)
+ Seeds abaxially with margined furrow for 1/3-1/2 of length; leaves adaxially glabrous, or pubescent on veins   (52)
51 (50) Leaf blade oblong or ovate-oblong.   54 R. wumingensis
+ Leaf blade ovate-elliptic, obovate-elliptic, or obovate.   50 R. rugulosa
52 (50) Seeds abaxially with margined furrow for nearly 1/2 of length, above furrow without fissured line   (53)
+ Seeds abaxially with basal rim shortly furrowed for less than 1/3 of length, above furrow with fissured line   (54)
53 (52) Leaf blade usually broadly elliptic, abaxially pubescent especially on veins, petiole 2-4 mm.   47 R. wilsonii
+ Leaf blade oblong or ovate-oblong, abaxially often turning reddish when dry, glabrous, petiole 4-9 mm.   55 R. brachypoda
54 (52) Branchlets gray-brown, not shiny, terminating in an obtuse spine; leaves often reflexed when dry, adaxially glabrous, abaxially veins or veins axils pilose, petioles 3-6 mm, glabrous or puberulent.   56 R. fulvotincta
+ Branchlets yellow-brown, shiny, terminating in a sharp spine; leaves not reflexed when dry, adaxially white adpressed hairy or pilose, abaxially veins or vein axils pilose or glabrous, petioles 6-25 mm, sparsely or densely pubescent.   57 R. schneideri

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