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from . The have in common is possession of so is the benchmark for name reflects the fact that 3 rather than 6 stamens. Maybe temperate . there is so much variegation on they do not need to produce so The inflorescence is open with the leaves that it seems as though much pollen where there is more well separated spikelets, and a yellow leaf has become striped wind. Unfortunately there are without basal sheaths or bracts. with green, rather than vice-versa. some temperate bamboos that To understand how we came to have 6 stamens, so even this be in such a mess with muddles distinction is not consistent. This and misapplications rife, we have is typical of bamboos--there are to look back at how our exceptions to every rule. knowledge of temperate Characters are rather plastic, bamboos has grown over the last often under physiological control, 200 years or so, and consider and boundaries are hard to pin some revelations from recent down, making the separation and Temperate recognition of groups somewhat Temperate molecular research. arbitrary at times. Bamboos are admittedly quite difficult to classify or identify, Why is this? From recent because groups are not always research (Triplett et al. 2014) it names separated clearly. Taxonomic seems that one reason may be bamboo names pervasive hybridisation. Even the groups at all levels can have

rather vague boundaries with major groups of bamboo, the Borinda - open panicles CHRIS STAPLETON surveys the long road overlapping characters. In tropicals, temperates, and addition, our understanding of herbaceous bamboos, originated Many cultivated temperate to nomenclatural stability and correct the characters has been poor. by hybridisation themselves. Our bamboo genera have very similar current polyploid groups flowers, all looking much the identification in cultivated bamboos aureosulcata This has often made it difficult to separate and define groups, even apparently arose as hybrids same. They include , 'Aureocaulis' in SW France when substantial differences were between previous diploid Chimonocalamus, Sarocalamus, and evident. lineages. Hybridisation has Borinda. They differ radically from probably obscured distinctions the compact spicate or capitate, Temperate bamboos are by far T IS NEARLY a quarter of a are in an era of much more The genus is now between groups of bamboos ever and densely bracteate the most important for century since David reliable and stable bamboo considered appropriate on the since they first evolved. inflorescences of tropical horticulture in Europe. These McClintock pondered the names. grounds of morphology and bamboos often obscured by bamboos have evolved out of the difficulties caused by what molecular data, and in that genus dense masses of stamens, and Generic discoveries heI called the 'shifting sands of it is now correctly known as windless and mild tropical forest Conflicting names bamboo nomenclature' in this Pleioblastus viridistriatus. The understorey. The main Looking at the temperate 6-stamened tropical flowers publication (McClintock 1992). As an example of the confusion generic name is a reflection of the morphological character that they bamboos, Tribe , that used to prevail, we have had Since then considerable progress different bud and branch there has clearly been a lot of multiple names in use in the UK 3-stamened temperate flowers has been made in our characters that distinguish this confusion at the generic level. It for the pretty little yellow-leaved understanding of relationships is interesting to see how our bamboo that the cognoscenti within the bamboos, and a knowledge of the variation and know to prune back hard in late relatively stable system of generic relationships within the tribe has winter to encourage a flush of names has now been in use for grown, and how attitudes to brilliant yellow spring shoots. several years, formalised in flora generic recognition have changed Genus, species & cultivar names accounts and online databases. as a consequence. have all varied on displayed and There have been a wealth of new Arundinaria was found in the marketed . I have seen species introductions from the USA by French collector genera Arundinaria, Bambusa, wild to try to identify, many of André Michaux, while looking for Pleioblastus and even Phyllostachys which continue to cause some American trees to reafforest used, with the species epithets, problems, but several France after long wars with viridistriatus auricoma, nana and misidentifications have been England took their toll on the aureospicata, and a few cultivar cleared up, and on the whole we forests. It was the first temperate those with dense spathes bearing names added for good measure. Pleioblastus viridistriatus genus to be named, in 1803, and small leaf blades, found in Japan by a German military doctor with also vary in other ways. Open missionary in Father Paul These bamboos with their dense was found to have dense the Dutch Navy, Phillip Siebold, inflorescences with bunching Guillaume Farges, and published toothbrush-like flowers are inflorescences when it eventually and named Phyllostachys in 1843. (fasciculation) at the nodes could usually found in the mountains of started to flower in the 1990s. have been seen by Munro in C China, while those of Borinda Thus use of the name khasianum from with open inflorescences come Sinarundinaria for the bamboos for example. from W China and the E with open inflorescences . is from including those now placed in Drepanostachyum - delicate even further west, right across the Yushania could no longer be flowers in bunches Himalayas. justified. Takenoshin Nakai was a Japanese botanist looking at bamboos in Vegetative characters the 1920s and 30s. He described In this way more characters were many of our cultivated running being studied, and more bamboo genera, including understanding of morphology Phyllostachys - flat culms Pleioblastus, Shibataea, was being gained. Rhizomes were , and looked at in more detail, four tropical, and named many new Sinobambusa, as well as the ill- distinct patterns of growth being bamboo species. From the fated Sinarundinaria. The - unilateral racemes discovered. Himalayas he described leptomorph-rhizomed genera that Thamnocalamus, a genus of in 1893. Even denser he described have been more Leptomorph rhizomes could lead temperate bamboos with inflorescences than those seen in useful than Sinarundinaria. This is to separate culms, as in Phyllostachys - leafy flowers compressed inflorescences, like Thamnocalamus are found in now a synonym of Fargesia, Phyllostachys in a favourable site, A Bristol soldier from the 39th those of Phyllostachys in having Thus there is substantial variation Fargesia, as a result of extreme because its type species, the or in clusters of culms as in (Dorsetshire) Regiment of Foot bracts, but without leafy blades, within the inflorescences of compression and confinement. Fountain Bamboo , Semiarundinaria fastuosa. was the next to describe new and the culms were round, not temperate bamboos, and bamboo genera, discovering flattened. By the time he had differences in vegetative another kind of compressed characteristics were also being temperate inflorescence in the discovered. Nevertheless a Himalayas. William Munro could conservative approach was taken be considered the Father of by institutional grass taxonomists Bamboo . He devoted when bamboos were included in his spare time to plants, especially the next major grass treatment, bamboos. He classified and putting the clock back. Bentham described all 219 bamboos & Hooker rejected Munro's known at the time, adding many Thamnocalamus, and it was back to new ones, in the first global 2 genera, Arundinaria and bamboo account in 1868. Phyllostachys for temperate Munro established botanic bamboos in their Genera gardens and vegetable gardens at Plantarum of 1883. Different the military stations where he opinions on bamboo genera are served, but more importantly for Thamnocalamus -spathed flowers nothing new. us he pioneered the revolutionary Moving on towards and through retired, as General Munro, he had use of vegetative characters to the 20th Century, many more described 14 new plant genera in separate bamboo genera--not just botanists stepped forward, total, not bad for an amateur their flowers. Munro was brave looking at more bamboos in taxonomist. enough to distinguish genus much more depth, describing Phyllostachys by a vegetative Within the temperate Asian further genera from a wider range character, its distinctive semi- bamboos there is still more of characters. The genus Fargesia, flattened internodes. variation. As well as differences which includes several of our in compression of the There are 4 different forms of rhizome in temperate bamboos He described several important horticulturally important species, inflorescence and the presence new bamboo genera, mostly was first collected by French and nature of spathes they can Pachymorph rhizomes could be was room for controversy over Molecular information to a more stable classification. - branchr shoots short-necked, forming tight which characters were most DNA has been most informative Development of this clumps as in Bambusa, important. at the level of tribes and collaborative and consensual Thamnocalamus, Borinda and By the mid 1980s 10 genera had subtribes, rather than genera, approach took a while, but use of Fargesia, or long-necked in been described in the because bamboo DNA is generic names for cultivated Yushania in a favourable site. pachymorph-rhizomed temperate particularly slow to differentiate, bamboos has become much more Just before and then after the bamboos. These have met with but it is useful to reveal stable over the past 15 years. Cultural Revolution, botanists in varying receptions however. convergent evolution, where Several important publications China from several universities Lumping of the genera has been similar species are not closely have helped along the way. described useful generic names common, but with different related. Species of large Bamboos of the World (Ohrnberger for temperate bamboos. First authorities prioritising different groupings such as broadly 1999) compiled all published Yushania was described for characters and lumping either defined Arundinaria, Sinarundinaria names, following advice from spreading bamboos differing according to the inflorescence-- and Thamnocalamus were shown to bamboo taxonomists around the from Arundinaria in having long- whether open or compressed, or be less closely related than world. It became a benchmark, necked pachymorph rhizomes by using other, vegetative expected. More, smaller genera and started a sense of stability. rather than leptomorph ones. characters that cut right across are now recognised. Isolated The bamboo Thamnocalamus - full sheathing Several other genera with has pendulous this distinction. Essentially a bamboos such as Thamnocalamus account (Li et al. 2006), pachymorph rhizomes were culm tips, long branches that can conflict was starting between tessellatus from S Africa, and supported by molecular described later, all forming scramble through trees, and those who favoured floral phylogeny, covered only half the clumps. Chimonocalamus was swollen nodes to catch on tree characters, especially grass bamboos of the world but added described for species with thorns branches for support. taxonomists and their followers, descriptions. The Species & at the nodes, just like those of Buds and branching patterns and those who favoured Sources List of the American the leptomorph-rhizomed have been found to show great vegetative characters, those for Bamboo Society has documented . These hardened variability. Several characters are whom field identification was bamboos for sale since 1980, like aerial roots develop through the involved, and the different more important. a Bamboo 'Plant Finder' for the base of the culm sheath in a ring patterns are still being The grass taxonomists' US, and now has nearly 500 around the node. investigated in depth. Progressive perspective prioritised flowers entries. It standardises names of loss of sheathing and the and avoided some vegetative bamboos in cultivation across the inclusion of a larger number of characters. Very large groups US. Our own RHS Plant Finder central branch internodes and were used based on whether the serves a similar function in the lateral branches produces inflorescence was open and UK. Many internet reference distinguishable patterns. Arundinaria-like (Sinarundinaria, sources are available, mostly Thamnocalamus has complete following standardised names. Fargesia - reduced sheathing including Yushania and other sheathing and relatively few genera) or compressed Bergbambos - fine bristles branches. Fargesia has lost some (Thamnocalamus, including Fargesia Arundinaria densifolia from Sri Misapplied species names sheaths and incorporated more and other genera). Not only was Lanka, were shown by their Clarifying the proper application laterals into the complement. this revealed as unnatural and DNA to be unrelated to others of species names is an important Himalayacalamus includes even unjustifiable lumping when DNA such as Thamnocalamus spathiflorus part of sorting and stabilising more laterals to produce a more evidence became available, it also from the Himalayas and bamboo nomenclature. Names complex layout. assumed that the type species of Arundinaria gigantea from the US. need to be tied to the right Sinarundinaria, Arundinaria nitida, New genera were required, e.g. Drepanostachyum and Chusquea lose species or there is no end of has open inflorescences. Sarocalamus, Bergbambos, Oldeania more sheaths still, and produce confusion. Bamboos in Chimonocalamus - root thorns Unfortunately, when it flowered and Kuruna. This explained some branches all around the culm. cultivation for a long time have in the 1990s, it was found to have unusual characters, such as the Two subtropical genera, often been incorrectly named. An compressed inflorescences particularly delicate oral setae of Drepanostachyum and the larger example was Himalayacalamus Character prioritisation instead, which is why it is now the widely cultivated Bergbambos Himalayacalamus, have multiple hookerianus. It was grown in the It is clear that in the temperate called Fargesia nitida and Yushania tessellata, formerly Thamnocalamus branches that break through the UK as Sinarundinaria falcata, while bamboos there is plenty of is used instead of Sinarundinaria tessellatus. culm sheath base, and thin the name Arundinaria hookeriana variation in vegetative for temperate spreading bamboos spikelet glumes. The former Combining the DNA data with was applied to Himalayacalamus morphology as well as with long-necked pachymorph having open inflorescences, while our better understanding of floral falconeri 'Damarapa' (Stapleton inflorescences. However, there Himalayacalamus - many branches rhizomes. the latter has condensed ones. and vegetative characters has led 1994). this cultivar of Dragon-head Bamboo 'Rufa'.

Conclusion Plenty of new species are yet to be discovered in the wild, and the prospects for finding exciting new horticultural introductions are very good. Much work remains to be done on recently introduced bamboos as well, but the road ahead looks much straighter now that the generic framework for cultivated bamboos has been stabilised, and An introduction with red sheaths, the sands are no longer shifting. misidentified as , has distinct . auricles that reveal its true identity as the CHRIS STAPLETON is a bamboo bamboo named F. dracocephala in China, taxonomist with a special interest Dragon-head Bamboo in Asian temperate bamboos. chris@bamboo- Fargesia rufa seem appropriate, but identification.co.uk Incorrect identification of that name really belongs to a recently introduced bamboos has different species altogether, best also caused considerable placed in Borinda. problems. As one example, a very REFERENCES Dragon-head bamboo has long handsome bamboo with orange Li, D.Z., Wang, Z.P., Zhu, Z.D., Xia, orange leaf sheath auricles with N.H., Jia, L.Z., Guo, Z.H., Yang, G.Y.; sheaths was introduced and sold twisted setae, all reminiscent of a as Fargesia rufa, but that was not dragon's head, complete with & Stapleton, C.M.A. (2006). the right name for this species. It flames. Bambuseae. In: Wu, Z. Y., Raven, P. H. was actually Fargesia dracocephala, & Hong, D. Y. (eds.) (2006). Flora of The name F. rufa should not have Dragon-head Bamboo. When it China, 22. : 7–180. Science been applied, but it was, and to was first introduced the red- Press, Beijing & Missouri Botanical keep the association we now call coloured sheaths made the name Garden Press, St. Louis. McClintock, D. (1992). The shifting ONLINE RESOURCES sands of bamboo genera. The Flora of China bamboo account Plantsman 14(3): 169-177. http://www.efloras.org/florataxon.aspx?flora_id=2&taxon_id=20753 Ohrnberger, D. (1999). The Bamboos of the World. Elsevier. American Bamboo Society Species List Stapleton, C.M.A. (1994). The blue- http://www.bamboo.org/BambooSourceList stemmed bamboo: Himalayacalamus Bamboo Identification website hookerianus. The New Plantsman 1(1): http://www.bamboo-identification.co.uk 1-9. TROPICOS database of plant names Triplett, J.K., Clark, L.G., Fisher, A.E., http://www.tropicos.org Wen, J. (2014). Independent allopolyploidization events preceded IPNI database of plant names speciation in the temperate and tropical http://www.ipni.org/ipni/plantnamesearchpage.do woody bamboos. New Phytol. 204 (1): 1469-8137.

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