生物多样性 ›› 2017, Vol. 25 ›› Issue (6): 654-674.doi: 10.17520/biods.2017050

• 研究报告 • 上一篇    下一篇

中国秋海棠属植物的自然杂交发生及其特点

田代科1, 2, *(), 李春1, 3, 肖艳1, 2, 付乃峰1, 2, 童毅1, 2, 吴瑞娟1, 2   

  1. 1 上海辰山植物园, 中国科学院上海辰山植物科学研究中心, 上海 201602
    2 上海市资源植物功能基因组学重点实验室, 上海 201602
    3 四川省芦山县林业局, 四川芦山 625600
  • 收稿日期:2017-02-20 接受日期:2017-06-28 出版日期:2017-06-20
  • 通讯作者: 田代科 E-mail:dktian@sibs.ac.cn
  • 基金项目:
    上海市绿化与市容管理局科技公关项目(F122416)和国家自然科学基金(3170199)

Occurrence and characteristics of natural hybridization in Begonia in China

Daike Tian1, 2, *(), Chun Li1, 3, Yan Xiao1, 2, Naifeng Fu1, 2, Yi Tong1, 2, Ruijuan Wu1, 2   

  1. 1 Shanghai Chenshan Plant Science Research Center, Chinese Academy of Sciences; Shanghai Chenshan Botanical Garden, Shanghai 201602
    2 Shanghai Key Laboratory of Plant Functional Genomics and Resources, Shanghai 201602
    3 Lushan Bureau of Forestry, Lushan, Sichuan 625600
  • Received:2017-02-20 Accepted:2017-06-28 Online:2017-06-20
  • Contact: Tian Daike E-mail:dktian@sibs.ac.cn

自然杂交在植物界中十分普遍, 一直是物种形成和进化研究的热点。秋海棠属(Begonia)为世界最大植物属之一, 其种类繁多, 但关于自然杂交的相关报道很少。系统调查秋海棠属的自然杂交不仅有利于理解该属植物的多样性, 也是弄清其资源和解决物种分类问题不可缺少的环节。本文通过对国产秋海棠属植物自然杂交现象的系统调查和分析, 发现总计29种(占国产已知200种秋海棠的约15%)参与了杂交, 产生了31个自然“杂种”约50个居群。其中, 掌叶秋海棠(B. hemsleyana)和粗喙秋海棠(B. longifolia)参与杂交程度最高, 分别与另外的8种和7种秋海棠发生了杂交; 而裂叶秋海棠(B. palmata)发生杂交的居群最多, 达16处。自然杂交以云南(尤其是南部)发生频率最高, 一共有20种参与, 产生了31处杂交居群; 台湾和广西次之。杂交多为单向发生, 个体以F1代个体为主, 尚未脱离亲本独立成种。野外传粉生物学初步观察发现: 秋海棠的主要访花昆虫为食蚜蝇类, 其次是蜜蜂科, 但各自传粉特点及效率尚待进一步研究。杂种及疑似亲本的鉴定可通过形态学、分子生物学和人工重复杂交等综合方法实现。杂种的基因组大小(C值)等于或接近两亲本的平均值, 误差通常小于10%, 有助于辅助鉴定杂种及亲本。我们总结后认为杂交发生及杂种形成必须满足5个条件: (1)重叠或邻近分布; (2)花期相遇; (3)有效传粉媒介; (4)杂交亲和; (5)适宜种子萌发幼苗生长的小生境及气候条件。据此, 我们预测了国产秋海棠属未来可能发生的约40个新杂交组合的可能亲本及地点。到目前为止, 尽管所有已知的秋海棠杂种尚未形成真正的新物种, 但是, 为了满足科学研究及实践交流的需要, 建议按照新分类群发表的严格要求, 将“杂种”逐一开展调查研究并整理发表, 并将其作为物种进化过程中的一个特殊分类单元对待。鉴于秋海棠属的杂种个体稀少且并未对亲本构成威胁, 反而在一定程度上增加了类群的多样性, 并为优良观赏品种的直接选育创造了机会, 因此, 国产秋海棠属种类丰富及杂种区应为就地保护的优先考虑范围之一。同时, 鉴于秋海棠属杂交亲和性高, 迁地保护栽培时应增大花期重叠种类的种植间距, 避免发生自然杂交。

关键词: 秋海棠属, 自然杂交, 发生频率, 特殊分类单元, 杂交方向, 传粉

Natural hybridization is a very common phenomenon in plants and has continuously been a hotspot in the scientific research of speciation and evolution. As one of the largest genera in plants, Begonia has a large number of species. However, very few cases have been reported on its natural hybrids. A systematic investigation of natural hybridization of Begonia can not only improve understanding of plant diversity and germplasm resources of this genus, but also be a step towards solving its taxonomical issues. Through a comprehensive field survey and experimental analysis of natural hybridization in Chinese begonias, we found nearly 50 populations of 31 natural hybrids occurred among 29 species, accounting for 15% of currently about 200 accepted species. Begonia hemsleyana and B. longifolia had the highest cross frequency and crossed with 8 and 7 other species, respectively. B. palmata contributed to 16 hybrid populations, the largest number in all individual species. The largest number of hybrids (31 hybrid populations from 20 parental species) occurred in Yunnan (mainly in the southern areas), followed by Taiwan and Guangxi. Hybridization between species was unidirectional in most cases, and the majority of hybrids were F1 individuals, which still rely on parents or hybridization zones and have not been established as a true species. Field observations showed that aphid flies, followed by members of the bee family (Apidae), were the most frequent visitors of begonia flowers, however, further studies are necessary to examine their behavior and the efficiency of pollination. The natural hybrids and their putative parents were identified using integrative methods of morphology, molecular research, and artificial crossing experiments. Genome size (C value) of hybrids was usually equal or close to the average of that of the two parents, with an error less than 10%, which could assist identification of hybrids and parents. In addition, we summarized the five conditions necessary for the occurrence of natural hybridization: overlapping or adjacent distributions, concurrent flowering (at least partially overlapping), efficient pollination media, cross compatibility, and suitable microhabitat and climate conditions for seed germination and seedling growth. Based on these, we also predicted future putative parents of about 40 new natural hybrids and possible hybridization sites. Thus far, although all the known natural begonia hybrids have not established as true species, it may be more appropriate to treat them as a special taxon of speciation and evolution, which should be thoroughly investigated and published referring to protocols for publishing a new taxon in the scientific research. Because the individuals of natural hybrids in Begonia are always sparsely distributed in small numbers, and hybrids are not harmful to its parents it increases plant diversity and the chances of obtaining a new ornamental cultivar by natural selection. Therefore, the priority of in situ conservation should be given to regions with natural hybrids and species-rich environments. Meanwhile, for the purpose of ex situ conservation, requiring sufficient planting distance between species should be considered in those species with overlapping flowering time to avoid possible natural hybridization due to high cross-compatibility, particularly in native begonias.

Key words: Begonia, natural hybridization, occurrence frequency, special taxon, cross direction, pollination

表1

中国秋海棠属的自然杂交后代及亲本鉴定材料(含不同居群)"

杂交居群可能亲本组合
Possible parents of natural hybrids
采集地
Location of collection
标本采集号
Voucher no.
分子鉴定 Molecular identification 基因组大小 Genome size
酸味秋海棠×勐养秋海棠
B. acetosella
× B. silletensis subsp. mengyangensis
云南勐腊国家级自然保护区
Mengla National Nature Reserve, Yunnan
TDK354
TDK358

老挝南塔至勐新县公路
Road between Nanta and Mengxin, Laos
TDK747*
歪叶秋海棠×粗喙秋海棠
B. augustinei × B. longifolia
云南勐养国家级自然保护区
Mengyang National Nature Reserve, Yunnan
ZPS001
周裂秋海棠×裂叶秋海棠
B. circumlobata × B. palmata

广东信宜县大雾山 Dawushan of Xinyi, Guangdong TDK868
TDK870
TDK873
广东惠州罗浮山 Luofushan of Huizhou, Guangdong TDK2064
大围山秋海棠×掌叶秋海棠
B. daweishanensis × B. hemsleyana
云南大围山国家级自然保护区
Daweishan National Nature Reserve, Yunnan
TDK1306
厚叶秋海棠×酸味秋海棠(粗喙秋海棠)
B. dryadis
× B. acetosella (B. longifolia)
云南勐腊县勐远乡
Mengyuan, Mengla,Yunnan
TDK396
厚叶秋海棠×粗喙秋海棠
B. dryadis × B. longifolia
云南勐腊县望天树景区
Wangtianshu Resort of Mengla,Yunnan
TDK368
全柱秋海棠×大理秋海棠
B. grandis subsp. holostyla × B. taliensis
云南丽江永胜县
Yongsheng, Lijiang, Yunnan
TDK2213
掌叶秋海棠×大裂秋海棠
B. hemsleyana × B. macrotoma
云南绿春县大水沟乡
Dashuigou, Lüchun, Yunnan
TDK264
掌叶秋海棠×裂叶秋海棠
B. hemsleyana × B. palmata

云南绿春县三勐乡巴东草果场
Badong, Sanmeng, Lüchun, Yunnan
TDK1252
TDK1257
云南绿春县平河乡 Pinghe, Lüchun, Yunnan TDK2697
TDK2697A
掌叶秋海棠×紫叶秋海棠
B. hemsleyana × B. purpureofolia
云南大围山国家级自然保护区
Daweishan National Nature Reserve, Yunnan
TDK1303
掌叶秋海棠×倒鳞秋海棠
B. hemsleyana × B. reflexisquamosa
云南绿春县三勐乡巴东草果场
Badong, Sanmeng, Lüchun, Yunnan
TDK1251
掌叶秋海棠×变色秋海棠
B. hemsleyana × B. versicolor
云南麻栗坡县豆豉店
Douchidian, Malipo, Yunnan
TDK1374
掌叶秋海棠×长纤秋海棠
B. hemsleyana × B. longiciliata
云南绿春县平河乡
Pinghe, Lüchun, Yunnan
TDK2689
蕺叶秋海棠×掌裂叶秋海棠
B. limprichtii × B. pedatifida
四川峨眉山
Mount Emei, Sichuan
TDK2984
蕺叶秋海棠×秀丽秋海棠
B. limprichtii × B. pulchrifolia
四川峨眉山
Mount Emei, Sichuan
TDK2976
蕺叶秋海棠×存疑种 B. limprichtii × Begonia sp. 四川峨眉山
Mount Emei, Sichuan
TDK2290
裂叶秋海棠×周裂秋海棠
B. palmata
× B. circumlobata
福建尤溪县坂面镇青坑村
Qingkeng, Banmian, Youxi, Fujian
TDK2542
广东惠州罗浮山
Luofushan of Huizhou, Guangdong
FNF006
裂叶秋海棠×掌叶秋海棠
B. palmata × B. hemsleyana
云南绿春县三勐乡巴东草果场
Badong, Sanmeng, Lüchun, Yunnan
TDK1255
杂交居群可能亲本组合
Possible parents of natural hybrids
采集地
Location of collection
标本采集号
Voucher no.
分子鉴定 Molecular identification 基因组大小
Genome size
云南屏边县至河口县
Between Pingbian and Hekou, Yunnan
TDK1310
云南绿春县平河乡 Pinghe, Lüchun, Yunnan TDK2677
掌裂叶秋海棠×蕺叶秋海棠
B. pedatifida × B. limprichtii
四川峨眉山 Mount Emei, Sichuan TDK2747
勐养秋海棠×粗喙秋海棠
B. silletensis subsp. mengyangensis ×
B. longifolia
云南勐腊县望天树景区
Wangtianshu Resort of Mengla,Yunnan
TDK362
多变秋海棠×癞叶秋海棠 B. variifolia ×
B. leprosa
广西东兰县新烟村 Xinyan, Donglan, Guangxi TDK1428
变色秋海棠×掌叶秋海棠
B. versicolor × B. hemsleyana
云南大围山国家级自然保护区
Daweishan National Nature Reserve, Yunnan
TDK1299
长毛秋海棠×变色秋海棠
B. villifolia × B. versicolor
云南麻栗坡县猛硐乡小坪寨
Xiaopingzhai, Mengdong, Malipo, Yunnan
TDK1348
云南麻栗坡县豆豉店 Douchidian, Malipo, Yunnan TDK1378
长毛秋海棠×变色秋海棠(反向杂交)
B. villifolia × B. versicolor (or reverse)
云南麻栗坡县猛硐乡小坪寨
Xiaopingzhai, Mengdong, Malipo, Yunnan
TDK1349
文山秋海棠×大围山秋海棠
B. wenshanensis × B. daweishanensis
云南大围山国家级自然保护区
Daweishan National Nature Reserve, Yunnan
TDK1296

图1

中国秋海棠的主要自然杂种及其亲本形态比较 A: 酸味秋海棠×勐养秋海棠: a. 杂种植株, b. 叶片形态比较(上: 酸味秋海棠; 中: 杂种; 下: 勐养秋海棠); B: 周裂秋海棠(a)×粗喙秋海棠(c)杂种(b); C: 周裂秋海棠×裂叶秋海棠杂种及其亲本叶片形态特征比较(a正面, b反面) (左: 周裂秋海棠, 右: 裂叶秋海棠, 下: 杂种); D: 大围山秋海棠×掌叶秋海棠: a. 杂种植株, b. 叶形比较(上: 掌叶秋海棠; 中: 杂种; 下: 大围山秋海棠); E: 厚叶秋海棠×粗喙秋海棠: a. 正交(左箭头所指: 粗喙秋海棠; 中: 杂种; 右: 厚叶秋海棠), b. 反交(上: 粗喙秋海棠; 下: 杂种); F: 全柱秋海棠×大理秋海棠: a. 亲本, b. 杂种, c. 叶片及花特征比较(上: 全柱秋海棠; 中: 杂种; 下: 大理秋海棠); G: 同一地点生长2个不同杂种群体: a.粗喙秋海棠(左上)×掌叶秋海棠(右上)杂种(左箭头所指)、掌叶秋海棠×长纤秋海棠杂种(右下), b. 掌叶秋海棠(左)×长纤秋海棠(右)叶片特征比较, 杂种(中)叶背颜色更深; H: 掌叶秋海棠×大裂秋海棠: a. 掌叶秋海棠(下箭头所指)、杂种(上箭头所指), b. 大裂秋海棠; I: 掌叶秋海棠×紫叶秋海棠: a. 杂种, b. 杂种(中)及亲本叶形比较(上: 掌叶秋海棠; 下: 紫叶秋海棠绿叶类型); J: 撕裂秋海棠及其亲本: a. 掌叶秋海棠, b. 撕裂秋海棠, c. 食用秋海棠; K: 掌叶秋海棠×裂叶秋海棠: a. 杂种植株, b. 叶形比较(上: 掌叶秋海棠; 中: 杂种; 下: 裂叶秋海棠); L: 蕺叶秋海棠×掌裂叶秋海棠: a. 蕺叶秋海棠及杂种(右上及箭头所指), b. 掌裂叶秋海棠(含反向杂交后代); M: 新种(右)×裂叶秋海棠杂种: 仅见1株小苗(箭头所指); N: 粗喙秋海棠(a)×裂叶秋海棠(c)杂种(b); O: 麻栗坡秋海棠(中)及其亲本掌叶秋海棠(左)、变色秋海棠(右)叶片形态比较, 正反交后代个体均可见; P: 倒鳞秋海棠×掌叶秋海棠: a. 杂种(上箭头所指)同母本倒鳞秋海棠(下箭头所指), b. 叶形比较(左: 倒鳞秋海棠; 左上: 掌叶秋海棠; 右下: 杂种); cde. 叶柄及茎被毛比较, 杂种d介于中间。除B中照片由谭运洪提供外, 其余由田代科提供。"

表2

已发现的中国秋海棠自然杂交组合(正反交算同组)"

杂种数
Hybrid no.
自然杂种或组合(母本×父本)
Natural hybrid (♀×♂)
杂种产地及居群数
Location and number of hybrid populations
发现(发表)时间 Time
discovered (published)
1 短葶秋海棠(多变秋海棠×癞叶秋海棠)
B. ×breviscapa (B. variifolia × B. leprosa)
广西东兰1处 Guangxi: Donglan 1 (Peng et al, 2010)
2 武威秋海棠(裂叶秋海棠×台湾秋海棠)
B. ×buimontana (B. palmata × B. taiwaniana)
台湾屏东2、高雄1、嘉义1
Taiwan: Pingdong 2, Gaoxiong 1, Jiayi 1
(Peng & Chen, 1991)
3 钟氏秋海棠(裂叶秋海棠×粗喙秋海棠)
B. ×chungii (B. palmata × B. longifolia)
台湾南投2 Taiwan: Nantou 2 (Peng & Ku, 2009)
4 台北秋海棠(水鸦脚×粗喙秋海棠)
B. ×taipeiensis (B. formosana × B. longfifolia)
台湾台北2 Taiwan: Taipei 2 (Peng & Sue, 2000)
5 玉柄秋海棠×掌裂叶秋海棠 B. rubinea × B. pedatifida 贵州习水1 Guizhou: Xishui 1 (Li et al, 2005)
6 酸味秋海棠×勐养秋海棠
B. acetosella × B. sillentensis subsp. mengyangensis
云南勐腊3 Yunnan: Mengla 3 1997, 2012, 2015
7 歪叶秋海棠×粗喙秋海棠 B. augustinei × B. longifolia 云南勐腊1 Yunnan: Mengla 1 2015
8 周裂秋海棠×粗喙秋海棠 B. circumlobata × B. longifolia 广西那坡1 Guangxi: Napo 1 2015
9 周裂秋海棠×裂叶秋海棠 B. circumlobata × B. palmata 广东信宜2、惠州1; 福建尤溪1 Guangdong: Xinyi 2, Huizhou 1; Fujian: Youxi 1 2014
10 大围山秋海棠×掌叶秋海棠 B. daweishanensis × B. hemsleyana 云南屏边1 Yunnan: Pingbian 1 2014
11 厚叶秋海棠×酸味秋海棠 B. dryadis × B. acetosella 云南勐腊1 Yunnan: Mengla 1 2012
12 厚叶秋海棠×粗喙秋海棠 B. dryadis × B. longifolia 云南勐腊1 Yunnan: Mengla 1 2012
13 全柱秋海棠×大理秋海棠 B. grandis subsp. holostyla × B. taliensis 云南永胜2 Yunnan: Yongsheng 2 2014
14 香花秋海棠×酸味秋海棠 B. handelii × B. acetosella 云南勐腊2、澜沧1、临沧1
Yunnan: Mengla 2, Lancang 1, Lincang 1
1997, 2012, 2015
15 掌叶秋海棠×食用秋海棠 B. hemsleyana × B. edulis 云南屏边1 Yunnan: Pingbian 1 1939*
16 掌叶秋海棠×长纤秋海棠 B. hemsleyana × B. longiciliata 云南绿春1 Yunnan: Lüchun 1 2015
17 掌叶秋海棠×粗喙秋海棠 B. hemsleyana × B. longifolia 云南绿春1 Yunnan: Lüchun 1 2015
18 掌叶秋海棠×大裂秋海棠 B. hemsleyana × B. macrotoma 云南绿春1 Yunnan: Lüchun 1 2012
19 掌叶秋海棠×裂叶秋海棠 B. hemsleyana × B. palmata 云南绿春2 Yunnan: Lüchun 2 2013, 2015
20 掌叶秋海棠×紫叶秋海棠 B. hemsleyana × B. purpureofolia 云南屏边1 Yunnan: Pingbian 1 2012
21 掌叶秋海棠×倒鳞秋海棠 B. hemsleyana × B. reflexisquamosa 云南绿春1 Yunnan: Lüchun 1 2013
22 蕺叶秋海棠×掌裂叶秋海棠 B. limprichtii × B. pedatifida 四川峨眉山2 Sichuan: Mount Emei 2 2014, 2016, 2017
23 蕺叶秋海棠×秀丽秋海棠 B. limprichtii × B. pulchrifolia 四川峨眉山1 Sichuan: Mount Emei 1 2016
24 长纤秋海棠×食用秋海棠 B. longiciliata × B. edulis 云南勐腊西双版纳热带植物园1
Yunnan: XSBN Tropical Botanical Garden 1
2015
25 长纤秋海棠×厚叶秋海棠 B. longiciliata × B. dryadis 云南勐腊西双版纳热带植物园1
Yunnan: XSBN Tropical Botanical Garden 1
2015
26 粗喙秋海棠×裂叶秋海棠 B. longifolia × B. palmata 广西上思1、武鸣1
Guangxi: Shangsi 1, Wuming 1
2014, 2016
27 粗喙秋海棠×勐养秋海棠 B. longifolia × B. silletensis subsp. mengyangensis 西藏墨脱1 Tibet: Motuo 1 2012 (存疑Possible)
28 新种×裂叶秋海棠 Begonia sp. nov. × B. palmata 广西防城港1 Guangxi: Fangchenggang 1 2013
29 变色秋海棠×掌叶秋海棠 B. versicolor × B. hemsleyana 云南屏边3、麻栗坡2
Yunnan: Pingbian 3, Malipo 2
1996
30 变色秋海棠×长毛秋海棠 B. versicolor × B. villifolia 云南麻栗坡2 Yunnan: Malipo 2 1997
31 文山秋海棠×大围山秋海棠 B. wenshanensis × B. daweishanensis 云南屏边1 Yunnan: Pingbian 1 2014

表3

中国秋海棠的自然杂交发生种类及频率"

编号
No.
种名 Species name 组名 Section 自然杂交发生地点及杂交居群数
Occurrence location and population number
1 酸味秋海棠 B. acetosella 无翅组 Sect. Sphenanthera 云南勐腊4、澜沧1、临沧1、绿春1
Yunnan: Mengla 4, Lancang 1, Lincang 1, Lüchun 1
2 歪叶秋海棠 B. augustinei 扁果组 Sect. Platycentrum 云南勐海1 Yunnan: Menghai 1
3 周裂秋海棠 B. circumlobata 扁果组 Sect. Platycentrum 广东信宜1、惠州1; 广西那坡1; 福建尤溪1
Guangdong: Xinyi 1, Huizhou 1; Guangxi: Napo 1; Fujian: Youxi 1
4 厚叶秋海棠 B. dryadis 扁果组 Sect. Platycentrum 云南勐腊3 Yunnan: Mengla 3
5 食用秋海棠 B. edulis 扁果组 Sect. Platycentrum 云南勐腊1、屏边1 Yunnan: Mengla 1, Pingbian 1
6 水鸭脚 B. formosana 扁果组 Sect. Platycentrum 台湾1 Taiwan: 1
7 全柱秋海棠 B. grandis subsp. holostyla 东亚秋海棠组 Sect. Diploclinium 云南永胜1 Yunnan: Yongsheng 1
8 香花秋海棠 B. handelii 无翅组 Sect. Sphenanthera 云南勐腊2、澜沧1、临沧1
Yunnan: Mengla 2, Lancang 1, Lincang 1
9 掌叶秋海棠 B. hemsleyana 扁果组 Sect. Platycentrum 云南屏边5、绿春2、麻栗坡2
Yunnan: Pingbian 5, Lüchun 2, Malipo 2
10 癞叶秋海棠 B. leprosa 棒果组 Sect. Leprosae 广西东兰1 Guangxi: Donglan 1
11 蕺叶秋海棠 B. limprichtii 扁果组 Sect. Platycentrum 四川峨眉山2 Sichuan: Mount Emei 2
12 长纤秋海棠 B. longiciliata 扁果组 Sect. Platycentrum 云南绿春1、勐腊1 Yunnan: Lüchun 1, Mengla 1
13 粗喙秋海棠 B. longifolia 东亚秋海棠组 Sect. Diploclinium 云南勐腊2、绿春2; 广西那坡1、上思1; 台湾2; 西藏墨脱1 Yunnan: Mengla 2, Lüchun 2; Guangxi: Napo 1, Shangsi 1; Taiwan: 2; Tibet: Motuo 1
14 大裂秋海棠 B. macrotoma 扁果组 Sect. Platycentrum 云南绿春2 Yunnan: Lüchun 2
15 裂叶秋海棠 B. palmata 扁果组 Sect. Platycentrum 台湾6; 云南绿春2、屏边1; 广东信宜2、惠州1; 广西上思1、防城港1、武鸣1; 福建尤溪1 Taiwan: 6; Yunnan: Lüchun 2, Pingbian 1; Guangdong: Xinyi 2, Huizhou 1; Guangxi: Shangsi 1, Fangchenggang 1, Wuming 1; Fujian: Youxi 1
16 掌裂秋海棠 B. pedatifida 扁果组 Sect. Platycentrum 四川峨眉山2 Sichuan: Mount Emei 2
17 秀丽秋海棠 B. pulchrifolia 扁果组 Sect. Platycentrum 四川峨眉山1 Sichuan: Mount Emei 1
18 紫叶秋海棠 B. purprueofolia 扁果组 Sect. Platycentrum 云南屏边1 Yunnan: Pingbian 1
19 倒鳞秋海棠 B. reflexisquamosa 扁果组 Sect. Platycentrum 云南绿春2 Yunnan: Lüchun 2
20 玉柄秋海棠 B. rubinea 扁果组 Sect. Platycentrum 贵州习水1 Guizhou: Xishui 1
21 厚壁秋海棠 B. silletensis subsp. silletensis 无翅组 Sect. Sphenanthera 西藏墨脱1 Tibet: Motuo 1
22 勐养秋海棠 B. silletensis
subsp. mengyangensis
无翅组 Sect. Sphenanthera 云南勐腊4 Yunnan: Mengla 4
23 台湾秋海棠 B. taiwaniana 东亚秋海棠组 Sect. Diploclinium 台湾1 Taiwan: 1
24 大理秋海棠 B. taliensis 东亚秋海棠组 Sect. Diploclinium 云南永胜1 Yunnan: Yongsheng 1
25 多变秋海棠 B. variifolia 侧膜组 Sect. Coelocentrum 广西东兰1 Guangxi: Donglan 1
26 变色秋海棠 B. versicolor 扁果组 Sect. Platycentrum 云南屏边3、麻栗坡2 Yunnan: Pingbian 3, Malipo 2
27 长毛秋海棠 B. villifolia 扁果组 Sect. Platycentrum 云南麻栗坡2 Yunnan: Malipo 2
28 文山秋海棠 B. wenshanensis 东亚秋海棠组 Sect. Diploclinium 云南屏边2、文山1 (疑似) Yunnan: Pingbian 2, Wenshan 1 (possible)
29 新种(待发表) Begonia sp. nov. 扁果组 Sect. Platycentrum 广西防城港1 Guangxi: Fangchenggang 1

表4

中国秋海棠自然杂交发生频率最高的种类"

种名
Species name
参与杂交种类及数量Species involved in natural cross 杂交发生地点
Location of hybridization occurrence
杂交发生频率高的理由
Reasons for high frequency of natural hybridization
掌叶秋海棠
B. hemsleyana
8 (1)云南、广西南部局部分布较广的种类, 所在分布区为国产秋海棠分布比较集中的地区, 种类多, 重叠概率大; (2)花期较长6-11月; (3)植株较高: 为直立茎类, 利于昆虫传粉。 (1) Widely distributed with higher chance of habitat overlapping in some regions of Yunnan and Guangxi provinces with rich begonia species; (2) Long flowering time from June to November; (3) Relatively taller plants benefiting visiting of pollinators.
食用秋海棠 B. edulis 云南屏边 Pingbian, Yunnan
粗喙秋海棠 B. longifolia 云南绿春 Lüchun, Yunnan
大裂秋海棠 B. macrotoma 云南绿春 Lüchun, Yunnan
裂叶秋海棠 B. palmata 云南绿春 Lüchun, Yunnan
紫叶秋海棠 B. purpureofolia 云南屏边 Pingbian, Yunnan
倒鳞秋海棠 B. reflexisquamosa 云南绿春 Lüchun, Yunnan
长纤秋海棠 B. longiciliata 云南绿春 Lüchun, Yunnan
变色秋海棠 B. versicolor 云南屏边、麻栗坡 Pingbian and Malipo, Yunnan
粗喙秋海棠
B. longifolia
7 (1)分布范围广, 为该属跨国分布最广种类之一, 仅次于裂叶秋海棠 B. plamata; (2)花期长: 5-12月; (3)植株高: 为直立茎、亚灌木类, 利于昆虫传粉。 (1) The second most widely distributed species only next to B. palmata; (2) Long flowering time from May to December; (3) Relatively taller subshrub benefiting pollinator visiting.
歪叶秋海棠 B. augustinei 云南勐腊 Mengla,Yunnan
周裂秋海棠 B. circumlobata 广西那坡 Napo, Guangxi
厚叶秋海棠 B. dryadis 云南勐腊 Mengla,Yunnan
水鸭脚 B. formosana 台湾 Taiwan
掌叶秋海棠 B. hemsleyana 云南绿春 Lüchun, Yunnan
裂叶秋海棠 B. palmata 台湾、云南绿春、广西上思Taiwan; Lüchun, Yunnan; Shangsi, Guangxi
厚壁秋海棠 B. silletensis 西藏墨脱 Motuo, Tibet
裂叶秋海棠
B. palmata
5 (1)分布范围最广, 为该属在国内和跨国分布最广种类; (2)该种适应性强, 居群多、范围及密度大、个体数多, 因此, 同其他种类重叠分布频率高; (3)花期长: 5-12月; (4)植株较高: 为直立茎类, 利于昆虫传粉。 (1) The most widely distributed species in Begonia; (2) High adaptation to environment, populations are large in number and usually in size and high density resulting in high frequency of habitat overlapping with other species; (3) Long flowering time from May to December; (4) Relatively taller plants benefiting pollinator visiting.
台湾秋海棠 B. taiwaniana 台湾 Taiwan
粗喙秋海棠 B. longifolia 台湾、广西上思 Taiwan; Shangsi, Guangxi
周裂秋海棠 B. circumlobata 广东信宜、惠州; 福建尤溪 Xinyi, Huizhou and Guangdong; Youxi, Fujian
掌叶秋海棠 B. hemsleyana 云南绿春 Lüchun, Yunnan
新种 Begonia sp. nov. 广西防城港 Fangchenggang, Guangxi
酸味秋海棠
B. acetosella
3 (1)分布范围相对窄, 限于西双版纳州及邻近地区, 但常同本组的香花秋海棠、勐养秋海棠和扁果组的厚叶秋海棠伴生; (2)为直立茎类、植株高, 利于昆虫传粉; (3)同属组花具有香味, 有利于吸引昆虫传粉。 (1) Narrowly distributed but frequently overlapped with habitats of other species (B. handelii, B. silletensis subsp. mengyangensis) in the same section and B. dryadis; (2) Shrub-like plants erect and tall to benefit pollinator visiting; (3) Flowers with strong fragrance to facilitate attracting pollinators.
厚叶秋海棠 B. dryadis 云南勐腊 Mengla,Yunnan
香花秋海棠 B. handelii 云南勐腊、澜沧、临沧Mengla, Lancang and Lincang, Yunnan
勐养秋海棠 B. sillentensis subsp. mengyangensis 云南勐腊 Mengla,Yunnan
长纤秋海棠
B. longiciliata
3 (1)分布范围相对狭窄, 限于云南绿春、江城等中越边境地区, 但该地区种类相对丰富; (2)西双版纳热带植物园栽培多种于热带雨林, 导致自然杂交。
(1) Narrowly distributed but frequently overlapped with relatively rich species in habitat in the border regions between Lüchun and Jiangcheng of Yunnan, China and Vietnam; (2) Natural cross also occurred in ex situ conservation site under the tropical rain forest of Xishuangbanna Tropical Botanical Garden.
掌叶秋海棠 B. hemsleyana 云南绿春 Lüchun, Yunnan
食用秋海棠 B. edulis 云南勐腊西双版纳热带植物园XishuangbannaTropical Botanical Garden, Mengla, Yunnan
厚叶秋海棠 B. dryadis 云南勐腊西双版纳热带植物园 Xishuangbanna Tropical Botanical Garden, Mengla, Yunnan

图2

中国秋海棠自然杂交发生频率及分布式样(实心点: 杂交居群) (藏南地区杂种未列入)"

表5

利用核基因ITS和叶绿体基因序列鉴别中国秋海棠的杂种及疑似亲本结果"

标本采集号 Voucher no. ITS 叶绿体基因 Chloroplast genes 标本采集号 Voucher no. ITS 叶绿体基因 Chloroplast genes
TDK244 TDK1296 × ×
TDK264 TDK1299
TDK354 × × TDK1303 ×
TDK358 × × TDK1306 ×
TDK362 TDK1310 ×
TDK368 × × TDK1348 × ×
TDK396 TDK1349 × ×
TDK868 × TDK1374
TDK870 × TDK1378 ×
TDK873 × TDK1428 ×
TDK1251 TDK2064
TDK1252 × TDK2213 ×
TDK1255 × TDK2290 × ×
TDK1257 × ZPS001

表6

中国秋海棠的自然杂种、人工杂交后代与其亲本的C值关系"

杂种采集号
Collect. no. of hybrid
母本 Mother species 父本Father species 2C值 2C value 相对误差 Relative error (%)
采集号 Collect. no. 物种
Species
采集号 Collect. no. 物种
Species
母本 Mother species 父本
Father species
杂种理论Theoretical of hybrid 杂种实测Measured of hybrid
TDK354 TDK355 酸味秋海棠
B. acetosella
TDK2645 勐养秋海棠
B. silletensis subsp.
mengyangensis
1.098 1.116 1.107 1.109 0.22
TDK368 TDK366 厚叶秋海棠
B. dryadis
TDK434 粗喙秋海棠
B. longifolia
1.204 1.122 1.163 1.263 8.59
TDK747 TDK355 酸味秋海棠
B. acetosella
TDK2645 勐养秋海棠
B. silletensis subsp.
mengyangensis
1.098 1.116 1.107 1.136 2.66
TDK2213 TDK2212 全柱秋海棠
B. grandis subsp. holostyla
TDK2214 大理秋海棠
B. taliensis
1.077 1.252 1.165 1.206 3.58
TDK2542 TDK2537 裂叶秋海棠
B. palmata
TDK1795 周裂秋海棠
B. circumlobata
0.993 0.999 0.996 1.006 1.08
TDK2677 TDK2669 裂叶秋海棠
B. palmata
TDK2696 掌叶秋海棠
B. hemsleyana
0.926 1.043 0.985 0.901 8.48
TDK2689 TDK2696 掌叶秋海棠
B. hemsleyana
TDK1283 长纤秋海棠
B. longiciliata
1.043 1.053 1.048 1.040 0.75
TDK2697 TDK2696 掌叶秋海棠
B. hemsleyana
TDK2669 裂叶秋海棠
B. palmata
1.043 0.926 0.985 0.891 9.50
TDK2747 TDK1924 掌裂叶秋海棠
B. pedatifida
TDK2250 蕺叶秋海棠
B. limprichtii
0.878 0.938 0.908 0.965 6.23
TDK2697A TDK2696 掌叶秋海棠
B. hemsleyana
TDK2669 裂叶秋海棠
B. palmata
1.043 0.926 0.985 0.940 4.49
TDK2976 TDK2983 蕺叶秋海棠
B. limprichtii
TDK2973 秀丽秋海棠
B. pulchrifolia
0.967 0.984 0.976 0.916 6.15
TDK2984 TDK2983 蕺叶秋海棠
B. limprichtii
TDK2981 掌裂叶秋海棠
B. pedatifida
0.967 1.285 1.126 1.345 19.45
FNF006 FNF005 裂叶秋海棠
B. palmata
TDK1795 周裂秋海棠
B. circumlobata
0.959 0.999 0.979 0.980 0.14
B7* B040-A 裂叶秋海棠
B. palmata
177 多毛秋海棠
B. polytricha
0.933 0.836 0.908 0.909 0.11
B9* B040-A 裂叶秋海棠
B. palmata
180 勐养秋海棠
B. silletensis subsp.
mengyangensis
0.933 1.116 0.979 1.079 10.24
B10* B040-A 裂叶秋海棠
B. palmata
177 多毛秋海棠
B. polytricha
0.933 0.836 0.908 0.932 2.64
B11* 278-C 花叶秋海棠
B. cathayana
180 勐养秋海棠
B. silletensis subsp.
mengyangensis
0.970 1.116 1.006 1.128 12.14
C4* B040-A 裂叶秋海棠
B. palmata
B022 秋海棠新种 Begonia sp. nov. 0.933 0.908 0.926 0.889 4.00
E5* 278-B 花叶秋海棠
B. cathayana
226 粗喙秋海棠
B. longifolia
0.970 1.105 1.003 1.103 9.89
E7* B040-A 裂叶秋海棠
B. palmata
226 粗喙秋海棠
B. longifolia
0.933 1.105 0.976 1.047 7.36
E8* B040-A 裂叶秋海棠
B. palmata
227 厚叶秋海棠
B. dryadis
0.933 1.204 1.001 1.151 15.03

表7

人工杂交实验验证中国秋海棠的自然杂交"

杂种组合
Hybrid
母本Mother plant (♀) 父本Father plant (♂) 是否成功(+/-) Success + /failure -
1 酸味秋海棠 B. acetosella 香花秋海棠 B. handelii +
香花秋海棠 B. handelii 酸味秋海棠 B. acetosella +
2 酸味秋海棠 B. acetosella 勐养秋海棠 B. silletensis subsp. mengyangensis +
勐养秋海棠 B. silletensis subsp. mengyangensis 酸味秋海棠 B. acetosella +
酸味秋海棠×勐养秋海棠
B. acetosella × B. silletensis subsp. mengyangensis
酸味秋海棠×勐养秋海棠 B. acetosella × B. silletensis subsp. mengyangensis -
3* 圆果秋海棠(=粗喙秋海棠) B. aptera (= B. longifolia) 水鸭脚 B. formosana -
水鸭脚 B. formosana 圆果秋海棠 B. aptera +
水鸭脚 B. formosana 台北秋海棠 B. ×taipeiensis (B. formosana × B. aptera) -
圆果秋海棠 B. aptera 台北秋海棠 B. ×taipeiensis -
台北秋海棠 B. ×taipeiensis 台北秋海棠 B. ×taipeiensis -
4 周裂秋海棠 B. circumlobata 裂叶秋海棠 B. palmata +
裂叶秋海棠 B. palmata 周裂秋海棠 B. circumlobata +
周裂秋海棠×裂叶秋海棠 B. circumlobata × B. palmata 周裂秋海棠 B. circumlobata -
周裂秋海棠×裂叶秋海棠 B. circumlobata × B. palmata 裂叶秋海棠 B. palmata -
5 厚叶秋海棠 B. dryadis 粗喙秋海棠 B. longifolia +
粗喙秋海棠 B. longifolia 厚叶秋海棠 B. dryadis +
厚叶秋海棠×粗喙秋海棠 B. dryadis × B. longifolia 厚叶秋海棠×粗喙秋海棠 B. dryadis × B. longifolia -
6 掌叶秋海棠 B. hemsleyana 变色秋海棠 B. versicolor +
变色秋海棠 B. versicolor 掌叶秋海棠 B. hemsleyana +
麻栗坡秋海棠 B. ×malipoensis (B. hemsleyana × B. versicolor) 麻栗坡秋海棠 B. ×malipoensis -
7 掌叶秋海棠 B. hemsleyana 大裂秋海棠 B. macrotoma +
掌叶秋海棠×大裂秋海棠 B. hemsleyana × B. macrotoma 掌叶秋海棠×大裂秋海棠 B. hemsleyana × B. macrotoma +
8* 粗喙秋海棠 B. longifolia 裂叶秋海棠 B. palmata +
钟氏秋海棠 B.×chungii (B. longifolia × B. plamata) 钟氏秋海棠 B.×chungii -

图3

连续三天不同时间段内上海辰山植物园秋海棠的主要访花昆虫的访花频率(次数)"

表8

中国秋海棠属植物未来可能发生新的自然杂交及其地点"

编号 No. 可能亲本A Possible parent A 可能亲本B Possible parent B 杂交可能发生地 Possible place of cross occurrence
1 大水沟秋海棠 B. ×dashuigouensis 掌叶秋海棠 B. hemsleyana; 大裂秋海棠 B. macrotoma 云南绿春 Lüchun, Yunnan
2 酸味秋海棠 B. acetosella 粗喙秋海棠 B. longifolia 云南勐腊 Mengla, Yunnan
3 点叶秋海棠 B. alveolata 裂叶秋海棠 B. palmata 云南马关、麻栗坡 Maguan & Malipo, Yunnan
4 树生秋海棠 B. arboreta 倒鳞秋海棠 B. reflexisquamosa 云南绿春 Lüchun, Yunnan
5 糙叶秋海棠 B. asperifolia 刺毛红孩儿B. difformis 云南贡山; 西藏墨脱 Gongshan, Yunnan; Motuo, Tibet
6 歪叶秋海棠 B. augustinei 裂叶秋海棠 B. palmata 云南勐腊、勐海、澜沧Mengla, Menghai & Lancang, Yunnan
7 巴马秋海棠 B. bamaensis 食用秋海棠 B. edulis 广西巴马 Bama, Guangxi
8 花叶秋海棠 B. cathayana 掌叶秋海棠 B. hemsleyana 云南屏边; 广西防城港 Pingbian, Yunnan; Fangchenggang, Guangxi
9 昌感秋海棠 B. cavaleriei 罗甸秋海棠 B. porteri; 卷毛秋海棠 B. cirrosa; 裂叶秋海棠 B. plamata 广西罗城 Luocheng, Guangxi
10 角果秋海棠 B. ceratocarpa 粗喙秋海棠 B. longifolia 云南马关 Maguan, Yunnan
11 凤山秋海棠 B. chingii 食用秋海棠 B. edulis 广西巴马、德保 Bama & Debao, Guangxi
12 赤水秋海棠 B. chishuiensis 掌裂叶秋海 B. pedatifida; 新种 Begonia sp. nov. 贵州赤水; 重庆江津四面山Chishui, Guizhou; Simianshan, Jiangjin, Chongqing; 四川纳溪Naxi, Sichuan
13 周裂秋海棠 B. circumlobata 裂叶秋海棠 B. palmata 贵州榕江Rongjiang, Guizhou
14 卷毛秋海棠 B. cirrosa 昌感秋海棠 B. cavaleriei 广西那坡 Napo, Guangxi
15 黄连山秋海棠 B. coptidimontana 多毛秋海棠 B. polytricha 云南绿春黄连山 Huanglianshan, Lüchun, Yunnan
16 水晶秋海棠 B. crystallina 光滑秋海棠 B. psyllophilla 云南麻栗坡 Malipo, Yunnan
17 瓜叶秋海棠 B. cucurbitifolia 裂叶秋海棠 B. palmata 云南马关 Maguan, Yunnan
18 大围山秋海棠 B. daweishanensis 掌叶秋海棠 B. hemsleyana; 裂叶秋海棠 B. palmata 云南屏边 Pingbian, Yunnan
19 德保秋海棠 B. debaoensis 食用秋海棠 B. edulis; 凤山秋海棠 B. chingii 广西德保 Debao, Guangxi
20 南川秋海棠 B. dielsiana 蕺叶秋海棠 B. limprichtii 四川峨眉山 Mount Emei, Sichuan
21 刺毛红孩儿B. difformis 宿苞秋海棠 B. yuii 云南景东无量山 Wuliangshan, Jingdong, Yunnan
22 槭叶秋海棠 B. digyna 裂叶秋海棠 B. palmata 福建长汀 Changting, Fujian
23 厚叶秋海棠 B. dryadis 小花秋海棠 B. peii 云南勐腊 Mengla, Yunnan
24 食用秋海棠 B. edulis 巴马秋海棠 B. bamaensis; 裂叶秋海棠 B. palmata 广西巴马Bama, Guangxi; 广东阳山Yangshan, Guangdong
25 紫背天葵B. fimbristipula 秋海棠 B. grandis 广东仁化; 湖南永兴Renhua, Guangdong; Yongxing, Hunan
26 金秀秋海棠 B. glechomifolia 裂叶秋海棠 B. pamlata 广西金秀 Jinxiu, Guangxi
27 圭山秋海棠 B. guishanensis 独牛B. henryi 云南石林 Shilin, Yunnan
28 古林箐秋海棠 B. gulinqingensis 裂叶秋海棠 B. pamlata 云南马关、古林箐 Gulinqing & Maguan, Yunnan
29 香花秋海棠 B. handelii 长果秋海棠 B. longicarpa 云南河口 Hekou, Yunnan
30 独牛 B. henryi 新种 Begonia sp. nov.; 圭山秋海棠 B. guishanensis 四川天全Tianquan, Sichuan; 云南石林Shilin, Yunnan
31 靖西秋海棠 B. jingxiensis 厚叶秋海棠 B. dryadis 广西靖西 Jingxi, Guangxi
32 缙云秋海棠 B. jinyunensis 裂叶秋海棠 B. palmata 贵州榕江 Rongjiang, Guizhou
33 心叶秋海棠 B. labordei 裂叶秋海棠 B. palmata; 中华秋海棠 B. grandis subsp. sinensis 云南多处; 广西Multi-places in Yunnan; Guangxi; 云南昆明、楚雄 Kunming & Chuxiong, Yunnan
34 大裂秋海棠 B. macrotoma 裂叶秋海棠 B. palmata 云南澜沧、绿春 Lancang & Lüchun, Yunnan
35 麻栗坡秋海棠 B. malipoensis 变色秋海棠 B. versicolor; 掌叶秋海棠 B. hemsleyana 云南屏边、麻栗坡 Pingbian & Malipo, Yunnan
36 不显秋海棠 B. obsolescens 裂叶秋海棠 B. palmata 云南马关 Maguan, Yunnan
37 掌裂叶秋海棠 B. pedatifida 赤水秋海棠 B. chishuiensis; 裂叶秋海棠 B. palmata; 新种Begonia sp. nov. 贵州赤水Chishui, Guizhou; 广西田林 Tianlin, Guangxi; 四川青城山 Qingchengshan, Sichuan
38 秀丽秋海棠 B. pulchrifolia 掌裂叶秋海棠 B. pedatifida 四川峨眉山 Mount Emei, Sichuan
39 长毛秋海棠 B. villifolia 裂叶秋海棠 B. palmata 云南麻栗坡 Malipo, Yunnan
40 一点血 B. wilsonii 新种Begonia sp. nov. 四川峨眉山、青城山 Mount Emei & Qingchengshan, Sichuan
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