被子植物隐性雌雄异株性系统的多样性、系统演化及进化意义

doi:10.17520/biods.2024149

摘要/Abstract

摘要：

隐性雌雄异株指植株表型为非雌雄异株, 但形态上两性花通常雄性不育或雌性不育, 植株个体实际仅行使雌性或雄性功能。两性花植株向雌雄异株的转变在被子植物中广泛存在, 隐性雌雄异株作为潜在的中间阶段对于理解植物性系统的演化具有重要意义。据APG IV分类系统, 这些物种在被子植物的22目36科65属约221种植物中有过报道, 分别约占被子植物总目数的34.4%、总科数的8.7%、总属数的0.5%和总种数的0.1%; 多为分布于热带或亚热带地区依靠生物传粉的木本植物, 且至少存在花蜜或花粉作为访花报酬。该性系统形态学的性表达类型多样: 可分为雄花两性花异株(类型I)、两性花植株(类型II)、雌花两性花异株(类型III)及其他类型(类型IV) 4种, 分别约占已报道隐性雌雄异株物种数的48.9%、47.5%、2.7%和0.9%。从系统发育看, 该性系统在被子植物木兰分支、单子叶植物分支及真双子叶植物分支均有分布, 且主要分布于较为进化的核心真双子叶植物类群中。隐性雌雄异株物种不育性器官的进化意义存在多种假说, 包括祖先假说、遗传约束假说、促进风媒花粉落置假说、传粉者吸引假说、拟态与欺骗假说等, 但相关实证研究较少。本文对隐性雌雄异株物种不同类群的性表达形式、不育性器官败育类型进行了归纳, 并对具该性系统的类群在被子植物中的分布与系统演化进行了分析与总结, 同时对有关隐性雌雄异株物种不育性器官进化意义的5个假说进行了介绍与评价, 最后对今后的相关研究方向进行了展望。以期为深入研究被子植物隐性雌雄异株性系统的进化式样与机制研究提供理论资料。

关键词: 隐性雌雄异株, 性表达, 性系统, 系统演化, 进化意义, 被子植物

Abstract

Background & Aim: Cryptic dioecy refers to a population that is phenotypically non-dioecious, but individuals actually function as males or females due to the morphologically hermaphroditic flower that typically exhibits female or male sterility. As a widespread transitional from hermaphroditism to dioecy, understandings of adaptive strategy of cryptic dioecy could help of illuminating the evolution of diverse sexual systems within angiosperms.
Progress: The morphological sex expressions of cryptic dioecy are diverse and can be classified into four types: androdioecy (type I), hermaphrodite (type II), gynodioecy (type III) and others (type IV), constituting 48.9%, 47.5%, 2.7%, and 0.9% of reported species, respectively. In the APG IV system, cryptic dioecy occurs across 22 orders, 36 families and 65 genera in around 221 species which accounts for approximately 34.4% of orders, 8.7% of families, 0.5% of genera and merely about 0.1% of angiosperms species. Most cryptic dioecious species are woody, biotic pollinated and distributed in tropics or subtropics. They have nectar or pollen as a floral reward. Phylogenetically, the sexual system appears in magnolia, monocotyledon and eudicots, particularly prevalent among more advanced core eudicots. Several hypotheses have been proposed to explain the evolutionary significance of sterile sex organs in cryptic dioecy, including ancestry hypothesis, genetic constraints hypothesis, promotion of wind-borne pollen dispersal hypothesis, pollinator attraction hypothesis, mimicry and deception hypothesis, but empirical studies remain little known. In this study, we provide a comprehensive summary of the forms of sex expression and types of sterile sex organs of cryptic dioecious in plant lineages. Families with cryptic dioecious species evolved are mapped into an updated phylogeny of angiosperms and systematic evolution of taxa with this sexual system in. Furthermore, five hypotheses regarding evolutionary significance of sterile sex organs in cryptic dioecious are also presented.
Prospects: The future trajectory of related research is presented, with the aim of providing insights into the empirical studies of selection underlying the evolution of cryptic dioecy in angiosperms.

Key words: cryptic dioecy, sex expression, sexual system, phylogeny, evolutionary significance, angiosperms

何花, 谭敦炎, 杨晓琛 (2024) 被子植物隐性雌雄异株性系统的多样性、系统演化及进化意义. 生物多样性, 32, 24149. DOI: 10.17520/biods.2024149.

Hua He, Dunyan Tan, Xiaochen Yang (2024) Cryptic dioecy in angiosperms: Diversity, phylogeny and evolutionary significance. Biodiversity Science, 32, 24149. DOI: 10.17520/biods.2024149.

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https://www.biodiversity-science.net/CN/Y2024/V32/I6/24149

图/表 4

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目 Order	科 Family	属 Genus	种 Species	参考文献 Reference
性表达类型I Sex expression type I
木兰目 Magnoliales	木兰科 Magnoliaceae	拟单性木兰属 Parakmeria	峨眉拟单性木兰 P. omeiensis	余道平等, 2019
棕榈目 Arecales	棕榈科 Arecaceae	海枣属 Phoenix	海枣 P. dactylifera	陆静等, 2021
毛茛目 Ranunculales	毛茛科 Ranunculaceae	唐松草属 Thalictrum	T. macrostylum, T. polygamum, 柔毛唐松草 T. pubescens	Melampy & Hayworth, 1980; Davis SL, 2002; Penny, 2018
昆栏树目 Trochodendrales	昆栏树科 Trochodendraceae	昆栏树属 Trochodendron	昆栏树 T. aralioides	Keng, 1959
五桠果目 Dilleniales	五桠果科 Dilleniaceae	锡叶藤属 Tetracera	锡叶藤属14种 Tetracera sp. 1-14	Kubitzki & Baretta-Kuipers, 1969
葡萄目 Vitales	葡萄科 Vitaceae	葡萄属 Vitis	夏葡萄 V. aestivalis, 河岸葡萄 V. riparia, V. vinifera ssp. silvestris	Brantjes, 1978; Kevan et al, 1985, 1988
金虎尾目 Malpighiales	藤黄科 Clusiaceae	藤黄属 Garcinia	G. brasiliensis	Leal et al, 2012
	红厚壳科 Calophyllaceae	南美杏属 Mammea	M. acuminata, M. africana, 马米杏 M. americana, M. sp. aff. bongo, M. eugenioides var. subsessilifolia, M. punctata, M. cf. punctata, M. siamensis	Dunthorn, 2004
	金虎尾科Malpighiaceae	折背藤属 Madagasikaria	M. andersonii	Davis CC, 2002
	金虎尾科Malpighiaceae	辐翅藤属 Rhynchophora	R. phillipsonii	Anderson WR, 2001
豆目 Fabales	豆科 Fabaceae	皂荚属 Gleditsia	南美皂荚 G. amorphoides	Cerino et al, 2018
桃金娘目 Myrtales	桃金娘科 Myrtaceae	子楝树属 Decaspermum	五瓣子楝树 D. parviflorum	Kevan & Lack, 1985
桃金娘目 Myrtales	桃金娘科 Myrtaceae	番樱桃属 Eugenia	E. albanensis, E. capensis, E. cf. mossambicensis, E. erythrophylla, E. natalitia, E. simii, E. umtamvunensis, E. verdoorniae, E. woodii, E. zeyheri, E.zuluensis, 番樱桃属3种 Eugenia sp. 1-3	van Wyk & Dedekind, 1985; van der Merwe et al, 2005
无患子目 Sapindales	无患子科 Sapindaceae	干果木属 Xerospermum	X. intermedium	Appanah, 1982
石竹目 Caryophyllales	石竹科 Caryophyllaceae	冰漆姑属 Honckenya	H. peploides var. major	Tsukui & Sugawara, 1992
山茱萸目 Cornales	蓝果树科 Nyssaceae	蓝果树属 Nyssa	云南蓝果树 N. yunnanensis	Sun et al, 2009
杜鹃花目 Ericales	柿科 Ebenaceae	柿属 Diospyros	君迁子 D. lotus	陆静等, 2021
	山矾科 Symplocaceae	山矾属 Symplocos	S. celastrinea, S. crenata, S. itatiaiae, S. lanceolata, S. mosenii, S. rhamnifolia, S. saxatilis, S. tetrandra, S. variabilis, S. inopinata, S. kleinii, S. occulta, 微毛山矾 S. wikstroemiifolia	Aranha Filho et al, 2009, 2013; Wang & Hu, 2011
	猕猴桃科 Actinidiaceae	猕猴桃属 Actinidia	软枣猕猴桃 A. arguta, 中华猕猴桃 A. chinensis, 美味猕猴桃 A. deliciosa var. deliciosa, 毛花猕猴桃 A. eriantha, 绵毛猕猴桃 A. fulvicoma var. lanata, 革叶猕猴桃 A. rubricaulis var. coriacea, 对萼猕猴桃 A. valvata, 葛枣猕猴桃 A. polygama	unpublished
	猕猴桃科 Actinidiaceae	水东哥属 Saurauia	S. bullosa, S. brachybotrys, S. excelsa, S. humboldtiana, S. tomentosa, S. putumayonis, S. ursina, S. veraguensis	Soejarto, 1969; Cane, 1993
伞形目 Apiales	毛柴木科 Pennantiaceae	毛柴木属 Pennantia	毛柴木属3种 Pennantia sp. 1-3	Baylis, 1977
伞形目 Apiales	五加科 Araliaceae	南洋参属 Polyscias	P. dzumacensis, P. microbotrys, P. pancheri, P. subincisa	Schlessman et al, 1990
紫草目 Boraginales	紫草科 Boraginaceae	破布木属 Cordia	C. crenata, C. cf. grandicalyx, C. sinensis	Pätzold et al, 2023
龙胆目 Gentianales	茜草科 Rubiaceae	木巴戟属 Morinda	鸡眼藤 M. parvifolia	Liu et al, 2012
唇形目 Lamiales	木樨科 Oleaceae	梣属 Fraxinus	维斯谷归化种群中的花梣 Fraxinus ornus in naturalized Vis Valley populations	Vernet et al, 2016
茄目 Solanales	茄科 Solanaceae	茄属 Solanum	S. asymmetriphyllum, S. carduiforme, S. cataphractum, S. cowiei, S. cunninghamii, S. dioicum, S. leopoldense, S. ossicruentum, S. petraeum, S. polygamum, S. tudununggae, S. vansittartensis, S. sejunctum	Anderson & Symon, 1989; Martine et al, 2013; Anderson et al, 2015; Ndem-Galbert et al, 2021
性表达类型II Sex expression type II
蔷薇目 Rosales	蔷薇科 Rosaceae	蔷薇属 Rosa	草原玫瑰 R. setigera	Kevan et al, 1990
桃金娘目 Myrtales	桃金娘科 Myrtaceae	多香果属 Pimenta	多香果 P. dioica	Boyd & Benkeblia, 2011
无患子目 Sapindales	无患子科 Sapindaceae	槭属 Acer	红花槭 A. rubrum	Batra, 1985
无患子目 Sapindales	楝科 Meliaceae	杜楝属 Turraea	T. dargentiana	Baider & Florens, 2016
锦葵目 Malvales	瑞香科 Thymelaeaceae	荛花属 Wikstroemia	夏威夷岛和可爱岛中荛花属种群 Wikstroemia populations on Kauai, Hawaii	Mayer & Charlesworth, 1991
锦葵目 Malvales	锦葵科 Malvaceae	非洲芙蓉属 Dombeya	D. ciliata, D. delislei, D. elegans, D. ficulnea, D. formosa, D. pilosa, D. punctata, D. reclinata	Humeau et al, 1999, 2000; Le Péchon et al, 2013
石竹目 Caryophyllales	仙人掌科 Cactaceae	旗号掌属 Consolea	C. corallicola, C. millspaughii, 鱼鳞掌 C. moniliformis, C. nashii, C. picardae, C. rubescens, C. spinosissima	Rebman & Pinkava, 2001; Strittmatter et al, 2006, 2008
		鹿角柱属 Echinocereus	赤花虾 E. coccineus, E. mombergerianus, E. pacificus, 多刺虾鹿角柱 E. polyacanthus	Hernández-Cruz et al, 2018
		仙人掌属 Opuntia	O. quitensis, O. robusta, O. stenopetala	Anderson EF, 2001; Hernández-Cruz et al, 2019
		摩天柱属 Pachycereus	武伦柱 P. pringlei	Fleming, 1989
		木麒麟属 Pereskia	P. portulacifolia, P. marcanoi, P. quisqueyana, P. zinniiflora	Areces-Mallea, 1992
杜鹃花目 Ericales	杜鹃花科 Ericaceae	南白珠属 Pernettya	P. rigida	Anderson et al, 2000
菊目 Asterales	菊科 Asteraceae	假泽兰属 Mikania	假泽兰属5种 Mikania 5 sp. 1-5	Holmes, 1991
茶茱萸目 Icacinales	茶茱萸科 Icacinaceae	假柴龙树属 Nothapodytes	臭味假柴龙树 N. nimmoniana	Sharma et al, 2010
龙胆目 Gentianales	茜草科 Rubiaceae	舒榄属 Bertiera	B. borbonica	Pailler et al, 1998b
		弯管花属 Chassalia	C. corallioides	Pailler et al, 1998a
		牛眼棠属 Cordiera	C. concolor	Judkevich et al, 2022
		靛榄属 Genipa	靛榄 G. americana	Judkevich et al, 2022
		蓝茜树属 Randia	R. calycina, R. heteromera	Judkevich et al, 2022
		木巴戟属 Morinda	须弥巴戟 M. villosa	Sugawara et al, 2011
		玉叶金花属 Mussaenda	短裂玉叶金花 M. breviloba, 尾裂玉叶金花 M. caudatiloba, 仁昌玉叶金花 M. chingii, 展枝玉叶金花 M. divaricata, 楠藤 M. erosa, 海南玉叶金花 M. hainanensis, 粗毛玉叶金花 M. hirsutula, 红毛玉叶金花 M. hossei, 广西玉叶金花 M. kwangsiensis, 广东玉叶金花 M. kwangtungensis, 疏花玉叶金花 M. laxiflora, 长瓣玉叶金花 M. longipetala, 多毛玉叶金花 M. mollissima, 小玉叶金花 M. parviflora, M. pilosissima, 屏边玉叶金花 M. pingbianensis, 玉叶金花 M. pubescens, 白花玉叶金花 M. pubescens var. alba, 贡山玉叶金花 M. treutleri, 玉叶金花属5种 Mussaenda sp. 1-5	邓小芳, 2007; Li et al, 2010; 陈湜, 2013
		沙九节属 Pagamea	P. duckei	Terra-Araujo et al, 2012
		九节属 Psychotria	九节 P. rubra	Watanabe et al, 2013
	马钱科 Loganiaceae	髯管花属 Geniostoma	G. borbonica	Humeau et al, 2003
	马钱科 Loganiaceae	顶髯管花属 Labordia	L. cyrtandrae, L. degeneri, L. fagraeoidea, L. hedyosmifolia, L. hirtella, L. hosakana, L. lydgatei, L. pumila, L. sessilis, L.venosa, L. waialealae, L. waiolani	Motley & Carr, 1998
	夹竹桃科 Apocynaceae	萝芙木属 Rauvolfia	R. sellowii	Souto & Koch, 2018
唇形目 Lamiales	马鞭草科 Verbenaceae	琴木属 Citharexylum	C. myrianthum	Rocca & Sazima, 2006
唇形目 Lamiales	唇形科 Lamiaceae	紫珠属 Callicarpa	C. glabra, C. nishimurae, C. subpubescens	Kawakubo, 1998
茄目 Solanales	茄科 Solanaceae	山盏茄属 Deprea	D. paneroi	Sawyer & Anderson, 2000
		巫灵木属 Dunalia	D. solanacea	Hepper, 1991
		枸杞属 Lycium	L. arenicolum, L. horridum, L. tetrandrum, L. villosum	Spies et al, 1993
茄目 Solanales	茄科 Solanaceae	茄属 Solanum	S. appendiculatum, S. campanulatum, S. confertiseriatum, S. crotonoides, S. conocarpum	Knapp et al, 1998; Anderson et al, 2015
		串钟花属 Symonanthus	S. aromaticus, S. bancroftii	Haegi, 1981
		睡茄属 Withania	W. aristata, W. coagulans	Sawyer & Anderson, 2000; Anderson et al, 2006
性表达类型III Sex expression type III
金虎尾目 Malpighiales	金虎尾科Malpighiaceae	异金英属 Spachea	S. membranacea	Steiner, 1985
石竹目 Caryophyllales	石竹科Caryophyllaceae	洋漆姑属 Schiedea	S. adamantis	Mayer & Charlesworth, 1991; Sakai et al, 1997
	仙人掌科 Cactaceae	圆柱掌属 Cylindropuntia	C. wolfii	Ramadoss et al, 2022
	仙人掌科 Cactaceae	裸萼球属 Gymnocalycium	罗星丸 G. bruchii	Delbón et al, 2024
杜鹃花目 Ericales	山榄科 Sapotaceae	星苹果属 Chrysophyllum	C. gonocarpum	Judkevich et al, 2023
杜鹃花目 Ericales	杜鹃花科 Ericaceae	岩梨属 Epigaea	岩梨 E. repens	Clay & Ellstrand, 1981
性表达类型IV Sex expression type IV (1) (2)
菊目 Asterales	菊科 Asteraceae	蜂斗菜属 Petasites	(1)蜂斗菜 P. japonicas; (2)毛裂蜂斗菜 P. tricholobus	Sakai et al, 2008; Yu et al, 2011

目 Order	科 Family	属 Genus	种 Species	参考文献 Reference
性表达类型I Sex expression type I
木兰目 Magnoliales	木兰科 Magnoliaceae	拟单性木兰属 Parakmeria	峨眉拟单性木兰 P. omeiensis	余道平等, 2019
棕榈目 Arecales	棕榈科 Arecaceae	海枣属 Phoenix	海枣 P. dactylifera	陆静等, 2021
毛茛目 Ranunculales	毛茛科 Ranunculaceae	唐松草属 Thalictrum	T. macrostylum, T. polygamum, 柔毛唐松草 T. pubescens	Melampy & Hayworth, 1980; Davis SL, 2002; Penny, 2018
昆栏树目 Trochodendrales	昆栏树科 Trochodendraceae	昆栏树属 Trochodendron	昆栏树 T. aralioides	Keng, 1959
五桠果目 Dilleniales	五桠果科 Dilleniaceae	锡叶藤属 Tetracera	锡叶藤属14种 Tetracera sp. 1-14	Kubitzki & Baretta-Kuipers, 1969
葡萄目 Vitales	葡萄科 Vitaceae	葡萄属 Vitis	夏葡萄 V. aestivalis, 河岸葡萄 V. riparia, V. vinifera ssp. silvestris	Brantjes, 1978; Kevan et al, 1985, 1988
金虎尾目 Malpighiales	藤黄科 Clusiaceae	藤黄属 Garcinia	G. brasiliensis	Leal et al, 2012
	红厚壳科 Calophyllaceae	南美杏属 Mammea	M. acuminata, M. africana, 马米杏 M. americana, M. sp. aff. bongo, M. eugenioides var. subsessilifolia, M. punctata, M. cf. punctata, M. siamensis	Dunthorn, 2004
	金虎尾科Malpighiaceae	折背藤属 Madagasikaria	M. andersonii	Davis CC, 2002
	金虎尾科Malpighiaceae	辐翅藤属 Rhynchophora	R. phillipsonii	Anderson WR, 2001
豆目 Fabales	豆科 Fabaceae	皂荚属 Gleditsia	南美皂荚 G. amorphoides	Cerino et al, 2018
桃金娘目 Myrtales	桃金娘科 Myrtaceae	子楝树属 Decaspermum	五瓣子楝树 D. parviflorum	Kevan & Lack, 1985
桃金娘目 Myrtales	桃金娘科 Myrtaceae	番樱桃属 Eugenia	E. albanensis, E. capensis, E. cf. mossambicensis, E. erythrophylla, E. natalitia, E. simii, E. umtamvunensis, E. verdoorniae, E. woodii, E. zeyheri, E.zuluensis, 番樱桃属3种 Eugenia sp. 1-3	van Wyk & Dedekind, 1985; van der Merwe et al, 2005
无患子目 Sapindales	无患子科 Sapindaceae	干果木属 Xerospermum	X. intermedium	Appanah, 1982
石竹目 Caryophyllales	石竹科 Caryophyllaceae	冰漆姑属 Honckenya	H. peploides var. major	Tsukui & Sugawara, 1992
山茱萸目 Cornales	蓝果树科 Nyssaceae	蓝果树属 Nyssa	云南蓝果树 N. yunnanensis	Sun et al, 2009
杜鹃花目 Ericales	柿科 Ebenaceae	柿属 Diospyros	君迁子 D. lotus	陆静等, 2021
	山矾科 Symplocaceae	山矾属 Symplocos	S. celastrinea, S. crenata, S. itatiaiae, S. lanceolata, S. mosenii, S. rhamnifolia, S. saxatilis, S. tetrandra, S. variabilis, S. inopinata, S. kleinii, S. occulta, 微毛山矾 S. wikstroemiifolia	Aranha Filho et al, 2009, 2013; Wang & Hu, 2011
	猕猴桃科 Actinidiaceae	猕猴桃属 Actinidia	软枣猕猴桃 A. arguta, 中华猕猴桃 A. chinensis, 美味猕猴桃 A. deliciosa var. deliciosa, 毛花猕猴桃 A. eriantha, 绵毛猕猴桃 A. fulvicoma var. lanata, 革叶猕猴桃 A. rubricaulis var. coriacea, 对萼猕猴桃 A. valvata, 葛枣猕猴桃 A. polygama	unpublished
	猕猴桃科 Actinidiaceae	水东哥属 Saurauia	S. bullosa, S. brachybotrys, S. excelsa, S. humboldtiana, S. tomentosa, S. putumayonis, S. ursina, S. veraguensis	Soejarto, 1969; Cane, 1993
伞形目 Apiales	毛柴木科 Pennantiaceae	毛柴木属 Pennantia	毛柴木属3种 Pennantia sp. 1-3	Baylis, 1977
伞形目 Apiales	五加科 Araliaceae	南洋参属 Polyscias	P. dzumacensis, P. microbotrys, P. pancheri, P. subincisa	Schlessman et al, 1990
紫草目 Boraginales	紫草科 Boraginaceae	破布木属 Cordia	C. crenata, C. cf. grandicalyx, C. sinensis	Pätzold et al, 2023
龙胆目 Gentianales	茜草科 Rubiaceae	木巴戟属 Morinda	鸡眼藤 M. parvifolia	Liu et al, 2012
唇形目 Lamiales	木樨科 Oleaceae	梣属 Fraxinus	维斯谷归化种群中的花梣 Fraxinus ornus in naturalized Vis Valley populations	Vernet et al, 2016
茄目 Solanales	茄科 Solanaceae	茄属 Solanum	S. asymmetriphyllum, S. carduiforme, S. cataphractum, S. cowiei, S. cunninghamii, S. dioicum, S. leopoldense, S. ossicruentum, S. petraeum, S. polygamum, S. tudununggae, S. vansittartensis, S. sejunctum	Anderson & Symon, 1989; Martine et al, 2013; Anderson et al, 2015; Ndem-Galbert et al, 2021
性表达类型II Sex expression type II
蔷薇目 Rosales	蔷薇科 Rosaceae	蔷薇属 Rosa	草原玫瑰 R. setigera	Kevan et al, 1990
桃金娘目 Myrtales	桃金娘科 Myrtaceae	多香果属 Pimenta	多香果 P. dioica	Boyd & Benkeblia, 2011
无患子目 Sapindales	无患子科 Sapindaceae	槭属 Acer	红花槭 A. rubrum	Batra, 1985
无患子目 Sapindales	楝科 Meliaceae	杜楝属 Turraea	T. dargentiana	Baider & Florens, 2016
锦葵目 Malvales	瑞香科 Thymelaeaceae	荛花属 Wikstroemia	夏威夷岛和可爱岛中荛花属种群 Wikstroemia populations on Kauai, Hawaii	Mayer & Charlesworth, 1991
锦葵目 Malvales	锦葵科 Malvaceae	非洲芙蓉属 Dombeya	D. ciliata, D. delislei, D. elegans, D. ficulnea, D. formosa, D. pilosa, D. punctata, D. reclinata	Humeau et al, 1999, 2000; Le Péchon et al, 2013
石竹目 Caryophyllales	仙人掌科 Cactaceae	旗号掌属 Consolea	C. corallicola, C. millspaughii, 鱼鳞掌 C. moniliformis, C. nashii, C. picardae, C. rubescens, C. spinosissima	Rebman & Pinkava, 2001; Strittmatter et al, 2006, 2008
		鹿角柱属 Echinocereus	赤花虾 E. coccineus, E. mombergerianus, E. pacificus, 多刺虾鹿角柱 E. polyacanthus	Hernández-Cruz et al, 2018
		仙人掌属 Opuntia	O. quitensis, O. robusta, O. stenopetala	Anderson EF, 2001; Hernández-Cruz et al, 2019
		摩天柱属 Pachycereus	武伦柱 P. pringlei	Fleming, 1989
		木麒麟属 Pereskia	P. portulacifolia, P. marcanoi, P. quisqueyana, P. zinniiflora	Areces-Mallea, 1992
杜鹃花目 Ericales	杜鹃花科 Ericaceae	南白珠属 Pernettya	P. rigida	Anderson et al, 2000
菊目 Asterales	菊科 Asteraceae	假泽兰属 Mikania	假泽兰属5种 Mikania 5 sp. 1-5	Holmes, 1991
茶茱萸目 Icacinales	茶茱萸科 Icacinaceae	假柴龙树属 Nothapodytes	臭味假柴龙树 N. nimmoniana	Sharma et al, 2010
龙胆目 Gentianales	茜草科 Rubiaceae	舒榄属 Bertiera	B. borbonica	Pailler et al, 1998b
		弯管花属 Chassalia	C. corallioides	Pailler et al, 1998a
		牛眼棠属 Cordiera	C. concolor	Judkevich et al, 2022
		靛榄属 Genipa	靛榄 G. americana	Judkevich et al, 2022
		蓝茜树属 Randia	R. calycina, R. heteromera	Judkevich et al, 2022
		木巴戟属 Morinda	须弥巴戟 M. villosa	Sugawara et al, 2011
		玉叶金花属 Mussaenda	短裂玉叶金花 M. breviloba, 尾裂玉叶金花 M. caudatiloba, 仁昌玉叶金花 M. chingii, 展枝玉叶金花 M. divaricata, 楠藤 M. erosa, 海南玉叶金花 M. hainanensis, 粗毛玉叶金花 M. hirsutula, 红毛玉叶金花 M. hossei, 广西玉叶金花 M. kwangsiensis, 广东玉叶金花 M. kwangtungensis, 疏花玉叶金花 M. laxiflora, 长瓣玉叶金花 M. longipetala, 多毛玉叶金花 M. mollissima, 小玉叶金花 M. parviflora, M. pilosissima, 屏边玉叶金花 M. pingbianensis, 玉叶金花 M. pubescens, 白花玉叶金花 M. pubescens var. alba, 贡山玉叶金花 M. treutleri, 玉叶金花属5种 Mussaenda sp. 1-5	邓小芳, 2007; Li et al, 2010; 陈湜, 2013
		沙九节属 Pagamea	P. duckei	Terra-Araujo et al, 2012
		九节属 Psychotria	九节 P. rubra	Watanabe et al, 2013
	马钱科 Loganiaceae	髯管花属 Geniostoma	G. borbonica	Humeau et al, 2003
	马钱科 Loganiaceae	顶髯管花属 Labordia	L. cyrtandrae, L. degeneri, L. fagraeoidea, L. hedyosmifolia, L. hirtella, L. hosakana, L. lydgatei, L. pumila, L. sessilis, L.venosa, L. waialealae, L. waiolani	Motley & Carr, 1998
	夹竹桃科 Apocynaceae	萝芙木属 Rauvolfia	R. sellowii	Souto & Koch, 2018
唇形目 Lamiales	马鞭草科 Verbenaceae	琴木属 Citharexylum	C. myrianthum	Rocca & Sazima, 2006
唇形目 Lamiales	唇形科 Lamiaceae	紫珠属 Callicarpa	C. glabra, C. nishimurae, C. subpubescens	Kawakubo, 1998
茄目 Solanales	茄科 Solanaceae	山盏茄属 Deprea	D. paneroi	Sawyer & Anderson, 2000
		巫灵木属 Dunalia	D. solanacea	Hepper, 1991
		枸杞属 Lycium	L. arenicolum, L. horridum, L. tetrandrum, L. villosum	Spies et al, 1993
茄目 Solanales	茄科 Solanaceae	茄属 Solanum	S. appendiculatum, S. campanulatum, S. confertiseriatum, S. crotonoides, S. conocarpum	Knapp et al, 1998; Anderson et al, 2015
		串钟花属 Symonanthus	S. aromaticus, S. bancroftii	Haegi, 1981
		睡茄属 Withania	W. aristata, W. coagulans	Sawyer & Anderson, 2000; Anderson et al, 2006
性表达类型III Sex expression type III
金虎尾目 Malpighiales	金虎尾科Malpighiaceae	异金英属 Spachea	S. membranacea	Steiner, 1985
石竹目 Caryophyllales	石竹科Caryophyllaceae	洋漆姑属 Schiedea	S. adamantis	Mayer & Charlesworth, 1991; Sakai et al, 1997
	仙人掌科 Cactaceae	圆柱掌属 Cylindropuntia	C. wolfii	Ramadoss et al, 2022
	仙人掌科 Cactaceae	裸萼球属 Gymnocalycium	罗星丸 G. bruchii	Delbón et al, 2024
杜鹃花目 Ericales	山榄科 Sapotaceae	星苹果属 Chrysophyllum	C. gonocarpum	Judkevich et al, 2023
杜鹃花目 Ericales	杜鹃花科 Ericaceae	岩梨属 Epigaea	岩梨 E. repens	Clay & Ellstrand, 1981
性表达类型IV Sex expression type IV (1) (2)
菊目 Asterales	菊科 Asteraceae	蜂斗菜属 Petasites	(1)蜂斗菜 P. japonicas; (2)毛裂蜂斗菜 P. tricholobus	Sakai et al, 2008; Yu et al, 2011