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国产龙血竭基原植物柬埔寨龙血树的分类学考证

花匠小妙招
2024-12-25 12:02

摘要:

“龙血竭”是龙血树属（Dracaena）植物树干产生的红色树脂，被称为“云南红药”。国药批准的国产“龙血竭”基原植物为天门冬科的柬埔寨龙血树（D. cambodiana Pierre ex Gagnep.）和剑叶龙血树（D. cochinchinensis (Lour.) S. C. Chen）。由于文献对两种龙血树的界定不清，在其名称上存在混淆使用。本研究对两种龙血树及其混淆种进行原始文献和模式标本考证，并利用龙血树属19种33个质体基因组数据开展系统发育重建。结果显示，19种龙血树可分成3个分支，其中，产龙血竭的物种位于分支Ⅰ和分支Ⅲ，柬埔寨龙血树样品分成两个独立的分支，即海南分支和云南分支。基于原始文献和模式标本考证，确认剑叶龙血树的形态与长花龙血树（D. angustifolia (Medik.) Roxb.）基本一致，可能是长花龙血树的一个新异名。本文结合龙血树质体基因组的系统发育和形态证据，明确了柬埔寨龙血树是蔡希陶等最早发现的国产龙血竭的基原植物，并确认岩棕（D. saposchnikowii Regel）和广西龙血树（未知种2）也是国产龙血竭的资源植物。

Abstract:

"Resina Draconis", also known as "Dragon's Blood Resin" or "Yunnan Hongyao", is red resin produced by the trunks of Dracaena species. Chinese "Resina Draconis" is derived from phenolic resins of D. cambodiana Pierre ex Gagnep. and D. cochinchinensis (Lour.) S. C. Chen. Due to taxonomic ambiguities between D. cambodiana and D. cochinchinensis in Chinese floras, there has been confusion in the literature regarding the source plant of Chinese "Resina Draconis". To resolve this, we examined the protologue and type materials of D. cambodiana, D. cochinchinensis, and D. angustifolia (Medik.) Roxb. and reconstructed the plastome phylogenies of 19 Dracaena species from 33 accessions. The plastome phylogenies strongly supported the division of these 19 Dracaena species into three clades. Species capable of producing phenolic resins were found in cladesⅠand Ⅲ, indicating their potential as source plants for "Resina Draconis". Notably, nine samples of D. cambodiana were not supported as monophyletic, forming two distinct groups: the Yunnan clade (D. cambodiana A) and the Hainan clade (D. cambodiana B). Morphological analysis of the protologue and neotype specimens indicated that D. cochinchinensis could not be differentiated from D. angustifolia, and thus should be considered as a synonym of D. angustifolia. Integrating phylogenetic analyses and morphological characteristics, we determined that D. cambodiana is the primary source plant of Chinese "Resina Draconis", with D. saposchnikowii Regel from South Yunnan and Dracaena sp. 2 from South Guangxi also contributing to its production. The plastome phylogenies clarified the source plants of "Resina Draconis" and elucidated the genetic relationships within the genus Dracaena, providing important information for the conservation and sustainable utilization of Dracaena germplasm resources.

图 1 龙血树属植物

A~C：柬埔寨龙血树模式标本。A：J.B.L. Pierre 660（GH00098599）；B：Clemens J. & Clemens M.S. 4057 （P00689821）； C：Poilane E. 4790 （P00689821）；D：剑叶龙血树新模式标本J. Clemens & M.S. Clemens 4048 （G00191146）；E：柬埔寨龙血树的软叶型（左）和硬叶型（右）；F：长花龙血树的模式图；G、H：岩粽的模式图和植物照片；I：引种于广西的龙血树（YX08）；J：引种于老挝的龙血树（YX05）。模式标本照片（A~D）下载于JSTOR全球植物网（https://plants.jstor.org/）。

Figure 1. Diversity of Dracaena species

A-C: Type specimens of D. cambodiana. A: J.B.L. Pierre 660 GH00098599); B: Clemens J. & Clemens M.S. 4057 (P00689821); C: Poilane E. 4790 (P00689821); D: Neotype of D. cochinchinensis J. Clemens & M.S. Clemens 4048 (G00191146); E: Soft leaf type (left) and hard leaf type (right) of D. cambodiana; F: Illustration of D. angustifolia; G: Illustration of D. saposchnikowii; H: Living plant of D. saposchnikowii; I: Dracaena sp. 2 (YX08); J: Dracaena sp.1 (YX05). Type specimen photos (A-D) were downloaded from Global Plants on JSTOR (https://plants.jstor.org/).

图 2 基于质体全基因组数据集的龙血树属植物贝叶斯推论（BI）系统发育树

Figure 2. Phylogenetic relationships of Dracaena species based on Bayesian Inference (BI) analysis using whole plastid genome dataset

**: MLBS=100, BIPP=1.00.

表 1 柬埔寨龙血树、剑叶龙血树和长花龙血树的原始文献与《中国植物志》特征描述的对比

Table 1 Comparison of characteristics between protologue and Flora Reipublicae Popularis Sinicae for Dracaena cambodiana, D. cochinchinensis, and D. angustifolia

性状特征
Traits 柬埔寨龙血树
D. cambodiana 剑叶龙血树
D. cochinchinensis 长花龙血树
D. angustifolia 原始文献《中国植物志》原始文献《中国植物志》原始文献《中国植物志》习性 Arbor 乔木状 Fruticosus 乔木状 Shrubby 灌木状株高 6~10 m 3~4 m 6-pedalis 5~15 m 8~10 ft 1~3 m 茎干 Rami divaricati 不分枝或分枝 Simplicissimus 分枝多 − 不分枝或稍分枝叶型 Lineari-laceolata 剑形 Laceolato-linearia 剑形 Linear 条状倒披针叶形叶长 60 cm ≥ 70 cm − 50~100 cm 30~45 cm 20~45 cm 叶宽 2~4 cm 1.5~3.0 cm − 2~5 cm 5 cm 1.5~5.5 cm [1] 蔡希陶,许再富. 国产血竭植物资源的研究[J]. 云南植物研究,1979,1(2):1−10.

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