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FOC | Family List | FOC Vol. 11 | Geraniaceae

1. Geranium Linnaeus, Sp. Pl. 2: 676. 1753.

老鹳草属 lao guan cao shu

Herbs, annual, biennial, or perennial [rarely shrublets or shrubs]. Leaves simple, usually palmately divided or cleft, sometimes entire or pinnately lobed, petiolate. Basal leaves usually forming a rosette. Cauline leaves opposite or alternate. Inflorescence terminal or axillary, cymose, bracteate. Flowers usually paired or occasionally solitary, actinomorphic, rarely somewhat zygomorphic. Sepals 5, imbricate, apex obtuse to caudate. Petals 5, distinct, sometimes clawed, apex rounded, retuse, or emarginate. Stamens 10, in 2 whorls, outer ones opposite to petals, inner ones alternating with petals, all with anthers or very rarely (in G. pusillum) 5 reduced to staminodes. Nectaries usually 5 and alternate with petals, rarely (in G. sinense) united into a ring. Ovary 5-locular, with 2 superposed ovules per locule; style distinctly 5-cleft. Fruit a schizocarp, long beaked, splitting into 5 1-seeded mericarps.

About 380 species: cosmopolitan, especially in temperate and tropical mountains; 50 species (18 endemic, three introduced) in China.

Primary leaf blade divisions are named segments and second ones lobes. The middle segment is measured from leaf blade apex to the insertion of the petiole with the leaf blade. The main sinus is measured from the leaf blade apex to the junction between the middle segment and the adjacent lateral segments. The second sinus is the deeper of the middle segment. Sepal length does not include mucro length. Fruit length is measured from the mericarp base to the apex of the stigmatic remains. The fruit rostrum length is measured from the mericarp apex to the base of the stigmatic remains including, when present, a narrowed apical part.

Five species recorded from China are of uncertain identity or are too poorly known to be treated here and are discussed unnumbered at the end of the generic treatment.

Geranium schrenkianum Trautvetter (Bull. Soc. Imp. Naturalistes Moscou 57: 53. 1882) is widespread in Kazakhstan and is expected to be found in Xinjiang. This species is a small annual, easily recognized by its palmately divided leaves, petals 3-4 mm with emarginate apex, and antrorsely appressed trichomes on stem, petioles, peduncle, and pedicels.

Species incertae sedis

The following species have been described from Chinese material, or recorded in China, but are imperfectly known as no specimens have been seen.

Geranium camaense C. C. Huang in S. Y. Jin, Cat. Type Spec. Herb. China, 301. 1994.

This species is only known from the type collected in Xizang (Dinggyê Xian) at ca. 4000 m. According to the original description, this species is similar to Geranium lucidum Linnaeus but differs in having leaves and flowers glabrous. Geranium lucidum is a European species that reaches the W Himalayas and Nepal. It is an annual and very easy to recognize by its calyx with lengthwise keels and transverse flaps between these. Unfortunately, it has not been possible to examine the type specimen of G. camaense.

Geranium platyrenifolium Z. M. Tan, Bull. Bot. Res., Harbin 6(2): 52. 1986.

Geranium tanii Aedo & Muñoz Garmendia; G. trifoliatum Z. M. Tan (1994), not Andrews (1805).

According to the original description, this species has roots bearing 1 or 2 tubers at distances up to several cm from their origin, opposite leaves deeply divided, stipules distinct, cymules 2-flowered, petals 1.5-1.8 cm, with rounded base and with trichomes, and indumentum nonglandular. It is found in meadows of SW Sichuan. Unfortunately, it has not been possible to examine any specimens belonging to this species.

Geranium rubifolium Lindley, Edwards’s Bot. Reg. 26: t. 67. 1840.

This species was reported in forests at ca. 2700 m in Xizang (Yadong) by L. R. Xu & C. C. Huang (Fl. Reipubl. Popularis Sin. 43(1): 72. 1998). This area is on the S side of the Himalayas between India (Sikkim) and Bhutan, but the species was previously considered endemic from N Pakistan and Kashmir far to the West. This Chinese report mentioned the plant as having petals with a basal spot. This feature does match Geranium rubifolium, which has uniform petals, and would correspond to G. procurrens Yeo (Bot. Mag. 179: t. 644. 1973), a species widespread in the Himalayas but not recorded from China. Unfortunately, it has not been possible to examine the specimen reported from China.

Geranium solitarium Z. M. Tan, Acta Phytotax. Sin. 33: 608. 1995.

According to the original description, this species has roots long and thickened, opposite leaves not deeply divided, stipules distinct, cymules 1-flowered, petals ca. 1.8 cm, with rounded apex and basally with trichomes, and indumentum nonglandular. The fruit and seed are unknown. It is found in alpine habitats of N Sichuan. Unfortunately, it has not been possible to examine any specimens belonging to this species.

Geranium yaanense Z. M. Tan, Acta Phytotax. Sin. 33: 611. 1995.

According to the original description, this species has opposite leaves not deeply divided, stipules distinct, cymules 2-flowered, petals ca. 9 mm, with rounded apex and basally with trichomes, and indumentum glandular. It has been considered a synonym of Geranium moupinense. However, the latter has alternate leaves and reticulate mericarps. Unfortunately, it has not been possible to examine any specimens belonging to this species.

1 Plants annual or rarely biennial   (2)
+ Plants perennial   (8)
       
2 (1) Leaves palmately divided.   1 G. robertianum
+ Leaves palmately cleft   (3)
       
3 (2) Petals purplish, with a dark basal spot.   2 G. ocellatum
+ Petals white, pale purple, or dark purple, without a basal spot   (4)
       
4 (3) Cymules umbelliform, in dense aggregates at apex of each branch; mericarps densely covered with 0.5-1.8 mm nonglandular and sometimes glandular trichomes.   12 G. carolinianum
+ Cymules solitary, axillary; mericarps glabrous or with 0.2-0.5 mm nonglandular trichomes   (5)
       
5 (4) Mericarps smooth   (6)
+ Mericarps with transverse ribs   (7)
       
6 (5) External whorl of stamens lacking anthers and with shorter filaments; petal apices emarginate.   10 G. pusillum
+ Both whorls of stamens with anthers and filaments of similar length; petal apices rounded.   13 G. rotundifolium
       
7 (5) Sepal mucros 0.7-1 mm; mericarps with to 0.5 mm appressed nonglandular trichomes.   8 G. divaricatum
+ Sepal mucros 0.1-0.2 mm; mericarps glabrous.   9 G. molle
       
8 (1) Mericarp reticulated   (9)
+ Mericarp smooth or with some transverse veins   (13)
       
9 (8) Basal cauline leaves opposite or whorled   (10)
+ Basal cauline leaves alternate   (11)
       
10 (9) Petals (1-)1.2-1.5 cm; leaves deeply divided, ratio of main sinus/middle segment length = (0.69-)0.8-0.88.   5 G. strictipes
+ Petals 0.9-1(-1.1) cm; leaves not deeply divided, ratio of main sinus/middle segment length = (0.55-)0.67-0.76(-0.8).   4 G. hispidissimum
       
11 (9) Flowers in solitary 2-flowered cymules; sepal mucros 1.7-1.8 mm.   6 G. moupinense
+ Flowers in several cymules at least partly arranged in pseudoumbels; sepal mucros 0.4-0.9(-1.4) mm   (12)
       
12 (11) Cymules 2(or 3)-flowered, in dense umbel-like aggregates at apex of each branch, peduncle usually absent; fruit rostrum without narrowed apex or rarely with narrowed apex to 1 mm.   3 G. polyanthes
+ Cymules (2 or)3(or 4)-flowered, in loose umbel-like aggregates at apex of each branch, peduncle 2.5-6.7 cm; fruit rostrum with narrowed apex ca. 2 mm.   7 G. umbelliforme
       
13 (8) Cymules usually with 1 flower   (14)
+ Cymules usually with 2(or 3) flowers   (17)
       
14 (13) Petals (1.2-)1.4-1.8(-2) cm; stipules broadly ovate, connate.   48 G. hayatanum
+ Petals (0.4-)0.5-0.8(-1.3) cm; stipules lanceolate to subulate, distinct   (15)
       
15 (14) Leaf blade middle segment obtriangular, 3-5-lobed; petals (5.2-)7.2-8.4(-12.7) mm.   47 G. suzukii
+ Leaf blade middle segment rhombic, 6-14-lobed; petals 4-5.9(-6.3) mm   (16)
       
16 (15) Leaf blade middle segment narrowly rhombic; sepal mucros long, ratio of mucro length/sepal length = (0.17-)0.23-0.28(-0.34).   15 G. sibiricum
+ Leaf blade middle segment broadly rhombic; sepal mucros short, ratio of mucro length/sepal length = (0.09-)0.14-0.21.   16 G. nepalense
       
17 (13) Petals reflexed   (18)
+ Petals not reflexed   (22)
       
18 (17) Pedicel glandular trichomes purple, coarse, dense.   18 G. refractum
+ Pedicel glandular trichomes colorless or with purple heads, fine, dense to sparse or absent   (19)
       
19 (18) Nectaries forming a ring around flower; petal basal margin with very few trichomes.   19 G. sinense
+ Nectaries 5, separate; petal basal margin with numerous trichomes   (20)
       
20 (19) Petals with trichomes only on basal margin.   20 G. shensianum
+ Petals with trichomes on basal margin and inside base   (21)
       
21 (20) Filaments almost glabrous or with a few 0.3-0.8(-1.2) mm trichomes; petals (0.6-)0.8-1(-1.2) cm, with an evident whitish basal zone.   21 G. delavayi
+ Filament base with numerous 1-1.8 mm trichomes; petals (1.2-)1.3-1.7 cm, without a whitish basal zone.   22 G. pogonanthum
       
22 (17) Stem arising from subglobose tubercles connected together by thin rootstock; cauline leaves alternate   (23)
+ Stem arising from rootstock without tubercles; cauline leaves opposite or alternate   (25)
       
23 (22) Petal apices deeply emarginate; leaves palmately divided; mericarps without a basal callus.   11 G. linearilobum
+ Petal apices rounded or rarely shallowly emarginate; leaves palmately cleft; mericarps with a basal callus   (24)
       
24 (23) Petals 0.6-1.2 cm; sepals 5.3-6.3 mm.   25 G. canopurpureum
+ Petals 1.6-1.8 cm; sepals 7.1-10.9 mm.   24 G. pylzowianum
       
25 (22) Cauline leaves alternate (upper usually opposite)   (26)
+ Cauline leaves opposite   (31)
       
26 (25) Staminal filaments with 1.5-3.2 mm nonglandular trichomes; rostrum narrowed apex 4.8-9.8 mm   (27)
+ Staminal filaments with 0.1-0.9 mm nonglandular trichomes; rostrum narrowed apex 0.9-3 mm   (28)
       
27 (26) Stem and petioles with retrorse appressed trichomes; leaves deeply divided.   29 G. erianthum
+ Stem and petioles with patent trichomes; leaves not deeply divided.   30 G. platyanthum
       
28 (26) Petals 4-6.2(-6.9) mm; leaf blade usually with 3 segments.   14 G. wilfordii
+ Petals 8.1-12.6(-15.6) mm; leaf blade with 5(or 7) segments   (29)
       
29 (28) Petals white, apex emarginate.   31 G. albiflorum
+ Petals purplish, apex rounded   (30)
       
30 (29) Leaf blade deeply divided, ratio of main sinus/middle segment length = (0.81-)0.87-0.91; leaf blade middle segment 8-22-lobed in distal half.   32 G. pseudosibiricum
+ Leaf blade not deeply divided, ratio of main sinus/middle segment length = 0.78-0.82; leaf blade middle segment 7-13-lobed in distal half.   50 G. franchetii
       
31 (25) Petals 0.4-0.8(-0.9) cm; anthers 0.3-1.1 mm   (32)
+ Petals (0.8-)1-2(-2.8) cm; anthers (0.9-)1.4-3.4 mm   (34)
       
32 (31) Leaf blade of cauline leaves with 3 segments.   14 G. wilfordii
+ Leaf blade of cauline leaves with 5 segments   (33)
       
33 (32) Sepals with glandular trichomes; petals (6.8-)7.3-8.3(-9) mm.   17 G. thunbergii
+ Sepals without glandular trichomes; petals 5.1-5.9(-6.3) mm.   16 G. nepalense
       
34 (31) Staminal filaments with a broadly triangular base and an abruptly narrowed apex   (35)
+ Staminal filaments lanceolate or sometimes slightly dilated at base   (38)
       
35 (34) Immature fruit reflexed   (36)
+ Immature fruit erect   (37)
       
36 (35) Leaf blade (4.2-)6.2-11.5 cm, deeply divided, ratio of main sinus/middle segment length = (0.81-)0.86-0.94; sepal with a (1.7-)2.2-3.9 mm mucro.   33 G. pratense
+ Leaf blade 2.2-3.8(-5.5) cm, not deeply divided, ratio of main sinus/middle segment length = 0.65-0.79(-0.86); sepal with a 0.7-1.3(-1.9) mm mucro.   34 G. himalayense
       
37 (35) Petals (1.6-)1.8-2.8 cm, apex emarginate; sepals (8.4-)9-10.3(-11.3) mm.   36 G. saxatile
+ Petals (0.8-)1-1.6(-1.8) cm, apex rounded; sepals 4.8-7.3(-9.3) mm.   35 G. collinum
       
38 (34) Staminal filaments and stigma blackish   (39)
+ Staminal filaments and stigma white or purplish   (40)
       
39 (38) Petals 1.2-1.4(-1.6) cm; staminal filaments with 0.2-0.8 mm trichomes.   38 G. christensenianum
+ Petals 2-2.4 cm; staminal filaments with 1.5-2 mm trichomes.   37 G. lambertii
       
40 (38) Stipules ovate, connate for most of their length   (41)
+ Stipules lanceolate or broadly lanceolate, distinct or sometimes connate for part of their length   (42)
       
41 (40) Petals with trichomes inside only at base, apex rounded; plants erect.   43 G. koreanum
+ Petals with trichomes inside on basal 1/3, apex emarginate; plants trailing to ascending.   39 G. wallichianum
       
42 (40) Dwarf alpine plants 4-20(-40) cm tall; stipules distinct   (43)
+ Plants with an erect stem (21-)50-129 cm tall; stipules connate or distinct   (45)
       
43 (42) Rootstock with several napiform roots clustered around it; leaf blade abaxially with glandular trichomes.   26 G. napuligerum
+ Rootstock without napiform roots clustered around it; leaf blade without glandular trichomes   (44)
       
44 (43) Petals very pale pink, claw 2.3-6.1 mm, apex rounded.   28 G. farreri
+ Petals deep pink, without claw, apex emarginate.   27 G. donianum
       
45 (42) Petal apices emarginate; stipules distinct.   49 G. rectum
+ Petal apices rounded; stipules usually connate or connate at least at base   (46)
       
46 (45) Rootstock horizontal or vertical, long, with thickened roots along it   (47)
+ Rootstock vertical, short, with a fascicle of long and thickened roots   (48)
       
47 (46) Petals (1.3-)1.6-1.9(-2.1) cm, outside with trichomes only at base; anthers blackish.   23 G. yunnanense
+ Petals (1-)1.2-1.4 cm, outside with trichomes on basal 1/3-1/2; anthers purplish.   46 G. rosthornii
       
48 (46) Leaf blade deeply divided, ratio of main sinus/middle segment length = (0.81-)0.86-0.96   (49)
+ Leaf blade not deeply divided, ratio of main sinus/middle segment length = 0.7-0.81(-0.84)   (51)
       
49 (48) Petals (1.3-)1.4-2 cm.   42 G. soboliferum
+ Petals (0.8-)0.9-1.3(-1.5) cm   (50)
       
50 (49) Stigmatic remains 4.6-6.6 mm; nectaries apically with a tuft of trichomes; sepals (6.2-)7-10 mm; middle leaf blade segment (4.3-)5.4-8.8 mm wide at base.   41 G. krameri
+ Stigmatic remains 2.8-4.5 mm; nectaries usually glabrous; sepals 5.5-6.9(-9) mm; middle leaf blade segment 1.6-4.8(-5.7) mm wide at base.   40 G. dahuricum
       
51 (48) Sepals (7.9-)9.4-11.2 mm; petals (1.5-)1.6-2.1 cm; leaf blade shallowly divided, ratio of second sinus/middle segment length = (0.13-)0.2-0.24(-0.25); nectaries apically with a tuft of trichomes; stigmatic remains 5-6.9 mm.   44 G. wlassovianum
+ Sepals 5.6-7.5(-7.9) mm; petals 1.1-1.4(-1.5) cm; leaf blade shallowly divided, ratio of second sinus/middle segment length = (0.11-)0.15-0.17(-0.21); nectaries glabrous; stigmatic remains 3.5-4.4 mm.   45 G. maximowiczii

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