Research Article
Research Article
Kengiochloa, a new bamboo genus to accommodate the morphologically unique species, Pseudosasa pubiflora (Poaceae)
expand article infoYi-Hua Tong§, Zheng-Yang Niu§|, Zhuo-Yu Cai§|, Jing-Bo Ni§, Nian-He Xia§
‡ South China Botanical Garden, Chinese Academy of Sciences, Guangzhou, China
§ South China National Botanical Garden, Guangzhou, China
| University of Chinese Academy of Sciences, Beijing, China
Open Access


Pseudosasa was confirmed as polyphyletic by recent phylogenetic analyses, with Chinese species of Pseudosasa distantly related to those from Japan. Among the Chinese species of Pseudosasa, Pseudosasa pubiflora is a morphologically unique as well as taxonomically problematic species endemic to South China, of which the generic designation is still uncertain. Molecular analyses based on both plastid and nuclear genomic data demonstrated that this species is closest to the recently published genus Sinosasa. Morphologically, the two are somewhat similar to each other in flowering branches developing at the nodes of every order of branches, raceme-like units of inflorescence with 3–5 short spikelets, each spikelet with few florets including a rudimentary one at the apex, and each floret with 3 stamens and 2 stigmas. However, P. pubiflora is very different from Sinosasa species in many reproductive and vegetative characters, such as the morphology of paracladia (lateral spikelet “pedicels”), the absence or existence of pulvinus at the base of paracladia, the relative length of the upper glume and the lowest lemma, the shape of lodicules and primary culm buds, the branch complement, the morphology of nodes, culm leaves and dried foliage leaf blades, and the number of foliage leaves per ultimate branchlet. The morphological and molecular evidence warrants recognition of a new genus to accommodate this unique species, which is here named Kengiochloa. After consulting related literature and examination of herbarium specimens or specimen photos, a taxonomic revision of K. pubiflora and its synonyms was made, and it was confirmed that four names, viz. P. gracilis, Yushania lanshanensis, Arundinaria tenuivagina and P. parilis, should be merged with K. pubiflora, while Indocalamus pallidiflorus and Acidosasa paucifolia are distinct species.


morphology, phylogeny, plastome, synonyms, taxonomy


Pseudosasa Makino ex Nakai, with ca. 20 species mainly distributed in East Asia and Vietnam, belongs to the subtribe Arundinariinae of the tribe Arundinarieae (Poaceae: Bambusoideae) (Zhang et al. 2020; Soreng et al. 2022; POWO 2022). Many molecular phylogenetic studies have shown that this genus is polyphyletic, as the Japanese species including the generic type, P. japonica (Siebold & Zucc. ex Steud.) Makino ex Nakai, and those from China are always distributed in different clades in the phylogenetic trees (Triplett and Clark 2010; Zhang et al. 2012; Guo et al. 2021; Triplett and Clark 2021). Morphologically, the two groups of species are also different: P. japonica has persistent culm leaf sheaths, a solitary branch at each node, or sometimes three branches at the top nodes of the culm, and three to five stamens (which also suggest that P. japonica has a hybrid origin between Japanese Pleioblastus Nakai and Sasa Makino & Shibata or Sasamorpha Nakai, as indicated by Triplett and Clark (2010, 2021)), while Chinese Pseudosasa species usually have deciduous culm leaf sheaths, one to three branches per node, and three stamens (Chen et al. 1996; Zhu et al. 2006; Zhang et al. 2012; Kobayashi 2017). Among those sampled Pseudosasa species, one species endemic to South China, viz. P. gracilis S. L. Chen & G. Y. Sheng, drew the attention of many researchers. This species was consistently associated with some Chinese Sasa species (which were recently transferred to the newly established genus, Sinosasa L. C. Chia ex N. H. Xia et al., see Qin et al. 2021), and far related with other Pseudosasa species in previous studies (Zhang et al. 2012; Guo et al. 2021) (Fig. 1). Zhang et al. (2012) argued that P. gracilis was not a typical member of Pseudosasa in terms of morphology, and suggested that more work was required to clarify the ascription of this species and its phylogenetic placement. In Flora of China, Zhu et al. (2006) considered that P. gracilis was possibly a synonym of P. pubiflora (Keng) Keng f. ex D. Z. Li & L. M. Gao, which they thought may not belong to Pseudosasa but Indocalamus Nakai.

Figure 1. 

Phylogenetic position of Pseudosasa gracilis, adapted from Guo et al. 2021.

Pseudosasa pubiflora has a relatively complicated taxonomic history. In 1936, Yi-Li Keng, the first Chinese botanist who studied the bamboo classification in China, described Arundinaria pubiflora Keng from northern Guangdong based on only one collection (To & Tsang 12284) with flowers (Keng 1936). His son, Pai-Chieh Keng then transferred it to Pseudosasa Makino ex Nakai without giving any reason (Keng 1957 & 1959). However, the new combination was invalid due to lack of a clear and direct reference to the basionym (Zhu et al. 2006). Yang and Chao (1994) recognized this species in Arundinaria Michx.., and considered several other names, including P. gracilis, Indocalamus pallidiflorus McClure, Yushania lanshanensis T. H. Wen and Acidosasa paucifolia W. T. Lin as well as several combinations based on these names, as synonyms of this species.

The P. pubiflora published in Flora of China (Zhu et al. 2006) was a combination largely based on Yang and Chao’s (1994) treatment, except that P. gracilis was suggested as “possibly a synonym of P. pubiflora”. Besides, two more names, viz. Arundinaria teuivagina W. T. Lin and P. parilis T. P. Yi & D. H. Hu, were further synonymized with P. pubiflora. Consequently, the distribution of this species was extended to include three provinces, i.e., Guangdong, Hunan and Jiangxi. Moreover, Zhu et al. (2006) argued that, for P. pubiflora, “the woolly internode apex is unusual in Pseudosasa and rather suggestive of Indocalamus Nakai”. Thus, the relationships between P. pubiflora and its synonyms and the generic designation of this species need a further study.

Materials and methods

Field trips to the type locality of P. pubiflora as well as its several synonyms including I. pallidiflorus, Arundinaria teuivagina and P. parilis were conducted from 2018 to 2022. The complete specimens including rhizomes, culms with culm leaves, branches and foliage leaves as well as fresh young foliage leaves used for molecular analyses were collected. The types and other specimens of the pertinent species housed at CANT, IBSC, N, SYS and ZJFI, as well as photographs of specimens housed at K, US and W, were examined. Herbaria acronyms follow Thiers (2022). The morphological description is based on specimens and referred to related literature (Keng 1936; Chen et al. 1983; Wen 1985, 1986; Lin and Wu 1990; Yi 1995; Chen et al. 1996; Zhu et al. 2006). General morphological terms follow Beentje (2016). The terms applied to the flowering structure in this study mainly follow the synflorescence concept applied to grasses (Vegetti and Anton 1996; Tivano et al. 2009; Cai and Xia 2021).

To study the phylogenetic position of P. pubiflora within the tribe Arundinarieae, the whole chloroplast genomes were used for building the phylogenetic tree. Five Pseudosasa species including one from Japan, also the type species of the genus, viz. P. japonica, and four from China were sampled. In total, there were 29 species represented by 30 samples of all the five subtribes in the tribe Arundinarieae (two samples for P. pubiflora) and one sample of Bambusa bambos (L.) Voss from the tribe Bambuseae was the outgroup. Species names, voucher information and GenBank accession numbers are provided in Table 1.

Table 1.

Species names, voucher information and GenBank accession numbers of the 30 whole chloroplast genomes used in this study.

Taxon Voucher information GenBank accession
Acidosasa glauca B. M. Yang CZY56 (IBSC) *OP850353
Ampelocalamus actinotrichus (Merr. & Chun) S. L. Chen T. H. Wen & G. Y. Sheng MPF10003 (KUN) MF066245
Chimonobambusa quadrangularis (Franceschi) Makino Not provided by the author MW928533
Ferrocalamus rimosivaginus T. H. Wen Zhang08019 (KUN) HQ337794
Ferrocalamus strictus Hsueh & Keng f. NH001 (IBSC) *OP850355
Gaoligongshania megalothyrsa (Hand.-Mazz.) D. Z. Li, Hsueh & N. H. Xia MPF10056 (KUN) JX513419
Gelidocalamus stellatus T. H. Wen BH102 (IBSC) *OP850347
Gelidocalamus tessellatus T. H. Wen & C. C. Chang MPF10049 (KUN) JX513420
Hsuehochloa calcarea (C. D. Chu & C. S. Chao) D. Z. Li & Y. X. Zhang MPF10050 (KUN) KJ496369
Indocalamus longiauritus Hand.-Mazz. MPF10168 (KUN) HQ337795
Indocalamus latifolius (Keng) McClure CZY76 (IBSC) *OP850354
Indocalamus sinicus (Hance) Nakai ZMY037 (KUN) MF066250
Indosasa crassiflora McClure BH58 (IBSC) *OK558536
Phyllostachys nidularia f. farcta H. R. Zhao & A. T. Liu 2020-JZ01 (NF) LC590826
Pleioblastus chino (Franch. & Sav.) Makino NH029 (IBSC) *OP850356
Pseudosasa amabilis (McClure) Keng f. NH032 (IBSC) *OP850358
Pseudosasa cantorii (Munro) Keng f. MPF10006 (KUN) MF066255
Pseudosasa japonica (Siebold & Zucc. ex Steud.) Makino ex Nakai Pjc-1 (ZJFC) KT428377
Pseudosasa nanunica (McClure) Z. P. Wang & G. H. Ye XNH36 (IBSC) *OP850361
Pseudosasa pubiflora (Keng) Keng f. ex D. Z. Li & L. M. Gao CZY146 (IBSC) *OP850350
Pseudosasa pubiflora (Keng) Keng f. ex D. Z. Li & L. M. Gao XNH63 (IBSC) *OP850362
Shibataea chinensis Nakai NH036 (IBSC) *OP850359
Sinobambusa tootsik (Makino) Makino ex Nakai NH031 (IBSC) *OP850357
Sinosasa fanjingshanensis N. H. Xia, Q. M. Qin & J. B. Ni BH124 (IBSC) *OP850348
Sinosasa sp. CZY173 (IBSC) *OP850352
Sinosasa longiligulata (McClure) N. H. Xia, Q. M. Qin & J. B. Ni CZY163 (IBSC) *OP850351
Ravenochloa wilsonii (Rendle) D. Z. Li & Y. X. Zhang MPF10146 (KUN) JX513421
Oligostachyum sulcatum Z. P. Wang & G. H. Ye CZY113 (IBSC) *OP850349
Yushania basihirsuta (McClure) Z. P. Wang & G. H. Ye XNH144 (IBSC) *OP850360
Bambusa bambos (L.) Voss B1-1 (ISC) KJ870988

Total genomic DNA was isolated from silica gel-dried leaves using TIANGEN Genomic DNA Extraction Kit (TIANGEN, Beijing, China). The extracted genomic DNA was fragmented randomly by Covaris M220 (Covaris, Woburn, MA), and the fragments with insert size of 350 bp were selected by using AxyPrep Mag PCR Clean Up Kit. The selected fragments were enriched by PCR after undergoing end repair, the addition of polyA tail and adaptor ligation. The paired-end (2 × 150 bp) libraries were constructed on NovaSeq 6000 platform. Finally, 20 Gb genome skimming data were generated for each sample.

After quality control of 20 Gb clean data by Trimmomatic v 0.39 (Bolger et al. 2014), the whole chloroplast (cp) genomes were assembled using the software GetOrganelle v 1.7.4 pipeline (Jin et al. 2018), with the reference cp genome sequence of Ampelocalamus sinovietnamensis (MW525255) and K-mer sizes of 45, 65, 85, 105 and 125. The generated whole cp genome was annotated using the program Plastid Genome Annotator (Qu et al. 2019) based on the annotation of A. sinovietnamensis. The transferred RNAs (tRNAs) were adjusted using the software tRNAscan-SE service (Schattner et al. 2005). The initiation and termination codons of all the coding genes were manually verified in Geneious Prime v 9.1.4 (Kearse et al. 2012).

All the cp genomes were aligned using MAFFT v. 7.450 (Katoh and Standley 2013). Maximum Likelihood (ML) analysis was conducted by RAxML v 8.2.12 (Stamatakis 2014) with the rapid bootstrap algorithm. GTRGAMMAI was selected as the best-fit model recommended by jModelTest v2.1.6 (Darriba et al. 2012). The number of 12345 was specified as the random seed of parsimony tree inference with 1000 replicates performed. For Bayesian Inference (BI), the data matrix was calculated using MrBayes v3.2.2 software (Ronquist et al. 2012). The best-fit model was selected as SYM+G under the Akaike information criterion (AIC) using MrModeltest v 2.3 (Nylander 2004). Rates of variations across sites were trimmed as gamma. For each analysis, two simultaneous runs of four Monte Carlo Markov Chains (three heated and one cold) were run for 20 million generations with a random tree as the starting point and saving trees every 1000 generations. After discarding the first 25% samples as burn-ins, the optimized topology was generated. The final results were visualized with Figtree 1.4.3 (


The chloroplast genomes ranged from 139,093 bp (Pleioblastus chino Makino) to 140,064 bp (Gaoligongshania megalothyrsa (Hand.-Mazz.) D. Z. Li, Hsueh & N. H. Xia) with an alignment length of 144,771 bp. The plastid data matrix was characterized by sequence divergence with 9,751 variable sites (6.74%), including 6,477 parsimony informative sites (4.47%) and 3,274 singleton variable sites (2.27%).

The reconstructed phylogenetic tree is shown in Fig. 2. Similar to some previous analyses on plastid sequence data (Triplett and Clark 2010; Zhang et al. 2012), Pseudosasa was resolved as polyphyletic in this analysis. Pseudosasa pubiflora was sister to the clade of Sinosasa with strong support (maximum likelihood bootstrap value/Bayesian inference posterior probability BS/PP = 100%/1.00). The other Pseudosasa taxa form a clade with five other genera of the subtribe Arundinariinae, namely Oligostachyum Z. P. Wang & G. H. Ye, Acidosasa B. M. Yang, Sinobambusa Makino, Indosasa Maclure and Pleioblastus (MLBS/PP = 100%/1.00).

Figure 2. 

Phylogeny of Arundinarieae based on whole chloroplast genome derived from maximum likelihood and Bayesian analyses. Only bootstrap values (BS) ≥ 70% and posterior probabilities (PP) ≥ 0.95 are indicated at each node.


Taxonomic revision of Pseudosasa pubiflora and its synonyms

After consulting related literature and examining the types and other herbarium specimens or specimen photographs, we agree with Yang and Chao (1994) that P. gracilis, Yushania lanshanensis should be merged into P. pubiflora, and with Zhu et al. (2006) that Arundinaria teuivagina and P. parilis are also synonyms of P. pubiflora, since all these species share the same key diagnostic characters, mainly including the leptomorph rhizome, the pluricaespitose, short (< 2 m) and thin (< 0.8 mm) culm (Fig. 3C), the flat node, the narrowly ovate culm buds (Fig. 4B), the branch complement with two to four branches at each mid-culm node, the persistent papery culm leaf sheath with fragile oral setae that are adnate at base (Fig. 3E), the erect and amplexicaul culm leaf blades that are usually longer than sheath (Fig. 3D), and the ultimate branchlet with one to four lanceolate or narrowly lanceolate foliage leaves (Fig. 3B). Among these characters, its oral setae and culm leaf blade are very special in Arundinarieae.

Figure 3. 

Kengiochloa pubiflora A habitat B habit C leptomorph rhizome and pluricaespitose culms, the arrow showing an internode with a bud D culm leaf, the arrow showing the junction of sheath and blade E culm leaf oral setae F intravaginal branching G prophyll of primary branch H three young branches at a node I foliage leaf branch. Photos by Y. H. Tong.

However, we do not agree that I. pallidiflorus and Acidosasa paucifolia are identical to P. pubiflora. Indocalamus pallidiflorus (≡ Pseudosasa pallidiflora (McClure) S. L. Chen & G. Y. Sheng) markedly differs from P. pubiflora in the woolly (vs. glabrous) infranodal region, the culm leaf sheath oral setae present (vs. absent) and the culm leaf blade much shorter (vs. usually longer) than sheath. Actually, the description of P. pubiflora in Flora of China is mainly based on that of I. pallidiflorus made by McClure (1940), which conflicts a lot with the protologue of P. pubiflora (Keng 1936). We cannot find the type specimens of Acidosasa paucifolia despite an exhaustive search in CANT where they should be preserved, as indicated by the author (Lin 1992). According to the description and illustration in the protologue (Lin 1992), this species significantly differs from P. pubiflora by the deciduous culm leaf with abaxially more or less hispid (vs. usually glabrous) sheaths and much longer ligules (2 mm vs less than 0.5 mm) and without (vs. with) oral setae, the absence (vs. existence) of foliage leaf oral setae and much longer foliage leaf inner ligules (3 mm vs. 0.3–1.5 mm). Thus, this species should not be synonymized with P. pubiflora either. The identities of these two taxa need further study.

Generic designation of Pseudosasa pubiflora

Recent research has proved that the formerly recognized Arundinaria in a broad sense should not be adopted any more, which at present is thought to be a small genus with only three species restricted to North America (Zhang et al. 2012, 2016, 2020; Guo et al. 2021; Soreng et al. 2022). Thus, Pseudosasa pubiflora should not be placed into Arundinaria. This species has many characteristics that are very unusual for Pseudosasa, such as the lanceolate (vs. narrowly terete or linear in other Pseudosasa species, the same below) and rather short (1.6–2 cm long vs. usually much longer and up to 20 cm) spikelets with robust and erect (vs. slender and porrect) pedicels and very few florets (2–4 vs. 3–30), each floret with two (vs. three) stigmas, the papery (vs. leathery) culm leaf sheath and the glabrous and nonpowdery (vs. conspicuously powdery) infranodal regions (Table 2). The phylogenetic analyses of both our study based on the chloroplast genomes and Guo et al. (2021) based on nuclear markers from ddRAD data demonstrated that this species had a distant relationship with other Pseudosasa. Thus, P. pubiflora should not be a member of Pseudosasa.

Table 2.

Comparison of key morphological characters of Pseudosasa pubiflora, Sinosasa, Indocalamus sinicus and Pseudosasa. The characters unique to P. pubiflora among these taxa are indicated in bold.

P. pubiflora Sinosasa Indocalamus sinicus Pseudosasa
Culm bud Narrowly ovate Trullate Triangle, lower half adnate to culm Trullate to narrowly trullate
Branches at mid-culm nodes 2 to 4 Solitary Solitary 1 to 3
Nodes Flat Prominent, supranodal ridge strongly raised Flat Flat or weakly prominent
Infranodal region Glabrous, nonpowdery With a sericeous or villous band Glabrous, nonpowdery Powdery
Culm leaf sheath Persistent, papery Persistent, papery Persistent, leathery-papery Persistent, usually leathery
Culm leaf oral setae Adnate at base Distinct Distinct or absent Distinct or absent
Culm leaf blades Erect, amplexicaul, usually longer than sheath Erect or slightly spreading, not amplexicaul, much shorter than sheath Erect or reflexed, not amplexicaul, narrowly triangular-lanceolate, much shorter than sheath Erect or reflexed, amplexicaul or not, narrowly triangular to strap-shaped, shorter than or as long as sheath
Number of foliage leaves per ultimate branchlet 1 to 4 5 to 17 7 to 14 2 to 10
Inner ligule Very short, 0.3–1.5 mm Long, (8–)10–20 mm Short, 2–3 mm Short to long, 1–5 mm
Foliage leaf blades Flat when dry Wavy when dry Flat when dry Flat when dry
Flowering branch Developing at the nodes of every order of branches Developing at the nodes of every order of branches or culm Terminal on culm or foliage leaf branches Developing at the nodes of every order of branches or culm
The unit of inflorescence of the synflorescence Raceme-like with 3–5 spikelets Raceme-like with 3–5 spikelets Panicle-like with many spikelets Raceme or panicle-like with several to many spikelets
Paracladia Robust and erect, appressed to the axis Slender, porrect to reflexed Slender, porrect Slender, porrect
Pulvinus at the base of pracladia Absent Present Present Absent
Spikelet Lanceolate, 1.6–2 cm long Lanceolate, 0.5–1.5 cm long Lanceolate, 1.5–2.5 cm long Narrowly terete or linear, 2.5–20 cm long
Number of floret per spikelet 2–4 2–3 3–4 3–30
Terminate rudimentary floret Exist Exist Exist Exist or not
The relative length of upper glume and the lowest lemma Upper glume shorter than the lowest lemma Upper glume nearly equal to the lowest lemma Upper glume shorter than the lowest lemma Upper glume shorter than the lowest lemma
Lodicule Oblong-obovate, spatulate or obovate, apex rounded Ovate or narrow oblong, apex acute Ovate to oblong, apex obtuse Oblong, obovate-lanceolate, spathulate or lanceolate, apex acute or obtuse
Stigma number 2 2 2 3 to 5

Interestingly, both of the two phylogenetic studies suggested a close relationship between Pseudosasa pubiflora and Sinosasa. In the analysis of Guo et al. (2021), Indocalamus sinicus (Hance) Nakai is closest to the Pseudosasa pubiflora + Sinosasa clade (called “Pseudosasa gracilis + Sasa subg. Sasa” clade (Fig. 1) in that paper) with a strong support. Morphologically, this small clade is characterized by having persistent papery culm leaf sheath, short (less than 2.5 cm) and lanceolate spikelet with few florets (2–4) including a terminate rudimentary one and each floret with two stigmas in the subtribe Arundinariinae. However, the flowering branch of Indocalamus sinicus is terminal on culm or foliage leaf branches with a large and panicle-like unit of inflorescence composed of many spikelets, while the flowering branches of the other two develop at the nodes of every order of branches with a raceme-like unit of inflorescence composed of only 3–5 spikelets. The culm buds of Indocalamus sinicus are also very special; they are triangular with lower half completely adnate to the culm (Fig. 4A).

Figure 4. 

Culm buds of three bamboos A Indocalamus sinicus B Kengiochloa pubiflora C Sinosasa sp. Photo A by Xi-Rong Zheng, B, C by Zhuo-Yu Cai. Scale bars: 1 cm (A, C); 5 mm (B).

Last, there are so many different vegetative and productive characters between Pseudosasa pubiflora and Sinosasa that they cannot belong to the same genus. Specifically, Pseudosasa pubiflora differs from Sinosasa in having narrowly ovate culm buds (vs. trullate) (Fig. 4), branch complement with 2–4 branches at mid-culm nodes (vs. solitary), flat nodes (vs. prominent with a strongly raised supranodal ridge), infranodal region being glabrous and nonpowdery (vs. with a sericeous or villous band), culm leaf with oral setae adnate at base (vs. distinct) and amplexicaul blade being usually longer than sheath (vs. not amplexicaul and much shorter than sheath), 1–4 foliage leaves per ultimate branchlet (vs. 5–17) with blades flat when dry (vs. wavy) and much shorter inner ligule (0.3–1.5 mm vs. (8–)10–20 mm), robust and erect (vs. slender and porrect to reflexed) paracladia being appressed to the axis and without (vs. with) pulvinus at base, upper glumes typically shorter than (vs. nearly equal to) the lowest lemma and lodicules with a rounded (vs. acute) apex (Detailed comparison is provided in Table 2). In conclusion, both the morphological and molecular phylogenetic evidence strongly supports the recognition of a new genus to accommodate Pseudosasa pubiflora.

Taxonomic treatment

Kengiochloa Y.H.Tong & N.H.Xia, gen. nov.


Kengiochloa pubiflora (Keng) Y. H. Tong & N. H. Xia (≡ Arundinaria pubiflora Keng).


This new genus is close to Sinosasa, but differs in having narrowly ovate culm buds, branch complement with 2–4 branches at mid-culm nodes, flat nodes, glabrous and nonpowdery infranodal region, culm leaf oral setae adnate at base, amplexicaul culm leaf blades that are usually longer than sheath, 1–4 foliage leaves per ultimate branchlet with blades flat when dry and short inner ligule, robust and erect paracladia (lateral spikelet pedicels) appressed to the axis and without pulvinus at base, upper lemma typically shorter than the lowest lemma and lodicules with a rounded apex.


Shrubby bamboo. Rhizomes leptomorph. Culms pluricaespitose, short and thin, less than 2 m tall and 8 mm in diam.; nodes flat. Culm buds narrowly ovate. Branches intravaginal, 2–4 at each mid-culm node. Culm leaf sheath persistent, papery; auricles absent; oral setae fragile, adnate at base; blade erect and amplexicaul, usually longer than sheath; ligule convex, short. Foliage leaves 1–4 per ultimate branchlet; auricles obscure; inner ligule short. Flowering branches developing at the nodes of every order of branches; the unit of inflorescence of synflorescence raceme-like with 3–5 spikelets; paracladia robust and erect, appressed to the axis and without pulvinus at base; florets 2–4 per spikelet including a rudimentary one at the apex; glumes 2, upper one shorter than the lowest lemma; lemma longer than palea; palea 2-keeled; lodicules 3, apex rounded; stamens 3, anthers pale yellow; styles 2, base slightly connate; stigmas 2, plumose. Caryopsis unknown.


Kengiochloa is named in honor of Professor Yi-Li Keng (1897–1975), a renowned botanist and the first Chinese who studied the bamboo taxonomy in China. The type species of this genus was first described by him, too. Its Chinese name is given as 以礼竹属 (pinyin: yĭ lĭ zhú shŭ).

Kengiochloa pubiflora (Keng) Y.H.Tong & N.H.Xia, comb. nov.

Arundinaria pubiflora Keng, Sinensia 7: 414, fig. 4. 1936; Yang & Chao, J. Bamb. Res. 13(1): 5. 1994, excl. syn. of Indocalamus pallidiflorus. ≡ Pseudosasa pubiflora (Keng) Keng f. ex D. Z. Li & L. M. Gao in Zhu et al., Fl. China 22: 118. 2006, excl. descr. and syn. of Indocalamus pallidiflorus and Acidosasa paucifolia; Keng f. in Keng, Clav. Gen. Sp. Gram. Prim. Sin. 154. 1957 & Fl. Ill. Pl. Prim. Sin. 32 & pl. 22. 1959, nom. inval. Type: China, Guangdong, on top of the hill at the rear of Iu [Yao] village, Lung Tau Mt. [Longtou Mountain], 30 May 1924, To & Tsang 12284 (holotype US00130296 (sheet 1) & US00065420 (sheet 2), image!, isotypes US0031360, image!, US00144993, image!, W19390008723, image!, SYS!). Figs 3, 4B; also see fig. 150 in Vol. 22 of Flora of China Illustrations (available at

=Pseudosasa gracilis S. L. Chen & G. Y. Sheng, Acta Phytotax. Sin. 21: 405. 1983; Chen et al., Fl. Reipubl. Popularis Sin. 9(1): 658. 1996; Zhu et al., Fl. China 22: 119. 2006. Type: China, Hunan, Yizhang County, Zeping, 6 May 1977, Z. P. Wang, H. R. Zhao & G. Y. Sheng 77004 (holotype NAS, not seen, isotypes N019022078!, N019022079!, N019022080!).

=Yushania lanshanensis T. H. Wen, J. Bamboo Res. 4(2): 13. 1985. ≡ Arundinaria lanshanensis (T. H. Wen) T. H. Wen, J. Bamboo Res. 5(1): 19. 1986. Type: China, Hunan, Lanshan County, Ziliang Township, 1125 m, 17 June 1984, S. Q. Chen CX84860 (holotype ZJFI!, isotypes ZJFI!).

=Arundinaria tenuivagina W. T. Lin & Z. M. Wu, J. S. China Agric. Univ. 11(3): 48. 1990. Type: China, Guangdong Province, Xinyi City, Dawuling, 1400 m, 12 April 1987, Z. M. Wu 0167 (holotype CANT00002410!, isotypes CANT00002408!, CANT00002409!, CANT00002411!, CANT00002412!, CANT00002413!, CANT00002414!, CANT00002415!).

=Pseudosasa parilis T. P. Yi & D. H. Hu in Yi, J. Bamboo Res. 14(1): 20. 1995. Type: China, Jiangxi, Suichuan County, Dabali, 1400 m, 21 November 1991, Q. Hu & D. H. Hu 032 (holotype SAUT, not seen, isotype Herbarium of Ji’an Forestry Institution, not seen).


Rhizomes amphipodial, internodes terete, 2–2.5 cm long, ca. 2.5 mm in diam. Culms 1.2–2 m tall, up to 8 mm in diam.; internodes terete, 8–24 cm long, green, not powdery, glabrous; wall thick, cavity with woolly or irregularly lamellate pith; supranodal ridge flat, remains of sheath base persistent, sheath scar pubescent or glabrous. Culm buds narrowly ovate. Branches intravaginal, 2–4 at each mid-culm node, base attached to culm. Culm leaf sheaths gradually deciduous or rather persistent, 1/2 to 4/7 as long as internodes, thickly papery, glabrous or sometimes sparsely brown appressed hispidulous, margins ciliolate; auricles absent; oral setae 5–11, lateral 1–4 curved and downward, others straight forward, 6–9 mm long, fragile, base usually adnate; ligule short, less than 0.5 mm, unevenly laciniate, glabrous; blade erect, amplexicaul, striate, broadly ovate-lanceolate, usually longer than sheath, both surfaces glabrous, margins ciliolate, apex acuminate. Foliage leaves 1 to 4 per ultimate branch; sheath 2–4 cm long, upper half more or less white-pilose abaxially, margins ciliolate, glabrescent; outer ligule ca. 0.5 mm long, margin white-pilose; auricles obscure; oral setae present, to 1.4 cm long; inner ligule short, 0.3–1.5 mm; pseudopetiole 1.5–3 mm long, slightly pilose adaxially, glabrous abaxially; blade lanceolate or narrowly lanceolate, 8–24 × 0.9–2 cm, slightly pilose at base adaxially, hispidulous abaxially, glabrescent, base cuneate, margins serrulate, apex acuminate; lateral veins 4–5 pairs, veinlets conspicuous. The unit of inflorescence of the synflorescence 3–9 cm long; main axis glabrous, flattened on the branching side, basal internode 0.8–1.8 cm long. Paracladia 2–4, 0.4–1 cm long, robust, close to the axis, puberulent or appressed-pubescent at the apex, base without a pulvinus. Spikelets 1.6–2 cm long, stramineous; developed (fertile) florets 1–3, uppermost one rudimentary; rachilla segments thickened upwards, 3–4 mm long, glabrous except the upper part white-tomentose; glumes 2, the lower one lanceolate to narrowly lanceolate, 7–10 mm long, glabrous at lower part, pubescent towards the apex, apex acuminate; the upper one with the same shape and indumentum as the lower one, 7–12 mm long; callus bearded with white- or grey-tomentose hairs; lemmas ovate, the lowest one 10–12 mm long, densely adpressed-pubescent abaxially, 9-veined, apex acuminate-mucronate; palea 7–8 mm long, apex obtuse, glabrous except the ciliate and strongly curved keels; lodicules 3, ca. 2 mm long, anterior 2 oblong-obovate, posterior 1 spatulate or obovate, rounded at apex, lower part brown-nerved, upper part hyaline and minutely ciliate; anthers 3, pale yellow, 5 mm long; ripe ovary 2 mm long, brownish when dry; styles 2, persistent, ca. 1 mm long, base slightly connate, stigmas very thin, plumose, ca. 3 mm long. Fruit unknown. Description of inflorescence follows Keng (1936).


Culm shoots produced in April to July, flowering in May.

Distribution and habitat

This species is distributed in Hunan, Jiangxi and Guangdong, China. It grows under evergreen broadleaved forests at an elevation from 1100–1600 m.

Chinese name

以礼竹 (pinyin: yĭ lĭ zhú).

Additional specimens examined

Kengiochloa pubiflora: China. Guangdong: Ruyuan County, Tianjingshan, 4 May 1978, Z. P. Wang & S. T. Liu 780047 (N019022073, N019022074, N019022075, N019022076); Shixing County, Longtou Moutain, elev. 1131 m, 25 April 2021, Z. Y. Cai & X. H. Ye czy-146 (IBSC); Xinyi City, Mianbeiding, 20 April 2020, N. H. Xia, Y. H. Tong, J. B. Ni, S. J. Zeng & B. M. Wang XNH-63 (IBSC). Hunan: Yizhang County, Mang Mountain, elev. 1600 m, 4 July 1964, B. M. Yang 026501 (N019022077). Jiangxi: Suichuan County, Dabali, 22 April 2022, C. Long BH-148 (IBSC); ibid. 13 November 2022, C. Long s.n. (IBSC). Indocalamus pallidiflorus: CHINA. Guangdong: Longmen County, Nankun Mountain, 12 April 1932, W. T. Tsang 20216 (SYS00095376, US00065467 (image), US00312706 (image), US00312707 (image)); ibid., 18 April 2018, X. R. Zheng, J. B. Ni & Y. Y. Zhang zxr-145 (IBSC). Indocalamus sinicus: CHINA. Hongkong: hill sides, C. Wright s.n. (K00092161 (image)); without precise locality, Hance s.n. (K00092162 (image), K00092163 (image)).


We are grateful to Dr. Si-Jin Zeng, Mr. Bing-Mou Wang, Mr. Yuan-Qiu Li, Ms. Xue-He Ye and Mr Chuan Long for help during field trips. Mr. Xi-Rong Zheng is thanked for providing the photo of Indocalamus sinicus. This work was supported by the Natural Science Foundation of Guangdong Province (no. 2114050002139) and National Natural Science Foundation of China (grant no. 32270227).


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