Interspecific relationship between Armeniaca hongpingensis T. T. Yu & C. L. Li and related species based on morphological characters
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摘要:
本研究通过野外调查获得洪平杏(Armeniaca hongpingensis T. T. Yu & C. L. Li)、杏(A. vulgaris Lam.)、梅(A. mume Siebold)和杏梅( A. mume var. bungo Makino)有关的叶、花、果、果核及枝条的21个定量性状和15个定性性状,并对这些数据进行了聚类分析。结果显示,无论是PAM聚类还是UPGMA聚类分析都可清晰地将4种杏属植物分开,说明物种之间具有明显的区分特征。主成分分析的前3个主成分中,一年生枝颜色、被毛情况,叶及果实被毛,叶形指标(叶形、叶尖、叶基)和果核表面特征的绝对权重值都在0.9以上,表明这些性状在4种杏属植物的分类中具有重要的作用。电镜扫描观察结果发现,供试花粉均是单粒花粉,等极,辐射对称,具3孔萌发沟。杏、梅、杏梅的花粉表面纹饰均为条纹状,但洪平杏的表面纹饰为间断条纹,且有不明显穿孔。4种植物叶上表皮细胞均具条纹状角质增厚。与其他种相比,洪平杏细胞表面条纹常弯曲、交叠,无固定走向。根据聚类分析结果和微形态特征,支持将洪平杏作为区别于梅和杏的一个独立种的处理。
Abstract:Armeniaca hongpingensis T. T. Yu & C. L. Li is a critically endangered species distributed only in Hongping Town, Shennongjia, Hubei Province, China. According to the Flora of China, A. hongpingensis and A. mume var. bungo Makino may be natural hybrids of A. mume Lam. and A. vulgaris Siebold. To date, however, no morphological study on the relationship between A. hongpingensis, A. vulgaris, A. mume, and A. mume var. bungo has been conducted. Through field investigations, 21 quantitative and 15 qualitative leaf, flower, fruit, kernel, and branch characters were obtained, followed by cluster analysis. Results showed that both PAM and UPGMA cluster analysis clearly divided the specimens into four species, indicating obvious distinguishing characters among them. Based on principal component analysis, the absolute weight values of annual branch color and hair-covering status, hair-covering status of leaf and fruit, leaf shape index (leaf shape, leaf tip, leaf base), and surface characters of kernel in the first three principal components were all above 0.9, indicating that these traits play important roles in the classification of the four Armeniaca species. Based on electron microscopy, the pollen grains were identified as single grain (monads), isopolar, radially symmetrical, and tri-colporate. The pollen surfaces of A. vulgaris, A. mume, and A. mume var. bungo were striate-ornamented, while the pollen of A. hongpingensis was discontinuously striped and showed inconspicuous perforation. The epidermal cells of the leaves of the four plants all showed striate keratinization. Compared to the other species, the stripes on the cell surface of A. hongpingensis were often curved, overlapped, and without fixed direction. According to cluster analysis and micromorphological characters, A. hongpingensis should be treated as an independent species from A. vulgaris and A. mume.
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Keywords:
- Armeniaca hongpingensis /
- Interspecific relationship /
- Micromorphology /
- Palynology
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图 3 叶背表皮毛 (A~D)、气孔(E~H) 、叶正面表皮细胞(I~L)及花粉 (M~T)电镜扫描图
A、E、I、M、Q:洪平杏(Armeniaca hongpingensis);B、F、J、N、R:杏(A. vulgaris);C、G、K、O、S:梅(A. mume);D、H、L、P、T:杏梅(A. mume var. bungo)。
Figure 3. Morphological characters of trichome (A–D), stomata (E–H) on leaf abaxial surface, epidermal cell on leaf adaxial surface (I–L), and pollen (M–T)
表 1 洪平杏及其近缘种定性性状及其赋值
Table 1 Selected characters and values for Armeniaca hongpingensis and related species
性状
Characters性状分类及赋值
Classification and values of characters一年生枝颜色 浅绿色—0 红褐色—1 紫红色—2 深紫色—3 一年生枝被毛 无—0 有—1 叶形 宽卵圆形—0 椭圆到椭圆状卵形—1 卵圆形—2 叶尖 短渐尖至急尖—0 长渐尖至尾尖—1 尾尖—2 叶基 宽楔形至圆形—0 圆形至近心形—1 叶缘 细锐锯齿—0 圆钝锯齿—1 叶柄腺体 无—0 有—1 叶片正面被毛 无—0 有—1 叶片背面被毛 无—0 有(密被浅黄褐色柔毛)—1 叶脉被毛 无或具短柔毛—0 密被浅黄褐色柔毛—1 果实被毛 微被短柔毛—0 密被浅黄褐色柔毛—1 核形 卵圆形—0 圆形—1 核顶端 圆钝—0 小钝尖—1 急尖—2 核表面 光滑或稍粗糙—0 蜂窝状孔穴—1 核腹面 龙骨状棱—0 纵沟—1 表 2 洪平杏、梅、杏梅及杏的数量性状
Table 2 Morphological quantitative characters of Armeniaca hongpingensis, A. mume, A. mume var. bungo, and A. vulgaris (mean±SD)
性状
Characters / mm洪平杏
A. hongpingensis杏
A. vulgaris梅
A. mume杏梅
A. mume var. bungo叶长 121.22±21.04a 64.49±11.52d 79.22±8.71c 108.86±6.22b 叶宽 57.34±11.37a 50.12±8.49b 41.52±4.63c 61.12±4.94a 叶柄长 17.15±1.74c 24.73±4.92a 11.14±1.58d 18.67±2.18b 叶长/宽 2.13±0.15a 1.29±0.10d 1.91±0.16b 1.79±0.12c 叶柄长/叶长 0.15±0.03c 0.39±0.06a 0.14±0.02d 0.17±0.02b 花冠直径 35.61±2.48a 25.95±1.20c 24.14±0.87d 31.46±2.21b 花梗长 4.19±0.72a 1.33±0.25d 3.47±0.39b 2.70±0.23c 花瓣长 19.09±1.18a 13.00±0.70c 12.14±1.06d 13.89±1.00b 花瓣宽 14.99±1.57a 12.23±0.94b 10.50±1.10c 14.58±1.11a 萼片长 5.11±0.40a 5.05±0.43a 4.43±0.32b 5.07±0.32a 萼片宽 3.55±0.31c 4.53±0.42a 3.73±0.29bc 3.84±0.33b 果长 33.52±1.71c 34.18±3.32a 33.9±1.64b 29.89±1.64c 果宽 33.52±1.70b 35.28±1.96a 28.1±2.13c 35.88±3.76a 果厚 30.55±1.53b 35.2±2.04a 26.5±1.65c 32.10±4.13b 果长宽比 1.09±0.04a 0.096±0.03c 1.04±0.05b 0.99±0.06c 果宽厚比 1.10±0.04a 1.00±0.04c 1.06±0.04b 1.12±0.06a 核长 25.12±1.66a 20.66±0.83c 20.85±2.16c 23.36±1.88b 核宽 17.36±1.19b 16.25±0.58c 14.22±1.32d 19.6±1.55a 核厚 10.27±0.68c 9.52±0.4d 11.23±0.73b 15.09±1.15a 核长宽比 1.45±0.10a 1.27±0.03b 1.48±0.18a 1.19±0.05c 核宽厚比 1.69±0.09a 1.71±0.05a 1.26±0.07b 1.30±0.04b 注:同行不同字母代表差异显著(P<0.05)。下同。 Note: Different letters on same line represent significant differences (P<0.05). Same below. 表 3 杏属4种植物的叶表面气孔及条纹数量特征
Table 3 Stomata and stripe quantitative characters on leaf abaxial surface of four Armeniaca species
性状Characters 洪平杏
A. hongpingensis杏
A. vulgaris梅
A. mume杏梅
A. mume var. bungo气孔长 / μm 20.96±2.02a 28.00±2.47b 20.92±2.03a 20.67±1.11a 气孔宽 / μm 15.82±1.04a 22.15±2.21c 17.06±1.41b 17.10±1.32b 气孔密度 / ind/mm2 549.25±114.02a 361.00±93.45c 809.63±159.25b 809.25±88.73b 条纹数 11.18±2.19a 8.61±2.40c 18.39±5.30b 9.32±2.30c 条纹宽 / μm 0.84±0.13a 1.29±0.16c 0.67±0.13b 0.87±0.12a 表 4 杏属4种植物的花粉形态、大小及表面条纹比较
Table 4 Comparison of pollen morphology, size, and stripes among four species of Armeniaca
性状
Characters洪平杏
A. hongpingensis杏
A. vulgaris梅
A. mume杏梅
A. mume var. bungo花粉形态 长球形 长球形 长球形 长球形 对称性与极性 等级辐射对称 等级辐射对称 等级辐射对称 等级辐射对称 外壁纹饰 间断粗条纹,走向不规则 细条纹,走向规则 粗条纹,走向规则 粗条纹,走向略不规则 极轴长(P)/ μm 33.08±1.62a 33.61±1.75a 31.18±1.13b 32.54±1.47ab 赤道轴长(E) / μm 24.59±0.76a 23.21±1.50c 20.50±1.56b 23.00±1.34c P/E 1.35±0.09a 1.45±0.13bc 1.53±0.12b 1.42±0.12ac 条纹宽 / μm 0.34±0.07a 0.16±0.03b 0.35±0.04a 0.34±0.04a -
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