Next Article in Journal
Solenopsis gutermannii (Campanulaceae), a New Species from Kefalonia (Ionian Islands, Greece)
Previous Article in Journal
A New Species in the Canthon indigaceus Species Group (Coleoptera: Scarabaeidae: Scarabaeinae) from the Mexican Pacific Coast
 
 
Search for Articles:
Title / Keyword
Author / Affiliation / Email
Journal
Article Type
 
 
Advanced Search
 
Section
Special Issue
Volume
Issue
Number
Page
 
Journals
Taxonomy
Volume 5
Issue 1
10.3390/taxonomy5010012
taxonomy-logo
Submit to this Journal Review for this Journal Propose a Special Issue
► Article Menu

Article Menu

share Share announcement Help format_quote Cite question_answer Discuss in SciProfiles
first_page
settings
Order Article Reprints
Font Type:
Arial Georgia Verdana
Font Size:
Aa Aa Aa
Line Spacing:
Column Width:
Background:
Article

New Combination and Two Synonyms of Indocalamus Nakai (Poaceae: Bambusoideae) from China Based on Morphological Characters and Phylogenetic Evidence

by
Liqin Gao
1,2,
Yonglong Li
2,
Chengkun Wang
2,
Jianqing Wang
2,
Guangyao Yang
2 and
Wengen Zhang
2,*
1
Jiangxi Academy of Forestry, Nanchang 330032, China
2
Jiangxi Provincial Key Laboratory of Improved Variety Breeding and Efficient Utilization of Native Tree Species, Jiangxi Agricultural University, Nanchang 330045, China
*
Author to whom correspondence should be addressed.
Taxonomy 2025, 5(1), 12; https://doi.org/10.3390/taxonomy5010012
Submission received: 20 January 2025 / Revised: 19 February 2025 / Accepted: 20 February 2025 / Published: 28 February 2025
Browse Figures
Review Reports Versions Notes

Abstract

:
Based on several sources of evidence from morphology, the micromorphology of the abaxial leaf epidermis, and a phylogenetic analysis using complete chloroplast genome data, a new combination and two new synonyms are reported in the genus Indocalamus Nakai. Indocalamus cordatus T. H. Wen et Y. Zou is synonymized with I. tessellatus (Munro) P. C. Keng, and I. chebalingensis W. T. Lin is synonymized with I. multinerus (W. T. Lin et Z. M. Wu) W. T. Lin. As a new combination, I. mollis (H. R. Zhao et Y. L. Yang) L. Q. Gao, W. G. Zhang et G. Y. Yang was created and is supplementally described. In addition, the lectotype of I. multinerus is designated herein.

1. Introduction

Indocalamus Nakai [1], ca. 23 species, is a frutescent genus of the temperate woody bamboos (i.e., the tribe Arundinarieae of Bambusoideae in Poaceae) with important economic and ecological value [2,3,4]. It is characterized by leptomorph rhizomes, solitary branches, persistent culm sheaths, relatively large leaves with distinct transverse veins, racemose or paniculate inflorescences, pedicellate spikelets, three stamens, and two stigmas [2,5,6,7]. The genus is mainly distributed in the southern Yangtze River in China, at elevations of 300–2400 m under evergreen broad-leaved forests, with culms used to make chopsticks and Chinese brush and leaves generally used for weaving bamboo hats and wrapping glutinous rice in China and some adjacent countries [2,5,8].
Molecular phylogenetic studies have recently indicated that Indocalamus is not monophyletic because its species were generally clustered into several lineages together with other genera, e.g., Ferrocalamus Hsueh & Keng f. in Keng & Hsueh [9], Gelidocalamus T. H. Wen [10], Yushania Keng [11], and Fargesia Franchet [3,12,13,14,15]. To accelerate the process of the taxonomic revision of the genus, taxonomical reassessment of some Indocalamus taxa is not only needed but obligatory. Recently, I. chouzhouensis Yi & Yang [16] was examined and synonymized with I. emeiensis C. D. Chu & C. S. Chao [5,15].
In the study, six taxa of Indocalamus, i.e., I. cordatus T. H. Wen et Y. Zou [17], I. tessellatus (Munro) P. C. Keng [11], I. multinerus (W. T. Lin et Z. M. Wu) W. T. Lin [18], I. chebalingensis W. T. Lin [19], I. guangdongensis H. R. Zhao et Y. L. Yang [20], and its variety I. guangdongensis var. mollis H. R. Zhao et Y. L. Yang [20], were re-evaluated. Based on several field works from 2019 to 2021, herbarium specimens, the micromorphological characteristics of abaxial leaf epidermis, and the phylogenetic tree were investigated comprehensively using complete chloroplast genome data, and the above taxa are treated herein.

2. Materials and Methods

2.1. Morphological Observation

Morphological data were obtained from filed investigations and based on the analysis of specimens deposited in the herbaria of Royal Botanical Gardens Kew (K), British Natural History Museum (BM), Nanjing University (N), Nanjing Forestry University (NFU), South China Agricultural University (CANT), Forestry School of Sichuan Province (SIFS), Sun Yat-Sen University (SYS), and Zhejiang Academy of Forestry (ZJFI); type specimens were particularly digitalized at BM and K. Six related taxa of Indocalamus were chosen from morphological observation (Supplementary Table S1). Some key features, e.g., culm leaf, including sheath, blade, auricles, and oral setae, mature culm appendage, node, and its longitudinal section, and foliage leaf, including sheath, auricles, and oral setae, were compared and photographed with the SLR camera EOS 60D using the macro lens EF 50 mm f/1.8 STM (Canon, Japan). Specimens were studied in the laboratory under a stereomicroscope. Voucher specimens were deposited in the herbarium of the College of Forestry, Jiangxi Agricultural University, China (JXAU).

2.2. Micromorphology of Abaxial Leaf Epidermis Study

Mature leaves were collected in the field and fixed in formalin–acetic acid–alcohol (FAA). We sampled 6 taxa of Indocalamus. We used the middle portion of foliage leaf blades for all samples. Leaves fixed in FAA were cleaned using the ultrasonic cleaner CPX2800H-C (Branson, CT, USA) with clean water and dried at room temperature before they were mounted on stubs. After gold sputtering with gold powder (3 nm), we photographed the samples using the scanning electron microscope NanoSEM 450 (Nova, OR, USA). The terminology for the epidermis appendages follows Ellis [21], Zhang et al. [22], and Leandro et al. [23]. Voucher specimens were deposited in the herbarium of the College of Forestry, Jiangxi Agricultural University, China (JXAU).

2.3. Molecular Phylogeny

2.3.1. Taxon Sampling

Leaf material was collected from living plants from type localities in China, including provinces Chongqing, Jiangxi, and Guangdong. A total of 61 individuals were sampled, including 7 new complete chloroplast genomes of Indocalamus in this study, as well as 51 complete chloroplast genomes to represent the 12 major clades of the tribe Arundinarieae (Supplementary Table S1), together with Bambusa emeiensis Chia & Fung [24], Buergersiochloa bambusoides Pilg. [25], and Chusquea culeou E. Desv. [26] as the outgroups, were obtained from NCBI (National Center for Biotechnology Information, USA) and the matrix of Guo et al. [4]. Fresh leaves were collected for all samples and then rapidly dried in silica gel. Voucher specimens were deposited at JXAU.

2.3.2. DNA Extraction, Library Preparation, and Data Assembly

Total genomic DNA was extracted from fresh leaves using a modified cetyl-trimethylammonium bromide (CTAB) method [27]. Two Illumina paired-end libraries were prepared and sequenced at Novogene Bioinformatics Technology Co., Ltd. (Beijing, China), and 6 GB of raw data for each taxon with an average read length of 150 bp were acquired. All contigs of two chloroplast genome sequences were spliced and assembled using SPAdes 3.13.0 [28]. Then, complete chloroplast genomes were annotated using CPGAVAS2 [29] and Geneious 9.1.4 [30], and they were finally uploaded to the National Center for Biotechnology Information.

2.3.3. Data Matrices and Phylogenetic Analyses

In order to understand the phylogenetic relationships among 5 species and 1 variant within Indocalamus, we selected and reannotated 51 complete chloroplast genomes to represent the 12 major clades of the tribe Arundinarieae, together with 3 outgroups (Supplementary Table S1).
Maximum likelihood (ML) and Bayesian inference (BI) analyses were performed to infer phylogenetic relationships. ML analyses with rapid bootstrapping and 1000 replications (IQ-TREE) [31] were conducted using complete chloroplast sequences under the TVM+I+G model, as recommended by jModelTest v2.1.4 [32]. With the same model, BI analyses were conducted in MrBayes v3.2.6 [33] using 2 runs of 4 chains each, and these analyses were run for ten million generations with the first 5000 trees discarded as burn-in.

3. Results and Discussion

3.1. Morphological Comparison

Species in Indocalamus have hollow culms, persistent culm sheathes, with the solitary branches sometimes as thick as the main culms, and large leaves with distinct transverse veins. We found other vegetative characteristics that provide useful information for identification in the field. Taxa in Indocalamus have some characters useful for identification in field populations, such as appendages on the infranodal regions of young culms and culm sheaths, the relationship between blades and the main culms, the shape and size of sheath auricles, and the number of leaves on the ultimate branches. The morphological characteristics of the six species were compared and analyzed as follows.
Indocalamus guangdongensis var. mollis was described by Huiru Zhao and Yaling Yang, and it was distinguished from I. guangdongensis mainly with a row of dense, short villi below the midrib side of leaves [20]. After studying specimens from both regions and field trips to type localities, we can conclude that the differences concerning the morphological characteristics between I. guangdongensis var. mollis and I. guangdongensis are significant.
The hairs on the mature culm are different. I. guangdongensis is densely covered with short, white tomentose (Figure 1A), and I. guangdongensis var. mollis is hairless (Figure 1G). The former’s culm node is elevated (Figure 1B,C), and the latter’s is slightly elevated (Figure 1H,I). The most obvious difference is the shape and form of the blade on the culm sheaths. The former has erect, ovate–lanceolate blades (Figure 1D), and the latter has recurved, subulate, or linear blades (Figure 1J). The leaf sheathes are also different. I. guangdongensis is dense with white tomenta and oral setae (Figure 1E), and I. guangdongensis var. mollis is smooth and hairless (Figure 1K). Moreover, the former has an abaxially glabrous leaf (Figure 1F), and the latter exhibits white pubescence on one side of the midrib or extending to the leaf base (Figure 1L). Therefore, there are significant differences in morphological characteristics between them.
Indocalamus cordatus is distinguished from other Indocalamus species mainly because of its cordate leaf base (Figure 2A,B), and both sides are asymmetrical along the midrib [17]. An observation of the paratypes of I. cordatus reveals that the character of the leaf base cordate is not stable (Figure 2C). An observation of the image of type and neotype specimens (Figure 3A,B) reveals that the mentioned differences fall within the variability encompassed by I. tessellatus. We investigated three populations of I. tessellatus in the Hunan, Chongqing, and Jiangxi provinces, and we carried out a further morphological comparison for I. tessellatus and I. cordatus. In this study, we selected the populations of Jiangxi Province with relatively complete morphological characteristics for the comparison herein. Some key characters showed no essential morphological differences between the two taxa above.
There are similarities in the habitats (Figure 4A,G). Annual mature sheathes are densely black–brown and strigose (Figure 4B,H). The infranodal culm region is covered with short, white tomenta (Figure 4C,I). The lanceolate blade is erect and variable in size (Figure 4D,J). Auricles and oral setae are absent, and the ligule is very short, with brown hirtellous (Figure 4E,K). The leaf sheaths are leathery and glabrous; the auricles and oral setae are rare or scarce (Figure 4F,L).
From the time of its original description until the present work, Indocalamus multinerus had been consistently unaccepted in previous studies, including in the Flora of China [2]. Yi et al. [34] placed I. multinerus in the incertae sedis taxon, but Vorontsova et al. [35] accepted it. Additionally, it had been synonymized with I. longiauritus Hand.-Mazz. [36]. After examining type specimens (Figure 5A,B) kept at CANT, we found that culm sheaths and leaf sheaths, blades, auricles, and oral setae showed no essential morphological differences between I. multinerus and I. chebalingensis. To figure out whether Indocalamus multinerus and I. chebalingensis are conspecific, we investigated the type localities, and we carried out a further morphological comparison (Figure 6). The results of some key characteristics showed no essential morphological differences between the above two taxa. Annual culms were uniformly distributed with pressed white fur–felt (Figure 6B,I). The culm sheaths of both taxa exhibit dense reddish brown bristles on the basal region of young culms (Figure 6A,H), the ovate–lanceolate blades are erect and hugging the culms, and the smaller falcate auricles show radiating oral setae (Figure 6C,J). The mature culms are nearly solid with a wall of about 3 mm in thickness and spongy filler (Figure 6F,G,M,N). The branches are usually solitary, with two to three upper branches (Figure 6E,L). Additionally, both taxa have three or four leaves on the ultimate branches with weak falcate auricles and radiating oral setae (Figure 6D,K).

3.2. Micromorphological Comparison of the Abaxial Leaf Epidermis

As a result of the scarcity of flowering collections, bamboo species identification mainly depends on vegetative features, and leaf epidermal micromorphology has proven useful in bamboo taxonomy [15,22,23].
Stomatal apparatuses, papillae, and various hairs are usually observed on the abaxial leaf epidermis. The leaf surface traits of all examined taxa are shown in Figure 7.
The abaxial leaf epidermis characteristics of I. guangdongensis var. mollis and I. guangdongensis are completely different. In the former one, there are two to four rows of stomatal apparatuses between the veins. Eight to ten elongated papillae overarch and have an apparent arciform shape that totally covers and conceals the stomata. Short papillae are abundant, and they are distributed in parallel with the veins and prickles. The microhairs are curved, and the silica bodies are inconspicuous (Figure 7A). A difference is that the former one, I. guangdongensis, has one to two rows of stomatal apparatuses between the veins. Most notable are the characteristics of the stomatal apparatuses and silica bodies. The elongated papillae, forming an inconspicuous, arciform shape, cover the stomata, and silica bodies are rare. Microhairs are straight and tiled on the veins (Figure 7B). As a result, I. guangdongensis var. mollis has entirely different micromorphological characteristics in its leaf epidermis than I. guangdongensis. The micromorphology of the abaxial leaf blade epidermis can separate these two taxa adequately.
Via scanning electron microscopy, the micromorphological features of the abaxial leaf epidermis of the population in the type locality of I. cordatus (Figure 7C) and three populations of I. tessellatus (Figure 7D–F) were investigated. Eight to ten elongated papillae overarch the stomatal apparatus, and short papillae are densely present. Macrohairs and microhairs are few in number, and they are scattered and distributed in the intercostal region. Saddle-shaped silica bodies are non-significant, and they are distributed in the costal and intercostal regions. As a result, I. cordatus has exactly the same micromorphological characteristics of the abaxial leaf epidermis as I. tessellatus.
Indocalamus chebalingensis and I. multinerus are characterized by the stomatal apparatuses, papillae, microhairs, prickles, and silica bodies (Figure 7G,H). We did not observe macrohairs. There are one to two rows of stomatal apparatuses between the veins. Eight to ten elongated papillae overarch, and an arciform structure totally covers and conceals the stomata. The short papillae are rare. Microhairs are distributed in the intercostal region and with equal lengths of the basal cells and apical cells (the apical cells are sleazy and brittle). Prickles occur on the costal and intercostal regions, and the direction of the sharp apices is irregular. The saddle-shaped silica bodies on the veins are also clearly identified. There are no significant differences between the two characteristics above.

3.3. Molecular Phylogenetic Relationships

In the phylogenetic tree, this study involved six taxa and eight populations of Indocalamus that are clustered into the Phyllostachys clade (V), which is consistent with most Indocalamus taxa.
Indocalamus guangdongensis var. mollis and I. guangdongensis are clustered with a distant clade, which shows a lower level of homology between the two taxa (Figure 8). The molecular phylogenetic relationships do not support the belief that the two taxa are the same. Therefore, Indocalamus guangdongensis var. mollis should be a separate taxon.
Indocalamus cordatus and I. tessellatus, I. multinerus, and I. chebalingensis are clustered together in the same clade with an almost negligible branch length, which shows a high level of homology between these taxa (a bootstrap value of 100% in the ML analysis and a posterior probability of 1.0 in the BI analysis) (Figure 8). The molecular phylogenetic relationships support the conclusions that I. cordatus is the same as I. tessellatus, that I. multinerus is the same as I. chebalingensis, and that the former should be merged with the latter.

4. Taxonomic Treatment

Based on the morphological, micromorphological, and molecular evidence, we here propose a new combination and two new synonyms of Indocalamus in China.
1. Indocalamus mollis (H. R. Zhao et Y. L. Yang) L. Q. Gao, W. G. Zhang et G. Y. Yang, comb. nov.
柔毛箬竹 (Rou Mao Ruo Zhu); Figure 9.
Basionym: Indocalamus guangdongensis var. mollis H. R. Zhao et Y. L. Yang in Acta Phytotaxon. Sin. 23 (6): 462. 1985.
Type: CHINA. Hunan: Yongzhou, Shuangpai, Yangming Shan, 14 May 1977, Z.P.Wang 77018 (holotype: N).
Description: Rhizomes leptomorph. Culms 0.9–2.2 m tall, 0.5–1.1 cm in diameter, hollow; internodes terete, 10–19 cm long, young culm puberulous, infranodal region white and light brown velutinous, glabrescent when old; wall 1.5–2.5 mm thick; supranodal ridge prominently slight or flat; intranodes 4–7 mm long. Culm sheaths tardily deciduous, oblong, thinly leathery or thickly papery, subequal to internodes, densely reddish brown strigose and aducous, margin densely reddish brown and ciliate; auricles not developed; oral setae none or several, caducous; ligule subtruncate, ca. 1–1.5 mm high, densely pubescent; blades recurved, subulate, or linear. Foliage leaves, 2–4 per ultimate branch; leaf sheaths smooth and hairless, margin without ciliate; leaf auricles not developed; oral setae, none; ligules truncate, 2–3 mm tall, margin without ciliate; blades oblong-lanceolate, 24.0–47.0 × 5.0–9.0 cm, adaxially glabrous, abaxially white and pubescent along one side of midrib and extending to leaf base; longitudinal veins, 8–14 pairs, base cuneate, apex long-acuminate, margin entire. Inflorescence unknown.
Additional specimens examined: —CHINA. Guangxi: Gongcheng, Y.K.Li 402992 (N); Hunan: Yongzhou, Shuangpai, Yangming Shan, Y.Tsiang et S.H.Chen 711 (SYS!); ibid., Y.Tsiang et S.H.Chen 484 (SYS!); ibid., L.Q.Gao et al. GXYM2014 (JXAU!); Hubei: Changyang, Duzhenwan, S.Y.Wang 660 (N).
2. Indocalamus tessellatus (Munro) Keng f., Acta Phytotaxon. Sin. 6(4): 355. 1957.
箬竹 (Ruo Zhu) Figure 2, Figure 3 and Figure 4.
Indocalamus cordatus Wen et Y. Zou, J. Bamboo Res. 10(1): 18~19. f. 3. 1991. Syn. Nov.—Type: CHINA. Jiangxi, Duchang, Dagang reservoir, 10 Sep 1982, Zou Yuan 286 (paratype: ZJFI!; NFU!); Duchang, Dagang, Fengshuxia, 2 Jun 1990, T. H. Wen et S. Jin 90661 (holotype: ZJFI).
Type: UK. London, Kew Bamboo Garden (cultivated, introduced from China), 18 Aug 1998, Stapleton 1122 (neotypes, designated by Stapleton (2000), K! [K000912169, K000912170, K000912171, K000912172]).
Additional specimens examined: —CHINA. Hunan: Ningyuan, Jiuyi Mountain, alt. 1350 m, 4 Sep. 1979, B.M.Yang 06329 (NYA!); Liuyang, Yangming Mountain, alt. 1050 m, 27 Aug. 2020, L.Q.Gao 2001 (JAXU!); Shimen, Jiashan town, alt. 650 m, 30 Aug. 2020, L.Q.Gao 2004 (JAXU!); Jiangxi: Xiushui, alt. 600 m, 13 Aug. 1994, C.M.Tan 94755 (PE!); ibid., alt. 800 m, 30 Apr. 1994, C.M.Tan 94342 (PE!); ibid., alt. 500, 5 Jun. 2016, C.M.Tan 1606985 (JJF!); Jing’an, alt. 150, 30 Jul. 2013, H.G.Ye et al. LXP10-31 (IBSC!); Fujian: Jiangle, alt. 600 m, 20 Sep. 1991, Longxi expedition 2515 (PE!); ibid., alt. 700, 16 Sep. 1991, Longxi expedition 2183 (PE!); Longyan, alt. 1185 m, 18 Jul. 2017, X.F.Zeng ZXF29072 (CZH!); ibid., alt. 929 m, 1 Jun. 2009, H.Z.Guo et al. 1822 (XMBG!); Zhejiang: Hangzhou, May 1980, Yao et Meng 80013 (HZ!); Anhui: Huoshan, alt. 200, 25 Sep. 1983, M.B.Deng 81671 (NAS!); Hubei: Jingshan, alt. 303, 6 Sep. 2005, J.Q.Wu et X.X.Yin 6736 (HIB!).
3. Indocalamus multinerus (W. T. Lin et Z. M. Wu) W. T. Lin, J. Bamboo Res. 19(4): 17. f. 1. 2000.
多脉箬竹 (Duo Mai Ruo Zhu); Figure 5 and Figure 6.
Indocalamus chebalingensis W. T. Lin, J. Bamboo Res. 19(1): 6. f. 1. 2000. Syn. Nov. Type: —CHINA. Guangdong, Shixing, Chebaling, 1 July 1989, C.L.Zhang 56102 (Syntypes: CANT! [CANT00002930, CANT00002931, CANT00002933, CANT00002934]).
Basionym: Arundinaria multinervis W. T. Lin et Z. M. Wu, J. South China Agr. Univ. 11(3):48. f. 3. 1990.
Type: —CHINA. Guangdong, Shixing, Chebaling, 300 m, 25 June 1989, Z.M.Wu 55876 (lectotype, designated here: CANT! [CANT00002994]; isolectotypes: CANT! [CANT00002992, CANT00002993, CANT00002995, CANT00002997, CANT00002998, CANT00002999]).
Additional specimens examined: —CHINA. Guangdong: Shixing, Chebaling, 29 July 1989, C. L. Zhang 56072; ibid., alt. 336 m, 26 Aug. 2020, L. Q. Gao et al. GDSX02 (JXAU!); ibid., alt. 384 m, 26 Aug. 2020, L. Q. Gao et al. GDSX02 (JXAU!); Hunan: Yongzhou, Dong’an, alt. 592 m, 27 Aug. 2020, L. Q. Gao et al. HN2032 (JXAU!).

Supplementary Materials

The following supporting information can be downloaded at https://www.mdpi.com/article/10.3390/taxonomy5010012/s1, Table S1: Data Matrices.

Author Contributions

Conceptualization: L.G. Investigation: L.G. and Y.L. Formal analysis: C.W. and J.W. Funding acquisition: G.Y. and W.Z. All authors have read and agreed to the published version of the manuscript.

Funding

This work was supported by the National Natural Science Foundation of China [Guangyao Yang, 31960335] and the Key R&D Planned Projects of Jiangxi Province [Wengen Zhang, 20192BBF60015].

Data Availability Statement

The data that support the findings of this study are openly available in the Science Data Bank at https://doi.org/10.57760/sciencedb.j00143.00093 (accessed on 26 January 2024).

Acknowledgments

We are grateful to the curators and officers of CANT, N, NJU, SIFS, SYS, and ZJFI for the permission to use their scanned images of specimens and research facilities. Thanks are also extended to Jiancheng Zhao (ZJFI), Lin Yang (SIFS), Xuehong Ma (NFU), Mingxuan Zhen (CANT), and Yanjun Pang (N) for facilitating some types of material from the respective herbaria and also to two anonymous reviewers for their valuable comments.

Conflicts of Interest

The authors have declared that no competing interests related to this work exist.

References

  1. Nakai, T. Two new genera of Bambusaceae, with special remarks on the related genera growing in eastern Asia. J. Arnold Arboretum 1925, 6, 145–153. [Google Scholar] [CrossRef]
  2. Wang, Z.P.; Stapleton, C. Indocalamus. In Flora of China; Wu, Z.Y., Raven, P.H., Hong, D.Y., Eds.; Science Press: Beijing, China; The Missouri Botanical Garden Press: St. Louis, Mo, USA, 2006; Volume 22, pp. 135–143. [Google Scholar]
  3. Guo, C.; Guo, Z.H.; Li, D.Z. Phylogenomic analyses reveal intractable evolutionary history of a temperate bamboo genus (Poaceae: Bambusoideae). Plant Divers. 2019, 41, 213–219. [Google Scholar] [CrossRef]
  4. Guo, C.; Ma, P.F.; Yang, G.Q.; Ye, X.Y.; Guo, Y.; Liu, J.X.; Liu, Y.L.; Eaton, D.A.R.; Guo, Z.H.; Li, D.Z. Parallel ddRAD and Genome Skimming Analyses Reveal a Radiative and Reticulate Evolutionary History of the Temperate Bamboos. Syst. Biol. 2021, 70, 756–773. [Google Scholar] [CrossRef] [PubMed]
  5. Chao, Q.S.; Chu, Z.D.; Hsiung, W.Y. A revision of some genera and species of Chinese bamboos. Acta Phytotaxon. 1980, 18, 20–36. [Google Scholar]
  6. Yang, Y.L. A revision of the genus Indocalamus of Bambusoideae from the world (I). J. Nanjing Univ. (Nat. Sci.) 1987, 23, 453–462. [Google Scholar]
  7. Zhao, H.R.; Yang, Y.L. Bambusoideae. In Flora Reipublicae Popularis Sinicae; Keng, P.C., Wang, Z.P., Eds.; Science Press: Beijing, China, 1996; Volume 9, pp. 676–704. [Google Scholar]
  8. Yang, Y.L. The endemic genus of China: Geographical distribution of Indocalamus Nakai. J. Plant Resour. Environ. 1993, 2, 41–44. [Google Scholar]
  9. Keng, P.C.; Hsueh, J.R. Ferrocalamus Hsueh & Keng f., a new bamboo genus in China. J. Bamboo Res. 1982, 1, 137–139. [Google Scholar]
  10. Wen, T.H. A new genus and some new species of Bambusoideae from China. J. Bamboo Res. 1982, 1, 20–45. [Google Scholar]
  11. Keng, P.C. One new genus and two new species of Chinese bamboos. J. Syst. Evol. 1957, 6, 355–360. [Google Scholar]
  12. Franchet, M.A. Fargesia, nouveau genre de Bambuseés de la Chine. Bull. Mens. Soc. Linn. Paris 1893, 2, 1067–1069. [Google Scholar]
  13. Zhou, Y.; Zhang, Y.Q.; Hou, X.Q.; Huang, L.; Kang, J.Q.; Zhang, J.Q.; Ren, Y. Phylogeny of Fargesia (Poaceae: Bambusoideae) and infrageneric adaptive divergence inferred from three cpDNA and nrITS sequence data. Plant Syst. Evol. 2019, 305, 61–75. [Google Scholar]
  14. Qin, Q.M.; Tong, Y.H.; Zheng, X.R.; Ni, J.B.; Xia, N.H. Sinosasa (Poaceae: Bambusoideae), a new genus from China. Taxon 2020, 70, 27–47. [Google Scholar] [CrossRef]
  15. Gao, L.Q.; Li, Y.L.; Guo, C.C.; Yang, G.Y.; Zhang, W.G. Indocalamus chongzhouensis (Poaceae: Bambusoideae), a new synonym of I. emeiensis: Evidence from morphology and complete chloroplast genome data. Phytotaxa 2022, 542, 53–63. [Google Scholar] [CrossRef]
  16. Yi, T.P.; Yang, L. A new species of Indocalamus Nakai from western Sichuan of China. J. Bamboo Res. 2004, 23, 13–15. [Google Scholar]
  17. Wen, T.H. Some ideas on the taxonomy of several Bambusoideae taxonss. J. Bamboo Res. 1991, 10, 11–25. [Google Scholar]
  18. Lin, W.T. A new species and two new combinations of Bambusoideae. J. Bamboo Res. 2000, 19, 6–8. [Google Scholar]
  19. Lin, W.T. Two new species and one new combinations of Bambusoideae. J. Bamboo Res. 2000, 19, 1–2, 17. [Google Scholar]
  20. Zhao, H.R.; Yang, Y.L. New taxa and new combinations of Indocalamus from China. J. Syst. Evol. 1985, 23, 460–465. [Google Scholar]
  21. Ellis, R.P. A procedure for standardizing comparative leaf anatomy in the Poaceae. II The epidermis as seen in surface view. Bothalia 1979, 12, 641–671. [Google Scholar] [CrossRef]
  22. Zhang, Y.X.; Zeng, C.X.; Li, D.Z. Scanning electron microscopy of the leaf epidermis in Arundinarieae (Poaceae: Bambusoideae): Evolutionary implications of selected micromorphological features. Bot. J. Linn. Soc. 2014, 176, 46–65. [Google Scholar] [CrossRef]
  23. Leandro, T.D.; Scatena, V.L.; Clark, L.G. Comparative leaf blade anatomy and micromorphology in the systematics and phylogeny of Bambusoideae (Poaceae: Poales). Bot. J. Linn. Soc. 2019, 192, 165–183. [Google Scholar] [CrossRef]
  24. Chia, L.Z.; Fung, X.L. On the validity of the genera Sinocalamus McClure and Lingnania McClure. J. Syst. Evol. 1980, 18, 211–216. [Google Scholar]
  25. Pilger, R. Neue und weniger bekannte Gramineen aus papuasien. Bot. Jahrb. Syst. 1914, 52, 168. [Google Scholar]
  26. Desvaux, Étienne-Émile. Flora Chilena 6; Paris, France, 1854; p. 450, t. 83. Available online: http://www.botanicus.org/page/1394960 (accessed on 26 January 2024).
  27. Doyle, J.J.; Doyle, J.L. A rapid DNA isolation procedure for small quantities of fresh leaf tissue. Phytochem. Bull. 1987, 19, 11–15. [Google Scholar]
  28. Bankevich, A.; Nurk, S.; Antipov, D.; Gurevich, A.A.; Dvorkin, M.; Kulikov, A.S.; Lesin, V.M.; Nikolenko, S.I.; Pham, S.; Prjibelski, A.D.; et al. SPAdes: A new genome assembly algorithm and its applications to single-cell sequencing. J. Comput. Biol. 2012, 19, 455–477. [Google Scholar] [CrossRef]
  29. Shi, L.C.; Chen, H.M.; Jiang, M.; Wang, L.; Wu, X.; Huang, L.; Liu, C. CPGAVAS2, an integrated plastome sequence annotator and analyzer. Nucl. Acid. Res. 2019, 47, 65–73. [Google Scholar] [CrossRef]
  30. Kearse, M.; Moir, R.; Wilson, A.; Stones-Havas, S.; Cheung, M.; Sturrock, S.; Buxton, S.; Cooper, A.; Markowitz, S.; Duran, C.; et al. Geneious Basic: An integrated and extendable desktop software platform for the organization and analysis of sequence data. Bioinformatics 2012, 12, 1647–1649. [Google Scholar] [CrossRef]
  31. Nguyen, L.T.; Schmidt, H.A.; Haeseler, A.; Minh, B.Q. IQ-TREE: A fast and effective stochastic algorithm for estimating maximum-likelihood phylogenies. Mol. Biol. Evol. 2014, 32, 268–274. [Google Scholar] [CrossRef]
  32. Darriba, D.; Taboada, G.L.; Doallo, R.; Posada, D. jModelTest 2: More models, new heuristics and parallel computing. Nat. Method 2012, 9, 772. [Google Scholar] [CrossRef]
  33. Ronquist, F.; Teslenko, M.; van der Mark, P.; Ayres, D.L.; Darling, A.; Höhna, S.; Larget, B.; Liu, L.; Suchard, M.A.; Huelsenbeck, J.P. MrBayes 3.2: Efficient Bayesian phylogenetic inference and model choice across a large model space. Syst. Biol. 2012, 61, 539–542. [Google Scholar] [CrossRef]
  34. Yi, T.P.; Shi, J.Y.; Ma, L.S.; Wang, H.T.; Yang, L. Iconographia Bambusoidearum Sinicarum; Science Press: Beijing, China, 2008; pp. 688–712, 728. [Google Scholar]
  35. Vorontsova, M.S.; Clark, L.G.; Dransfield, J.; Govaerts, R.; Baker, W.J. World Checklist of Bamboos and Rattans; Science Press: Beijing, China, 2016; pp. 106–110. [Google Scholar]
  36. Xia, N.H.; Lin, R.S.; Wang, R.H. Taxonomic notes of Bambusoideae (Poaceae) from Guangdong. J. Trop. Subtrop. Bot. 2009, 17, 351–354. [Google Scholar]
Taxonomy 05 00012 g001
Figure 1. Morphological comparison between Indocalamus guangdongensis (A–F) and I. guangdongensis var. mollis (G–L). (A,G) Part of annual mature culm, showing appendages on the surface of the culm; (B,H) Part of the mature biennial culm, showing the supranodal branch ridge; (C,I) Part of the annual mature culm, showing basal warty setae on the culm sheath and appendages on the infranodal region; (D,J) Blade; (E,K) Part of the leaf sheath, showing a ligule and oral setae; (F,L) Part of the annual mature blade, showing the leaf base, back, and hair. (scale bars = 1 cm).
Figure 1. Morphological comparison between Indocalamus guangdongensis (A–F) and I. guangdongensis var. mollis (G–L). (A,G) Part of annual mature culm, showing appendages on the surface of the culm; (B,H) Part of the mature biennial culm, showing the supranodal branch ridge; (C,I) Part of the annual mature culm, showing basal warty setae on the culm sheath and appendages on the infranodal region; (D,J) Blade; (E,K) Part of the leaf sheath, showing a ligule and oral setae; (F,L) Part of the annual mature blade, showing the leaf base, back, and hair. (scale bars = 1 cm).
Taxonomy 05 00012 g001
Taxonomy 05 00012 g002
Figure 2. Indocalamus cordatus. (A) Holotype illustration; (A1) Branches, leaves and culms; (A2) Outer surface of culm sheaths; (A3) Inner surface of culm sheaths; Illustrated by Jin S. (from specimen Wen T.H. et Jin S. no. 90661, ZJFI). (B,C). Paratypes (Y. Zou 286, ZJFI! and NF!). (scale bars = 1 cm).
Figure 2. Indocalamus cordatus. (A) Holotype illustration; (A1) Branches, leaves and culms; (A2) Outer surface of culm sheaths; (A3) Inner surface of culm sheaths; Illustrated by Jin S. (from specimen Wen T.H. et Jin S. no. 90661, ZJFI). (B,C). Paratypes (Y. Zou 286, ZJFI! and NF!). (scale bars = 1 cm).
Taxonomy 05 00012 g002
Taxonomy 05 00012 g003
Figure 3. Indocalamus tessellatus. (A) Type (Munro s.n., BM000959209 [image!]); (B) Neotype (Stapleton 1122, K000912169 [image!]). (scale bars = 1 cm).
Figure 3. Indocalamus tessellatus. (A) Type (Munro s.n., BM000959209 [image!]); (B) Neotype (Stapleton 1122, K000912169 [image!]). (scale bars = 1 cm).
Taxonomy 05 00012 g003
Taxonomy 05 00012 g004
Figure 4. Morphological comparison between Indocalamus cordatus (A–F) and I. tessellatus (G–L). (A,G) Habitat; (B,H) Culm sheath, showing basal warty setae; (C,I) Part of annual culm, showing white, felted wool infranodal region; (D,J) Sheath auricle and oral setae; (E,K) Blade and ligule; (F,L) Part of the ultimate leafy branch, showing leaf sheath, micro auricles, and oral setae. (scale bars = 1 cm).
Figure 4. Morphological comparison between Indocalamus cordatus (A–F) and I. tessellatus (G–L). (A,G) Habitat; (B,H) Culm sheath, showing basal warty setae; (C,I) Part of annual culm, showing white, felted wool infranodal region; (D,J) Sheath auricle and oral setae; (E,K) Blade and ligule; (F,L) Part of the ultimate leafy branch, showing leaf sheath, micro auricles, and oral setae. (scale bars = 1 cm).
Taxonomy 05 00012 g004
Taxonomy 05 00012 g005
Figure 5. (A) Syntype of Indocalamus chebalingensis (C. L. Zhang 56102, CANT!); (B) Lectotype of I. multinerus (Z. M. Wu 55876, CANT!). (scale bars = 1 cm).
Figure 5. (A) Syntype of Indocalamus chebalingensis (C. L. Zhang 56102, CANT!); (B) Lectotype of I. multinerus (Z. M. Wu 55876, CANT!). (scale bars = 1 cm).
Taxonomy 05 00012 g005
Taxonomy 05 00012 g006
Figure 6. Morphological comparison between Indocalamus chebalingensis (A–G) and I. multinerus (H–N). (A,H) Part of annual young culm, showing white, felted wool infranodal region; (B,I) Part of annual mature culm, showing powdery, short, white hair; (C,J) Blade, ligule, sheath auricle, and oral setae; (D,K) Part of ultimate leafy branch, showing leaf sheath, auricles, and oral setae; (E,L) Part of mature culm, showing branch; (F,M) Part of mature culm, cross section; (G,N) Part of mature culm, longitudinal section. (scale bars = 1 cm).
Figure 6. Morphological comparison between Indocalamus chebalingensis (A–G) and I. multinerus (H–N). (A,H) Part of annual young culm, showing white, felted wool infranodal region; (B,I) Part of annual mature culm, showing powdery, short, white hair; (C,J) Blade, ligule, sheath auricle, and oral setae; (D,K) Part of ultimate leafy branch, showing leaf sheath, auricles, and oral setae; (E,L) Part of mature culm, showing branch; (F,M) Part of mature culm, cross section; (G,N) Part of mature culm, longitudinal section. (scale bars = 1 cm).
Taxonomy 05 00012 g006
Taxonomy 05 00012 g007
Figure 7. Micromorphology of the abaxial leaf epidermis of Indocalamus (scanning electron microscopy). (A) I. guangdongensis var. mollis (Yangmingshan, Hunan, China); (B) I. guangdongensis (Futang, Guangdong, China); (C) I. cordatus (DuChang, Jiangxi, China); (D) I. tessellatus (Daweishan, Hunan, China); (E) I. tessellatus (Xiushan, Chongqing, China); (F) I. tessellatus (Nanchang, Jiangxi, China); (G) I. chebalingensis (Chebaling, Guangdong, China); (H) I. multinerus (Chebaling, Guangdong, China). Abbreviations: ep, elongated papillae; sp, short papillae; ma, macrohairs; mi, microhairs; pr, prickles; sb, silica bodies; st, stomatal apparatuses. (scale bar = 10 μm).
Figure 7. Micromorphology of the abaxial leaf epidermis of Indocalamus (scanning electron microscopy). (A) I. guangdongensis var. mollis (Yangmingshan, Hunan, China); (B) I. guangdongensis (Futang, Guangdong, China); (C) I. cordatus (DuChang, Jiangxi, China); (D) I. tessellatus (Daweishan, Hunan, China); (E) I. tessellatus (Xiushan, Chongqing, China); (F) I. tessellatus (Nanchang, Jiangxi, China); (G) I. chebalingensis (Chebaling, Guangdong, China); (H) I. multinerus (Chebaling, Guangdong, China). Abbreviations: ep, elongated papillae; sp, short papillae; ma, macrohairs; mi, microhairs; pr, prickles; sb, silica bodies; st, stomatal apparatuses. (scale bar = 10 μm).
Taxonomy 05 00012 g007
Taxonomy 05 00012 g008
Figure 8. Phylogeny of Arundinarieae inferred from a maximum likelihood (ML) analysis based on the complete chloroplast genomes of 61 representative bamboos. The branches indicate 11 accepted lineages of Arundinarieae (I–XI) [4]. The numbers associated with branches indicate the bootstrap value and the posterior probability, respectively. The asterisks indicate 100% bootstrap support or a posterior probability of 1.0.
Figure 8. Phylogeny of Arundinarieae inferred from a maximum likelihood (ML) analysis based on the complete chloroplast genomes of 61 representative bamboos. The branches indicate 11 accepted lineages of Arundinarieae (I–XI) [4]. The numbers associated with branches indicate the bootstrap value and the posterior probability, respectively. The asterisks indicate 100% bootstrap support or a posterior probability of 1.0.
Taxonomy 05 00012 g008
Taxonomy 05 00012 g009
Figure 9. Indocalamus mollis. (A,B) Habitat and plants; (C,D) rhizome and new shoot; (E) infranodal region of mature culm; (F) branch; (G,H) culm sheath, blade, and ligule; (I,J) leaf; (K) branching node and bud; (L) longitudinal section of branching node. Scale bars = 1 m (A,B); 1 cm (C–L).
Figure 9. Indocalamus mollis. (A,B) Habitat and plants; (C,D) rhizome and new shoot; (E) infranodal region of mature culm; (F) branch; (G,H) culm sheath, blade, and ligule; (I,J) leaf; (K) branching node and bud; (L) longitudinal section of branching node. Scale bars = 1 m (A,B); 1 cm (C–L).
Taxonomy 05 00012 g009
Disclaimer/Publisher’s Note: The statements, opinions and data contained in all publications are solely those of the individual author(s) and contributor(s) and not of MDPI and/or the editor(s). MDPI and/or the editor(s) disclaim responsibility for any injury to people or property resulting from any ideas, methods, instructions or products referred to in the content.

Share and Cite

MDPI and ACS Style

Gao, L.; Li, Y.; Wang, C.; Wang, J.; Yang, G.; Zhang, W. New Combination and Two Synonyms of Indocalamus Nakai (Poaceae: Bambusoideae) from China Based on Morphological Characters and Phylogenetic Evidence. Taxonomy 2025, 5, 12. https://doi.org/10.3390/taxonomy5010012

AMA Style

Gao L, Li Y, Wang C, Wang J, Yang G, Zhang W. New Combination and Two Synonyms of Indocalamus Nakai (Poaceae: Bambusoideae) from China Based on Morphological Characters and Phylogenetic Evidence. Taxonomy. 2025; 5(1):12. https://doi.org/10.3390/taxonomy5010012

Chicago/Turabian Style

Gao, Liqin, Yonglong Li, Chengkun Wang, Jianqing Wang, Guangyao Yang, and Wengen Zhang. 2025. "New Combination and Two Synonyms of Indocalamus Nakai (Poaceae: Bambusoideae) from China Based on Morphological Characters and Phylogenetic Evidence" Taxonomy 5, no. 1: 12. https://doi.org/10.3390/taxonomy5010012

APA Style

Gao, L., Li, Y., Wang, C., Wang, J., Yang, G., & Zhang, W. (2025). New Combination and Two Synonyms of Indocalamus Nakai (Poaceae: Bambusoideae) from China Based on Morphological Characters and Phylogenetic Evidence. Taxonomy, 5(1), 12. https://doi.org/10.3390/taxonomy5010012

Article Metrics

Back to TopTop