花对称性与植物花大小的变异性: 在高寒草甸植物群落检验Berg的假说

doi:10.3724/SP.J.1258.2014.00042

摘要/Abstract

摘要：

植物花对称性与传粉系统密切相关, 花特征的变异性受到传粉者的选择作用。由于特化的传粉者对两侧对称的花的稳定选择, Berg假说认为两侧对称植物花大小的变异性比辐射对称植物的更低; 而且, 在传粉者的选择作用下花特征比植物营养特征明显有更低的变异性, 因为后者更易受环境影响。该文对青藏高原东部高山草甸植物群落的50种开花植物(包括19种两侧对称植物和31种辐射对称植物)的花和叶特征进行了测定和分析。结果表明不论是两侧对称植物还是辐射对称植物, 花大小的变异性都显著低于叶片大小的变异性, 表明传粉者对花施加的稳定选择有利于花的稳定性。但是, 辐射对称物种花大小的变异程度和两侧对称物种的相似, 即使在控制物种系统发育的影响后, 也没有发现显著的差异, 这与Berg假说不一致。高山生态系统中传粉者种类相对较少, 以熊蜂和蝇类为主, 传粉者的活动受局域气候环境影响较大, 因此传粉昆虫对植物花的选择作用强度可能有较大的变异性。

关键词: Berg假说, 两侧对称, 花对称性, 系统发育, 辐射对称

Abstract:

Aims Flower symmetry is closely related to pollination system and variations in flower traits are under the selection role of pollinators. Berg predicted that flower size varies much less in bilateral species than in radial species because of the stable and/or directional selection by pollinators, and that it should also show lower variability than vegetative organs which have relatively higher sensitivity to environments. We examined these so-called Berg’s hypotheses in an alpine meadow.
Methods By measuring the traits of flowers and leaves in 50 flowering plants in an alpine meadow in the eastern Qinghai-Xizang Plateau, we compared the coefficients of variations (CV) in flower size and leaf size between 31 radial and 19 bilateral species and also performed phylogenetically independent comparisons.
Important findings Our results were not completely consistent with the Berg’s hypotheses. The CV of flower size was significantly smaller than that of leaf size in both radial and bilateral species as predicted by Berg, indicating that pollinator-mediated stabilizing selection would benefit flower stability. But the CV of flower size did not differ between the bilateral and the radial species, which is inconsistent with the Berg’s hypothesis even if we controlled confounding effects of phylogenetic relatedness. The role of pollinators in alpine ecosystem where bumble bees and flies predominate is likely more affected by local climatic environments, which leads to considerable variability in the selection of pollinators on flowers.

Key words: Berg’ hypothesis, bilateral symmetry, flower symmetry, phylogeny, radial symmetry

路宁娜,赵志刚. 花对称性与植物花大小的变异性: 在高寒草甸植物群落检验Berg的假说. 植物生态学报, 2014, 38(5): 460-467. DOI: 10.3724/SP.J.1258.2014.00042

LU Ning-Na,ZHAO Zhi-Gang. Flower symmetry and flower size variability: an examination of Berg’s hypotheses in an alpine meadow. Chinese Journal of Plant Ecology, 2014, 38(5): 460-467. DOI: 10.3724/SP.J.1258.2014.00042

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https://www.plant-ecology.com/CN/Y2014/V38/I5/460

图/表 3

参考文献 36

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	科 Family	特征 Trait	变异系数 Mean CV
辐射对称物种 Radial species
喉毛花 Comastoma pulmonarium⁴	龙胆科 Gentianaceae	叶大小(长、宽) Leaf size (length, width)	0.242
喉毛花 Comastoma pulmonarium⁴	龙胆科 Gentianaceae	花大小(花直径、瓣长、瓣宽) Flower size (diameter, length, width)	0.144
湿生扁蕾 Gentianopsis paludosa⁴	龙胆科 Gentianaceae	叶大小(长、宽) Leaf size (length, width)	0.355
湿生扁蕾 Gentianopsis paludosa⁴	龙胆科 Gentianaceae	花大小(花直径、瓣长、瓣宽) Flower size (diameter, length, width)	0.134
瞿麦 Dianthus superbus⁶	石竹科 Caryophyllaceae	叶大小(长、宽) Leaf size (length, width)	0.262
瞿麦 Dianthus superbus⁶	石竹科 Caryophyllaceae	花大小(花直径、瓣长、瓣宽) Flower size (diameter, length, width)	0.118
沼生柳叶菜 Epilobium palustre⁶	柳叶菜科 Onagraceae	叶大小(长、宽) Leaf size (length, width)	0.280
沼生柳叶菜 Epilobium palustre⁶	柳叶菜科 Onagraceae	花大小(花直径、瓣长、瓣宽) Flower size (diameter, length, width)	0.136
柳兰 Chamerion angustifolium⁶	柳叶菜科 Onagraceae	叶大小(长、宽) Leaf size (length, width)	0.200
柳兰 Chamerion angustifolium⁶	柳叶菜科 Onagraceae	花大小(花直径、瓣长、瓣宽) Flower size (diameter, length, width)	0.055
花荵 Polemonium caeruleum⁶	花荵科 Polemoniaceae	叶大小(长、宽) Leaf size (length, width)	0.239
花荵 Polemonium caeruleum⁶	花荵科 Polemoniaceae	花大小(花直径、瓣长、瓣宽) Flower size (diameter, length, width)	0.082
莓叶委陵菜 Potentilla fragarioides²	蔷薇科 Rosaceae	叶大小(长、宽) Leaf size (length, width)	0.198
莓叶委陵菜 Potentilla fragarioides²	蔷薇科 Rosaceae	花大小(花直径、瓣长、瓣宽) Flower size (diameter, length, width)	0.119
银露梅 Potentilla glabra²	蔷薇科 Rosaceae	叶大小(长、宽) Leaf size (length, width)	0.185
银露梅 Potentilla glabra²	蔷薇科 Rosaceae	花大小(花直径、瓣长、瓣宽) Flower size (diameter, length, width)	0.119
金露梅 Potentilla fruticosa²	蔷薇科 Rosaceae	叶大小(长、宽) Leaf size (length, width)	0.222
金露梅 Potentilla fruticosa²	蔷薇科 Rosaceae	花大小(花直径、瓣长、瓣宽) Flower size (diameter, length, width)	0.064
丝叶毛茛 Ranunculus tanguticus var. capillaceus¹	毛茛科 Ranunculaceae	叶大小(长、宽) Leaf size (length, width)	0.575
丝叶毛茛 Ranunculus tanguticus var. capillaceus¹	毛茛科 Ranunculaceae	花大小(花直径、瓣长、瓣宽) Flower size (diameter, length, width)	0.136
钝裂银莲花 Anemone obtusiloba¹	毛茛科 Ranunculaceae	叶大小(长、宽) Leaf size (length, width)	0.464
钝裂银莲花 Anemone obtusiloba¹	毛茛科 Ranunculaceae	花大小(花直径、瓣长、瓣宽) Flower size (diameter, length, width)	0.140
高山唐松草 Thalictrum alpinum¹	毛茛科 Ranunculaceae	叶大小(长、宽) Leaf size (length, width)	0.113
高山唐松草 Thalictrum alpinum¹	毛茛科 Ranunculaceae	花大小(花直径、瓣长、瓣宽) Flower size (diameter, length, width)	0.154
小花草玉梅 Anemone rivularis var. flore-minore¹	毛茛科 Ranunculaceae	叶大小(长、宽) Leaf size (length, width)	0.223
小花草玉梅 Anemone rivularis var. flore-minore¹	毛茛科 Ranunculaceae	花大小(花直径、瓣长、瓣宽) Flower size (diameter, length, width)	0.143
毛茛状金莲花 Trollius ranunculoides¹	毛茛科 Ranunculaceae	花大小(花直径、瓣长、瓣宽) Flower size (diameter, length, width)	0.137
甘青老鹳草 Geranium pylzowianum⁶	牻牛儿苗科 Geraniaceae	叶大小(长、宽) Leaf size (length, width)	0.127
甘青老鹳草 Geranium pylzowianum⁶	牻牛儿苗科 Geraniaceae	花大小(花直径、瓣长、瓣宽) Flower size (diameter, length, width)	0.096
鼠掌老鹳草 Geranium sibiricum⁶	牻牛儿苗科 Geraniaceae	叶大小(长、宽) Leaf size (length, width)	0.281
鼠掌老鹳草 Geranium sibiricum⁶	牻牛儿苗科 Geraniaceae	花大小(花直径、瓣长、瓣宽) Flower size (diameter, length, width)	0.155
水团花 Adina pilulifera³	茜草科 Rubiaceae	叶大小(长、宽) Leaf size (length, width)	0.236
水团花 Adina pilulifera³	茜草科 Rubiaceae	花大小(花直径、瓣长、瓣宽) Flower size (diameter, length, width)	0.114
直立点地梅 Androsace erecta⁶	报春花科 Primulaceae	花大小(花直径、瓣长、瓣宽) Flower size (diameter, length, width)	0.067
黄帚橐吾 Ligularia virgaurea⁵	菊科 Asteraceae	叶大小(长、宽) Leaf size (length, width)	0.233
黄帚橐吾 Ligularia virgaurea⁵	菊科 Asteraceae	花大小(花直径、瓣长、瓣宽) Flower size (diameter, length, width)	0.161
毛连菜 Picris hieracioides⁶	菊科 Asteraceae	叶大小(长、宽) Leaf size (length, width)	0.223
毛连菜 Picris hieracioides⁶	菊科 Asteraceae	花大小(花直径、瓣长、瓣宽) Flower size (diameter, length, width)	0.153
沼生苦苣菜 Sonchus palustris⁶	菊科 Asteraceae	花大小(花直径、瓣长、瓣宽) Flower size (diameter, length, width)	0.100
黑褐千里光 Senecio atrofuscus⁶	菊科 Asteraceae	花大小(花直径、瓣长、瓣宽) Flower size (diameter, length, width)	0.127
缘毛紫菀 Aster souliei⁶	菊科 Asteraceae	花大小(花直径、瓣长、瓣宽) Flower size (diameter, length, width)	0.132
高山紫菀 Aster alpinus⁶	菊科 Asteraceae	叶大小(长、宽) Leaf size (length, width)	0.172
高山紫菀 Aster alpinus⁶	菊科 Asteraceae	花大小(花直径、瓣长、瓣宽) Flower size (diameter, length, width)	0.172
	科 Family	特征 Trait	变异系数 Mean CV
甘青微孔草 Microula pseudotrichocarpa³	紫草科 Boraginaceae	花大小(花直径、瓣长、瓣宽) Flower size (diameter, length, width)	0.105
刺芒龙胆 Gentiana aristata⁴	龙胆科 Gentianaceae	花大小(花冠筒长、花冠直径、花冠裂片长、花冠裂片宽、花冠筒直径) Flower size (corolla tube length, corolla diameter, corolla lobe length, corolla lobe width, corolla tube diameter)	0.093
鳞叶龙胆 Gentiana squarrosa⁴	龙胆科 Gentianaceae	叶大小(长、宽) Leaf size (length, width)	0.223
鳞叶龙胆 Gentiana squarrosa⁴	龙胆科 Gentianaceae	花大小(花冠筒长、花冠直径、花冠裂片长、花冠裂片宽、花冠筒直径) Flower size (corolla tube length, corolla diameter, corolla lobe length, corolla lobe width, corolla tube diameter)	0.240
狼毒 Stellera chamaejasme⁶	瑞香科 Thymelaeaceae	花大小(花直径、瓣长、瓣宽) Flower size (diameter, length, width)	0.128
夏河缬草 Valeriana xiaheensis⁵	败酱科 Valerianaceae	花大小(花冠筒长、花冠筒直径、花直径、花瓣长、花瓣宽) Flower size (corolla tube length, corolla tube diameter, corolla diameter, petal length and width)	0.220
高山韭 Allium sikkimense	百合科 Liliaceae	花大小(花瓣长、宽) Flower size (length and width)	0.115
高原鸢尾 Iris collettii	鸢尾科 Iridaceae	花大小(外花被直径、内花被直径、外花瓣长、外花瓣宽、内花瓣长、内花瓣宽) Flower size (outside perianth diameter, inside perianth diameter, outside petal length and width, inside petal length and width )	0.080
两侧对称物种 Bilateral species
碎米蕨叶马先蒿 Pedicularis cheilanthifolia⁵	列当科 Orobanchaceae	叶大小(长、宽) Leaf size (length and width)	0.248
碎米蕨叶马先蒿 Pedicularis cheilanthifolia⁵	列当科 Orobanchaceae	花大小(花冠筒长、下唇长、下唇宽、花冠与下唇间距) Flower size (corolla tube length, Lower lip length and width, distance between corolla and lower lip)	0.102
甘肃马先蒿 Pedicularis kansuensis⁵	列当科 Orobanchaceae	叶大小(长、宽) Leaf size (length and width)	0.101
甘肃马先蒿 Pedicularis kansuensis⁵	列当科 Orobanchaceae	花大小(花冠筒长、下唇长、下唇宽、花冠与下唇间距) Flower size (corolla tube length, Lower lip length and width, distance between corolla and lower lip)	0.077
四川马先蒿 Pedicularis szetschuanica⁵	列当科 Orobanchaceae	叶大小(长、宽) Leaf size (length and width)	0.286
四川马先蒿 Pedicularis szetschuanica⁵	列当科 Orobanchaceae	花大小(花冠筒长、下唇长、下唇宽、花冠与下唇间距) Flower size (corolla tube length, Lower lip length and width, distance between corolla and lower lip)	0.124
露蕊乌头 Gymnaconitum gymnandrum¹	毛茛科 Ranunculaceae	叶大小(长、宽) Leaf size (length and width)	0.259
露蕊乌头 Gymnaconitum gymnandrum¹	毛茛科 Ranunculaceae	花大小(盔高、萼片直径、侧萼片宽、侧萼片高、侧萼片间距) Flower size (galea height, sepal diameter, Lateral sepal width and length, distance between lateral sepals)	0.122
展毛翠雀花 Delphinium kamaonense¹	毛茛科 Ranunculaceae	叶大小(长、宽) Leaf size (length and width)	0.124
展毛翠雀花 Delphinium kamaonense¹	毛茛科 Ranunculaceae	花大小(花冠直径、距长、上萼片宽、下萼片宽) Flower size (corolla diameter, spur length, upper sepal width, under sepal width)	0.068
黄花棘豆 Oxytropis ochrocephala²	豆科 Fabaceae	叶大小(长、宽) Leaf size (length and width)	0.247
黄花棘豆 Oxytropis ochrocephala²	豆科 Fabaceae	花大小(翼瓣长、龙骨瓣长、旗瓣长) Flower size (wing petal length, keel length, flag petal length)	0.158
甘肃米口袋 Gueldenstaedtia gansuensis²	豆科 Fabaceae	叶大小(长、宽) Leaf size (length and width)	0.092
甘肃米口袋 Gueldenstaedtia gansuensis²	豆科 Fabaceae	花大小(龙骨瓣长、旗瓣长、龙骨瓣宽、翼瓣长、旗瓣宽) Flower size (keel length and width, wing petal length, flag petal length and width)	0.199
青海苜蓿 Medicago archiducis-nicolai²	豆科 Fabaceae	叶大小(长、宽) Leaf size (length and width)	0.292
青海苜蓿 Medicago archiducis-nicolai²	豆科 Fabaceae	花大小(龙骨瓣长、旗瓣长、龙骨瓣宽、翼瓣长、旗瓣宽) Flower size (keel length and width, wing petal length and width, flag petal length and width)	0.09
歪头菜 Vicia unijuga²	豆科 Fabaceae	叶大小(长、宽) Leaf size (length and width)	0.198
歪头菜 Vicia unijuga²	豆科 Fabaceae	花大小(翼瓣长、龙骨瓣长、旗瓣长) Flower size (wing petal length, keel length, flag petal length)	0.208
苦马豆 Sphaerophysa salsula²	豆科 Fabaceae	叶大小(长、宽) Leaf size (length and width)	0.064
苦马豆 Sphaerophysa salsula²	豆科 Fabaceae	花大小(翼瓣长、龙骨瓣长、旗瓣长) Flower size (wing petal length, keel length, flag petal length)	0.05
山黧豆 Lathyrus quinquenervius²	豆科 Fabaceae	叶大小(长、宽) Leaf size (length and width)	0.276
山黧豆 Lathyrus quinquenervius²	豆科 Fabaceae	花大小(翼瓣长、龙骨瓣长和宽、旗瓣长) Flower size (wing petal length, keel length and width, flag petal length)	0.124
	科 Family	特征 Trait	变异系数 Mean CV
救荒野豌豆 Vicia sativa²	豆科 Fabaceae	花大小(翼瓣长、龙骨瓣长、旗瓣长) Flower size (wing petal length, keel length, flag petal length)	0.060
大叶鼠尾草 `Salvia grandifolia⁴	唇形科 Lamiaceae	叶大小(长、宽) Leaf size (length and width)	0.181
大叶鼠尾草 `Salvia grandifolia⁴	唇形科 Lamiaceae	花大小(花冠筒长、下唇长、下唇宽、上下唇间距) Flower size (corolla tube length, lower lip length and width, distance between lips)	0.099
藏荆芥 Nepeta angustifolia⁴	唇形科 Lamiaceae	叶大小(长、宽) Leaf size (length and width)	0.187
藏荆芥 Nepeta angustifolia⁴	唇形科 Lamiaceae	花大小(花萼筒长、花萼筒直径、上唇长、上唇宽、下唇长、下唇宽) Flower size (corolla tube length, corolla tube diameter, upper lip length and width, lower lip length and width)	0.076
异色黄芩 Scutellaria discolor⁴	唇形科 Lamiaceae	叶大小(长、宽) Leaf size (length and width)	0.225
异色黄芩 Scutellaria discolor⁴	唇形科 Lamiaceae	花大小(花萼筒长、花萼筒直径、上唇长、上唇宽、下唇长、下唇宽) Flower size (corolla tube length, corolla tube diameter, upper lip length and width, lower lip length and width)	0.080
毛果婆婆纳 Veronica eriogyne³	玄参科 Scrophulariaceae	叶大小(长、宽) Leaf size (length and width)	0.111
毛果婆婆纳 Veronica eriogyne³	玄参科 Scrophulariaceae	花大小(花萼筒直径、上唇宽、下唇宽) Flower size (corolla tube diameter, upper lip width, lower lip width)	0.129
短腺小米草 Euphrasia regelii³	玄参科 Scrophulariaceae	叶大小(长、宽) Leaf size (length and width)	0.204
短腺小米草 Euphrasia regelii³	玄参科 Scrophulariaceae	花大小(上唇长、上唇宽、下唇长、下唇宽、上下唇间距) Flower size (upper lip length and width, lower lip length and width, distance between lips)	0.151
灯笼草 Clinopodium polycephalum⁴	唇形科 Lamiaceae	叶大小(长、宽) Leaf size (length and width)	0.167
灯笼草 Clinopodium polycephalum⁴	唇形科 Lamiaceae	花大小(花萼筒长、上唇长、上唇宽、下唇长、下唇宽) Flower size (corolla tube length, upper lip length and width, lower lip length and width)	0.118
陕西紫堇 Corydalis shensiana⁶	罂粟科 Papaveraceae	叶大小(长、宽)Leaf size (length and width)	0.264
陕西紫堇 Corydalis shensiana⁶	罂粟科 Papaveraceae	花大小(上花瓣长、距长、下花瓣长、下花瓣宽、上下瓣间距) Flower size (upper petal length, lower petal length and width, distance between petals, spur length)	0.180

	科 Family	特征 Trait	变异系数 Mean CV
辐射对称物种 Radial species
喉毛花 Comastoma pulmonarium⁴	龙胆科 Gentianaceae	叶大小(长、宽) Leaf size (length, width)	0.242
喉毛花 Comastoma pulmonarium⁴	龙胆科 Gentianaceae	花大小(花直径、瓣长、瓣宽) Flower size (diameter, length, width)	0.144
湿生扁蕾 Gentianopsis paludosa⁴	龙胆科 Gentianaceae	叶大小(长、宽) Leaf size (length, width)	0.355
湿生扁蕾 Gentianopsis paludosa⁴	龙胆科 Gentianaceae	花大小(花直径、瓣长、瓣宽) Flower size (diameter, length, width)	0.134
瞿麦 Dianthus superbus⁶	石竹科 Caryophyllaceae	叶大小(长、宽) Leaf size (length, width)	0.262
瞿麦 Dianthus superbus⁶	石竹科 Caryophyllaceae	花大小(花直径、瓣长、瓣宽) Flower size (diameter, length, width)	0.118
沼生柳叶菜 Epilobium palustre⁶	柳叶菜科 Onagraceae	叶大小(长、宽) Leaf size (length, width)	0.280
沼生柳叶菜 Epilobium palustre⁶	柳叶菜科 Onagraceae	花大小(花直径、瓣长、瓣宽) Flower size (diameter, length, width)	0.136
柳兰 Chamerion angustifolium⁶	柳叶菜科 Onagraceae	叶大小(长、宽) Leaf size (length, width)	0.200
柳兰 Chamerion angustifolium⁶	柳叶菜科 Onagraceae	花大小(花直径、瓣长、瓣宽) Flower size (diameter, length, width)	0.055
花荵 Polemonium caeruleum⁶	花荵科 Polemoniaceae	叶大小(长、宽) Leaf size (length, width)	0.239
花荵 Polemonium caeruleum⁶	花荵科 Polemoniaceae	花大小(花直径、瓣长、瓣宽) Flower size (diameter, length, width)	0.082
莓叶委陵菜 Potentilla fragarioides²	蔷薇科 Rosaceae	叶大小(长、宽) Leaf size (length, width)	0.198
莓叶委陵菜 Potentilla fragarioides²	蔷薇科 Rosaceae	花大小(花直径、瓣长、瓣宽) Flower size (diameter, length, width)	0.119
银露梅 Potentilla glabra²	蔷薇科 Rosaceae	叶大小(长、宽) Leaf size (length, width)	0.185
银露梅 Potentilla glabra²	蔷薇科 Rosaceae	花大小(花直径、瓣长、瓣宽) Flower size (diameter, length, width)	0.119
金露梅 Potentilla fruticosa²	蔷薇科 Rosaceae	叶大小(长、宽) Leaf size (length, width)	0.222
金露梅 Potentilla fruticosa²	蔷薇科 Rosaceae	花大小(花直径、瓣长、瓣宽) Flower size (diameter, length, width)	0.064
丝叶毛茛 Ranunculus tanguticus var. capillaceus¹	毛茛科 Ranunculaceae	叶大小(长、宽) Leaf size (length, width)	0.575
丝叶毛茛 Ranunculus tanguticus var. capillaceus¹	毛茛科 Ranunculaceae	花大小(花直径、瓣长、瓣宽) Flower size (diameter, length, width)	0.136
钝裂银莲花 Anemone obtusiloba¹	毛茛科 Ranunculaceae	叶大小(长、宽) Leaf size (length, width)	0.464
钝裂银莲花 Anemone obtusiloba¹	毛茛科 Ranunculaceae	花大小(花直径、瓣长、瓣宽) Flower size (diameter, length, width)	0.140
高山唐松草 Thalictrum alpinum¹	毛茛科 Ranunculaceae	叶大小(长、宽) Leaf size (length, width)	0.113
高山唐松草 Thalictrum alpinum¹	毛茛科 Ranunculaceae	花大小(花直径、瓣长、瓣宽) Flower size (diameter, length, width)	0.154
小花草玉梅 Anemone rivularis var. flore-minore¹	毛茛科 Ranunculaceae	叶大小(长、宽) Leaf size (length, width)	0.223
小花草玉梅 Anemone rivularis var. flore-minore¹	毛茛科 Ranunculaceae	花大小(花直径、瓣长、瓣宽) Flower size (diameter, length, width)	0.143
毛茛状金莲花 Trollius ranunculoides¹	毛茛科 Ranunculaceae	花大小(花直径、瓣长、瓣宽) Flower size (diameter, length, width)	0.137
甘青老鹳草 Geranium pylzowianum⁶	牻牛儿苗科 Geraniaceae	叶大小(长、宽) Leaf size (length, width)	0.127
甘青老鹳草 Geranium pylzowianum⁶	牻牛儿苗科 Geraniaceae	花大小(花直径、瓣长、瓣宽) Flower size (diameter, length, width)	0.096
鼠掌老鹳草 Geranium sibiricum⁶	牻牛儿苗科 Geraniaceae	叶大小(长、宽) Leaf size (length, width)	0.281
鼠掌老鹳草 Geranium sibiricum⁶	牻牛儿苗科 Geraniaceae	花大小(花直径、瓣长、瓣宽) Flower size (diameter, length, width)	0.155
水团花 Adina pilulifera³	茜草科 Rubiaceae	叶大小(长、宽) Leaf size (length, width)	0.236
水团花 Adina pilulifera³	茜草科 Rubiaceae	花大小(花直径、瓣长、瓣宽) Flower size (diameter, length, width)	0.114
直立点地梅 Androsace erecta⁶	报春花科 Primulaceae	花大小(花直径、瓣长、瓣宽) Flower size (diameter, length, width)	0.067
黄帚橐吾 Ligularia virgaurea⁵	菊科 Asteraceae	叶大小(长、宽) Leaf size (length, width)	0.233
黄帚橐吾 Ligularia virgaurea⁵	菊科 Asteraceae	花大小(花直径、瓣长、瓣宽) Flower size (diameter, length, width)	0.161
毛连菜 Picris hieracioides⁶	菊科 Asteraceae	叶大小(长、宽) Leaf size (length, width)	0.223
毛连菜 Picris hieracioides⁶	菊科 Asteraceae	花大小(花直径、瓣长、瓣宽) Flower size (diameter, length, width)	0.153
沼生苦苣菜 Sonchus palustris⁶	菊科 Asteraceae	花大小(花直径、瓣长、瓣宽) Flower size (diameter, length, width)	0.100
黑褐千里光 Senecio atrofuscus⁶	菊科 Asteraceae	花大小(花直径、瓣长、瓣宽) Flower size (diameter, length, width)	0.127
缘毛紫菀 Aster souliei⁶	菊科 Asteraceae	花大小(花直径、瓣长、瓣宽) Flower size (diameter, length, width)	0.132
高山紫菀 Aster alpinus⁶	菊科 Asteraceae	叶大小(长、宽) Leaf size (length, width)	0.172
高山紫菀 Aster alpinus⁶	菊科 Asteraceae	花大小(花直径、瓣长、瓣宽) Flower size (diameter, length, width)	0.172
	科 Family	特征 Trait	变异系数 Mean CV
甘青微孔草 Microula pseudotrichocarpa³	紫草科 Boraginaceae	花大小(花直径、瓣长、瓣宽) Flower size (diameter, length, width)	0.105
刺芒龙胆 Gentiana aristata⁴	龙胆科 Gentianaceae	花大小(花冠筒长、花冠直径、花冠裂片长、花冠裂片宽、花冠筒直径) Flower size (corolla tube length, corolla diameter, corolla lobe length, corolla lobe width, corolla tube diameter)	0.093
鳞叶龙胆 Gentiana squarrosa⁴	龙胆科 Gentianaceae	叶大小(长、宽) Leaf size (length, width)	0.223
鳞叶龙胆 Gentiana squarrosa⁴	龙胆科 Gentianaceae	花大小(花冠筒长、花冠直径、花冠裂片长、花冠裂片宽、花冠筒直径) Flower size (corolla tube length, corolla diameter, corolla lobe length, corolla lobe width, corolla tube diameter)	0.240
狼毒 Stellera chamaejasme⁶	瑞香科 Thymelaeaceae	花大小(花直径、瓣长、瓣宽) Flower size (diameter, length, width)	0.128
夏河缬草 Valeriana xiaheensis⁵	败酱科 Valerianaceae	花大小(花冠筒长、花冠筒直径、花直径、花瓣长、花瓣宽) Flower size (corolla tube length, corolla tube diameter, corolla diameter, petal length and width)	0.220
高山韭 Allium sikkimense	百合科 Liliaceae	花大小(花瓣长、宽) Flower size (length and width)	0.115
高原鸢尾 Iris collettii	鸢尾科 Iridaceae	花大小(外花被直径、内花被直径、外花瓣长、外花瓣宽、内花瓣长、内花瓣宽) Flower size (outside perianth diameter, inside perianth diameter, outside petal length and width, inside petal length and width )	0.080
两侧对称物种 Bilateral species
碎米蕨叶马先蒿 Pedicularis cheilanthifolia⁵	列当科 Orobanchaceae	叶大小(长、宽) Leaf size (length and width)	0.248
碎米蕨叶马先蒿 Pedicularis cheilanthifolia⁵	列当科 Orobanchaceae	花大小(花冠筒长、下唇长、下唇宽、花冠与下唇间距) Flower size (corolla tube length, Lower lip length and width, distance between corolla and lower lip)	0.102
甘肃马先蒿 Pedicularis kansuensis⁵	列当科 Orobanchaceae	叶大小(长、宽) Leaf size (length and width)	0.101
甘肃马先蒿 Pedicularis kansuensis⁵	列当科 Orobanchaceae	花大小(花冠筒长、下唇长、下唇宽、花冠与下唇间距) Flower size (corolla tube length, Lower lip length and width, distance between corolla and lower lip)	0.077
四川马先蒿 Pedicularis szetschuanica⁵	列当科 Orobanchaceae	叶大小(长、宽) Leaf size (length and width)	0.286
四川马先蒿 Pedicularis szetschuanica⁵	列当科 Orobanchaceae	花大小(花冠筒长、下唇长、下唇宽、花冠与下唇间距) Flower size (corolla tube length, Lower lip length and width, distance between corolla and lower lip)	0.124
露蕊乌头 Gymnaconitum gymnandrum¹	毛茛科 Ranunculaceae	叶大小(长、宽) Leaf size (length and width)	0.259
露蕊乌头 Gymnaconitum gymnandrum¹	毛茛科 Ranunculaceae	花大小(盔高、萼片直径、侧萼片宽、侧萼片高、侧萼片间距) Flower size (galea height, sepal diameter, Lateral sepal width and length, distance between lateral sepals)	0.122
展毛翠雀花 Delphinium kamaonense¹	毛茛科 Ranunculaceae	叶大小(长、宽) Leaf size (length and width)	0.124
展毛翠雀花 Delphinium kamaonense¹	毛茛科 Ranunculaceae	花大小(花冠直径、距长、上萼片宽、下萼片宽) Flower size (corolla diameter, spur length, upper sepal width, under sepal width)	0.068
黄花棘豆 Oxytropis ochrocephala²	豆科 Fabaceae	叶大小(长、宽) Leaf size (length and width)	0.247
黄花棘豆 Oxytropis ochrocephala²	豆科 Fabaceae	花大小(翼瓣长、龙骨瓣长、旗瓣长) Flower size (wing petal length, keel length, flag petal length)	0.158
甘肃米口袋 Gueldenstaedtia gansuensis²	豆科 Fabaceae	叶大小(长、宽) Leaf size (length and width)	0.092
甘肃米口袋 Gueldenstaedtia gansuensis²	豆科 Fabaceae	花大小(龙骨瓣长、旗瓣长、龙骨瓣宽、翼瓣长、旗瓣宽) Flower size (keel length and width, wing petal length, flag petal length and width)	0.199
青海苜蓿 Medicago archiducis-nicolai²	豆科 Fabaceae	叶大小(长、宽) Leaf size (length and width)	0.292
青海苜蓿 Medicago archiducis-nicolai²	豆科 Fabaceae	花大小(龙骨瓣长、旗瓣长、龙骨瓣宽、翼瓣长、旗瓣宽) Flower size (keel length and width, wing petal length and width, flag petal length and width)	0.09
歪头菜 Vicia unijuga²	豆科 Fabaceae	叶大小(长、宽) Leaf size (length and width)	0.198
歪头菜 Vicia unijuga²	豆科 Fabaceae	花大小(翼瓣长、龙骨瓣长、旗瓣长) Flower size (wing petal length, keel length, flag petal length)	0.208
苦马豆 Sphaerophysa salsula²	豆科 Fabaceae	叶大小(长、宽) Leaf size (length and width)	0.064
苦马豆 Sphaerophysa salsula²	豆科 Fabaceae	花大小(翼瓣长、龙骨瓣长、旗瓣长) Flower size (wing petal length, keel length, flag petal length)	0.05
山黧豆 Lathyrus quinquenervius²	豆科 Fabaceae	叶大小(长、宽) Leaf size (length and width)	0.276
山黧豆 Lathyrus quinquenervius²	豆科 Fabaceae	花大小(翼瓣长、龙骨瓣长和宽、旗瓣长) Flower size (wing petal length, keel length and width, flag petal length)	0.124
	科 Family	特征 Trait	变异系数 Mean CV
救荒野豌豆 Vicia sativa²	豆科 Fabaceae	花大小(翼瓣长、龙骨瓣长、旗瓣长) Flower size (wing petal length, keel length, flag petal length)	0.060
大叶鼠尾草 `Salvia grandifolia⁴	唇形科 Lamiaceae	叶大小(长、宽) Leaf size (length and width)	0.181
大叶鼠尾草 `Salvia grandifolia⁴	唇形科 Lamiaceae	花大小(花冠筒长、下唇长、下唇宽、上下唇间距) Flower size (corolla tube length, lower lip length and width, distance between lips)	0.099
藏荆芥 Nepeta angustifolia⁴	唇形科 Lamiaceae	叶大小(长、宽) Leaf size (length and width)	0.187
藏荆芥 Nepeta angustifolia⁴	唇形科 Lamiaceae	花大小(花萼筒长、花萼筒直径、上唇长、上唇宽、下唇长、下唇宽) Flower size (corolla tube length, corolla tube diameter, upper lip length and width, lower lip length and width)	0.076
异色黄芩 Scutellaria discolor⁴	唇形科 Lamiaceae	叶大小(长、宽) Leaf size (length and width)	0.225
异色黄芩 Scutellaria discolor⁴	唇形科 Lamiaceae	花大小(花萼筒长、花萼筒直径、上唇长、上唇宽、下唇长、下唇宽) Flower size (corolla tube length, corolla tube diameter, upper lip length and width, lower lip length and width)	0.080
毛果婆婆纳 Veronica eriogyne³	玄参科 Scrophulariaceae	叶大小(长、宽) Leaf size (length and width)	0.111
毛果婆婆纳 Veronica eriogyne³	玄参科 Scrophulariaceae	花大小(花萼筒直径、上唇宽、下唇宽) Flower size (corolla tube diameter, upper lip width, lower lip width)	0.129
短腺小米草 Euphrasia regelii³	玄参科 Scrophulariaceae	叶大小(长、宽) Leaf size (length and width)	0.204
短腺小米草 Euphrasia regelii³	玄参科 Scrophulariaceae	花大小(上唇长、上唇宽、下唇长、下唇宽、上下唇间距) Flower size (upper lip length and width, lower lip length and width, distance between lips)	0.151
灯笼草 Clinopodium polycephalum⁴	唇形科 Lamiaceae	叶大小(长、宽) Leaf size (length and width)	0.167
灯笼草 Clinopodium polycephalum⁴	唇形科 Lamiaceae	花大小(花萼筒长、上唇长、上唇宽、下唇长、下唇宽) Flower size (corolla tube length, upper lip length and width, lower lip length and width)	0.118
陕西紫堇 Corydalis shensiana⁶	罂粟科 Papaveraceae	叶大小(长、宽)Leaf size (length and width)	0.264
陕西紫堇 Corydalis shensiana⁶	罂粟科 Papaveraceae	花大小(上花瓣长、距长、下花瓣长、下花瓣宽、上下瓣间距) Flower size (upper petal length, lower petal length and width, distance between petals, spur length)	0.180