被子植物翅果的多样性及演化

doi:10.17521/cjpe.2018.0053

摘要/Abstract

摘要：

翅果能够依靠风力进行传播, 可能是被子植物快速散布和物种分化的一个重要因素。狭义的翅果是指果皮延伸成翅且不开裂的干果; 广义的翅果则包括果皮、花被片或苞片形成果翅的所有果实。根据果翅形态及其生长方式的不同, 广义的翅果可分为单侧翅果、周位翅果(圆翅果与蝶翅果)、棱翅果、披针翅果、翼状萼翅果、叶状苞翅果6种类型, 其空中运动方式有自旋式(单侧翅果、翼状萼翅果)、波浪式(周位翅果、叶状苞翅果)、翻滚自旋式(周位翅果)、直升机式(披针翅果、翼状萼翅果)和滚筒式(棱翅果)。棱翅果与圆翅果在被子植物基部类群樟目就有发生, 并同时出现在单子叶植物和双子叶植物中, 可能是最早出现的翅果类型。翅果的演化过程呈现出果翅数量增加、果翅偏向单侧和果翅负荷(果实质量与果翅面积之比)降低的趋势, 以利于适应较小的风并增加传播距离。果翅除了促进果实与种子的风力传播外, 还具有物理防御、调节种子萌发和促进二次传播等作用。泛热带分布的金虎尾科有着极其丰富的翅果类型, 与其多次跨洋长距离扩散密切相关, 可以作为研究翅果适应与演化的一个模式类群。结合生态和演化-发育生物学方法, 研究不同类型翅果在适应风力传播方面的差异、萼片或苞片发育成翅的分子与遗传机制、翅果不同类型的演化历史及其对被子植物物种多样性的影响等是今后值得探讨的重要问题。

关键词: 翅果, 长距离扩散, 金虎尾路线, 适应演化

Abstract:

Samara (winged fruit) can be dispersed easily by wind and may be a crucial factor for angiosperm spread and diversification. In a narrow sense, a samara is an indehiscent dry fruit with wing(s) developed from fruit pericarp, while in a broad sense samaras also include all winged fruits with wings developed from both pericarp and perianth or bracts. According to the wing shape and growth patterns of samaras, we divided samaras into six types, i.e. single-winged, lanceolate-winged, rib-winged, sepal-winged, bract-winged, and perigynous samaras. Perigynous samaras can be further classified into two forms, i.e. round-winged and butterfly-winged samaras. Accordingly, the aerodynamic behavior of samaras can be classified into five types, autogyro, rolling autogyro, undulator, helicopter, and tumbler. The rib-winged and round-winged samaras can be found in Laurales, a basal angiosperm, and may represent the primitive type of early samaras. In the derived clades, samaras evolved enlarged but unequal wings and decreased wing loading (the ratio of fruit weight to wing size), which is likely an adaptation to gentle wind and secondary dispersal through water or ground wind. The wings of some samaras (such as sepal-winged and bract-winged samaras) may have multiple functions including wind dispersal, physical defense for the seeds, and adjust seed germination strategy. The pantropical family Malpighiaceae is extraordinarily rich in samara types, which is likely related to its multiple inter-continent dispersal in history, which is known as “Malpighiaceae Route”. Therefore, Malpighiaceae can be used as a model system for the studies on samara adaptation and evolution. We identified the following issues that deserve further examination in future studies using both ecological and evo-devo methods: 1) the adaption of different types of samaras in dispersal processes, 2) the molecular and developmental mechanism of sepal- and bract-wings, and 3) the evolution of samara types and their effects on angiosperm diversification.

Key words: samara, long-distance dispersal, Malpighiaceae Route, adaptive evolution

谭珂, 董书鹏, 卢涛, 张亚婧, 徐诗涛, 任明迅. 被子植物翅果的多样性及演化. 植物生态学报, 2018, 42(8): 806-817. DOI: 10.17521/cjpe.2018.0053

TAN Ke, DONG Shu-Peng, LU Tao, ZHANG Ya-Jing, XU Shi-Tao, REN Ming-Xun. Diversity and evolution of samara in angiosperm. Chinese Journal of Plant Ecology, 2018, 42(8): 806-817. DOI: 10.17521/cjpe.2018.0053

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https://www.plant-ecology.com/CN/Y2018/V42/I8/806

图/表 5

参考文献 63

[1]	Al-Mayah AA, Stace CA ( 1989). New species, names, and combinations in American Combretaceae. Annals of the Missouri Botanical Garden, 76, 1125-1128. DOI URL
[2]	Al-Shehbaz IA ( 1986). The genera of Lepidieae(Cruciferae; Brassicaceae) in the southeastern United States. Journal of the Arnold Arboretum, 67, 265-311. DOI URL
[3]	Anderson WR, Anderson C, Davis CC ( 2006). Malpighiaceae. . Cited: 2018-03-01.
[4]	Augspurger CK ( 1986). Morphology and dispersal potential of wind-dispersed diaspores of neotropical trees. American Journal of Botany, 73, 353-363. DOI URL
[5]	Augspurger CK ( 1988). Mass allocation, moisture content, and dispersal capacity of wind-dispersed tropical diaspores. New Phytologist, 108, 357-368. DOI URL
[6]	Baskin JM, Lu JJ, Baskin CC, Tan DY, Wang L ( 2014). Diaspore dispersal ability and degree of dormancy in heteromorphic species of cold deserts of northwest China: A review. Perspectives in Plant Ecology, Evolution and Systematics, 16, 93-99. DOI URL
[7]	Burrows CJ ( 1994). Fruit types and seed dispersal modes of woody plants in Ahuriri summit Bush, Port Hills, western Banks Peninsula, Canterbury, New Zealand. New Zealand Journal of Botany, 32, 169-181. DOI URL
[8]	Burrows FM ( 1975). Wind-borne seed and fruit movement. New Phytologist, 75, 405-418. DOI URL
[9]	Chang MC, Qiu LQ, Wei Z, Green PS ( 1996). Oleaceae. In: Wu ZY, Raven PH, Hong DY eds. Flora of China. Vol. 15. Science Press & Missouri Botanical Garden Press, Beijing.
[10]	Chen SL, Gilbert MG ( 1994). Verbenaceae. In: Wu ZY, Raven PH, Hong DY eds. Flora of China. Vol. 17. Science Press & Missouri Botanical Garden Press, Beijing.
[11]	Corlett RT ( 2009). Seed dispersal distances and plant migration potential in tropical East Asia. Biotropica, 41, 592-598. DOI URL
[12]	Cremer KW ( 1977). Distance of seed dispersal in Eucalyptus estimated from seed weights. Australian Forest Research, 7, 225-228.
[13]	Dallwitz MJ, Paine TA, Zurcher EJ ( 2000). Principles of interactive keys. . Cited: 2018-03-01.
[14]	Davis CC, Anderson WR ( 2010). A complete generic phylogeny of Malpighiaceae inferred from nucleotide sequence data and morphology. American Journal of Botany, 97, 2031-2048. DOI URL PMID
[15]	Davis CC, Bell CD, Mathews S, Donoghue MJ ( 2002). Laurasian migration explains Gondwanan disjunctions: Evidence from Malpighiaceae. Proceedings of the National Academy of Sciences of the United States of America, 99, 6833-6837. DOI URL PMID
[16]	Davis CC, Schaefer H, Xi ZX, Baum DA, Donoghue MJ, Harmon LJ ( 2014). Long-term morphological stasis maintained by a plant-pollinator mutualism. Proceedings of the National Academy of Sciences of the United States of America, 111, 5914-5919. DOI URL PMID
[17]	Davis CC, Anderson WR, Donoghue MJ ( 2001). Phylogeny of Malpighiaceae: Evidence from chloroplast ndhF and trnL-F nucleotide sequences. American Journal of Botany, 88, 1830-1846. DOI URL PMID
[18]	Delgado JA, Jimenez MD, Gomez A ( 2009). Samara size versus dispersal and seedling establishment in Ailanthus altissima(Miller) Swingle. Journal of Environmental Biology, 30, 183-186. URL PMID
[19]	Deng LL, Wei KY, Fan GS ( 2003). A new species of Acer from Yunnan. Acta Botanica Yunnanica, 25, 197-198. DOI URL
	[ 邓莉兰, 魏开云, 樊国盛 ( 2003). 云南槭属植物一新种. 云南植物研究, 25, 197-198.] DOI URL
[20]	Eriksson O, Bremer B ( 1992). Pollination systems, dispersal modes, life forms, and diversification rates in angiosperm families. Evolution, 46, 258-266. DOI URL PMID
[21]	Eriksson O, Friis EM, Löfgren P ( 2000). Seed size, fruit size, and dispersal systems in angiosperms from the Early Cretaceous to the Late Tertiary. The American Naturalist, 156, 47-58. DOI URL PMID
[22]	Fang RC, Staples G ( 1995). Convolvulaceae. In: Wu ZY, Raven PH, Hong DY eds. Flora of China. Vol. 16. Science Press & Missouri Botanical Garden Press, Beijing.
[23]	Friis EM, Crane PR, Pedersen KR ( 2011). Early Flowers and Angiosperm Evolution. Cambridge University Press. Cambridge, UK.
[24]	Green DS ( 1980). The terminal velocity and dispersal of spinning samaras. American Journal of Botany, 67, 1218-1224. DOI URL
[25]	Greene DF, Johnson EA ( 1990). The dispersal of winged fruits and seeds differing in autorotative behaviour. Canadian Journal of Botany, 68, 2693-2697. DOI URL
[26]	Gunasekara N ( 2004). Phylogenetic and Molecular Dating Analyses of The Tropical Tree Family Dipterocarpaceae Based on Chloroplast matK Nucleotide Sequence Data. Master degree dissertation, Concordia University, Montreal.
[27]	Harris JG, Harris MW ( 2001). Plant Identification Terminology: An Illustrated Glossary. Spring Lake Publishing, Spring Lake, Utah.
[28]	Herrera CM ( 1989). Seed dispersal by animals: A role in angiosperm diversification? The American Naturalist, 133, 309-322. DOI URL
[29]	Hong DY, Barrie FR, Bell CD ( 2011). Valerianaceae. In: Wu ZY, Raven PH, Hong DY eds. Flora of China. Vol. 19. Science Press & Missouri Botanical Garden Press, Beijing.
[30]	Huang YJ, Zhu H, Chen WY, Zhou ZK ( 2013). Intraspecific variation in samara morphology of Acer and its implication in fossil identification. Plant Diversity and Resources, 35, 295-302. DOI URL
	[ 黄永江, 朱海, 陈文允, 周浙昆 ( 2013). 槭属翅果种内形态变异性及对其化石鉴定的意义. 植物分类与资源学报, 35, 295-302.] DOI URL
[31]	Jurado E, Westoby M, Nelson D ( 1991). Diaspore weight, dispersal, growth form and perenniality of Central Australian plants. Journal of Ecology, 79, 811-830. DOI URL
[32]	Lansdown RV ( 2006). The genus Callitriche(Callitrichaceae) in Asia. Novon, 16, 354-361. DOI URL
[33]	Li AR, Bao BJ, Grabovskaya-Borodina AE, Hong SP, McNeill J, Mosyakin SL, Ohba H, Park CW (2003). Polygonaceae. In: Wu ZY, Raven PH, Hong DY eds. Flora of China. Vol. 5. Science Press & Missouri Botanical Garden Press, Beijing.
[34]	Li PQ, Skvortsov AK ( 1999). Betulaceae. In: Wu ZY, Raven PH, Hong DY eds. Flora of China. Vol. 4. Science Press & Missouri Botanical Garden Press, Beijing.
[35]	Li XW, Li J, Ashton PS ( 2007). Dipterocarpaceae. In: Wu ZY, Raven PH, Hong DY eds. Flora of China. Vol. 13. Science Press & Missouri Botanical Garden Press, Beijing.
[36]	Lord J, Egan J, Clifford T, Jurado E, Leishman M, Williams D, Westoby M ( 1997). Larger seeds in tropical floras: Consistent patterns independent of growth form and dispersal mode. Journal of Biogeography, 24, 205-211. DOI URL
[37]	Lu AM, Stone DE, Grauke LJ ( 1999). Juglandaceae. In: Wu ZY, Raven PH, Hong DY eds. Flora of China. Vol. 4. Science Press & Missouri Botanical Garden Press, Beijing.
[38]	Ma JS, Zhang ZX, Liu QR, Peng H, Funston AM ( 2008). Celastraceae. In: Wu ZY, Raven PH, Hong DY eds. Flora of China. Vol. 11. Science Press & Missouri Botanical Garden Press, Beijing.
[39]	Manchester SR, O’Leary EL ( 2010). Phylogenetic distribution and identification of fin-winged fruits. The Botanical Review, 76, 1-82. DOI URL
[40]	Norberg RA ( 1973). Autorotation, self-stability, and structure of single-winged fruits and seeds (samaras) with comparative remarks on animal flight. Biological Review, 48, 561-596. DOI URL
[41]	Peck CJ, Lersten NR ( 1991). Samara development of black maple (Acer saccharum ssp. nigrum) with emphasis on the wing. Canadian Journal of Botany, 69, 1349-1360.
[42]	Planchuelo G, Catalán P, Delgado JA ( 2016). Gone with the wind and the stream: Dispersal in the invasive species Ailanthus altissima. Acta Oecologica, 73, 31-37.
[43]	Qian ZN, Ren MX ( 2016). Floral evolution and pollination shifts of the “Malpighiaceae route” taxa, a classical model for biogeographical study. Biodiversity Science, 24, 95-101. DOI URL
	[ 钱贞娜, 任明迅 ( 2016). “金虎尾路线”植物的花进化与传粉转变. 生物多样性, 24, 95-101.] DOI URL
[44]	Qin HN, Gilbert MG ( 2007). Elaeagnaceae. In: Wu ZY, Raven PH, Hong DY eds. Flora of China. Vol. 13. Science Press & Missouri Botanical Garden Press, Beijing.
[45]	Ren MX ( 2015). The upper reaches of the largest river in Southern China as an “evolutionary front” of tropical plants: Evidences from Asia-endemic genus Hiptage(Malpighiaceae). Collectanea Botanica, 34, e003. DOI: 10.3989/collectbot.2015.v34.003.
[46]	Seter D, Rosen A ( 1992). Study of the vertical autorotation of a single winged samara. Biological Reviews, 67, 175-197. DOI URL
[47]	Shang XL, Xu XZ, Fang SZ ( 2011). Seed dormancy mechanism of Cyclocarya paliurus. Scientia Silvae Sinica, 47(3), 68-74. DOI URL
	[ 尚旭岚, 徐锡增, 方升佐 ( 2011). 青钱柳种子休眠机制. 林业科学, 47(3), 68-74.] DOI URL
[48]	Shi Z, Chen YL, Chen YS, Lin YR, Liu SW, Ge XJ, Gao TG, Zhu SX, Liu Y, Yang QE, Humphries CJ, von Raab-Straube E, Gilbert MG, Nordenstam B, Kilian N, Brouillet L, Illarionova ID, Hind DJN, Jeffrey C, Bayer RJ, Kirschner J, Greuter W, Anderberg AA, Semple JC, Štepánek J, Freire SE, Martins L, Koyama H, Kawahara T, Vincent L, Sukhorukov AP, Mavrodiev EV, Gottschlich G (2011). Asteraceae. In: Wu ZY, Raven PH, Hong DY eds. Flora of China. Vol. 20-21. Science Press & Missouri Botanical Garden Press, Beijing.
[49]	Takeno K, Yamaguchi H ( 1991). Diversity in seed germination behavior in relation to heterocarpy in Salsola komarovii Iljin. The Botanical Magazine, 104, 207-215. DOI URL
[50]	Tanai T ( 1978). Taxonomical investigation of the living species of the genus Acer L., based on vein architecture of leaves. Journal of Faculty of Science, 18, 243-282.
[51]	Tanai T ( 1983). Revisions of tertiary Acer from East Asia. Journal of Faculty of Science, 20, 291-390.
[52]	Tang Y, Gilbert MG, Dorr LJ ( 2007). Tiliaceae. In: Wu ZY, Raven PH, Hong DY eds. Flora of China. Vol. 12. Science Press & Missouri Botanical Garden Press, Beijing.
[53]	The Angiosperm Phylogeny Group ( 2016). An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG IV. Botanical Journal of the Linnean Society, 181, 1-20. DOI URL
[54]	van der Niet T, Johnson SD ( 2012). Phylogenetic evidence for pollinator-driven diversification of angiosperms. Trends in Ecology & Evolution, 27, 353-361. DOI URL PMID
[55]	Wang J, Yang C, Liu SR ( 2002). A study of ecological significance of wind-rolling plants in steppe of middle reach of Xilin River, Inner Mongolia. Grassland of China, 24(6), 43-46. DOI URL
	[ 王静, 杨持, 刘书润 ( 2002). 锡林河中游草原风滚植物区系分析及生态意义. 中国草地, 24(6), 43-46.] DOI URL
[56]	Wang L, Xie SP, Liu KN, Wen WW, Zhang LR, Sun BN ( 2012). Late miocene samara of Fraxinus(Oleaceae) from the Lincang in Yunnan Province and its paleophytogeographical significance. Journal of Jilin University (Earth Science Edition), 42(suppl. 2), 331-342.
	[ 王磊, 解三平, 刘珂男, 温雯雯, 张利锐, 孙柏年 ( 2012). 云南临沧晚中新世梣属翅果化石及其古植物地理学意义. 吉林大学学报(地球科学版), 42(suppl. 2), 331-342.]
[57]	Wang YZ, Gereau R ( 2007). Ancistrocladaceae. In: Wu ZY, Raven PH, Hong DY eds. Flora of China. Vol. 13. Science Press & Missouri Botanical Garden Press, Beijing. DOI URL
[58]	Wei Y, Dong M, Huang ZY, Tan DY ( 2008). Factors influencing seed germination of Salsola affinis(Chenopodiaceae), a dominant annual halophyte inhabiting the deserts of Xinjiang, China. Flora, 203, 134-140. DOI URL
[59]	Willis JC ( Revised by Airy Shaw HK)( 1973). A Dictionary of the Flowering Plants and Ferns. 8th edn. Cambridge University Press, Cambridge, UK.
[60]	Xia NH, Gilbert MG ( 2003). Santalaceae. In: Wu ZY, Raven PH, Hong DY eds. Flora of China. Vol. 5. Science Press & Missouri Botanical Garden Press, Beijing.
[61]	Xu TZ, Chen YS, de Jong PC, Oterdoom HJ, Chang CS ( 2008). Aceraceae. In: Wu ZY, Raven PH, Hong DY eds. Flora of China. Vol. 11. Science Press & Missouri Botanical Garden Press, Beijing.
[62]	Yamaguchi H, Ichihara K, Takeno K, Hori Y, Saito T ( 1990). Diversities in morphological characteristics and seed germination behavior in fruits of Salsola komarovii Iljin. The Botanical Magazine, 103, 177-190. DOI URL
[63]	Zhang JQ, Wang LZ ( 1985). The geographic distribution of Dipterocarpaceae plants. Chinese Bulletin of Botany, 3, 1-8.
	[ 张金泉, 王兰州 ( 1985). 龙脑香科植物的地理分布. 植物学通报, 3, 1-8.]

翅果类型 Samara type	主要分布类群 Systematic distribution
1 单侧翅果 Single-winged samara (SWS)
1.1 单翅果 Mono-SWS	金虎尾科(异翅藤属, Acridocarpus, Banisteriopsis, Barnebya, Bronwenia, Cordobia, Cottsia, Dinemagonum, Diplopterys, Ectopopterys, Janusia, Peixotoa, Sphedamnocarpus, Stigmaphyllon), 木犀科陉属, 远志科蝉翼藤属, 鼠李科翼核果属, 榆科刺榆属, 梧桐科银叶树属 Malpighiaceae (Heteropterys, Acridocarpus, Banisteriopsis, Barnebya, Bronwenia, Cordobia, Cottsia, Dinemagonum, Diplopterys, Ectopopterys, Janusia, Peixotoa, Sphedamnocarpus, Stigmaphyllon), Fraxinus (Oleaceae), Securidaca (Polygalaceae), Ventilago (Rhamnaceae), Hemiptelea (Ulmaceae), Heritiera (Sterculiaceae)
1.2 双聚单翅果 Di-SWS	槭树科槭树属, 木犀科陉属 Acer (Aceraceae), Fraxinus (Oleaceae)
1.3 三聚单翅果 Tri-SWS	金虎尾科翅实藤属, 异翅藤属 Malpighiaceae (Ryssopterys, Heteropterys)
2 周位翅果 Perigynous samara
2.1 圆翅果 Round-winged samara	金虎尾科(盾翅藤属, Alicia, Amorimia, Calcicola, Caucanthus, Christianella, Diaspis, Excentradenia, Flabellaria, Madagasikaria, Malpighiodes, Mascagnia, Mezia), 胡桃科(青钱柳属, 化香树属), 豆科(紫檀属, 黄檀属), 十字花科(菘蓝属, 屈曲花属, 菥属), 桦木科(桤木属, 桦木属), 榆科(榆属, 青檀属), 蓼科山蓼属, 木樨科雪柳属, 槭树科金钱槭属, 苦木科臭椿属, 杜仲科杜仲属, 芸香科榆橘属, 蓝果树科喜树属, 莲叶桐科青藤属, 鼠李科马甲子属, 马尾树科马尾树属 Malpighiaceae (Aspidopterys, Alicia, Amorimia, Calcicola, Caucanthus, Christianella, Diaspis, Excentradenia, Flabellaria, Madagasikaria, Malpighiodes, Mascagnia, Mezia), Juglandaceae (Cyclocarya, Platycarya), Fabaceae (Pterocarpus, Dalbergia), Cruciferae (Isatis, Iberis, Thlaspi), Betulaceae (Alnus, Betula), Ulmaceae (Ulmus, Pteroceltis), Oxyria (Polygonaceae), Fontanesia (Oleaceae), Dipteronia (Aceraceae), Ailanthus (Simaroubaceae), Eucommia (Eucommiaceae), Ptelea (Rutaceae), Camptotheca (Nyssaceae), Illigera (Hernandiaceae), Paliurus (Rhamnaceae), Rhoiptelea (Rhoipteleaceae)
2.2 蝶翅果 Butterfly-winged samara	金虎尾科(Adelphia, Aenigmatanthera, Hiraea, Amorimia, Callaeum, Carolus, Christianella, Dinemandra等), 菊科(金鸡菊属, 偶雏菊属, 蟛蜞菊属), 木犀科(雪柳属, 六道木叶属), 使君子科(诃子属, 榆绿木属, 假榆橘属) Malpighiaceae (Adelphia, Christianella, Aenigmatanthera, Hiraea, Amorimia, Callaeum, Carolus, Dinemandra, etc.), Asterceae (Coreopsis, Boltonia, Sphagneticola), Oleaceae (Fontanesia, Abeliophyllum), Combretaeeae (Terminalia, Anogeissus, Pteleopsis)
3 棱翅果 Rib-winged samara	金虎尾科(Aspicarpa, Calcicola, Digoniopterys), 蓼科(翅果蓼属, 红药子属, 大黄属, 沙拐枣属, 等), 卫矛科(雷公藤属, 卫矛属, 等), 使君子科(风车子属, 诃子属), 无患子科(车桑子属, 黄梨木属), 椴树科(滇桐属, 蚬木属, 柄翅果属, 一担柴属), 鼠李科咀签属, 莲叶桐科青藤属, 茶茱萸科心翼果属, 薯蓣科薯蓣属, 藜科四翅滨藜, 胡颓子科翅果油树 Malpighiaceae (Aspicarpa, Calcicola, Digoniopterys), Polygonaceae (Parapteropyrum, Pteroxygonum, Rheum, Calligonum etc.), Celastraceae (Tripterygium, Euonymus etc.), Sapindaceae (Dodonaea, Boniodendron), Tiliaceae (Craigia, Excentrodendron, Burretiodendron, Colona), Gouania (Rhamnaceae), Illigera (Hernandiaceae), Cardiopteris (Icacinaceae), Dioscorea (Dioscoreaceae), Atriplex canescens (Chenopodiaceae), Elaeagnus mollis (Elaeagnaceae)
4 披针翅果 Lanceolate-winged samara	金虎尾科(风筝果属, 三星果属, Lophopterys, Dicella, Microsteira, Niedenzuella, Rhynchophora, Tetrapterys), 椴树科六翅木属, 卫矛科斜翼属, 十字花科沙芥属, 使君子科萼翅藤属 Malpighiaceae (Hiptage, Tristellateia, Lophopterys, Dicella, Microsteira, Niedenzuella, Rhynchophora, Tetrapterys), Berrya (Tiliaceae), Plagiopteron (Celastraceae), Pugionium (Crucificeae), Getonia (Combretaeeae)
5 翼状萼翅果 Sepal-winged samara	龙脑香科, 钩枝藤科钩枝藤属, 旋花科(飞蛾藤属, 三翅藤属, 白花叶属, 地旋花属) Dipterocarpaceae, Ancistrocladus (Ancistrocladaceae), Convolvulaceae (Dinetus, Tridynamia, Poranopsis, Xenostegia)
6 叶状苞翅果 Bract-winged samara	桦木科(鹅耳枥属, 铁木属, 桦木属, 桤木属, 榛属), 胡桃科(黄杞属, 枫杨属), 败酱科败酱属, 旋花科盾苞藤属, 马鞭草科楔翅藤属, 檀香科米面蓊属 Betulaceae (Carpinus, Ostrya, Betula, Alnus, Corylus), Juglandaceae (Engelhardia, Pterocarya), Patrinia (Valerianaceae), Neuropeltis (Convolvulaceae), Sphenodesme (Verbenaceae), Buckleya (Santalaceae)

翅果类型 Samara type	主要分布类群 Systematic distribution
1 单侧翅果 Single-winged samara (SWS)
1.1 单翅果 Mono-SWS	金虎尾科(异翅藤属, Acridocarpus, Banisteriopsis, Barnebya, Bronwenia, Cordobia, Cottsia, Dinemagonum, Diplopterys, Ectopopterys, Janusia, Peixotoa, Sphedamnocarpus, Stigmaphyllon), 木犀科陉属, 远志科蝉翼藤属, 鼠李科翼核果属, 榆科刺榆属, 梧桐科银叶树属 Malpighiaceae (Heteropterys, Acridocarpus, Banisteriopsis, Barnebya, Bronwenia, Cordobia, Cottsia, Dinemagonum, Diplopterys, Ectopopterys, Janusia, Peixotoa, Sphedamnocarpus, Stigmaphyllon), Fraxinus (Oleaceae), Securidaca (Polygalaceae), Ventilago (Rhamnaceae), Hemiptelea (Ulmaceae), Heritiera (Sterculiaceae)
1.2 双聚单翅果 Di-SWS	槭树科槭树属, 木犀科陉属 Acer (Aceraceae), Fraxinus (Oleaceae)
1.3 三聚单翅果 Tri-SWS	金虎尾科翅实藤属, 异翅藤属 Malpighiaceae (Ryssopterys, Heteropterys)
2 周位翅果 Perigynous samara
2.1 圆翅果 Round-winged samara	金虎尾科(盾翅藤属, Alicia, Amorimia, Calcicola, Caucanthus, Christianella, Diaspis, Excentradenia, Flabellaria, Madagasikaria, Malpighiodes, Mascagnia, Mezia), 胡桃科(青钱柳属, 化香树属), 豆科(紫檀属, 黄檀属), 十字花科(菘蓝属, 屈曲花属, 菥属), 桦木科(桤木属, 桦木属), 榆科(榆属, 青檀属), 蓼科山蓼属, 木樨科雪柳属, 槭树科金钱槭属, 苦木科臭椿属, 杜仲科杜仲属, 芸香科榆橘属, 蓝果树科喜树属, 莲叶桐科青藤属, 鼠李科马甲子属, 马尾树科马尾树属 Malpighiaceae (Aspidopterys, Alicia, Amorimia, Calcicola, Caucanthus, Christianella, Diaspis, Excentradenia, Flabellaria, Madagasikaria, Malpighiodes, Mascagnia, Mezia), Juglandaceae (Cyclocarya, Platycarya), Fabaceae (Pterocarpus, Dalbergia), Cruciferae (Isatis, Iberis, Thlaspi), Betulaceae (Alnus, Betula), Ulmaceae (Ulmus, Pteroceltis), Oxyria (Polygonaceae), Fontanesia (Oleaceae), Dipteronia (Aceraceae), Ailanthus (Simaroubaceae), Eucommia (Eucommiaceae), Ptelea (Rutaceae), Camptotheca (Nyssaceae), Illigera (Hernandiaceae), Paliurus (Rhamnaceae), Rhoiptelea (Rhoipteleaceae)
2.2 蝶翅果 Butterfly-winged samara	金虎尾科(Adelphia, Aenigmatanthera, Hiraea, Amorimia, Callaeum, Carolus, Christianella, Dinemandra等), 菊科(金鸡菊属, 偶雏菊属, 蟛蜞菊属), 木犀科(雪柳属, 六道木叶属), 使君子科(诃子属, 榆绿木属, 假榆橘属) Malpighiaceae (Adelphia, Christianella, Aenigmatanthera, Hiraea, Amorimia, Callaeum, Carolus, Dinemandra, etc.), Asterceae (Coreopsis, Boltonia, Sphagneticola), Oleaceae (Fontanesia, Abeliophyllum), Combretaeeae (Terminalia, Anogeissus, Pteleopsis)
3 棱翅果 Rib-winged samara	金虎尾科(Aspicarpa, Calcicola, Digoniopterys), 蓼科(翅果蓼属, 红药子属, 大黄属, 沙拐枣属, 等), 卫矛科(雷公藤属, 卫矛属, 等), 使君子科(风车子属, 诃子属), 无患子科(车桑子属, 黄梨木属), 椴树科(滇桐属, 蚬木属, 柄翅果属, 一担柴属), 鼠李科咀签属, 莲叶桐科青藤属, 茶茱萸科心翼果属, 薯蓣科薯蓣属, 藜科四翅滨藜, 胡颓子科翅果油树 Malpighiaceae (Aspicarpa, Calcicola, Digoniopterys), Polygonaceae (Parapteropyrum, Pteroxygonum, Rheum, Calligonum etc.), Celastraceae (Tripterygium, Euonymus etc.), Sapindaceae (Dodonaea, Boniodendron), Tiliaceae (Craigia, Excentrodendron, Burretiodendron, Colona), Gouania (Rhamnaceae), Illigera (Hernandiaceae), Cardiopteris (Icacinaceae), Dioscorea (Dioscoreaceae), Atriplex canescens (Chenopodiaceae), Elaeagnus mollis (Elaeagnaceae)
4 披针翅果 Lanceolate-winged samara	金虎尾科(风筝果属, 三星果属, Lophopterys, Dicella, Microsteira, Niedenzuella, Rhynchophora, Tetrapterys), 椴树科六翅木属, 卫矛科斜翼属, 十字花科沙芥属, 使君子科萼翅藤属 Malpighiaceae (Hiptage, Tristellateia, Lophopterys, Dicella, Microsteira, Niedenzuella, Rhynchophora, Tetrapterys), Berrya (Tiliaceae), Plagiopteron (Celastraceae), Pugionium (Crucificeae), Getonia (Combretaeeae)
5 翼状萼翅果 Sepal-winged samara	龙脑香科, 钩枝藤科钩枝藤属, 旋花科(飞蛾藤属, 三翅藤属, 白花叶属, 地旋花属) Dipterocarpaceae, Ancistrocladus (Ancistrocladaceae), Convolvulaceae (Dinetus, Tridynamia, Poranopsis, Xenostegia)
6 叶状苞翅果 Bract-winged samara	桦木科(鹅耳枥属, 铁木属, 桦木属, 桤木属, 榛属), 胡桃科(黄杞属, 枫杨属), 败酱科败酱属, 旋花科盾苞藤属, 马鞭草科楔翅藤属, 檀香科米面蓊属 Betulaceae (Carpinus, Ostrya, Betula, Alnus, Corylus), Juglandaceae (Engelhardia, Pterocarya), Patrinia (Valerianaceae), Neuropeltis (Convolvulaceae), Sphenodesme (Verbenaceae), Buckleya (Santalaceae)