中国秋海棠属植物的自然杂交发生及其特点

doi:10.17520/biods.2017050

摘要/Abstract

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

田代科, 李春, 肖艳, 付乃峰, 童毅, 吴瑞娟. (2017) 中国秋海棠属植物的自然杂交发生及其特点. 生物多样性, 25(6), 654-674.

Daike Tian, Chun Li, Yan Xiao, Naifeng Fu, Yi Tong, Ruijuan Wu. (2017) Occurrence and characteristics of natural hybridization in Begonia in China. Biodiversity Science, 25(6), 654-674.

图/表 11

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	√
	云南绿春县三勐乡巴东草果场 Badong, Sanmeng, Lüchun, Yunnan	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	√

表1

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。"

图1

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2

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3

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

表4

图2

表5

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 (%)
杂种采集号 Collect. no. of hybrid	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	相对误差 Relative error (%)
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6

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	-

表7

图3

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

表8

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