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FOC | Family List | FOC Vol. 24 | Liliaceae

11. Smilax Linnaeus, Sp. Pl. 2: 1028. 1753.

菝葜属 ba qia shu

Chen Xinqi (陈心启 Chen Sing-chi); Tetsuo Koyama1

Vines climbing or shrubs, woody, less often suberect or herbs, dioecious, usually with short, thick rhizomes. Stems and branches usually prickly. Petiole usually narrowly winged proximally, with an abscission zone between winged portion and apex; tendrils often present. Leaf blade usually ovate to lanceolate, main veins 3--7, connected by a network of cross veins and veinlets. Inflorescence borne in axil of leaf or scalelike bract, of 1(--3) umbels or a panicle, raceme, or spike of umbels; peduncle sometimes with a scalelike prophyll at base. Flowers small; tepals 6, usually free (connate in Smilax synandra). Male flowers: stamens 6, rarely 8 or more, inserted at base of tepals; anthers mostly 1-loculed. Female flowers: ovary 3-loculed; ovules 1 or 2 per locule; style very short; stigmas 3; staminodes absent to 6. Fruit a berry, red to black, usually 1- or 2-seeded. Seeds dark brown.

About 300 species: tropical, subtropical, and temperate regions of both hemispheres; 79 species (39 endemic) in China.


1 Rhizomes stoloniferous, creeping, elongate.   (2)
+ Rhizomes not stoloniferous, usually tuberous, short.   (4)
       
2 (1) Peduncle with a prophyll at base; leaves not glaucous abaxially   65 Smilax pinfaensis
+ Peduncle without a prophyll at base; leaves glaucous abaxially.   (3)
       
3 (2) Petiole 1--2 mm, narrowly winged for ca. 2/3 its length; stem much branched   26 Smilax nana
+ Petiole 4--13 mm, scarcely winged; stem slightly branched or not   25 Smilax pachysandroides
       
4 (1) Inflorescence a spike of umbels with 5--25 sessile umbels on a common axis.   (5)
+ Inflorescence a solitary umbel or a raceme of umbels with 2 to many stalked umbels on a common axis.   (6)
       
5 (4) Branches terete, ridged-angled, sparsely prickly; leaf blade 6--11 cm   78 Smilax aspera
+ Branches 4-angled, not prickly; leaf blade 20--30 cm   79 Smilax elegantissima
       
6 (4) Branches verruculose or bristly.   (7)
+ Branches neither verruculose nor bristly.   (16)
       
7 (6) Branches densely bristly.   (8)
+ Branches verruculose, sometimes young branches minutely bristly.   (9)
       
8 (7) Petiole not bristly; peduncle without a prophyll at base   19 Smilax horridiramula
+ Petiole densely bristly; peduncle with a prophyll at base   74 Smilax setiramula
       
9 (7) Peduncle without a prophyll at base.   (10)
+ Peduncle with a prophyll at base.   (11)
       
10 (9) Branches minutely verruculose, not prickly; inflorescence borne in axil of well-developed leaf   49 Smilax nervomarginata
+ Branches verruculose or spinulose-verruculose, ± sparsely prickly; inflorescence borne in axil of young leaf on new branches   13 Smilax lebrunii
       
11 (9) Young branches, petiole, and leaf base minutely bristly   57 Smilax kwangsiensis
+ Young branches, petiole, and leaf base not bristly.   (12)
       
12 (11) Leaf blade orbicular or nearly so, thickly leathery   70 Smilax bauhinioides
+ Leaf blade not orbicular, leathery or papery.   (13)
       
13 (12) Petiole wingless, tendrils absent; receptacle suboblong   67 Smilax fooningensis
+ Petiole winged, tendrils usually present; receptacle subglobose.   (14)
       
14 (13) Inflorescence 1-umbellate; branches usually 2- or 3-angled   54 Smilax chapaensis
+ Inflorescence 2--5-umbellate, forming a raceme of umbels; branches not angled.   (15)
       
15 (14) Inflorescence with 3--7 umbels; petiole 10--15 mm, tendrils usually present   61 Smilax aspericaulis
+ Inflorescence with 2 umbels; petiole 7--12 mm, tendrils usually absent   63 Smilax malipoensis
       
16 (6) Inflorescence of 2 to many umbels (if a solitary umbel, then peduncle articulate in proximal part with a prophyll at base).   (17)
+ Inflorescence a solitary umbel; peduncle not articulate, usually without a prophyll at base.   (36)
       
17 (16) Branches ± 4-angled.   (18)
+ Branches terete, not angled.   (19)
       
18 (17) Branches not zigzagged, angles narrowly winged   72 Smilax gagnepainii
+ Branches strongly zigzagged, angles wingless   68 Smilax quadrata
       
19 (17) Tepals of male flowers longitudinally wrinkled; stamens 8--10   50 Smilax luei
+ Tepals of male flowers not wrinkled; stamens 6.   (20)
       
20 (19) Petiole wings well developed, each 4--13 mm wide.   (21)
+ Petiole wings weakly developed, each less than 4 mm wide.   (23)
       
21 (20) Petiole wings not clasping node at base   75 Smilax griffithii
+ Petiole wings deeply clasping node at base.   (22)
       
22 (21) Inflorescence of 2--7 solitary umbels; filaments proximally connate, forming a short column   76 Smilax ocreata
+ Inflorescence of 10--30 umbels, umbels 2- or 4-subwhorled or nearly clustered; filaments free   77 Smilax perfoliata
       
23 (20) Filaments proximally connate, forming a short column.   (24)
+ Filaments free.   (25)
       
24 (23) Receptacle suboblong, 3--6 × 2--4.5 mm; outer tepals of male flowers ca. 5 mm   56 Smilax hemsleyana
+ Receptacle subglobose, 2--3 mm in diam.; outer tepals of male flowers 7--8 mm   55 Smilax vanchingshanensis
       
25 (23) Branches densely prickly; petiole verruculose-prickly   73 Smilax densibarbata
+ Branches smooth or sparsely prickly; petiole not verruculose-prickly.   (26)
       
26 (25) Leaf blade 8--14(--20) cm wide   62 Smilax ovalifolia
+ Leaf blade 1--5(--7) cm wide.   (27)
       
27 (26) Petiole laterally compressed, winged portion 1--3 cm   71 Smilax planipes
+ Petiole subterete, winged portion less than 0.9 cm.   (28)
       
28 (27) Umbels 2- or 3-flowered; peduncle 1--3 mm; petiole 2.5--5 mm; stem and branches 2- or 4-angled   66 Smilax basilata
+ Umbels 3--15-flowered; peduncle and petiole both more than 5 mm; stem and branches not or indistinctly obtusely angled.   (29)
       
29 (28) Stamens very short, less than 1 mm, 1/5--1/3 as long as tepals   64 Smilax cocculoides
+ Stamens 3--6 mm, more than 1/2 as long as tepals.   (30)
       
30 (29) Leaf blade thickly leathery, orbicular to elliptic, with 3 main veins concave adaxially and veinlets indistinct; petiole scarcely winged   69 Smilax lunglingensis
+ Leaf blade leathery or papery, variable in shape, with 3--5 main veins raised or sometimes midvein slightly concave adaxially; petiole winged.   (31)
       
31 (30) Inflorescence with 3--7 umbels.   (32)
+ Inflorescence with 1 umbel.   (34)
       
32 (31) Leaf blade leathery, apex acuminate; raceme with terminal umbel abortive; receptacle ellipsoid   59 Smilax yunnanensis
+ Leaf blade papery, apex mucronate; raceme with terminal umbel well developed; receptacle subglobose.   (33)
       
33 (32) Berries 1.5--2 cm in diam.; petiole 1.5--5 cm; inflorescence of 2 or 3 umbels; male flowers with tepals greenish yellow   58 Smilax megacarpa
+ Berries 5--7 mm in diam.; petiole 1--2 cm; inflorescence of 3--6(--10) umbels; male flowers with tepals dull red   60 Smilax bracteata
       
34 (31) Berries 1.5--2 cm in diam.   58 Smilax megacarpa
+ Berries 0.6--1 cm in diam.   (35)
       
35 (34) Seeds smooth; leaf blade becoming dark green or sometimes tinged black when dried, not wrinkled adaxially   52 Smilax lanceifolia
+ Seeds deeply 5- or 6-furrowed; leaf blade becoming grayish when dried, wrinkled along main veins adaxially   53 Smilax astrosperma
       
36 (16) Stems usually annual, herbaceous.   (37)
+ Stems perennial, becoming woody.   (39)
       
37 (36) Leaf blade abaxially pale green, usually glabrous or papillose-puberulent, not white powdery   2 Smilax riparia
+ Leaf blade abaxially glaucous, white powdery.   (38)
       
38 (37) Petiole scarcely winged, tendrils (when developed) borne in proximal part   1 Smilax nipponica
+ Petiole narrowly winged, tendrils borne at middle or in distal part   3 Smilax pottingeri
       
39 (36) Petiole wings lacerate-fimbriate at margin.   (40)
+ Petiole wings subentire at margin.   (41)
       
40 (39) Branchlets usually 4-angled; leaf blade usually rhombic-ovate, with 3(--5) main veins raised adaxially, base cuneate; prickles sparse or absent   44 Smilax myrtillus
+ Branchlets 2- or 3-angled; leaf blade cordate to ovate, with (3--)5 main veins slightly concave adaxially, base truncate to cordate; prickles rather dense   45 Smilax munita
       
41 (39) Petiole with abscission zone below apex, hence leaf blade abscising together with distal part of petiole.   (42)
+ Petiole with abscission zone at apex, hence leaf blade abscising with almost no part of petiole.   (61)
       
42 (41) Leaf blade abaxially pubescent, papillose-pubescent, or powdery-scabrous.   (43)
+ Leaf blade abaxially glabrous.   (44)
       
43 (42) Peduncle 0.5--3 cm; receptacle thickened and elongate   12 Smilax chingii
+ Peduncle 3--5 cm; receptacle scarcely thickened, not elongate   37 Smilax aberrans
       
44 (42) Peduncle 0.1--0.3 cm   46 Smilax bapouensis
+ Peduncle 0.4--5 cm.   (45)
       
45 (44) Petiole wings semicircular, each 3--5 mm wide, occupying entire length of petiole   16 Smilax discotis
+ Petiole wings not as above.   (46)
       
46 (45) Filaments ± connate, forming a column.   (47)
+ Filaments free.   (48)
       
47 (46) Outer tepals free; filaments connate in proximal part   24 Smilax emeiensis
+ Outer tepals connate for ca. 1/2 their length, forming a tube; filaments connate throughout   33 Smilax synandra
       
48 (46) Inflorescence arising from branches or branchlets with leaves fully developed.   (49)
+ Inflorescence arising from young branches or branchlets with leaves not fully developed.   (52)
       
49 (48) Leaf blade abaxially glaucous.   (50)
+ Leaf blade abaxially green or pale green.   (51)
       
50 (49) Inflorescence usually racemose; pedicel erect in fruit   36 Smilax austrozhejiangensis
+ Inflorescence umbellate; pedicel bent downward in fruit   38 Smilax retroflexa
       
51 (49) Peduncle slightly shorter or longer than petiole; female flowers with 6 staminodes   4 Smilax sieboldii
+ Peduncle much shorter than petiole (usually less than 1/2 as long); female flowers with 3 staminodes   5 Smilax scobinicaulis
       
52 (48) Petiole with abscission zone 2--3 mm above winged portion.   (53)
+ Petiole with abscission zone immediately above winged portion.   (54)
       
53 (52) Leaf blade elliptic; petiole winged for ca. 1/2 its length   17 Smilax glaucochina
+ Leaf blade ovate-elliptic to oblong-lanceolate; petiole winged for ca. 2/3 its length   18 Smilax elongatoumbellata
       
54 (52) Umbels 1- or 2-flowered or racemes 3--5-flowered   9 Smilax trinervula
+ Umbels 3- to many flowered.   (55)
       
55 (54) Receptacle oblong or ellipsoid, ± elongate.   (56)
+ Receptacle subglobose.   (57)
       
56 (55) Petiole winged for 2/3--4/5 its length   10 Smilax ferox
+ Petiole winged for 1/3--1/2 its length   11 Smilax megalantha
       
57 (55) Leaf blade herbaceous, becoming membranous or thinly papery when dried; tendrils partly developed.   (58)
+ Leaf blade papery or leathery; tendrils well developed.   (59)
       
58 (57) Leaf blade abaxially glaucous; berries red at maturity   14 Smilax polycolea
+ Leaf blade abaxially green or pale green; berries purplish black at maturity   15 Smilax outanscianensis
       
59 (57) Petiole wings each 2--4 mm wide, wider than petiole; tendrils short, slender   8 Smilax davidiana
+ Petiole wings each 0.5--1 mm wide, about as wide as petiole; tendrils long, rather thick.   (60)
       
60 (59) Tepals of male flowers greenish to yellowish green, recurved at anthesis, outer ones 1.8--2.5 mm wide   6 Smilax china
+ Tepals of male flowers yellow, tinged pinkish red, spreading at anthesis, outer ones 1.3--1.5 mm wide   7 Smilax nantoensis
       
61 (41) Leaves and inflorescences becoming blackish or blackish brown when dried.   (62)
+ Leaves and inflorescences not becoming blackish or blackish brown when dried.   (64)
       
62 (61) Branches slightly ridged, smooth or prickly   20 Smilax nigrescens
+ Branches neither ridged nor prickly.   (63)
       
63 (62) Petiole 3--7 mm; umbels 1--3-flowered   39 Smilax darrisii
+ Petiole 5--5.5 cm; umbels densely 50--100-flowered   48 Smilax jiankunii
       
64 (61) Petiole wings each with a lanceolate auricle at apex.   (65)
+ Petiole wings without auricle at apex.   (70)
       
65 (64) Outer tepals connate for ca. 1/2 their length, inner ones adnate to filament column   33 Smilax synandra
+ All tepals free.   (66)
       
66 (65) Plants suberect or subscandent; tendrils absent.   (67)
+ Vines climbing; tendrils wholly or partly developed.   (68)
       
67 (66) Leaf blade rhombic-elliptic or ovate-rhombic; peduncle 7--22 mm; auricles of petiole wings 1--1.5 mm   35 Smilax hayatae
+ Leaf blade ovate; peduncle 3--4 mm; auricles of petiole wings 0.3--0.5 mm   40 Smilax mairei
       
68 (66) Leaf blade thinly papery, ovate-lanceolate to broadly lanceolate; branchlets zigzagged; female flowers with 6 staminodes   31 Smilax elegans
+ Leaf blade leathery, ovate to elliptic; branchlets not zigzagged; female flowers with 3 staminodes.   (69)
       
69 (68) Peduncle 4--15 mm, 2/3--4/5 as long as petiole   32 Smilax corbularia
+ Peduncle 1--5 mm, less than 1/2 as long as petiole   34 Smilax hypoglauca
       
70 (64) Shrubs erect or suberect; tendrils absent.   (71)
+ Vines climbing, shrubby; tendrils wholly or partly developed.   (74)
       
71 (70) Leaf blade lanceolate to oblong-lanceolate, more than 5 × as long as wide; stamens very short, ca. 1/8 as long as tepals   30 Smilax tsinchengshanensis
+ Leaf blade ovate to elliptic, 2--3 × as long as wide; stamens much longer, 1/3--1/2 as long as tepals.   (72)
       
72 (71) Filaments connate in proximal part, forming a column   23 Smilax cyclophylla
+ Filaments free.   (73)
       
73 (72) Petiole and leaf blade glabrous   21 Smilax stans
+ Distal 1/2 of petiole and abaxial base of leaf blade (particularly on veins) papillose-puberulent   22 Smilax trachypoda
       
74 (70) Peduncle shorter than or subequaling petiole; receptacle much thickened, with many persistent bracteoles.   (75)
+ Peduncle longer than petiole; receptacle not or only slightly thickened, without persistent bracteoles   (77)
       
75 (74) Male flowers slightly 6-angled, scarcely open, ca. 3 mm in diam.; outer tepals cucullate, abaxially deeply channeled   43 Smilax glabra
+ Male flowers not angled, fully open, 1--1.5 mm in diam.; outer tepals neither cucullate nor abaxially channeled.   (76)
       
76 (75) Branches minutely prickly; peduncle slightly shorter than petiole; female flowers with 3 staminodes   42 Smilax microphylla
+ Branches not prickly; peduncle much shorter than petiole; female flowers with 6 staminodes   41 Smilax longebracteolata
       
77 (74) Leaf blade abaxially green or pale green.   (78)
+ Leaf blade abaxially ± glaucous.   (80)
       
78 (77) Leaf blade with 3--5 main veins, outermost veins separate from leaf margins; petiole winged for 1/2--3/4 its length, tendrils borne near middle   51 Smilax arisanensis
+ Leaf blade with 5--7 main veins, outermost veins nearly combined with leaf margins; petiole winged for ca. 1/4 its length, tendrils borne near base.   (79)
       
79 (78) Leaf blade papery or thinly leathery, adaxially ± concave along midvein; anthers shorter than filaments   47 Smilax biumbellata
+ Leaf blade leathery, adaxially not concave along midvein; anthers slightly longer than filaments   49 Smilax nervomarginata
       
80 (77) Flowers large, tepals 6--7 mm   29 Smilax lushuiensis
+ Flowers much smaller, tepals 2--3.5 mm.   (81)
       
81 (80) Leaf blade with 3 main veins; petiole 3--7 mm, winged for 1/3--1/2 its length; stamens ca. 1/2 as long as tepals   39 Smilax darrisii
+ Leaf blade with 5--7 main veins; petiole (5--)8--12 mm, winged for 2/3--3/4 its length; stamens ca. 1/3 as long as tepals.   (82)
       
82 (81) Leaf blade with 5 main veins; filaments basally connate, forming a short column   27 Smilax menispermoidea
+ Leaf blade with 7 main veins; filaments free   28 Smilax pygmaea

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