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FOC | Family List | FOC Vol. 19 | Rubiaceae

28. Galium Linnaeus, Sp. Pl. 1: 105. 1753.

拉拉藤属 la la teng shu

Authors: Tao Chen & Friedrich Ehrendorfer

Galium verum

Credit: Harvard University Herbaria

Subshrubs to perennial or annual herbs. Stems often weak and clambering, often notably prickly or "sticky" (i.e., retrorsely aculeolate, "velcro-like"). Raphides present. Leaves opposite, mostly with leaflike stipules in whorls of 4, 6, or more, usually sessile or occasionally petiolate, without domatia, abaxial epidermis sometimes punctate- to striate-glandular, mostly with 1 main nerve, occasionally triplinerved or palmately veined; stipules interpetiolar and usually leaflike, sometimes reduced. Inflorescences mostly terminal and axillary (sometimes only axillary), thyrsoid to paniculiform or subcapitate, cymes several to many flowered or infrequently reduced to 1 flower, pedunculate to sessile, bracteate or bracts reduced especially on higher order axes [or bracts sometimes leaflike and involucral], bracteoles at pedicels lacking. Flowers mostly bisexual and monomorphic, hermaphroditic, sometimes unisexual, andromonoecious, occasionally polygamo-dioecious or dioecious, pedicellate to sessile, usually quite small. Calyx with limb nearly always reduced to absent; hypanthium portion fused with ovary. Corolla white, yellow, yellow-green, green, more rarely pink, red, dark red, or purple, rotate to occasionally campanulate or broadly funnelform; tube sometimes so reduced as to give appearance of free petals, glabrous inside; lobes (3 or)4(or occasionally 5), valvate in bud. Stamens (3 or)4(or occasionally 5), inserted on corolla tube near base, exserted; filaments developed to ± reduced; anthers dorsifixed. Inferior ovary 2-celled, ± didymous, ovoid, ellipsoid, or globose, smooth, papillose, tuberculate, or with hooked or rarely straight trichomes, 1 erect and axile ovule in each cell; stigmas 2-lobed, exserted. Fruit on pedicels sometimes elongating during development, green, gray, or infrequently white (to red, orange, or black), mostly dry to leathery schizocarps, infrequently spongy, rarely ± fleshy and berrylike, ellipsoid to subglobose; schizocarps separating into 2 indehiscent mericarps, each with 1 seed, subglobose, ellipsoid-oblong, or reniform, smooth and glabrous to tuberculate and/or covered with trichomes often hooked and clinging; seeds small, grooved ventrally (i.e., adaxially); testa membranous; endosperm corneous; embryo curved; cotyledons leaflike; radicle terete, inferior.

More than 600 species: worldwide, mostly in meridional to temperate but also in alpine and arctic regions or in subtropical and tropical zones at higher elevations; 63 species (23 endemic, four of unconfirmed occurrence) in China.

Galium is by far the largest and most widespread genus within the tribe Rubieae (subfamily Rubioideae). According to the most recent contributions (Natali et al., Opera Bot. Belg. 7: 193-203. 1996; Ehrendorfer et al., Fl. Iranica 176: 1-287. 2005; Bremer & Eriksson, Int. J. Pl. Sci. 170: 766-793. 2009; Soza & Olmstead, Taxon 59: 755-771. 2010), this tribe is closest to Theligoneae, Putorieae, and Paederieae, and includes the following genera treated (or mentioned) in the present flora: Asperula, Cruciata Miller, Galium, Kelloggia, Leptunis, Microphysa, Phuopsis, Rubia, and Sherardia Linnaeus.

So far, the genera Cruciata and Sherardia have not been found in China yet but may be expected there because of their partly weedy character and widely adventive occurrence. They are included in the key below for future reference but not among the full generic presentations. Sherardia arvensis Linnaeus is widely distributed in warm temperate and high-elevation tropical regions and can be separated from Asperula, Phuopsis, Leptunis, or Galium by its terminal capitate inflorescences enclosed by leaflike bracts, its clearly developed calyx with 6 acute lobes, and its pink or violet corollas with well-developed funnelform tubes and 4 lobes.

Among the few Cruciata species, the W Eurasiatic C. pedemontana (Bellardi) Ehrendorfer appears occasionally as an adventive in warm temperate regions. It is common, e.g., in SE North America, and could be found in China too. Cruciata can be separated from Galium by its flowering stems with vegetative apices and the inflorescences consisting only of lateral axillary cymes on middle and lower stem nodes. These cymes are equal to or shorter than the subtending leaves when fully developed. In contrast, the inflorescences are mostly terminal and axillary and longer than the leaves in Galium.

The characters relevant for the taxonomy of Galium and other Rubieae deserve some comments. Life and growth forms are important, particularly with respect to the differentiation into half-shrubs, herbaceous perennials, and annuals. Stem and leaf posture, consistency, shape, and indumentum (e.g., pubescent or retrorsely aculeolate with recurved microhairs) are often quite diverse and may vary within species or even populations. The true leaves are always opposite and 2, but interpetiolar stipules may vary from inconspicuous and divided or simple to enlarged and leaflike, forming whorls of 4 or up to 6 and more. During seedling and shoot development all these taxa pass through the 2- and 4-whorl stage, but some taxa remain at this stage, while others continue to develop more numerous whorl elements toward the middle of their stems. This is a most informative differential character within Rubieae. Other relevant features relate to leaf shape, venation, texture, and particularly indumentum. Here, the presence of longer or shorter microhairs (use a lens) on surfaces and particularly margins as well as their forward or backward direction is of taxonomic importance.

Other morphological characters decisive for Rubieae taxonomy concern the inflorescences (e.g., the position and structure of the cymes). Flower shape is essential for the traditional separation of the genera Asperula (with salverform, funnelform, or cup-shaped corollas) and Galium (with ± rotate corollas). It is now clear that there are transitions between these character states and that even closely related taxa may differ in this respect. So far, it has been possible to provisionally maintain Asperula and Galium by the transfer of obviously misplaced taxa and by using the presence or absence of bracts and bracteoles as a differential character for the two genera (see Ehrendorfer et al., loc. cit. 2005).

The indumentum of ovaries and fruit as well as fruit consistency also vary strongly within Rubieae. Informative are, for example, ± fleshy berries (as in Relbunium Bentham & J. D. Hooker, Rubia, and certain Galium taxa) vs. dry schizocarps or the presence vs. absence of hairiness and whether the trichomes are hooked (i.e., the fruit disperse as "stick-tights" on animals) vs. straight. However, the distinction between all these structures is arbitrary, and there are even transitions between trichomes and tuberculate protuberances of various shapes as well as between hairy and glabrous. All this is well illustrated by Yang and Li (Bull. Natl. Mus. Nat. Sci., Taichung 11: f. 1. 1998). Furthermore, ovary and fruit indumentum and surface structures may change during development and sometimes vary genetically within species or even within populations, as in several Galium species. In general, authors in other regions have documented infraspecific variation from glabrous to densely hairy or tuberculate fruit but traditionally have only separated plants with hooked trichomes into different species. However, intrepid Chinese authors have easily combined these latter morphotypes, e.g., in G. dahuricum sensu W. C. Chen (in FRPS 71(2): 255. 1999), whereas Fl. Japan (3a: 238-239. 1993) distinguished G. manshuricum on the basis of this character. Only careful studies and field observations can clarify such cases, as in G. spurium, where the infraspecific variation of fruit, either smooth, tuberculate, or covered with hooked hairs, has been proven.

Further differential characters for the taxonomy of Rubieae come from the fields of palynology (e.g., number of colpi), karyology (e.g., deviations from the normal chromosome base number x = 11 in Asperula sect. Cynanchicae (Candolle) Boissier with x = 10 or in Galium sect. Aparinoides (Jordan) Grenier with x = 12; common occurrence of polyploidy), and reproductive biology. Most of the perennial Rubieae taxa have conspicuous hermaphroditic or andromonoecious flowers and inflorescences and are insect-pollinated and self-incompatible outbreeders (e.g., Phuopsis or G. boreale and G. verum). Nevertheless, for several annuals with small and inconspicuous flower aggregates selfing and autogamy have been documented (e.g., G. aparine, G. spurium, and Sherardia arvensis). Furthermore, polygamodioecy and dioecy occur in some groups (e.g., G. elegans). Up to now, only few and insufficient data from all these fields are available for Asian Rubieae species and have not been mentioned in FRPS. Nevertheless, such data are significant and will have to be addressed in more detailed future systematic Rubieae studies from this region.

The α-taxonomy of Rubieae in E Asia is still in a problematic state. A general survey of the collections at the herbaria KUN, MO, PE, W, and WU has revealed the existence of many very polymorphic, complex, and insufficiently understood species groups. Therefore, the present treatment has to be regarded as provisional.

A particularly critical case concerns several Galium species described by H. Léveillé from 1904-1917 (see Lauener & Ferguson, Notes Roy. Bot. Gard. Edinburgh 32: 103-115. 1973). These descriptions are most fragmentary and the relevant types are not yet studied sufficiently (but see Mill, Edinburgh J. Bot. 53: 193-213. 1996). Relevant taxa in alphabetical order are G. blinii (see under that name), G. bodinieri (see under G. blinii and G. rebae), G. cavaleriei (see under G. asperifolium), G. comarii (see under G. dahuricum), G. esquirolii (see under G. asperifolium), G. hongnoense (see under G. spurium), G. mairei (see under G. elegans), G. martini (see under G. bungei), G. quinatum (see under G. blinii), G. remotiflorum (see under G. bungei), and G. venosum (see under G. bungei).

The treatment of Galium for the Flora of Taiwan by Yang and Li (Bull. Natl. Mus. Nat. Sci., Taichung 11: 101-117. 1998; Fl. Taiwan, ed. 2, 4: 254-259. 1998) is not satisfactory in several aspects: keys and descriptions are rather idealized and lack carefully observed ranges of morphological variation for the taxa; species are circumscribed more narrowly and based on different characters than used by other authors in the region (e.g., presence vs. absence of leaf indumentum is considered variable within species by most other authors); the treatment is not well reconciled with continental Galium taxonomy (e.g., there are no references to the Russian floras, and names synonymized by others are used without explanation); and at least two names based on types from Taiwan are missing.

With respect to a more "natural" and general taxonomic classification of the Rubieae and Galium, a number of recent morphological, karyological, palynological, and particularly DNA-analytical studies (e.g., Natali et al., loc. cit.; Robbrecht & Manen, Syst. & Geogr. Pl. 76: 85-146. 2006; Bremer & Eriksson, loc. cit.; Soza & Olmstead, loc. cit.) are available. They show that Theligonum should be placed into a separate tribe (Theligoneae), that the tribe Rubieae is monophyletic, and that Kelloggia (as subtribe Kelloggiinae, still with normal Rubiaceae stipules, calyx teeth, and 3-colpate pollen, but already with hooked trichomes on the dry mericarps) occupies a basal position in Rubieae. The Central American genus Didymaea J. D. Hooker (still with normal stipules but with the calyx already lacking, 5-coplate pollen, and seeds separating from the fleshy pericarp) represents a link to the genus Rubia in the true Rubiinae. Their stipules are nearly always leaflike, the pollen is polycolpate, and the seeds never separate from the pericarp. Rubia, a well-circumscribed and certainly monophyletic genus, is always perennial, has 5-lobed corollas, and berrylike fruit.

The remaining Rubiinae are also monophyletic as a whole, but their traditional genera Asperula, Bataprine Nieuwland, Callipeltis Steven, Crucianella Linnaeus, Cruciata, Galium, Leptunis, Mericarpaea Boissier, Microphysa, Phuopsis, Relbunium, Sherardia, Valantia Linnaeus, and Warburgina Eig are all essentially interdigitated. They are difficult to separate and can hardly be brought into concordance with available phylogenetic data. These advanced Rubiinae tend to develop more and more apomorphic character profiles, i.e., change from perennial to annual, increase in numbers of leaflike stipules from 4 to numerous, loss of bracts and prophylls in the inflorescences, reduction from 5-lobed to (3 or)4-lobed corollas, specialization of mericarps, etc. As shown by the most comprehensive phylogram available so far (Natali et al., loc. cit.: f. 2; Soza & Olmstead, loc. cit.: f. 1, 2) and new findings (unpubl.), these more apomorphic Rubiinae form a polytomy or a grade with seven parallel clades. The most basal clade (1) consists of the monotypic Galium sect. Cymogalia Pobedimova only. The following Sherardia clade (2) includes Crucianella, Phuopsis, Sherardia, and several sections of Asperula together with Leptunis. Separate clades are formed by G. sect. Depauperata Pobedimova (3), A. sect. Glabella Grisebach, including G. sect. Aparinoides (4), and A. sect. Asperula (5). The Cruciata clade (6) consists not only of the genera Cruciata and Valantia but also of all sections of Galium (including the traditional genera Bataprine, Microphysa, and Relbunium) that form whorls of 2 leaves and normally not more than 2 even-sized leaflike stipules. Finally, the G. sect. Galium clade (7) comprises this and various other sections of Galium, which regularly develop whorls of leaves and leaflike stipules with 5 to more elements.

From the above data and the fact that a number of major groups of Rubiinae have not been DNA-analyzed yet, it is obvious that it is still difficult and partly impossible to harmonize DNA-supported clades with the traditional genera and sections. Thus, extensive changes are expected for generic and sectional circumscriptions within Rubiinae in the future. Therefore, we refrain from taxonomic changes for the present flora, list taxa in alphabetical order, and only supplement phylogenetic comments. Thus, the present treatment in principle follows FRPS (71(2): 216-286. 1999), mainly based on Pobedimova et al. (Fl. URSS 23: 287-381. 1958), but also considers Ehrendorfer et al. (loc. cit. 2005). In order to make comparison with available phylogenetic data and present infrageneric classification easier, relevant information is inserted as a "Taxonomic Conspectus" before the individual species descriptions. It was not until this volume was ready for the press that the need for the nomen novum, Galium glabriusculum, was discovered; therefore, this species alone is outside of the alphabetical order.

Here the key to species of Galium is extensively revised from that of FRPS. It includes all of the Chinese Galium species with full ranges of differential character variation. Furthermore, it keys out all other Rubieae genera which are easily confused with Galium and are documented or can be expected in China. Details on the genera Asperula, Leptunis, Microphysa, Phuopsis and Rubia can be found where they are listed in alphabetical order, references to Cruciata and Sherardia appear in the comments above.

Several species are keyed out more than once in the present key because they are circumscribed by combinations of characters rather than by unique features. Furthermore, many Galium species are markedly variable because of genetic differentiation (e.g., G. bungei, G. elegans) but also because of phenetic plasticity due to different environmental conditions. References to the number of leaves and leaflike stipules in whorls as well as leaf measurements refer to middle stem regions. Measurements of organs with hairy surfaces (e.g., leaves, fruit, and mericarps) here apply to the solid surface of the structure and do not include the trichomes. The terms "leaf whorl," "ovary," and "uncinate trichome" follow common usage in Galium. Infraspecific taxa are adopted from FRPS in order to facilitate future and more detailed work on this group and comparison with other floras. They are not included in the following main key but are subordinated under the relevant species in alphabetical order and keyed out there.

Taxonomic conspectus of the Rubieae (excluding Kelloggia and Rubia)

In FRPS (71(2): 216-286. 1999) the taxa of Galium were placed in the following sections (designated here by capital letters): G. sect. Depauperata (A), G. sect. Aparine (B), G. sect. Pseudaparine Lange (C), G. sect. Cymogalia (D), G. sect. Trachygalium (E), G. sect. Leptogalium Lange (F), G. sect. Platygalium (G), G. sect. Galium (H), G. sect. Leiogalium (I), G. sect. Trichocarpa (Pobedimova) Pobedimova (J), G. sect. Asperuloides Pobedimova (K), and G. sect. Brachyantha (Boissier) Pobedimova (L). Some species of uncertain position were classified as dubious (M). Species accepted in the present treatment but lacking in FRPS are designated as (Z). For comparison, a current (e.g., Jelenevsky et al., Novosti Sist. Vyssh. Rast. 35: 174-187. 2003; Ehrendorfer et al., Fl. Iranica 176: 1-287. 2005), DNA-analytically supported (e.g., Natali et al., Opera Bot. Belg. 7: 193-203. 1996; Soza & Olmstead, Taxon 59: 755-771. 2010; Ehrendorfer, unpubl.) but still provisional taxonomic conspectus is presented below. It lists all species here accepted under their clades and sections. For the clades 1-6 one should compare the comments above, for the sections compare Ehrendorfer et al. (loc. cit.). The placement of Galium species into sections (or lack of placement) by FRPS is indicated by showing the relevant letters used above in parentheses after the species names.

Clade 1

Galium sect. Cymogalia Pobedimova s.s.

39. Galium paradoxum (D)

Clade 2

Phuopsis stylosa

Sherardia arvensis

Leptunis trichodes

Asperula oppositifolia

Clade 3

Galium sect. Depauperata Pobedimova

16. Galium exile (A; incl. G. songaricum sensu FRPS)

Clade 4

Galium sect. Aparinoides (Jordan) Grenier

27. Galium karakulense (E)

25. Galium innocuum (as G. trifidum: E; incl. "G. palustre")

Clade 5

Asperula sect. Asperula

Asperula orientalis

Clade 6

Galium sect. Platygalium W. D. J. Koch s.l.

9. Galium bungei (E; incl. G. martini: M)

45. Galium salwinense (E)

12. Galium crassifolium (A)

31. Galium linearifolium (E)

21. Galium hirtiflorum (Z)

20. Galium glandulosum (A)

18. Galium forrestii (D)

44. Galium rupifragum (Z)

35. Galium morii (D)

54. Galium tarokoense (A)

34. Galium minutissimum (M)

36. Galium nankotaizanum (M; incl. G. maborasense)

47. Galium serpylloides (A)

29. Galium kinuta (G)

24. Galium hupehense (M)

30. Galium kunmingense (Z)

17. Galium formosense (as G. kwanzanense: M)

15. Galium elegans (D)

63. Galium yunnanense (M)

26. Galium kamtschaticum (D)

10. Galium chekiangense (as G. nakaii: G)

Microphysa elongata

40. Galium platygalium (K)

32. Galium maximoviczii (K)

7. Galium boreale s.l. (G)

60. Galium turkestanicum (M)

Clade 7

[Galium s.s.]

Galium sect. Hylaea (Grisebach) Ehrendorfer s.l.

37. Galium odoratum (J)

4. Galium asperuloides (A)

22. Galium hoffmeisteri (as subsp. of G. asperuloides: A)

14. Galium echinocarpum (A)

53. Galium takasagomontanum (M)

59. Galium triflorum (A)

58. Galium trifloriforme (Z)

Galium sect. Trachygalium K. Schumann s.l.

48. Galium sichuanense (Z)

13. Galium dahuricum (as "G. davuricum": F; incl. G. comarii, G. niewerthii, G. pseudoasprellum)

56. Galium tokyoense (as var. of "G. davuricum": F)

41. Galium prattii (M)

52. Galium taiwanense (M)

3. Galium asperifolium (I)

6. Galium blinii (as syn. of G. asperifolium var.: I; incl. G. quinatum: M)

51. Galium sungpanense (A)

42. Galium pusillosetosum (A)

1. Galium acutum (M)

43. Galium rebae (Z)

33. Galium megacyttarion (Z)

5. Galium baldensiforme (A)

49. Galium glabriusculum (A)

28. Galium karataviense (as G. rivale s.l.: K)

61. Galium uliginosum (F)

Galium sect. Leiogalium Ledebour

38. Galium paniculatum (M; incl. G. xinjiangense: J)

Galium sect. Orientigalium Ehrendorfer

8. Galium bullatum (I)

Galium sect. Galium

23. Galium humifusum (L)

62. Galium verum (H)

11. Galium consanguineum (as G. majmechense: H)

46. Galium saurense (M)

Galium sect. Aparine W. D. J. Koch s.s.

50. Galium spurium (as G. aparine var. tenerum: B)

2. Galium aparine (B)

Galium sect. Kolgyda Dumortier s.s.

57. Galium tricornutum (as G. tricorne: B)

Galium sect. Microgalium Grisebach

19. Galium ghilanicum (Z)

55. Galium tenuissimum (C)

Key to species of Galium and to related genera of the Rubieae

1 (1) Interpetiolar stipules inconspicuous, multifid or fimbriate, not leaflike and not forming whorls with true leaves; corolla funnelform, (4 or)5-lobed; ovary and dry mericarps with hooked trichomes.   Kelloggia (see p. 183)
+ Interpetiolar stipules mostly leaflike and in whorls with true leaves, rarely reduced   (2)
2 (2) Corolla lobes regularly 5; fruit fleshy, mericarps berrylike, 2(or 1, by non-development), often dispersed together.   Rubia (see p. 305)
+ Corolla lobes usually 4 (rarely 3); fruit dry or leathery, mericarps mostly 2, nearly always separating for dispersal   (3)
3 (2) Leaves in middle stem region opposite, with stipules reduced or ± leaflike and in whorls of 4 but then always clearly smaller than true leaves   (4)
+ Leaves in middle stem region opposite and with very similar leaflike stipules in whorls of 4-16   (6)
4 (4) Corolla pink, funnelform, with well-developed tube longer than lobes; fruit smooth Asperula oppositifolia (see p. 78)   (4)
+ Corolla white, rotate, with tube shorter than lobes; fruit with uncinate trichomes   (5)
5 (4) Perennial herbs; leaves 5-30 × 5-23 mm, obtuse to truncate at base, on petioles 1.5-10 mm; flowers 3-11 in cymes; corolla with 4 lobes.   39 G. paradoxum
+ Annual herbs; leaves 2-12 × 1-4 mm, acute to cuneate at base, subsessile or on short petioles; flowers solitary at each node; corolla mostly with 3 lobes.   16 G. exile
6 (3) Leaf apex rounded, obtuse, or ± blunt, never acute or with a hyaline mucro; leaves in whorls of 4-6, linear to broadly oblong, 1-nerved, dried blackening; ripe mericarps globose, didymous and only with a short zone of contact, glabrous; corolla cup-shaped to slightly campanulate, 3- or 4-merous   (7)
+ Leaf apex mostly ± acute, often with a hyaline mucro; leaves in whorls of 4-16, sometimes broader and with 3-5 palmate nerves; ripe mericarps ovoid to subglobose, with a longer zone of contact and with diverse surface structures; corolla diverse, but often rotate and always 4-merous   (8)
7 (6) Inflorescences with many-flowered cymes; corolla 4-lobed, 2.5-4 mm in diam.; leaves mostly 15-20 × 5-8 mm.   27 G. karakulense
+ Inflorescences with 1-3(or 4)-flowered cymes; corolla mostly 3-lobed, 1-1.8 mm in diam.; leaves mostly 3-8 × 1-2 mm.   25 G. innocuum
8 (6) Leaves and leaflike stipules in middle stem region never in whorls of more than 4 (if rarely in whorls of up to 6 then leaves with 3-5 palmate principal veins), from linear to broadly ovate   (9)
+ Leaves and leaflike stipules in middle stem region regularly in whorls of more than 4, i.e., in whorls of 5-16, with only 1 principal vein, linear to broadly lanceolate or elliptic, but never ovate or with a length/breadth index of less than 2.5   (44)
9 (9) Stem apex vegetative, with few- to several-flowered lateral cymes only in leaf axils and shorter than or ± equal to subtending leaves, nodding in fruit.  

Cruciata (see comments above)

+ Stem apex usually floriferous, with terminal and axillary cymes, often longer than subtending leaves and mostly not nodding in fruit   (10)
10 (9) Condensed plants of rocks or high elevations; stems usually less than 10 cm tall, glabrous or with spreading (but never retrorse) hairs; leaves mostly ± ovate, (1-)3-8(-20) × (0.8-)2-4(-10) mm, with 1-3 main veins; corolla rotate, often only 1.5-2 mm in diam.; mericarps with spreading (rarely appressed) hooked or ± straight trichomes   (11)
+ Taller plants, usually of lower elevations with larger leaves (if plants ± condensed then stem hairs retrorsely curved or fruit hairs appressed but not hooked)   (18)
11 (10) Mericarps with ± straight hairs, 2-2.5 mm in diam.; stems mostly pilose or hirtellous   (12)
+ Mericarps with weakly to strongly curved and uncinate trichomes; stems partly glabrous   (13)
12 (12) Fruiting pedicels straight; Xizang.   47 G. serpylloides
+ Fruiting pedicels nodding; Taiwan.   36 G. nankotaizanum
13 (11) Corolla ca. 3 mm in diam.; stems glabrous.   53 G. takasagomontanum
+ Corolla 1.2-2 mm in diam.; stems ± hairy or glabrous   (14)
14 (13) Stems ± hairy   (15)
+ Stems glabrous; Taiwan   (16)
15 (14) Leaves ovate to broadly lanceolate, acute, up to 3.5 mm wide; Yunnan.   44 G. rupifragum
+ Leaves broadly elliptic to obovate, obtuse and mucronate, up to 10 mm wide; Taiwan.   17 G. formosense
16 (14) Leaves very small, only 0.8-1 mm wide, with 1 main vein only; fruit hairs spreading.   34 G. minutissimum
+ Leaves wider, with 1 or 3 main veins; fruit hairs appressed   (17)
17 (16) Leaves with 3 main veins; corolla only ca. 1.2 mm in diam.   35 G. morii
+ Leaves with 1 main vein; corolla ca. 2 mm in diam.   54 G. tarokoense
18 (10) Leaves with 1 principal vein or 2 lateral veins only weakly visible and not extending past middle of blade   (19)
+ Leaves with 3-5 palmate principal veins, lateral veins well marked and extending for more than half of blade length   (29)
19 (19) Open corollas funnelform, 2.5-3 mm, tube somewhat shorter than lobes; fruit with pericarp smooth to granular, becoming slightly inflated, enclosing both mericarps at dispersal.   Microphysa elongata (see p. 216)
+ Open corollas rotate, fused basal part much shorter than lobes; mericarps clearly separated   (20)
20 (19) Stems ± strigose-hirsute, with hairs ± retrorse (but not retrorsely aculeolate); leaves ovate or elliptic to linear-lanceolate, broadest ± in middle, at lower side usually with glandlike spots; flowers unisexual, usually yellowish, ± greenish, or reddish; fruit normally with uncinate trichomes   (21)
+ Stems glabrous or with indumentum, but not with retrorse hairs; leaves often broadest above middle and thinner, without glandlike spots; flowers usually bisexual   (23)
21 (20) Plants usually less than 15 cm tall, strongly branched from base; leaves often less than 8 mm, mostly glabrescent or glabrous, subleathery; inflorescences with few-flowered, bracteate cymes.   20 G. glandulosum
+ Plants usually more than 15 cm tall, little branched; leaves usually longer than 8 mm, hairy on both sides; inflorescences paniculate to corymbiform, little bracteate   (22)
22 (21) Leaves linear-elliptic to narrowly lanceolate, mostly 8-17 × 1-2.5 mm, dried rather papery; inflorescence paniculate.   21 G. hirtiflorum
+ Leaves ovate-elliptic, mostly 8-12 × 3-5 mm, dried rather subleathery; inflorescence corymbiform.   18 G. forrestii
23 (20) Leaves ± linear, often longer than 20 mm, in addition to 1 principal, with 2 weaker lateral veins; corolla 4-5 mm in diam   (24)
+ Leaves not linear, mostly shorter than 20 mm; corolla smaller   (25)
24 (23) Leaves linear-spatulate, 1-4 mm wide; inflorescences loose, broadly paniculiform; ovaries and fruit glabrous and smooth.   31 G. linearifolium
+ Leaves linear-lanceolate, 3-9 mm wide; inflorescences dense, elongate-paniculate; ovaries and fruit with sparse hooked trichomes or glabrous.   60 G. turkestanicum
25 (23) Leaves ovate, length/breadth index 2 or less, in addition to 1 principal, with 2 weaker lateral veins; corolla larger; fruit with spreading hooked or straight hairs; Taiwan   (26)
+ Leaves ovate-oblong to lanceolate, length/breadth index 2 or more, with only 1 principal vein; corolla 1.5-2 mm in diam.; ovaries and fruit glabrous or with various indumentum   (27)
26 (25) Mericarps with straight trichomes; corolla 2-2.5 mm in diam.; stems pilose or glabrescent.   36 G. nankotaizanum
+ Mericarps with hooked trichomes; corolla ca. 3 mm in diam.; stems glabrous.   53 G. takasagomontanum
27 (25) Leaves dried subleathery; fruit with appressed, ± curved (but not uncinate) hairs; Shanxi.   12 G. crassifolium
+ Leaves dried papery; fruit with various indumentum   (28)
28 (27) Plants ascending, weak, sparsely hairy or glabrous; inflorescence few flowered, peduncles and pedicels very thin and elongated, latter mostly 4-8 mm; fruit with spreading uncinate trichomes.   45 G. salwinense
+ Plant erect, more robust, indumentum diverse; inflorescences ± many flowered, peduncles and pedicels thicker and shorter, latter mostly 2-4 mm; fruit tuberculate, with appressed or spreading hooked trichomes, or more rarely smooth.   9 G. bungei
29 (18) Corolla funnelform or cup-shaped, 2-5 mm in diam., with fused lower part ± as long as lobes; ovaries and fruit glabrous   (30)
+ Corolla rotate, (1-)2-5 mm in diam., with fused base much shorter than lobes   (32)
30 (29) Corolla cup-shaped or campanulate, 2-2.7 mm in diam.; cauline leaves broadly lanceolate, always in whorls of 4.   30 G. kunmingense
+ Corolla funnelform or campanulate, 2.5-5 mm in diam.; middle stem leaves ovate to elliptic, in whorls of 4-6   (31)
31 (30) Open corollas 3.5-5 mm in diam.; cauline leaves usually in whorls of 4, 12-28 mm.   40 G. platygalium
+ Open corollas 2.5-3.5 mm in diam.; cauline leaves in whorls of 4-6(-8), 23-53 mm.   32 G. maximoviczii
32 (29) Leaves linear to linear-lanceolate, 27-40 × 3-9 mm, in addition to 1 principal, with 2 weaker lateral veins; corolla 4-5 mm in diam.; ovaries and fruit with sparse uncinate trichomes or glabrous.   60 G. turkestanicum
+ Leaves lanceolate to ovate, shorter, 3 principal veins mostly readily visible   (33)
33 (32) Fruit glabrous, smooth to granular-papillose, or with appressed and ± hooked or with spreading and straight (but never with spreading and hooked) trichomes   (34)
+ Fruit with ± spreading and uncinate trichomes   (39)
34 (33) Open corollas 3-4 mm in diam.; stems (except nodes) mostly glabrous and smooth   (35)
+ Open corollas (1-)2-2.5 mm in diam.; stems glabrous or ± hairy   (36)
35 (34) Leaves ovate-lanceolate to ovate, papillose, length/breadth index mostly 2.5 or less; cymes rather few flowered; ovaries and fruit with ± appressed, apically somewhat bent trichomes.   10 G. chekiangense
+ Leaves mostly rather narrowly lanceolate, smooth or ± hairy, length/breadth index mostly 3 or more; cymes many flowered; ovaries and fruit glabrous or with various indumentum.   7 G. boreale
36 (34) Cauline leaves broadly to narrowly lanceolate, length/breadth index often 3.5 or more   (37)
+ Cauline leaves narrowly to broadly ovate-lanceolate, length/breadth index usually 3 or less   (38)
37 (36) Stems hairy throughout; leaves lanceolate; fruit with straight hairs or rarely glabrous.   24 G. hupehense
+ Stems (except nodes) glabrous; leaves ovate-lanceolate (sometimes also broader), apex subacute to acuminate, striate-punctate glandular below; fruit glabrous and smooth.   29 G. kinuta
38 (36) Leaves 6-30 × 3-20 mm; fruiting pedicels straight; fruit glabrous or scaberulous; mainland.   15 G. elegans
+ Leaves 4-10 × 2-5 mm; fruiting pedicels nodding; fruit with ± straight trichomes; Taiwan.   36 G. nankotaizanum
39 (33) Open corollas 3 mm or more in diam.; stems (except nodes) often glabrous and smooth   (40)
+ Open corollas 2.5 mm or less in diam.; stems often ± hairy   (42)
40 (39) Leaves lanceolate to ovate-lanceolate or elliptic, length/breadth index 3.5 or more, apex acute to acuminate.   7 G. boreale
+ Leaves ovate, length/breadth index less than 3   (41)
41 (40) Leaf apex obtuse to rounded, usually mucronate; mainland.   26 G. kamtschaticum
+ Leaf apex acuminate; Taiwan.   53 G. takasagomontanum
42 (39) Leaves ovate-lanceolate to narrowly elliptic, with acute apex, length/breadth index normally more than 2.5.   63 G. yunnanense
+ Leaves ovate to broadly elliptic, with obtuse to rounded apex, entire or shortly mucronate, length/breadth index normally less than 2.5   (43)
43 (42) Leaves up to 20 mm wide; plants slender to usually rather robust; mainland.   15 G. elegans
+ Leaves up to 10 mm wide; slender low plants; Taiwan.   17 G. formosense
44 (8) Inflorescences capitate and enclosed by leaflike bracts; corolla funnelform or salverform, 4-15 mm, with 4 or 5 lobes   (45)
+ Inflorescences branched, not enclosed by bracts; corolla rotate, campanulate, or funnelform, 0.5-13 mm, mostly with 4 (rarely 3) lobes   (47)
45 (45) Plants perennial, 20-70 cm tall; calyx limb obsolete; corolla 5-lobed, 12-14 mm.   Phuopsis stylosa (see p. 291)
+ Plants annual; corolla 4-lobed, shorter   (46)
46 (46) Calyx teeth well developed; corolla pink, tube 4-5 mm.  

Sherardia arvensis (see comments above)

+ Calyx lacking; corolla bluish, tube 7-13(-15) mm.   Asperula orientalis (see p. 78)
47 (44) Medium stem leaves marginally (and often on upper side) with microhairs directed forward (use 20× lens), thus antrorsely ciliolate or aculeolate; stems mostly not retrorsely aculeolate   (48)
+ Medium stem leaves marginally (not on leaf surface) with microhairs directed backward (use 20× lens), thus retrorsely aculeolate or completely glabrous and smooth; stems often retrorsely aculeolate   (61)
48 (47) Ovaries and fruit densely covered with uncinate trichomes; plants perennial with smooth stems   (49)
+ Ovaries and fruit glabrous or hairy, but never with uncinate trichomes; plants perennial or annual   (53)
49 (48) Corolla funnelform, with tube ± as long as lobes.   37 G. odoratum
+ Corolla rotate, with tube markedly shorter than lobes   (50)
50 (49) Robust, procumbent to ascending plants often longer than 50 cm; inflorescences terminal and lateral with cymes in upper 2-4 nodes.   59 G. triflorum
+ Slender erect plants, less than 50 cm tall; inflorescences predominantly terminal   (51)
51 (50) Leaves 6-25 × 2-7 mm; open corollas ca. 2 mm in diam.; Taiwan (cf. also 53. G. takasagomontanum).   14 G. echinocarpum
+ Leaves often larger; open corollas 1.5-3.5 mm in diam.; mainland   (52)
52 (51) Leaves mostly in whorls of 7 or 8, (ob)lanceolate, length/breadth index mostly 3.5-4.5, subpetiolate; fruit with uncinate trichomes 0.6-0.8 mm; expected in Xizang.   4 G. asperuloides
+ Leaves mostly in whorls of 6, narrowly obovate to broadly oblanceolate, length/breadth index mostly 2.5-3.5, clearly petiolate; fruit with uncinate trichomes 0.8-1.2 mm or longer; widespread.   22 G. hoffmeisteri
53 (48) Plants annual, slender; flowers on pedicels and peduncles often longer than 5 mm and in lax, diffuse inflorescences   (54)
+ Plants perennial, slender to robust; flowers on pedicels and peduncles 0.5-5 mm, in lax to ± congested inflorescences   (56)
54 (54) Leaves filiform, 20-30 mm, ascending; corolla funnelform, pink to red; ovary and mericarps with dense, short and curved hairs.   Leptunis trichodes (see p. 213)
+ Leaves linear to oblanceolate, 4-20 mm, spreading to reflexed; corolla ± rotate, whitish, yellowish, or greenish; ovary and fruit glabrous or slightly tuberculate   (55)
55 (54) Inflorescences broadly ovate, diffuse and intricate, with fruiting pedicels elongated to 20 mm.   55 G. tenuissimum
+ Inflorescences rather narrowly thyrsoid, not diffuse and intricate, with fruiting pedicels only up to 4 mm.   19 G. ghilanicum
56 (53) Corolla funnelform, lobed for 1/2-2/3, white   (57)
+ Corolla rotate, lobed for 3/4 or more, often yellowish; plants erect to ascending   (58)
57 (56) Inflorescences lax, ± ebracteate; stems erect, smooth; main stem leaves 15-65 × 3-12 mm.   38 G. paniculatum
+ Inflorescences congested, strongly bracteate; stems procumbent, mostly ± hairy; main stem leaves 5-23 × 1-2(-5) mm.   23 G. humifusum
58 (56) Leaves in middle stem region in whorls of not more than 6; plants of (sub)alpine region, not taller than 30 cm.   46 G. saurense
+ Leaves in middle stem region in whorls of more than 6 and up to 12   (59)
59 (58) Open corollas 3.5-5 mm in diam., white; fruit somewhat spongy or fleshy, 3-3.5 mm, with a dry pericarp separating from rest of fruit.   8 G. bullatum
+ Open corollas ca. 3 mm in diam., yellow to whitish; fruit with dry mericarps, 1.5-2 mm, with pericarp dark and firmly attached to rest of fruit   (60)
60 (59) Leaves mostly 2.5-5 mm wide, glabrous abaxially; fruit ca. 1.5 mm.   11 G. consanguineum
+ Leaves 1-2.5 mm wide, glabrous to densely pubescent abaxially; fruit 1.5-2 mm.   62 G. verum
61 (47) Annuals, often in ± disturbed, weedy habitats; stems and leaf margins retrorsely aculeolate; fully developed mericarps subspherical, 2-6 mm; open corollas 1-2 mm in diam   (62)
+ Perennials, in ± natural habitats; fully developed mericarps ellipsoid, 1.5-3 mm; open corollas 1-4 mm in diam   (64)
62 (61) Fruit becoming pendulous on arching peduncles and pedicels, verrucose to spinulose; leaves glabrescent above.   57 G. tricornutum
+ Fruit on divaricate straight peduncles and pedicels (only latter sometimes bent just beneath fruit), mostly with uncinate trichomes; leaves ± hairy above   (63)
63 (62) Open corollas 1.5-2 mm in diam.; individual mature mericarps 2.5-5 mm in diam., with trichomes arising from tuberculate bases.   2 G. aparine
+ Open corollas 1-1.5 mm in diam.; individual mature mericarps 1-3 mm in diam., with trichomes straight from base.   50 G. spurium
64 (61) Middle stem leaves narrowly obovate to oblanceolate, mostly 18-28 × 5-10 mm; stems slightly retrorsely aculeolate; inflorescences of medium size, with terminal and lateral, few- to several-flowered, rather loose cymes with small bracts, in fruit stiffly divaricate; corolla rotate, 1.5-2 mm in diam.; fruit with hooked trichomes.   58 G. trifloriforme
+ Middle stem leaves mostly smaller; inflorescences different, usually more bracteate; fruit smooth, verrucose, or with hooked trichomes   (65)
65 (64) Corolla funnelform to subcampanulate, with tube ± as long as or slightly shorter than lobes, whitish; ovaries and fruit glabrous, smooth or verrucose; leaves papery to subleathery and glossy; stems rough, retrorsely aculeolate, procumbent to clambering   (66)
+ Corolla rotate, fused basal part much shorter than lobes; ovaries and fruit with hooked trichomes, tuberculate, or glabrous; stems glabrous, rough, or hairy   (67)
66 (65) Corolla funnelform; plants robust, 0.6-1.2 m, often forming mats; main stem leaves 8-50 × 2-8 mm.   28 G. karataviense
+ Corolla subcampanulate; plants slender and ± erect, 10-60 cm tall; main stem leaves 3-16 × 1-3 mm.   61 G. uliginosum
67 (65) Middle stem leaves larger, (5-)10-35(-50) × (1-)2.5-10 mm, mostly ± hairy, at least margins retrorsely aculeolate; cymes lateral and terminal, many flowered; ovaries and fruit glabrous or with diverse indumentum; plants from lower elevations, relatively robust, erect or clambering, stems up to 0.7 m tall   (68)
+ Middle stem leaves uniformly small, 2-12(-15) × 0.3-3.5 mm, glabrous and smooth to ± hairy; cymes lateral and terminal, few flowered; ovaries and fruit with uncinate (very rarely ± straight) trichomes or glabrous; plants from high elevations, usually reduced and weak, caespitose to procumbent, stems only up to 0.3 m   (74)
68 (67) Stems branched from base; cymes leafy, with bracts to last branches; stems and leaves (nodes and margins excepted) glabrous and smooth; mericarps with spreading uncinate trichomes.   48 G. sichuanense
+ Stems normally branched from middle; cymes usually less leafy; stems and leaves mostly with more indumentum; fruit glabrous, papillose, tuberculate, or with uncinate trichomes   (69)
69 (68) Peduncles and pedicels slender, filiform and often ± flexuose, with inconspicuous bracts; pedicels up to 5 mm, in fruit elongated to 10 mm or more; flowers never reddish; plants usually clambering; ovary and fruit surfaces diverse   (70)
+ Peduncles and pedicels rather stiff and often divaricate and ± bracteate; pedicels shorter than 5 mm and hardly elongated in fruit; ovary and fruit surfaces diverse, but often glabrous   (71)
70 (69) Fruit often with uncinate trichomes; mainland.   13 G. dahuricum
+ Fruit glabrous; Taiwan.   52 G. taiwanense
71 (69) Corolla red to purple (very rarely maroon or white), 1.5-2.5 mm in diam   6 G. blinii
+ Corolla whitish, yellowish, or greenish   (72)
72 (71) Corolla small, 1.5-2 mm in diam., yellow to greenish white, lobes aristate; inflorescences divaricate and regularly bracteate with bracts similar to but smaller than leaves, giving a diffuse miniature aspect; ovary and fruit surface variable; plants often clambering.   3 G. asperifolium
+ Corolla larger, mostly more than 2 mm in diam.; inflorescences ebracteate or with bracts ± reduced and irregularly distributed; ovary and fruit surface smooth to tuberculate; plants erect, hardly clambering   (73)
73 (72) Leaves lanceolate, gradually narrowed into acute apex.   41 G. prattii
+ Leaves subspatulate to obovate, apex rounded and abruptly narrowed into a mucro.   56 G. tokyoense
74 (67) Ovaries and fruit with uncinate (very rarely ± straight) trichomes   (75)
+ Ovaries and fruit glabrous, smooth, papillose, or verrucose; Himalaya   (78)
75 (74) Leaves and stems ± densely hispid and often retrorsely aculeolate; stems with 4 conspicuous whitish angles.   42 G. pusillosetosum
+ Leaves completely glabrous and smooth or only slightly hairy and/or retrorsely aculeolate; stems with 4 inconspicuous angles   (76)
76 (75) Leaves dried blackening, papery and thin, oblanceolate to narrowly obovate, with flat margins, hardly longer than 7 mm.   5 G. baldensiforme
+ Leaves dried greenish-brownish, with ± revolute margins, often longer than 7 mm   (77)
77 (76) Plants nearly always smooth; leaves ± subleathery; ovary in flower ca. 1 mm.   49 G. glabriusculum
+ Plants retrorsely aculeolate at least on margins and lower side of papery leaves; ovary in flower 0.5-0.8 mm.   51 G. sungpanense
78 (74) Plants weak to procumbent but not mat-forming; cells of adaxial leaf surface relatively large, readily visible individually with 20× lens; corolla mostly whitish.   33 G. megacyttarion
+ Plants procumbent and often mat-forming; cells of adaxial leaf surface small, not or hardly visible individually with 20× lens   (79)
79 (78) Stems ± densely hairy and/or retrorsely aculeolate, with 4 conspicuous whitish angles.   42 G. pusillosetosum
+ Stems glabrous or only slightly retrorsely aculeolate, with inconspicuous angles   (80)
80 (79) Leaves on main stems 2-8.5 mm; inflorescence cymes 1- to few flowered, fascicled; corolla white, pale green, or pale yellow, with upper surface of lobes papillose.   1 G. acutum
+ Leaves on main stems 5-10.5 mm; inflorescence cymes 1-6-flowered; corolla nearly always red or purple, with upper surface of lobes glabrous and smooth except ± puberulent on margins and central vein.   43 G. rebae

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