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Taxonomy and biogeography of the Bambuseae (Gramineae: Bambusoideae) - D.Z. Li

Deputy Director, Research Program, Kunming Institute of Botany, Kunming, Yunnan, China

Introduction

Bamboo is one of the most important plants in the daily life of people in Asia, Africa and Latin America. In this paper, bamboo is used in a strict sense, including only the woody members of the Bambusoideae or the Bambuseae. In Asia, bamboo is of important economic value amounting to US$ 7 billion per year (Stevens 1995). In China, bamboo is called the “one of the three friends of the hardy winter”, together with the pines and the winterplum, and has long been the main subject of artists, poets and traditional Chinese painters. Bamboo yields are high, they grow fast and are used widely for construction, food and cooking, hunting, gathering material, and paper-making material. Bamboo is the only food for giant panda and also the favourite food for many animals, such as the red panda (Ailurus fulgens) in China.

Importance of morphological characters in bamboo taxonomy

The morphological characters of bamboo are not clearly understood. The following attempts to explain certain criteria in defining a working list of genera and their relationships based on current understanding.

Inflorescence type: Using inflorescence type to define genera was first suggested by McClure (1934). Based on his study of the inflorescences of Schizostachyum species in China, he proposed the term 'pseudospikelet'. McClure (1966) further developed his ideas on the different types of inflorescences in bamboo, based on the two different types of spikelets (spikelets vs. pseudospikelets) by creating another two terms, semelauctant and iterauctant, or determinate and indeterminate inflorescence. McClure's concept was widely accepted. Keng (1982) proposed a system of bamboo classification, including two supertribes in his concept of Bambusoideae (the woody bamboos), one with semelauctant, and the other with iterauctant inflorescences. This was adopted in the FRPS (Keng and Wang 1996). Although recent study shows that both semelauctant and iterauctant inflorescence may occur in most subtribes (Soderstrom and Ellis 1987), it is generally agreed that types of inflorescence are good criteria in defining various bamboo genera.

It is not always easy to determine inflorescence types in some bamboos. The first example is the Racemobambos group. Holttum (1956b) stated that this genus was allied to Bambusa, because of the same ovary type. The inflorescence of Racemobambos was interpreted as semelauctant by Dransfield (1983, 1992), but as iterauctant by Chao and Renvoize (1989). The point is that in the inflorescence of Racemobambos each spikelet is subtended by tiny bracts. In an allied species originally called Arundinaria prainii, the bracts are well developed and it was sometimes treated under the genus, Neomicrocalamus Keng f. (Keng 1983). Keng himself first described inflorescence of N. prainii as semelauctant, but later as iterauctant (in Wen 1986). Dransfield (1992) holds the same view that in Neomicrocalamus it is iterauctant, while Stapleton (1994c) regarded it as semelauctant. The other group is usually treated in a separate subtribe, Shibataeinae (Soderstrom and Ellis 1987; Keng 1992), including Phyllostachys, Semiarundinaria, Sinobambusa, Shibataea, Chimonobambusa and Indosasa, all with bracteate inflorescence. It is easy to determine axillary buds in the bracts during the developing stages, like those in P. nidularia (McClure 1966: p. 99), but not so easy in herbarium specimens. Soderstrom and Ellis (1987, p. 237) followed McClure (1966) who pointed out that the inflorescences of Phyllostachys were iterauctant, but they placed Chimonobambusa in subtribe Arundinariinae, in which all other genera have semelauctant inflorescences. Stapleton (1994c) recognized the close relationship between Chimonobambusa and Phyllostachys, thus correctly treading both of them in Shibataeinae, but he described the inflorescence as 'semelauctant' (Stapleton 1994c, p. 327). Other authors treated it as 'weakly iterauctant' (Clayton and Renvoize 1986, p. 48). In my opinion, these bracts represent an intermediate state between the two types of inflorescences. Evidence from leaf anatomy, chromosome numbers and ovary structure support a close relationship between the Shibataeinae and the Arundinariinae, although they produce different types of inflorescences. Further study is needed to elucidate this relationship between them, as there are many species which are superficially similar to one another at vegetative state, in Acidosasa and Arundinaria of the Arundinariinae and Indosasa and Sinobambusa of the Shibataeinae. Wang and Ye (1980) pointed out that Indosasa shibateoides and Sinobambusa might link the two inflorescence types. Keng supported this idea although he still weighted this character with great importance in his classification system (Keng 1992).

Spikelet structure: As members of the grass family, the bamboos have reduced flowers called 'florets' as part of a spikelet. The spikelet proper is a basic unit of a bamboo flower. There is no proper perianth in the floret, being substituted by lodicules. The origin of lodicules and other bracts (lemma and palea) is not clear. It is widely accepted that it is a monochlamydeous flower and the lodicules form perianth. This is largely on the basis of comparative morphology. However, the 'pseudospikelets' of Streptochaeta, as interpreted by Soderstrom (1981), suggest it is reduced with a single terminal achlamydeous flower. The presence or absence of lodicules is generally used as a generic character, as in Melocalamus vs Dinochloa and Bambusa vs Dendrocalamus. It is suggested that modification of stamens may give rise to irregular number of lodicules and intermediate structures as in Schizostachyum (Arber 1934).

Many tropical bamboos have six stamens. Arundinaria and its close allies have three. The number of stamens may be the major difference between some bamboo genera, e.g. Sinobambusa vs Indosasa, Indocalamus vs Sasa and Arundinaria vs Acidosasa. It is reported that in Oligostachyum there are ususally three or four stamens, lying between Arundinaria and Acidosasa (Wang and Ye 1980). This may be the reason why Soderstrom and Ellis (1987) did not place it properly in the subtribes. Generally speaking, other detailed structures, such as state of the uppermost floret, presence or absence of rachilla extension, shape of uppermost palea and number of stigmas, may also be considered as generic characters, but these characters should be used more critically.

Ovary and fruit: The fruit is among the most poorly known structures of the bamboos. Typical grass fruits or caryopses can be found in species of the Arundinariinae. They have a short appendage, except those of Ferrocalamus whose fruit is berry-like (Keng and Hsueh 1982). The subtribe Bambusinae usually has fruit with a conspicuous appendage (persistent style) which is a growth of the pericarp at the tip of the ovary formed after fertilization. The pear-like fruit of Melocanna has attracted the attention of many botanists (e.g. Stapf 1904; McClure 1966). However, it is suggested that fleshy fruit may have been derived independently in several subtribes (Soderstrom and Ellis 1987).

Holttum (1956a) drew special attention to the anatomy of ovary structure. His work is of fundamental importance in bamboo classification. This was the beginning of the revision of the traditional, Bentham-Munro system. Early support was received from culm anatomy (Grossor and Liese 1971, 1973). A marked change at the generic level was the delimitation of Oxytenanthera. Holttum restricted the genus to its only African member (the type of the genus), O. abbyssinica, having a hollow style (ovary appendage) which is unlike that found in Asiatic species of that genus, all of which were reassigned to Dendrocalamus or Gigantochloa (Holttum 1956a).

Underground system: Riviere and Riviere (1879) were the first to publish a clear account of 'two basic forms' of bamboo rhizome. They noticed the differences in the growth of Gigantochloa and Phyllostachys one leading to caespitose habit, the other to spreading habit. Gamble (1897) gave good descriptions and excellent illustrations of rhizomes of some species but he did not pay much attention to the underground parts of bamboos. McClure (1925) proposed the terms sympodial and monopodial to describe the growth habit of rhizomes, corresponding to Riviere and Riviere's terms 'caespitose' and 'spreading'. This was followed and modified by Japanese and Chinese botanists, notably by Nakai (1935) and Keng (1948), the latter creating a new term 'amphipodial'. McClure (1966, 1973) redefined them as pachymorph and leptomorph and used them extensively in his revision of bamboo genera of the New World. The terms sympodial, amphipodial and monopodial were used with reference to the manner of the origin of culms rather than the habit of the associated 'rhizomes', although these terms were widely used in describing the underground system (the rhizomes) (Holttum 1958; Keng 1959; Clayton and Renvoize 1986; Soderstrom and Ellis 1987; Chao 1989).

It is not always easy to distinguish the two basic forms since those with pachymorph or sympodial rizomes sometimes produce well developed culm neck, up to several metres long in some species, which gives rise to culms not tufted at all and appear like leptomorph or monopodial rhizomes. The leptomorph rhizomes produce buds and roots at each node while the long-necked pachymorph ones, or McClure's (1966 p. 29) metamorph I, do not. Regardless of the terms, sympodial vs monopodial, or pachymorph vs leptomorph, there is a fundamental difference between them. The intermediate type, amphipodial, should be regarded as a subform of the monopodial or leptomorph rhizomes. It was suggested that the condition in Chusquea fendleri and the amphipodial condition of Arundinaria and Indocalamus are not quite the same ('tiller' in the latter genera) (Wong 1986). In my opinion, there is no fundamental difference between the 'culm base' of Arundinaria and the rhizome proper in McClure's (1966) sense. The answer may be simple: those with sympodial or pachymorph 'rhizomes' are in fact different from the underground stems of leptomorph 'rhizomes'. With or without running underground stem is a very distinctive character of bamboo plants. Every bamboo genus has a particular type of underground system. It is generally accepted that the exception is Chusquea of the New World. The underground system of this genus is still poorly understood. The species with both pachymorph and leptomorph rhizomes, like C. fendleri, illustrated in McClure's (1966 p. 33) book, in my opinion, true rhizomes. Such species are very different from those with sympodial 'rhizome', e.g. Chusquea culeou and C. quila.

Buds and branching: No attention was paid to either the underground or the branching system in distinguishing bamboo genera until 1925. Several genera were identified or validated by Nakai (1925), primarily based on vegetative characters such as rhizomes and branching. Among them, Indocalamus, Pseudosasa and Pleioblastus were distinguished mainly by different branching systems from Arundinaria, although later studies do not really support the distinction of Pleioblastus (Chao and Chu 1979; Soderstrom and Ellis 1987). Later detailed study demonstrated the use of branching in separating bamboo genera in the New World (McClure 1966, 1973). Chao et al. (1980) revised some Asian bamboo genera by branching characters. Famous examples are Phyllostachys and Sasa with a fixed number of branches at each culm node, while usually only the mid-culm branch complement can be used since in most bamboos the branching at the uppermost culm nodes is not very distinctive.

Efforts have been made to explore the taxonomic value of prophylls of mid-culm buds and the inflorescence buds and variation in prophyll structure may be taxonomically useful at the species level (McClure 1966; Stapleton 1991, 1994a). A larger number of genera should be examined before this is used as a generic character.

Culm foliage: The unusually developed branching and fast growth make bamboos unique in the plant kingdom. During development, the culm foliage has become specialized for protection rather than a photosynthetic organ. It is widely accepted that culm foliage is of great taxonomic importance in bamboos. According to Holttum (1958), Gamble (1897) was the first to describe and illustrate the culm foliage (culm-sheath) for diagnostic purposes who also used it as a specific character in descriptions. Nakai (1925) used it in generic diagnosis. In a given species of bamboo, the characters of mid-culm sheaths are constant and may be used as a tool to distinguish species and even genera. Chimonobambusa and Qiongzhuea are among those in which the blades of culm-sheaths are reduced to needle-like points.

Bamboo taxonomy at the subtribal and generic levels

The taxonomy of bamboos, or the science of documenting bamboo diversity at species level, is the most difficult field (Table 1). One reason is that the classification of flowering plants largely depends on the floral characters, but flowering is rare in most bamboo species. Some species flower at intervals of as long as 120 years. In addition, information on underground parts, branching and culm sheaths is also important to understand the differences among bamboos (Soderstrom and Young 1981) and earlier herbarium collections of bamboos were mostly incomplete. All this has made bamboo taxonomy a puzzle for a long time. Because of these reasons modern botanists have different opinions on bamboo taxonomy. Although Keng and Wang (1996) treated the woody bamboo in several supertribes, most people tended to accept that the woody bamboos should be treated in a tribe because of the fact that they all share a set symapomorphies. At the subtribal and generic levels, Clayton and Renvoize (1986) recognized only 49 genera in three subtribes, i.e., Arundinariinae (20 genera), Bambusinae (25 genera) and Melocanniae (4 genera), while Soderstrom and Ellis (1987) accepted 59. The estimated number of bamboo genera in recently-published systems of bamboo classification ranges from 69 (Dransfield and Widjaja 1995) to 78 (Stapleton 1994c, and pers. comm). A combination of morphological characters, as discussed above, and anatomical characters make the grouping of bamboo genera and subtribes more reasonable. One of the most interesting findings using molecular markers and gene sequences, such as ndhF of the cpDNA, was that woody bamboos are monophyletic (Watanabe et al. 1994; Clark et al. 1995; Xia, pers. comm.).

Table 1. The subtribes and genera of woody bamboos (tribe Bambuseae)

Subtribe

Clayton and Renvoize (1986)

Soderstrom and Ellis (1987)

Dransfield and Widjaja (1995)

Stapleton (1994 and 1997)

The present paper

Subtribe Arthrostylidiinae


11 genera, New World tropics

13 genera, confined to the New World

13 genera, New World

13 genera, New World

Subtribe Arundinariinae

20 genera, Pan-tropic

12 genera, essentially Old World

14 genera, Old World (1 in N. Am.)

19 genera, Old World (1 in N. am.)

14 genera, essentially Old World

Subtribe Bambusinae

25 genera, Pan-tropic

6 genera, Old World tropics

13 genera, mostly Tropical Asia

11 genera, Old World

9 genera, Old World tropics

Subtribe Chusqueinae


1 genus, New World tropics

2 genera, New World tropics

2 genera, New World

2 genera, New World tropics

Subtribe Guaduinae


5 genera, New World tropics

5 genera, New World tropics

5 genera, New World tropics

5 genera, New World tropics

Subtribe Melocanninae

4 genera, Old World tropics

7 genera, Old World tropics

8 genera, mostly Asia

10 genera, mostly Asia

9 genera, Old World tropics

Subtribe Nastinae


4 genera, Tanzania and Madagascar to Indonesia and New Guinea

6 genera, southern hemisphere of the Old World tropics

6 genera, southern hemisphere of the Old World tropics

6 genera, Tropical Africa and Madagascar to Tropical Asia

Subtribe Neurolepidinae


1 genus, New World high elevations




Subtribe Racemobambosinae



3 genera, Himalayas and Tropical Asia

3 genera

1 genus, Tropical Asia

Subtribe Shibataeinae


7 genera, Japan to Himalayas

5 genera, temperate Asia

5 genera

7 genera, E. Asia forests

Unclear placement


5 genera


3 (2 not yet published plus Phyllosasa)

(1 Temburongia)

Total

49 genera

59 genera

69 genera

77 genera

67 genera


According to our present understanding of the Chinese bamboos (Li 1997), plus the knowledge of bamboo taxonomy based mainly on the systems of Soderstrom and Ellis (1987) and Dransfield and Widjaja (1995), we accept 67 genera represented in different subtribes of bamboo in the present paper. Following is a conspectus of bamboo genera and subtribes of the world.

Conspectus of bamboo genera of the world

I. Subtribe Arthrostylidiinae:

1. Actinocladum, 2. Alvimia, 3. Apoclada, 4. Arthrostylidium, 5. Athroostachys, 6. Atractantha, 7. Aulonemia (Matudacalamus), 8. Colanthelia, 9. Elytrostachys, 10. Glaziophyton, 11. Merostachys, 12. Myriocladus, 13. Rhipidocladum.
II. Subtribe Arundinariinae:
14. Acidosasa, 15. Ampelocalamus, 16. Arundinaria, 17. Chimonocalamus, 18. Drepanostachyum (Himalayacalamus), 19. Fargesia (Borinda, Yushania), 20. Ferrocalamus, 21. Gaoligongshania, 22. Gelidocalamus, 23. Indocalamus, 24. Oligostachyum, 25. Pseudosasa, 26. Sasa, 27. Thamnocalamus
III. Subtribe Bambusinae:
28. Bambusa (Dendrocalamopsis), 29. Bonia (Monocladus), 30. Dendrocalamus (Klemachloa, Oreobambos, Oxynanthera, Sinocalamus), 31. Gigantochloa, 32. Dinochloa, 33. Holttumochloa, 34. Kinabaluchloa (Maclurochloa, Soejatmia), 35. Melocalamus, 36. Sphaerobambos, 37. Thyrsostachys
IV. Subtribe Chusqueinae:
38. Chusquea, 39. Nerolepis
V. Subtribe Guaduinae:
40. Criciuma, 41. Eremocaulon, 42. Guadua, 43. Olmeca, 44. Otatea
VI. Subtribe Melocanninae:
45. Cephalostachyum, 46. Davidsea, 47. Leptocanna, 48. Melocanna, 49. Neohouzeaua, 50. Ochlandra, 51. Pseudostachyum, 52. Schizostachyum, 53. Teinostachyum
VII. Subtribe Nastinae:
54. Decaryochloa, 55. Greslania, 56. Hickelia, 57. Hitchcockella (?), 58. Nastus, 59. Perrierbambus (?)
VIII. Subtribe Racemobambodinae:
60. Racemobambos (Neomicrocalamus, Vietnamosasa)
IX. Subtribe Shibataeinae:
61. Chimonobambusa, 62. Indosasa, 63. Phyllostachys, 64. Qiongzhuea, 65. Semiarundianria (Brachystachyum), 66. Shibataea, 67. Sinobambusa, 68. Temburongia (whereabout uncertain)
Bamboo taxonomy in China

As the most diverse flora in the North Temperate Zone, the flora of China is among the richest on earth including bamboo diversity. Chinese civilization has had a close relationship with bamboo since ancient times. In contrast, the taxonomy of bamboo, or the science of documenting bamboo diversity, is one of the most difficult fields. Earlier books on bamboo in Chinese history can be traced back to the Jing Dynaty (265-316 AD), during that time Dai Kai-Zhi wrote Zhu Pu, a treatise on bamboo, in which 61 species of bamboo were described (of which 34 species or varieties remain valid even today). Modern taxonomy was not used for bamboo in China until the 1930s. Since the 1950s, botanists in many botanical institutions and universities throughout China have made extensive collections of bamboo. The volume for Bambusoideae of the Flora Reipublicae Popularis Sinicae (FRPS) was eventually published in 1996. The estimated number of the Chinese genera varies between 22 and 51, and the FRPS includes 37, along with several new genera. At the specific level, some 479 new species have been published (figures from Chao and Tang 1993, updated) since 1976, together with 199 species published between 1753 and 1975, and 515 species were recognized in the FRPS. The Flora of China Project was initiated to revise the FRPS and to provide an English edition for it, For this purpose, we revised the taxonomy of this subfamily based on the morphological characters used in defining bamboo genera, i.e. inflorescence types, spikelet structure, ovary and fruit, underground parts, branching and culm foliage. According to our present understanding there are some 450 species of 33 genera of bamboo in China, accounting for about 45% and 48.5% of the world's totals, respectively. The diversity of bamboo at community and specific levels in China represents one of the very richest in the world, with potentially high infraspecific diversity. Most of the bamboo vegetation types, ranging from tropical, to subtropical, temperate and alpine, are found in China.

Conspectus of Bambusoideae in China

Subfamily Bambusoideae Ascherson & Graebner, Syn. Mittleleurop. Fl. 2:769. 1902.

Supertribe Bambusodae Keng & Keng f. ex L. Liou in Acta Phytotax. Sin. 18:323. 1980.

Tribe Bambuseae Nees, Agrost. Bras. 520. 1829.

Subtribe Bambusinae Agardh, Aphor., g. 153. 1823.: Bambusa, Thyrsostachys, Dendrocalamus, Gigantochloa, Melocalamus, Bonia.

Subtribe Melocanninae Reichenbach, Deutsch. Fl. 6:6. 1846.: Schizostachyum, Leptocanna, Cephalostachyum, Pseudostachyum, Melocanna.

Subtribe Racemobambosinae Stapleton in Edinb. J. Bot. 51:323. 1994: Racemobambos.

Subtribe Arundinariinae Bentham in J. Linn. Soc. Bot. 19:31. 1881.: Sinarundinaria, Thamnocalamus, Drepanostachyum, Ampelocalamus, Chimonocalamus, Gaoligongshania, Acidosasa, Sasa, Oligostachyum, Arundinaria, Gelidocalamus, Ferrocalamus, Pseudosasa, Indocalamus.

Subtribe Shibateainae (Naka) Soderstrom & Ellis in Grass. Syst. Evol. 238. 1987.: Indosasa, Sinobambusa, Semiarundinaria, Chimonobambusa, Qiongzhuea, Shibataea, Phyllostachys.

Biogeography

Clearly, Asia is the main centre of distribution of bamboos. According to incomplete statistics, there are some 600 species of 44 genera of bamboo in Asia, representing 6 subtribes (Table 2). Because Asia not only possesses the highest number of species, but also those of the genera and subtribes, it is also the centre of diversification of the bamboos. Little information is available on bamboo cytogeography. Soderstrom pointed out that mainland Southeast Asia is a centre of distribution for tetraploid and hexaploid bamboos, but one species of Arundinaria and one species of Phyllostachys with 2n=24 (diploids) are also reported from China (Hsu 1967, 1972). The diploids are mostly found in the herbaceous bamboo species of central and southern America. The chromosome numbers for most of the Bambusa species are 2n=70, while 2n=24 and 2n=64 are found in some Chinese Bambusa (Hsu 1972; Zhang 1985). These indicate that mainland Southeast Asia including the southern part of China may be the centre of origin for the woody bamboos. However, molecular evidence shows that the woody bamboos of the New World and those of the Old World developed quite independently. Further study from various evidences including molecular, anatomical and cytological aspects is needed to elucidate the relationships of the bamboos in the two hemispheres.

Table 2. Geographical distribution of bamboo subtribes, genera and species

Geographical area

Subtribes

Genera

Species

Chromosome No.

Asia

6

44

ca. 600

46, 48, 64, 72

Africa

2

3

5 (endemic)

72

Madagascar

2

6

20 (endemic)


Australia

2

2

3


Pacific

2

2

4


America

4

21

ca. 400

40, 46, 48

Total

9

68

ca. 1000



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