被子植物雌全同株性系统: 系统演化、性表达与进化意义

doi:10.3724/SP.J.1258.2014.00008

摘要/Abstract

摘要：

雌全同株是指雌花和两性花共同发生在同一植株上的性表达形式。作为被子植物从雌雄同花(两性花)向雌雄同株异花进化的一个重要阶段, 雌全同株性系统在减少昆虫对雌性的取食和伤害、提高异交率以减少近交衰退、减少雌/雄功能干扰、提高雌/雄性功能间资源分配的灵活性, 以及吸引传粉者等方面具有重要的进化适应意义。根据APG III分类系统, 雌全同株性系统在被子植物木兰分支(magnoliids)的短蕊花科、单子叶植物分支(monocots)的天南星科和禾本科, 以及核心真双子叶植物分支(core eudicots)中的菊科、苋科、唇形科和石竹科等23科中均有报道, 且以菊科植物中最多。雌全同株植物不同类群的雌花和两性花在位置、形态、大小及开花时间等性表达特征上表现出多样化, 且这些特征不仅受遗传因子的调控, 还受可获得资源(如营养、光照、温度和水分等条件)的制约。该文针对我国对雌全同株性系统的研究还相对较少的现状, 重点对具雌全同株性系统的类群在被子植物中的分布与系统演化、性表达与环境的关系等方面进行了分析与总结, 并对有关其进化适应意义的5个假说进行了介绍和评价, 对今后的研究方向进行了展望, 以期为推动我国对被子植物雌全同株性系统的进化式样与机制研究提供理论资料。

关键词: 被子植物, 进化意义, 雌全同株, 系统演化, 性表达, 性系统

Abstract:

Gynomonoecy is the mode of sex expression in which female and bisexual flowers occur on the same plants; it is an important step in the evolution of monoecy from hermaphroditism in angiosperms. The sexual system of gynomonoecy is considered to play several important roles, including reducing herbivore damage to pistils, reducing inbreeding depression by favoring out-crossing and pollen-pistil interference, enhancing the flexibility of resource allocation to male and female functions in two flower types, and attracting pollinators. According to the classification of APG III system, gynomonoecy occurs in about 23 families in angiosperms including Lactoridaceae in magnoliids, Araceae and Poaceae in monocots, and Asteraceae, Amaranthaceae, Lamiaceae and other families in eudicots. Gynomonoecy is most common in Asteraceae. Female and bisexual flowers from different gynomonoecious taxa show diversified sex expressions, such as their position in inflorescence, morphs, size and flowering time. These traits are regulated not only by genetic factors, but also by resource availability (e.g., nutrition, light, temperature, and moisture conditions). Because reports on gynomonoecy in China are rare, this paper emphasizes our analysis and summary of the phylogeny of gynomonoecious taxa and their evolution in angiosperm and the relationship between their sex expression and environmental conditions. In addition, five hypotheses on the evolutionary significance of gynomonoecy are introduced and evaluated in this paper. Finally, we discuss prospects for further research in this topical area. Our purpose is to provide a theoretical framework for research on evolutionary patterns and mechanisms of gynomonoecy for angiosperms in China.

Key words: angiosperms, evolutionary significance, gynomonoecy, phylogeny, sex expression, sexual systems

吉乃提汗·马木提, 谭敦炎. 被子植物雌全同株性系统: 系统演化、性表达与进化意义. 植物生态学报, 2014, 38(1): 76-90. DOI: 10.3724/SP.J.1258.2014.00008

Jannathan MAMUT, TAN Dun-Yan. Gynomonoecy in angiosperms: phylogeny, sex expression and evolutionary significance. Chinese Journal of Plant Ecology, 2014, 38(1): 76-90. DOI: 10.3724/SP.J.1258.2014.00008

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图/表 2

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分类群 Taxon	性系统类型 Type of sexual system		参考文献 Reference
短蕊花科 Lactoridaceae Lactoris	雌全同株 Gynomonoecy		Bernardello et al., 1999
禾本科 Gramineae 小丽草属 Coelachne	雌全同株 Gynomonoecy		Yampolsky & Yampolsky, 1922; Lu & Huang, 2006
Tripsacum, Aulonemia	雌全同株 Gynomonoecy		Dewald et al., 1987; Grombone-Guaratini et al., 2011
刺叶树科 Xanthorrhoeaceae 独尾草属 Eremurus	雌全同株、两性花植株 Gynomonoecy, hermaphroditism		Mamut et al., 2014
天南星科 Araceae 麒麟尾属 Epipremnum, Heteropsis, 崖角藤属 Rhaphidophora, Rhodospatha	雌全同株 Gynomonoecy		Yampolsky & Yampolsky, 1922; Lu & Huang, 2006
洋二仙草科 Gunneraceae Gunnera	雌全同株 Gynomonoecy		Palkovic, 1978; Lowrey & Robinson, 1988
胡桃科 Juglandaceae 马尾树属 Rhoiptelea	雌全同株 Gynomonoecy		Sun et al., 2006; Lu & Huang, 2006
荨麻科 Urticaceae 墙草属 Parietaria	雌全同株 Gynomonoecy		Franchi et al., 2007
桃金娘科 Myrtaceae 白千层属 Melaleuca	雌全同株、雌全异株 Gynomonoecy, gynodioecy		Barlow & Forrester, 1984
柳叶菜科 Onagraceae 倒挂金钟属 Fuchsia	雌全同株 Gynomonoecy		Lu & Huang, 2006
牻牛儿苗科 Geraniaceae 牻牛儿苗属 Erodium, 老鹳草属 Geranium	雌全同株 Gynomonoecy		Lu & Huang, 2006
景天科 Crassulaceae 八宝属 Hylotelephium, 景天属 Sedum	雌全同株 Gynomonoecy		Lu & Huang, 2006
石竹科 Caryophyllaceae 蝇子草属 Silene	雌全同株 Gynomonoecy		Hermanutz & Innes, 1994; Maurice, 1999; Lu & Huang, 2006
蝇子草属 Silene,石竹属 Dianthus,繁缕属 Stellaria	雌全同株、雌全异株Gynomonoecy, gynodioecy		Maurice, 1999; Collin et al., 2009; Dufay et al., 2010 ; Casimiro-Soriguer et al., 2013
麦仙翁属 Agrostemma, 蚤缀属 Arenaria, 卷耳属 Cerastium, 石竹属 Dianthus, 剪秋罗属 Lychnis, 高山漆姑草属 Minuartia,漆姑草属 Sagina, 牛漆姑草属 Spergularia,繁缕属 Stellaria	雌全同株 Gynomonoecy		Lu & Huang, 2006
苋科 Amaranthaceae (包括藜科 Chenopodiaceae) Arthrocnemum, Didymantlius, Heterostachys, Microcnemum, Monolepis, Osteocarpum, Spirostachys, Secticornia, Threlkeldia	雌全同株 Gynomonoecy		Yampolsky & Yampolsky, 1922
藜属 Chenopodium	雌全同株 Gynomonoecy		Bhargava et al., 2007
盐节木属 Halocnemum, 盐千屈菜属 Halopeplis, 盐爪爪属 Kalidium, 盐角草属 Salicornia, 碱蓬属 Suaeda	雌全同株 Gynomonoecy		Yampolsky & Yampolsky, 1922; Lu & Huang, 2006
刺藜属 Dysphania, 棉藜属 Kirilowia, 地肤属 Kochia, 绒藜属 Londesia, 兜藜属 Panderia	雌全同株 Gynomonoecy		Lu & Huang, 2006
菠菜属 Spinacia	雌株、雄株、雌全同株、雄全同株 Female, male, gynomonoecy, andromonoecy		Komai & Masuda, 2004; Onodera et al., 2008
花荵科 Polemoniaceae 花荵属 Polemonium	雌全同株 Gynomonoecy		Yampolsky & Yampolsky, 1922; Lu & Huang, 2006
山榄科 Sapotaceae 山榄属 Planchonella	雌全同株 Gynomonoecy		Méndez & Munzinger,2010
龙胆科 Gentianaceae 假龙胆属 Gentianella	雌全同株 Gynomonoecy		Pringle, 2011
分类群 Taxon	性系统类型 Type of sexual system		参考文献 Reference
爵床科 Acanthaceae Brachystephanus	雌全同株 Gynomonoecy		Champluvier, 1997
木犀科 Oleaceae 木犀榄属 Olea, 木犀属 Osmanthus, 丁香属 Syringa	雌全同株 Gynomonoecy		Lu & Huang, 2006
唇形科 Lamiaceae 新风轮属 Calamintha,青兰属 Dracocephalum, 活血丹属 Glechoma,神香草属 Hyssopus, 地笋属 Lycopus, 蜜蜂花属 Melissa, 薄荷属 Mentha,荆芥属 Nepeta, 牛至属 Origanum, 夏枯草属 Prunella, 鼠尾草属 Salvia,水苏属 Stachys, 百里香属 Thymus	雌全同株 Gynomonoecy		Lu & Huang, 2006
车前草科 Plantaginaceae 车前草属 Plantago	雌全同株 Gynomonoecy		Lu & Huang, 2006
菊科 Asteraceae Abrotanella, Acamptopappus, Achaetogeron, Achmophora, Achyrachaena, Achyrocline, Adelostigma, Aganippea, Aiolotheca, Allendea, Aniauria, Am- blyocarpum, Amblyopappus, Amellus, Amphidoxa, Amphiglossa, Amphori- carpus, Anaglypha, Anaxeton, Antithrixia, Aphanactis, Aphanostephus, Agyroxiphium, Arnica, Arnicastrum, Arrowsmithia, Asteropsis, Astephania, Asteromoea, Athrixia, Athroisma, Axiniphyllum, Baeria, Balleya, Balsam- orhiza, Bartlettia, Bellida, Bellium, Bellis, Berlandiera, Bi-Leveillea, Blennosperma, Blepharispermum, Blepharizonia, Boltonia, Borrichia, Br- achychaeta, Brachyclados, Brachycome, Brachyglottis, Bradburia, Bryo- morphe, Burrielia, Cacosmia, Cadiscus, Calyptrocarpus, Cassinia, Cel- misia, Ceratogyne, Ceruana, Chaetanthira, Chaetopappa, Chaptalia, Chardinia, Charieis, Chiliophyllum, Chiliotrichium, Chionolaena, 被菊属 Chlamydites, Chroilema, Chromolepis, Chrysactinia, Chrysanthellum, Chrysogonum, Chrysoma, Chrysopsis, Clappia, Clibadium, Cnicothamnus, Coleocoma, Commidendron, Corethrogyne,秋英属 Cosmos, Crocidium, Crupina, Cylindrocline, Cypselodontia, Delamerea, Denekia, Dichaeto- phora, Dicranocarpus, Dimorphocoma, Dimorphotheca, Diplostephium, Dipterocome, Disparago, Dugesia, Egletes, Elachanthus, Elvira, Engel- mannia, Epallage, Eremothamnus, Eriachaenium, Ericameria, Erioce- phalus, Eriothrix, Ermeiastrum, Espeletia,梳黄菊属 Euryops, Eutetras, Evax, Faceles, Faujasia, Felicia, Fresenia, Gamolepis, Garcilassa, Garuleum, Geigeria, Glossocardia, Glycideras, Goldmanella, Golionema, Grantia, Gratwickia, Greenella, Grypocarpha, Guardiola,小葵子属 Guizotia, Gundlachia, Gundelia, Gutierrezia, Gymnarrhena, Gymnodiscus, Gymnosperma, Gymnostephium, Haastia, Hazardia, Heliopsis, Hemi- zonella, Hemizonia, Henricia, Heptanthus, Hertia, Heterolepis, Heteros- permum, Heterotheca, Hidalgoa, Hinterhubera, Hippia, Homochaete, Homochroma, Homogyne, Hysterionica, Ichthyothere, Inulopsis, Ischnea, Isocarpha, Isoetopsis, Isostigma, Ixiolaena, Jaegena, Jaegeria, Jasonia, Keerlia, Koehneola, Lachnophyllum, Laestadia,瓶头草属 Lagenophora, Lagophylla, Lantanopsis, Lasiocoma, Lasiopogon, Lasthenia, Lecocarpus, Leontonyx, Lepidophyllum, Lepidostephium, Leptocarpha, Leptorhynchus, Lessingia, Leucopholis, Leyssera, Liabum, Lindheimera, Lipochaeta, Lucilia, Macowania, Madia, Mairia, Melalema, Melampodium, Mela- nodendron, Melanthera, Merrittia, Micractis, Microlecane, Micropsis, Micropus, Microtrichia, Milleria, Minuria, Minurothamnus, Monactis, Monarrhenus, Monolopia, Monoptilon, Moonia, Montanoa,帚菊木属 Mutisia, Nananthea,毛冠菊属 Nannoglottis, Nanothamnus, Narvalina, Nerrittia, Nestlera, Nicolasia, Nidorella, Nolletia, Odontospermum, Oedera, Olearia, Oligocarpus, Olivaea, Onoseris, Oreochrysum, Osmites, Osmitopsis, Osteospermum, Othonna, Otochlamys, Oxypappus, Pach- ylaena, Pachyrhynchus, Parthenice, Parantennaria, Pectis, Pentachaeta, Perityle, Perralderia, Perymenium, Petalacte, Petalactella, Petrollinia, Phacellothrix, Phaenocoma, Phalacrocarpum, Phemmera, Philactis, Philoglossa, Philyrophyllum, Phymaspermum, Pinillosia, Plagiocheilus, Plazia, Pleurophyllum, Podochaenium, Podocoma, Podolepis, Polygyne, Polymnia, Postia, Printzia, Psiadia, Psila, Psilactis, Psilocarphus, Pterigeron, Raillardella, Raoulia, Relhania, Remya, Rhantherium, Rhodogeron, Riddellia, Riencourtia, Rigiopappus, Rochonia, Ruckeria, Rumfordia, Sabazia, Sanvitalia, Sartwellia, Schistocarpha, Schist- ostephium, Schizoptera, Schizotrichia, Schkuhria, Schoenia, Sclerocarpus, Scyphocoronis, Selloa, Sericocarpus, Siebera, Silphium, Sommerfeltia, Sphacophyllum, Sphaeromorphaea, Sphagneticola, Spilanthus, Stachy- cephalum, Staurochlamys, Steirodiscus, Stemmatella, Stenocline, Stephan- odoria, Stilpnogyne, Stuartina, Styloclyne, Syncephalantha, Syntrichop- appus, Tessaria, Tetracanthus, Tetragonotheca, Tetramolopium,	雌全同株 Gynomonoecy		Yampolsky & Yampolsky, 1922
分类群 Taxon	性系统类型 Type of sexual system		参考文献 Reference
Tetranthus, Tetraperone, Thelesperma, Thespidium, Thymopsis, Tolbonia,地菊属 Townsendia, Tragoceros, Trichocline, Trigonospermum, Triploce- phalum, Tripteris, Urbinella, Vanclevea, Varilla, Venegazia, Vieraea, Villanova, Vittadinia, Waitzia, Welwitschiella, Whitneya, Wyomingia, Xanthisma,旱花属 Xeranthemum, Zaluzania, Zexmenia, Zoegea
Lasianthaea, Liabum,羽芒菊属 Tridax, Verbesina	雌全同株 Gynomonoecy		Bullock, 1985
Baltimora,鬼针草属 Bidens,鳢肠属 Eclipta	雌全同株 Gynomonoecy		Bullock, 1985; Yampolsky & Yampolsky, 1922
Ambylocarpum, Asteriscus, Chrysophthalmum, Cylindrocline, Dittrichia, Doellia, Duhaldea, Geigeria, Ighermia, Limbarda, Pseudoconyza, Rhan- terium, Schizogyne, Streptoglossa, Telekia, Vierea, Xerolekia, Zoutpan- sbergia	雌全同株 Gynomonoecy		Torices & Anderberg, 2009
Callilepis, Calostephane, Lifago, Mollera, Ondetia, Pallenis, Pechuel- L?schea, Porphyrostemma, Rosenia, Sachsia, Stenachaenium	雌全同株 Gynomonoecy		Yampolsky & Yampolsky, 1922; Torices & Anderberg, 2009
Ifloga	雌全同株 Gynomonoecy		Yampolsky & Yampolsky, 1922; Bergh & Verboom, 2011
假莲属 Conyza	雌全同株 Gynomonoecy		Yampolsky & Yampolsky, 1922; Urdampilleta et al., 2005; Lu & Huang, 2006
旋覆花属 Inula, 山黄菊属 Anisopappus, 牛眼菊属 Buphthalmum, 天门精属 Carpesium, 六棱菊属 Laggera, 蚤草属 Pulicaria	雌全同株 Gynomonoecy		Yampolsky & Yampolsky, 1922; Lu & Huang, 2006; Torices & Anderberg, 2009
艾纳香属 Blumea, 阔苞菊属 Pluchea	雌全同株 Gynomonoecy		Lu & Huang, 2006; Torices & Anderberg, 2009
蟛蜞菊属 Wedelia, 百日菊属 Zinnia	雌全同株 Gynomonoecy		Yampolsky & Yampolsky, 1922; Bullock, 1985; Lu & Huang, 2006
紫菀属 Aster	雌全同株 Gynomonoecy		Bertin & Kerwin, 1998; Lu & Huang, 2006
山芫荽属 Cotula	两性花植株、雌全同株、雌雄同株、亚雌雄异株、雌雄异株Hermaphroditism, gynomonoecy, monoecy, subdioecy, dioecy		Lloyd, 1972a, 1972b; Lu & Huang, 2006
千里光属 Senecio	雌全同株 Gynomonoecy		Abbott & Schmitt, 1985; Lu & Huang, 2006
一枝黄花属 Solidago	雌全同株 Gynomonoecy		Bertin & Gwisc, 2002; Lu & Huang, 2006
刺苞果属 Acanthospermum, 蓍属 Achillea,腺梗菜属 Adenocaulon,异芒菊属 Blainvillea, 金盏花属 Cal- endula, 翠菊属 Callistephus, 刺冠菊属 Calotis, 石胡荽属 Centipeda, 蕲艾属 Crossostephium,杯菊属 Cyathocline, 鱼眼草属 Dichrocephala, 多榔菊属 Doronicum, 鳢肠属 Eclipta, 沼菊属 Enydra, 菊芹属 Erechtites,飞蓬属 Erigeron, 絮菊属 Filago, 大丁草属 Gerbera, 香茹属 Glossogyne, 鼠麴草属 Gnaphalium, 田基黄属 Grangea,蜡菊属 Helichrysum, 狗娃花属 Heteropappus,小舌菊属 Microglossa, 齿冠草属 Myriactis, 银胶菊属 Parthenium, 瓜叶菊属 Pericallis,棉毛草属 Phagnalon, 秋分草属 Rhynchospermum, 虾须草属 Sheareria, 豨莶属 Siegesbeckia, 裸柱菊属 Soliva, 戴星草属 Sphaeranthus, 合苞菊属 Symphyllocarpus, 歧伞菊属 Thespis, 款冬属 Tussilago	雌全同株 Gynomonoecy		Yampolsky & Yampolsky, 1922; Lu & Huang, 2006
拟艾脑属 Blumeopsis, 苇谷草属 Pentanema	雌全同株 Gynomonoecy		Lu & Huang, 2006; Torices & Anderberg, 2009
橐吾属 Ligularia	雌全同株 Gynomonoecy		Lu & Huang, 2006; Zhang et al., 2012
亚菊属 Ajania, 画笔菊属 Ajaniopsis, 香青属 Anaphalis, 春黄菊属 Anthemis, 莎菀属 Arctogeron, 木筒蒿属 Argyranthemum, 蒿属 Artemisia, 紫菀木属 Asterothamnus, 雏菊属 Bellis, 短舌菊属 Bra- chanthemum, 短星菊属 Brachyactis, 葶菊属 Cavea, 果香菊属 Chamaemelum, 茼蒿属 Chrysanthemum, 鞘冠菊属 Coleostephus, 金鸡菊属 Coreopsis, 垂头菊属 Cremanthodium, 大丽花属 Dahlia, 菊属 Dendranthema, 东风菜属 Doellingeria, 鹅不食草属 Epaltes, 大吴风	雌全同株 Gynomonoecy		Lu & Huang, 2006
分类群 Taxon		性系统类型 Type of sexual system		参考文献 Reference
草属 Farfugium, 线叶菊属 Filifolium, 复芒菊属 Formania, 天人菊属 Ga- illardia, 牛膝菊属 Galinsoga, 裸菀属 Gymnaster, 异裂菊属 Heteroplexis, 马兰属 Kalimeris, 花花紫属 Karelinia, 喀什菊属 Kaschgaria, 岩菊属 Krylovia, 瓶头菊属 Lagenophora, 滨菊属 Leucanthemum, 母菊属Matricaria, 羽叶千里光属 Nemosenecio, 栉叶蒿属 Neopallasia, 太行菊属 Opisthop- appus, 蜂斗菜属 Petasites, 寒莲属 Psychrogeton, 小黄菊属 Pyrethrum, 金光菊属 Rudbeckia, 华蟹甲草属 Sinacalia, 蒲儿根属 Sinosenecio, 金钮扣属 Spilanthes, 金腰箭属 Synedrella, 合耳菊属 Synotis, 万寿菊属 Tagetes, 菊蒿属 Tanacetum, 狗舌草属 Tephroseris, 肿柄菊属 Tithonia, 羽芒菊属 Tridax, 三肋果属 Tripleurospermum, 碱菀属 Tripolium, 女菀属 Turcza- ninovia, 扁芒菊属 Waldheimia
桔梗科 Campanulaceae 半边莲属 Lobelia		雌全同株 Gynomonoecy		Molano-flores & Danderson, 2009
忍冬科 Caprifoliaceae (包括败酱科 Valerianaceae) 缬草属 Valeriana		雌全同株 Gynomonoecy		Lu & Huang, 2006
缬草属 Valeriana		雌全同株、雌全异株 Gynomonoecy, gynodioecy		Duarte-Silva et al., 2010

分类群 Taxon	性系统类型 Type of sexual system		参考文献 Reference
短蕊花科 Lactoridaceae Lactoris	雌全同株 Gynomonoecy		Bernardello et al., 1999
禾本科 Gramineae 小丽草属 Coelachne	雌全同株 Gynomonoecy		Yampolsky & Yampolsky, 1922; Lu & Huang, 2006
Tripsacum, Aulonemia	雌全同株 Gynomonoecy		Dewald et al., 1987; Grombone-Guaratini et al., 2011
刺叶树科 Xanthorrhoeaceae 独尾草属 Eremurus	雌全同株、两性花植株 Gynomonoecy, hermaphroditism		Mamut et al., 2014
天南星科 Araceae 麒麟尾属 Epipremnum, Heteropsis, 崖角藤属 Rhaphidophora, Rhodospatha	雌全同株 Gynomonoecy		Yampolsky & Yampolsky, 1922; Lu & Huang, 2006
洋二仙草科 Gunneraceae Gunnera	雌全同株 Gynomonoecy		Palkovic, 1978; Lowrey & Robinson, 1988
胡桃科 Juglandaceae 马尾树属 Rhoiptelea	雌全同株 Gynomonoecy		Sun et al., 2006; Lu & Huang, 2006
荨麻科 Urticaceae 墙草属 Parietaria	雌全同株 Gynomonoecy		Franchi et al., 2007
桃金娘科 Myrtaceae 白千层属 Melaleuca	雌全同株、雌全异株 Gynomonoecy, gynodioecy		Barlow & Forrester, 1984
柳叶菜科 Onagraceae 倒挂金钟属 Fuchsia	雌全同株 Gynomonoecy		Lu & Huang, 2006
牻牛儿苗科 Geraniaceae 牻牛儿苗属 Erodium, 老鹳草属 Geranium	雌全同株 Gynomonoecy		Lu & Huang, 2006
景天科 Crassulaceae 八宝属 Hylotelephium, 景天属 Sedum	雌全同株 Gynomonoecy		Lu & Huang, 2006
石竹科 Caryophyllaceae 蝇子草属 Silene	雌全同株 Gynomonoecy		Hermanutz & Innes, 1994; Maurice, 1999; Lu & Huang, 2006
蝇子草属 Silene,石竹属 Dianthus,繁缕属 Stellaria	雌全同株、雌全异株Gynomonoecy, gynodioecy		Maurice, 1999; Collin et al., 2009; Dufay et al., 2010 ; Casimiro-Soriguer et al., 2013
麦仙翁属 Agrostemma, 蚤缀属 Arenaria, 卷耳属 Cerastium, 石竹属 Dianthus, 剪秋罗属 Lychnis, 高山漆姑草属 Minuartia,漆姑草属 Sagina, 牛漆姑草属 Spergularia,繁缕属 Stellaria	雌全同株 Gynomonoecy		Lu & Huang, 2006
苋科 Amaranthaceae (包括藜科 Chenopodiaceae) Arthrocnemum, Didymantlius, Heterostachys, Microcnemum, Monolepis, Osteocarpum, Spirostachys, Secticornia, Threlkeldia	雌全同株 Gynomonoecy		Yampolsky & Yampolsky, 1922
藜属 Chenopodium	雌全同株 Gynomonoecy		Bhargava et al., 2007
盐节木属 Halocnemum, 盐千屈菜属 Halopeplis, 盐爪爪属 Kalidium, 盐角草属 Salicornia, 碱蓬属 Suaeda	雌全同株 Gynomonoecy		Yampolsky & Yampolsky, 1922; Lu & Huang, 2006
刺藜属 Dysphania, 棉藜属 Kirilowia, 地肤属 Kochia, 绒藜属 Londesia, 兜藜属 Panderia	雌全同株 Gynomonoecy		Lu & Huang, 2006
菠菜属 Spinacia	雌株、雄株、雌全同株、雄全同株 Female, male, gynomonoecy, andromonoecy		Komai & Masuda, 2004; Onodera et al., 2008
花荵科 Polemoniaceae 花荵属 Polemonium	雌全同株 Gynomonoecy		Yampolsky & Yampolsky, 1922; Lu & Huang, 2006
山榄科 Sapotaceae 山榄属 Planchonella	雌全同株 Gynomonoecy		Méndez & Munzinger,2010
龙胆科 Gentianaceae 假龙胆属 Gentianella	雌全同株 Gynomonoecy		Pringle, 2011
分类群 Taxon	性系统类型 Type of sexual system		参考文献 Reference
爵床科 Acanthaceae Brachystephanus	雌全同株 Gynomonoecy		Champluvier, 1997
木犀科 Oleaceae 木犀榄属 Olea, 木犀属 Osmanthus, 丁香属 Syringa	雌全同株 Gynomonoecy		Lu & Huang, 2006
唇形科 Lamiaceae 新风轮属 Calamintha,青兰属 Dracocephalum, 活血丹属 Glechoma,神香草属 Hyssopus, 地笋属 Lycopus, 蜜蜂花属 Melissa, 薄荷属 Mentha,荆芥属 Nepeta, 牛至属 Origanum, 夏枯草属 Prunella, 鼠尾草属 Salvia,水苏属 Stachys, 百里香属 Thymus	雌全同株 Gynomonoecy		Lu & Huang, 2006
车前草科 Plantaginaceae 车前草属 Plantago	雌全同株 Gynomonoecy		Lu & Huang, 2006
菊科 Asteraceae Abrotanella, Acamptopappus, Achaetogeron, Achmophora, Achyrachaena, Achyrocline, Adelostigma, Aganippea, Aiolotheca, Allendea, Aniauria, Am- blyocarpum, Amblyopappus, Amellus, Amphidoxa, Amphiglossa, Amphori- carpus, Anaglypha, Anaxeton, Antithrixia, Aphanactis, Aphanostephus, Agyroxiphium, Arnica, Arnicastrum, Arrowsmithia, Asteropsis, Astephania, Asteromoea, Athrixia, Athroisma, Axiniphyllum, Baeria, Balleya, Balsam- orhiza, Bartlettia, Bellida, Bellium, Bellis, Berlandiera, Bi-Leveillea, Blennosperma, Blepharispermum, Blepharizonia, Boltonia, Borrichia, Br- achychaeta, Brachyclados, Brachycome, Brachyglottis, Bradburia, Bryo- morphe, Burrielia, Cacosmia, Cadiscus, Calyptrocarpus, Cassinia, Cel- misia, Ceratogyne, Ceruana, Chaetanthira, Chaetopappa, Chaptalia, Chardinia, Charieis, Chiliophyllum, Chiliotrichium, Chionolaena, 被菊属 Chlamydites, Chroilema, Chromolepis, Chrysactinia, Chrysanthellum, Chrysogonum, Chrysoma, Chrysopsis, Clappia, Clibadium, Cnicothamnus, Coleocoma, Commidendron, Corethrogyne,秋英属 Cosmos, Crocidium, Crupina, Cylindrocline, Cypselodontia, Delamerea, Denekia, Dichaeto- phora, Dicranocarpus, Dimorphocoma, Dimorphotheca, Diplostephium, Dipterocome, Disparago, Dugesia, Egletes, Elachanthus, Elvira, Engel- mannia, Epallage, Eremothamnus, Eriachaenium, Ericameria, Erioce- phalus, Eriothrix, Ermeiastrum, Espeletia,梳黄菊属 Euryops, Eutetras, Evax, Faceles, Faujasia, Felicia, Fresenia, Gamolepis, Garcilassa, Garuleum, Geigeria, Glossocardia, Glycideras, Goldmanella, Golionema, Grantia, Gratwickia, Greenella, Grypocarpha, Guardiola,小葵子属 Guizotia, Gundlachia, Gundelia, Gutierrezia, Gymnarrhena, Gymnodiscus, Gymnosperma, Gymnostephium, Haastia, Hazardia, Heliopsis, Hemi- zonella, Hemizonia, Henricia, Heptanthus, Hertia, Heterolepis, Heteros- permum, Heterotheca, Hidalgoa, Hinterhubera, Hippia, Homochaete, Homochroma, Homogyne, Hysterionica, Ichthyothere, Inulopsis, Ischnea, Isocarpha, Isoetopsis, Isostigma, Ixiolaena, Jaegena, Jaegeria, Jasonia, Keerlia, Koehneola, Lachnophyllum, Laestadia,瓶头草属 Lagenophora, Lagophylla, Lantanopsis, Lasiocoma, Lasiopogon, Lasthenia, Lecocarpus, Leontonyx, Lepidophyllum, Lepidostephium, Leptocarpha, Leptorhynchus, Lessingia, Leucopholis, Leyssera, Liabum, Lindheimera, Lipochaeta, Lucilia, Macowania, Madia, Mairia, Melalema, Melampodium, Mela- nodendron, Melanthera, Merrittia, Micractis, Microlecane, Micropsis, Micropus, Microtrichia, Milleria, Minuria, Minurothamnus, Monactis, Monarrhenus, Monolopia, Monoptilon, Moonia, Montanoa,帚菊木属 Mutisia, Nananthea,毛冠菊属 Nannoglottis, Nanothamnus, Narvalina, Nerrittia, Nestlera, Nicolasia, Nidorella, Nolletia, Odontospermum, Oedera, Olearia, Oligocarpus, Olivaea, Onoseris, Oreochrysum, Osmites, Osmitopsis, Osteospermum, Othonna, Otochlamys, Oxypappus, Pach- ylaena, Pachyrhynchus, Parthenice, Parantennaria, Pectis, Pentachaeta, Perityle, Perralderia, Perymenium, Petalacte, Petalactella, Petrollinia, Phacellothrix, Phaenocoma, Phalacrocarpum, Phemmera, Philactis, Philoglossa, Philyrophyllum, Phymaspermum, Pinillosia, Plagiocheilus, Plazia, Pleurophyllum, Podochaenium, Podocoma, Podolepis, Polygyne, Polymnia, Postia, Printzia, Psiadia, Psila, Psilactis, Psilocarphus, Pterigeron, Raillardella, Raoulia, Relhania, Remya, Rhantherium, Rhodogeron, Riddellia, Riencourtia, Rigiopappus, Rochonia, Ruckeria, Rumfordia, Sabazia, Sanvitalia, Sartwellia, Schistocarpha, Schist- ostephium, Schizoptera, Schizotrichia, Schkuhria, Schoenia, Sclerocarpus, Scyphocoronis, Selloa, Sericocarpus, Siebera, Silphium, Sommerfeltia, Sphacophyllum, Sphaeromorphaea, Sphagneticola, Spilanthus, Stachy- cephalum, Staurochlamys, Steirodiscus, Stemmatella, Stenocline, Stephan- odoria, Stilpnogyne, Stuartina, Styloclyne, Syncephalantha, Syntrichop- appus, Tessaria, Tetracanthus, Tetragonotheca, Tetramolopium,	雌全同株 Gynomonoecy		Yampolsky & Yampolsky, 1922
分类群 Taxon	性系统类型 Type of sexual system		参考文献 Reference
Tetranthus, Tetraperone, Thelesperma, Thespidium, Thymopsis, Tolbonia,地菊属 Townsendia, Tragoceros, Trichocline, Trigonospermum, Triploce- phalum, Tripteris, Urbinella, Vanclevea, Varilla, Venegazia, Vieraea, Villanova, Vittadinia, Waitzia, Welwitschiella, Whitneya, Wyomingia, Xanthisma,旱花属 Xeranthemum, Zaluzania, Zexmenia, Zoegea
Lasianthaea, Liabum,羽芒菊属 Tridax, Verbesina	雌全同株 Gynomonoecy		Bullock, 1985
Baltimora,鬼针草属 Bidens,鳢肠属 Eclipta	雌全同株 Gynomonoecy		Bullock, 1985; Yampolsky & Yampolsky, 1922
Ambylocarpum, Asteriscus, Chrysophthalmum, Cylindrocline, Dittrichia, Doellia, Duhaldea, Geigeria, Ighermia, Limbarda, Pseudoconyza, Rhan- terium, Schizogyne, Streptoglossa, Telekia, Vierea, Xerolekia, Zoutpan- sbergia	雌全同株 Gynomonoecy		Torices & Anderberg, 2009
Callilepis, Calostephane, Lifago, Mollera, Ondetia, Pallenis, Pechuel- L?schea, Porphyrostemma, Rosenia, Sachsia, Stenachaenium	雌全同株 Gynomonoecy		Yampolsky & Yampolsky, 1922; Torices & Anderberg, 2009
Ifloga	雌全同株 Gynomonoecy		Yampolsky & Yampolsky, 1922; Bergh & Verboom, 2011
假莲属 Conyza	雌全同株 Gynomonoecy		Yampolsky & Yampolsky, 1922; Urdampilleta et al., 2005; Lu & Huang, 2006
旋覆花属 Inula, 山黄菊属 Anisopappus, 牛眼菊属 Buphthalmum, 天门精属 Carpesium, 六棱菊属 Laggera, 蚤草属 Pulicaria	雌全同株 Gynomonoecy		Yampolsky & Yampolsky, 1922; Lu & Huang, 2006; Torices & Anderberg, 2009
艾纳香属 Blumea, 阔苞菊属 Pluchea	雌全同株 Gynomonoecy		Lu & Huang, 2006; Torices & Anderberg, 2009
蟛蜞菊属 Wedelia, 百日菊属 Zinnia	雌全同株 Gynomonoecy		Yampolsky & Yampolsky, 1922; Bullock, 1985; Lu & Huang, 2006
紫菀属 Aster	雌全同株 Gynomonoecy		Bertin & Kerwin, 1998; Lu & Huang, 2006
山芫荽属 Cotula	两性花植株、雌全同株、雌雄同株、亚雌雄异株、雌雄异株Hermaphroditism, gynomonoecy, monoecy, subdioecy, dioecy		Lloyd, 1972a, 1972b; Lu & Huang, 2006
千里光属 Senecio	雌全同株 Gynomonoecy		Abbott & Schmitt, 1985; Lu & Huang, 2006
一枝黄花属 Solidago	雌全同株 Gynomonoecy		Bertin & Gwisc, 2002; Lu & Huang, 2006
刺苞果属 Acanthospermum, 蓍属 Achillea,腺梗菜属 Adenocaulon,异芒菊属 Blainvillea, 金盏花属 Cal- endula, 翠菊属 Callistephus, 刺冠菊属 Calotis, 石胡荽属 Centipeda, 蕲艾属 Crossostephium,杯菊属 Cyathocline, 鱼眼草属 Dichrocephala, 多榔菊属 Doronicum, 鳢肠属 Eclipta, 沼菊属 Enydra, 菊芹属 Erechtites,飞蓬属 Erigeron, 絮菊属 Filago, 大丁草属 Gerbera, 香茹属 Glossogyne, 鼠麴草属 Gnaphalium, 田基黄属 Grangea,蜡菊属 Helichrysum, 狗娃花属 Heteropappus,小舌菊属 Microglossa, 齿冠草属 Myriactis, 银胶菊属 Parthenium, 瓜叶菊属 Pericallis,棉毛草属 Phagnalon, 秋分草属 Rhynchospermum, 虾须草属 Sheareria, 豨莶属 Siegesbeckia, 裸柱菊属 Soliva, 戴星草属 Sphaeranthus, 合苞菊属 Symphyllocarpus, 歧伞菊属 Thespis, 款冬属 Tussilago	雌全同株 Gynomonoecy		Yampolsky & Yampolsky, 1922; Lu & Huang, 2006
拟艾脑属 Blumeopsis, 苇谷草属 Pentanema	雌全同株 Gynomonoecy		Lu & Huang, 2006; Torices & Anderberg, 2009
橐吾属 Ligularia	雌全同株 Gynomonoecy		Lu & Huang, 2006; Zhang et al., 2012
亚菊属 Ajania, 画笔菊属 Ajaniopsis, 香青属 Anaphalis, 春黄菊属 Anthemis, 莎菀属 Arctogeron, 木筒蒿属 Argyranthemum, 蒿属 Artemisia, 紫菀木属 Asterothamnus, 雏菊属 Bellis, 短舌菊属 Bra- chanthemum, 短星菊属 Brachyactis, 葶菊属 Cavea, 果香菊属 Chamaemelum, 茼蒿属 Chrysanthemum, 鞘冠菊属 Coleostephus, 金鸡菊属 Coreopsis, 垂头菊属 Cremanthodium, 大丽花属 Dahlia, 菊属 Dendranthema, 东风菜属 Doellingeria, 鹅不食草属 Epaltes, 大吴风	雌全同株 Gynomonoecy		Lu & Huang, 2006
分类群 Taxon		性系统类型 Type of sexual system		参考文献 Reference
草属 Farfugium, 线叶菊属 Filifolium, 复芒菊属 Formania, 天人菊属 Ga- illardia, 牛膝菊属 Galinsoga, 裸菀属 Gymnaster, 异裂菊属 Heteroplexis, 马兰属 Kalimeris, 花花紫属 Karelinia, 喀什菊属 Kaschgaria, 岩菊属 Krylovia, 瓶头菊属 Lagenophora, 滨菊属 Leucanthemum, 母菊属Matricaria, 羽叶千里光属 Nemosenecio, 栉叶蒿属 Neopallasia, 太行菊属 Opisthop- appus, 蜂斗菜属 Petasites, 寒莲属 Psychrogeton, 小黄菊属 Pyrethrum, 金光菊属 Rudbeckia, 华蟹甲草属 Sinacalia, 蒲儿根属 Sinosenecio, 金钮扣属 Spilanthes, 金腰箭属 Synedrella, 合耳菊属 Synotis, 万寿菊属 Tagetes, 菊蒿属 Tanacetum, 狗舌草属 Tephroseris, 肿柄菊属 Tithonia, 羽芒菊属 Tridax, 三肋果属 Tripleurospermum, 碱菀属 Tripolium, 女菀属 Turcza- ninovia, 扁芒菊属 Waldheimia
桔梗科 Campanulaceae 半边莲属 Lobelia		雌全同株 Gynomonoecy		Molano-flores & Danderson, 2009
忍冬科 Caprifoliaceae (包括败酱科 Valerianaceae) 缬草属 Valeriana		雌全同株 Gynomonoecy		Lu & Huang, 2006
缬草属 Valeriana		雌全同株、雌全异株 Gynomonoecy, gynodioecy		Duarte-Silva et al., 2010