园艺学报 ›› 2021, Vol. 48 ›› Issue (3): 539-552.doi: 10.16420/j.issn.0513-353x.2020-0474
虞钦岚1, 刘守赞2,*(), 徐韧析谋1, 潘晨航1, 颜忆铭1, 王义英1, 夏国华1,**()
收稿日期:
2020-08-12
出版日期:
2021-03-25
发布日期:
2021-04-02
通讯作者:
刘守赞,夏国华
E-mail:szliu@zafu.edu.cn;zjfc_ghxia@126.com
基金资助:
YU Qinlan1, LIU Shouzan2,*(), XU Renximou1, PAN Chenhang1, YAN Yiming1, WANG Yiying1, XIA Guohua1,**()
Received:
2020-08-12
Online:
2021-03-25
Published:
2021-04-02
Contact:
LIU Shouzan,XIA Guohua
E-mail:szliu@zafu.edu.cn;zjfc_ghxia@126.com
摘要:
野外调查发现,珍稀濒危植物江南牡丹草(Gymnospermium kiangnanense)资源量日益萎缩,其药用和园艺观赏开发受到较大限制。为揭示江南牡丹草的种群结构和繁育系统特征,对分布于浙江淳安临岐和诸暨璜山的2个天然种群开展了样地调查,编制了种群特定时间生命表,分析了存活曲线、生存函数,运用时间序列模型预测种群数量动态,对杂交指数、花粉/胚珠比、花粉活力、柱头可授性以及传粉昆虫等繁育系统特性进行测定。结果表明:(1)淳安种群为中龄级比例较大的纺锤形结构,诸暨种群幼龄级和老龄级比例较大、中龄级严重缺乏;2个种群均表现为衰退型,存活曲线均趋于Deevey-Ⅱ型;(2)花部表现出泛化传粉类型的特征,P/O值为7 920,繁育系统类型为异交,需要传粉者;主要传粉昆虫为中华蜜蜂、黑带食蚜蝇等;其通过短期内高密度集中开花的开花式样、泛化传粉系统以及早春空白生态位来提高传粉效率,保障传粉成功。
中图分类号:
虞钦岚, 刘守赞, 徐韧析谋, 潘晨航, 颜忆铭, 王义英, 夏国华. 珍稀濒危植物江南牡丹草种群结构和繁育系统研究[J]. 园艺学报, 2021, 48(3): 539-552.
YU Qinlan, LIU Shouzan, XU Renximou, PAN Chenhang, YAN Yiming, WANG Yiying, XIA Guohua. Population Structure and Breeding Systems of Rare and Endangered Gymnospermium kiangnanense(Berberidaceae)[J]. Acta Horticulturae Sinica, 2021, 48(3): 539-552.
图2 江南牡丹草种群龄级结构 Ⅰ~Ⅹ:10个径级,级差5 mm。下同。
Fig. 2 The age-class structure of Gymnospermium kiangnanense population Ⅰ~Ⅹ:There are 10 diameter classes and the grade difference is 5 mm. The same below.
种群 Population | 龄级 Age class | 径级/mm(TD) Tuber diameter | Ax | ax | lx | ln(lx) | dx | qx | Lx | Tx | ex | Kx | |||||||||||||
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
淳安 Chun'an | Ⅰ | TD < 10 | 29 | 33 | 1 000 | 6.908 | 30 | 0.030 | 985 | 8 270 | 8.270 | 0.030 | |||||||||||||
Ⅱ | 10 ≤ TD < 15 | 24 | 32 | 970 | 6.878 | 29 | 0.030 | 956 | 7 285 | 7.506 | 0.030 | ||||||||||||||
Ⅲ | 15 ≤ TD < 20 | 39 | 32 | 942 | 6.848 | 28 | 0.030 | 928 | 6 328 | 6.720 | 0.030 | ||||||||||||||
Ⅳ | 20 ≤ TD < 25 | 33 | 31 | 914 | 6.818 | 27 | 0.030 | 900 | 5 401 | 5.909 | 0.030 | ||||||||||||||
Ⅴ | 25 ≤ TD < 30 | 30 | 30 | 887 | 6.788 | 26 | 0.030 | 874 | 4 500 | 5.074 | 0.030 | ||||||||||||||
Ⅵ | 30 ≤ TD < 35 | 30 | 29 | 861 | 6.758 | 25 | 0.030 | 848 | 3 626 | 4.213 | 0.030 | ||||||||||||||
Ⅶ | 35 ≤ TD < 40 | 40 | 28 | 835 | 6.728 | 25 | 0.030 | 823 | 2 778 | 3.326 | 0.030 | ||||||||||||||
Ⅷ | 40 ≤ TD < 45 | 39 | 27 | 811 | 6.698 | 24 | 0.030 | 799 | 1 955 | 2.412 | 0.030 | ||||||||||||||
Ⅸ | 45 ≤ TD < 50 | 20 | 26 | 787 | 6.668 | 23 | 0.030 | 775 | 1 157 | 1.470 | 0.030 | ||||||||||||||
Ⅹ | TD ≥ 50 | 18 | 26 | 763 | 6.638 | 763 | — | 382 | 382 | 0.500 | — | ||||||||||||||
诸暨 Zhuji | Ⅰ | TD < 10 | 45 | 25 | 1 000 | 6.908 | 227 | 0.227 | 886 | 4 543 | 4.543 | 0.258 | |||||||||||||
Ⅱ | 10 ≤ TD < 15 | 17 | 19 | 773 | 6.650 | 133 | 0.172 | 706 | 3 657 | 4.734 | 0.189 | ||||||||||||||
Ⅲ | 15 ≤ TD < 20 | 2 | 16 | 639 | 6.461 | 94 | 0.148 | 592 | 2 951 | 4.615 | 0.160 | ||||||||||||||
Ⅳ | 20 ≤ TD < 25 | 3 | 14 | 545 | 6.301 | 73 | 0.134 | 508 | 2 359 | 4.327 | 0.144 | ||||||||||||||
Ⅴ | 25 ≤ TD < 30 | 2 | 12 | 472 | 6.157 | 60 | 0.127 | 442 | 1 850 | 3.921 | 0.136 | ||||||||||||||
Ⅵ | 30 ≤ TD < 35 | 4 | 10 | 412 | 6.021 | 51 | 0.123 | 387 | 1 408 | 3.418 | 0.131 | ||||||||||||||
Ⅶ | 35 ≤ TD < 40 | 1 | 9 | 361 | 5.890 | 44 | 0.121 | 339 | 1 022 | 2.827 | 0.129 | ||||||||||||||
Ⅷ | 40 ≤ TD < 45 | 1 | 8 | 318 | 5.761 | 39 | 0.122 | 298 | 682 | 2.148 | 0.130 | ||||||||||||||
Ⅸ | 45 ≤ TD < 50 | 4 | 7 | 279 | 5.631 | 35 | 0.124 | 262 | 384 | 1.376 | 0.132 | ||||||||||||||
Ⅹ | TD ≥ 50 | 48 | 6 | 244 | 5.499 | 244 | — | 122 | 122 | 0.500 | — |
表1 江南牡丹草淳安种群和诸暨种群静态生命表
Table 1 Life table of Chun'an population and Zhuji population of Gymnospermium kiangnanense
种群 Population | 龄级 Age class | 径级/mm(TD) Tuber diameter | Ax | ax | lx | ln(lx) | dx | qx | Lx | Tx | ex | Kx | |||||||||||||
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
淳安 Chun'an | Ⅰ | TD < 10 | 29 | 33 | 1 000 | 6.908 | 30 | 0.030 | 985 | 8 270 | 8.270 | 0.030 | |||||||||||||
Ⅱ | 10 ≤ TD < 15 | 24 | 32 | 970 | 6.878 | 29 | 0.030 | 956 | 7 285 | 7.506 | 0.030 | ||||||||||||||
Ⅲ | 15 ≤ TD < 20 | 39 | 32 | 942 | 6.848 | 28 | 0.030 | 928 | 6 328 | 6.720 | 0.030 | ||||||||||||||
Ⅳ | 20 ≤ TD < 25 | 33 | 31 | 914 | 6.818 | 27 | 0.030 | 900 | 5 401 | 5.909 | 0.030 | ||||||||||||||
Ⅴ | 25 ≤ TD < 30 | 30 | 30 | 887 | 6.788 | 26 | 0.030 | 874 | 4 500 | 5.074 | 0.030 | ||||||||||||||
Ⅵ | 30 ≤ TD < 35 | 30 | 29 | 861 | 6.758 | 25 | 0.030 | 848 | 3 626 | 4.213 | 0.030 | ||||||||||||||
Ⅶ | 35 ≤ TD < 40 | 40 | 28 | 835 | 6.728 | 25 | 0.030 | 823 | 2 778 | 3.326 | 0.030 | ||||||||||||||
Ⅷ | 40 ≤ TD < 45 | 39 | 27 | 811 | 6.698 | 24 | 0.030 | 799 | 1 955 | 2.412 | 0.030 | ||||||||||||||
Ⅸ | 45 ≤ TD < 50 | 20 | 26 | 787 | 6.668 | 23 | 0.030 | 775 | 1 157 | 1.470 | 0.030 | ||||||||||||||
Ⅹ | TD ≥ 50 | 18 | 26 | 763 | 6.638 | 763 | — | 382 | 382 | 0.500 | — | ||||||||||||||
诸暨 Zhuji | Ⅰ | TD < 10 | 45 | 25 | 1 000 | 6.908 | 227 | 0.227 | 886 | 4 543 | 4.543 | 0.258 | |||||||||||||
Ⅱ | 10 ≤ TD < 15 | 17 | 19 | 773 | 6.650 | 133 | 0.172 | 706 | 3 657 | 4.734 | 0.189 | ||||||||||||||
Ⅲ | 15 ≤ TD < 20 | 2 | 16 | 639 | 6.461 | 94 | 0.148 | 592 | 2 951 | 4.615 | 0.160 | ||||||||||||||
Ⅳ | 20 ≤ TD < 25 | 3 | 14 | 545 | 6.301 | 73 | 0.134 | 508 | 2 359 | 4.327 | 0.144 | ||||||||||||||
Ⅴ | 25 ≤ TD < 30 | 2 | 12 | 472 | 6.157 | 60 | 0.127 | 442 | 1 850 | 3.921 | 0.136 | ||||||||||||||
Ⅵ | 30 ≤ TD < 35 | 4 | 10 | 412 | 6.021 | 51 | 0.123 | 387 | 1 408 | 3.418 | 0.131 | ||||||||||||||
Ⅶ | 35 ≤ TD < 40 | 1 | 9 | 361 | 5.890 | 44 | 0.121 | 339 | 1 022 | 2.827 | 0.129 | ||||||||||||||
Ⅷ | 40 ≤ TD < 45 | 1 | 8 | 318 | 5.761 | 39 | 0.122 | 298 | 682 | 2.148 | 0.130 | ||||||||||||||
Ⅸ | 45 ≤ TD < 50 | 4 | 7 | 279 | 5.631 | 35 | 0.124 | 262 | 384 | 1.376 | 0.132 | ||||||||||||||
Ⅹ | TD ≥ 50 | 48 | 6 | 244 | 5.499 | 244 | — | 122 | 122 | 0.500 | — |
龄级 Age class | 淳安种群Chun'an population | 诸暨种群Zhuji population | ||||||
---|---|---|---|---|---|---|---|---|
原始数据 Primary data | M2[ | M5[ | M8[ | 原始数据 Primary data | M2[ | M5[ | M8[ | |
Ⅰ | 29 | 45 | ||||||
Ⅱ | 24 | 27 | 17 | 31 | ||||
Ⅲ | 39 | 32 | 2 | 10 | ||||
Ⅳ | 33 | 36 | 3 | 3 | ||||
Ⅴ | 30 | 32 | 31 | 2 | 3 | 14 | ||
Ⅵ | 30 | 30 | 31 | 4 | 3 | 6 | ||
Ⅶ | 40 | 35 | 34 | 1 | 3 | 2 | ||
Ⅷ | 39 | 40 | 34 | 33 | 1 | 1 | 2 | 9 |
Ⅸ | 20 | 30 | 32 | 32 | 4 | 3 | 2 | 4 |
Ⅹ | 18 | 19 | 29 | 31 | 48 | 26 | 12 | 8 |
总计Total | 302 | 279 | 192 | 96 | 127 | 81 | 38 | 22 |
表2 江南牡丹草淳安种群和诸暨种群个体数量动态变化的时间序列分析
Table 2 The time sequence analysis of Gymnospermium kiangnanense population
龄级 Age class | 淳安种群Chun'an population | 诸暨种群Zhuji population | ||||||
---|---|---|---|---|---|---|---|---|
原始数据 Primary data | M2[ | M5[ | M8[ | 原始数据 Primary data | M2[ | M5[ | M8[ | |
Ⅰ | 29 | 45 | ||||||
Ⅱ | 24 | 27 | 17 | 31 | ||||
Ⅲ | 39 | 32 | 2 | 10 | ||||
Ⅳ | 33 | 36 | 3 | 3 | ||||
Ⅴ | 30 | 32 | 31 | 2 | 3 | 14 | ||
Ⅵ | 30 | 30 | 31 | 4 | 3 | 6 | ||
Ⅶ | 40 | 35 | 34 | 1 | 3 | 2 | ||
Ⅷ | 39 | 40 | 34 | 33 | 1 | 1 | 2 | 9 |
Ⅸ | 20 | 30 | 32 | 32 | 4 | 3 | 2 | 4 |
Ⅹ | 18 | 19 | 29 | 31 | 48 | 26 | 12 | 8 |
总计Total | 302 | 279 | 192 | 96 | 127 | 81 | 38 | 22 |
图5 江南牡丹草精细解剖图 A1:花序;A2:花枝;B:花离析;C1 ~ C3:花苞;C4 ~ C5:花正面观;C6:花背面观;D:子房纵剖;E:小苞片;F:花(除萼片);G:花瓣(蜜腺状);H:雄蕊;I:萼片和花瓣。
Fig. 5 Fine structure and morphology of Gymnospermium kiangnanense A1:Inflorescence;A2:Flower branch;B:Anatomy of the flower;C1-C3:Alabastrum;C4-C5:Frontal view of the flower;C6:Dorsal view of the flower;D:Longitudinal profile of ovary;E:Bractlet;F:Flower(except calyx);G:Petal(nectary-like);H:Stamen;I:Sepal and petal.
花器官 Flower characteristics | 淳安种群 Chun'an population | 诸暨种群 Zhuji population | 花器官 Flower characteristics | 淳安种群 Chun'an population | 诸暨种群 Zhuji population |
---|---|---|---|---|---|
花序长度/mm Length of inflorescence | 91.50 ± 17.37 | 140.60 ± 26.12 ** | 花瓣数 Number of petal | 5.96 ± 0.19 | 5.93 ± 0.25 |
小花数 Number of flower | 15.07 ± 1.98 | 15.77 ± 3.13 | 花瓣长/cm Length of petal | 2.34 ± 0.46 | 2.03 ± 0.56 * |
花梗长/cm Length of pedicel | 33.34 ± 4.79 | 12.90 ± 4.87 ** | 花瓣宽/cm Width of petal | 1.66 ± 0.34 | 1.64 ± 0.25 |
小苞片长/cm Length of bractlet | 6.65 ± 0.90 | 5.15 ± 1.04 ** | 雄蕊数 Number of stamen | 5.93 ± 0.27 | 6.00 ± 0.00 |
小苞片宽/cm Width of bractlet | 5.29 ± 1.37 | 4.39 ± 1.26 * | 花丝长/cm Length of filament | 2.86 ± 0.37 | 2.83 ± 0.54 |
花冠口直径/cm Diameter of corolla | 17.30 ± 1.68 | 16.36 ± 2.18 | 花药长/cm Length of anther | 2.43 ± 0.42 | 2.18 ± 0.35 * |
花萼数 Number of calyx | 6.56 ± 0.58 | 6.90 ± 0.31 ** | 子房长/cm Length of ovary | 2.86 ± 0.45 | 3.71 ± 1.00 ** |
花萼长/cm Length of calyx | 9.23 ± 0.98 | 8.30 ± 1.02 ** | 子房直径/cm Diameter of ovary | 1.71 ± 0.36 | 1.98 ± 0.37 ** |
花萼宽/cm Width of calyx | 5.29 ± 0.90 | 4.57 ± 0.71 ** | 子房托长度/cm Length of ovary base | 1.45 ± 0.28 | 0.71 ± 0.26 ** |
表3 花部器官特征统计值
Table 3 Statistical value of flower characteristics
花器官 Flower characteristics | 淳安种群 Chun'an population | 诸暨种群 Zhuji population | 花器官 Flower characteristics | 淳安种群 Chun'an population | 诸暨种群 Zhuji population |
---|---|---|---|---|---|
花序长度/mm Length of inflorescence | 91.50 ± 17.37 | 140.60 ± 26.12 ** | 花瓣数 Number of petal | 5.96 ± 0.19 | 5.93 ± 0.25 |
小花数 Number of flower | 15.07 ± 1.98 | 15.77 ± 3.13 | 花瓣长/cm Length of petal | 2.34 ± 0.46 | 2.03 ± 0.56 * |
花梗长/cm Length of pedicel | 33.34 ± 4.79 | 12.90 ± 4.87 ** | 花瓣宽/cm Width of petal | 1.66 ± 0.34 | 1.64 ± 0.25 |
小苞片长/cm Length of bractlet | 6.65 ± 0.90 | 5.15 ± 1.04 ** | 雄蕊数 Number of stamen | 5.93 ± 0.27 | 6.00 ± 0.00 |
小苞片宽/cm Width of bractlet | 5.29 ± 1.37 | 4.39 ± 1.26 * | 花丝长/cm Length of filament | 2.86 ± 0.37 | 2.83 ± 0.54 |
花冠口直径/cm Diameter of corolla | 17.30 ± 1.68 | 16.36 ± 2.18 | 花药长/cm Length of anther | 2.43 ± 0.42 | 2.18 ± 0.35 * |
花萼数 Number of calyx | 6.56 ± 0.58 | 6.90 ± 0.31 ** | 子房长/cm Length of ovary | 2.86 ± 0.45 | 3.71 ± 1.00 ** |
花萼长/cm Length of calyx | 9.23 ± 0.98 | 8.30 ± 1.02 ** | 子房直径/cm Diameter of ovary | 1.71 ± 0.36 | 1.98 ± 0.37 ** |
花萼宽/cm Width of calyx | 5.29 ± 0.90 | 4.57 ± 0.71 ** | 子房托长度/cm Length of ovary base | 1.45 ± 0.28 | 0.71 ± 0.26 ** |
图8 江南牡丹草开花动态观察 A:蕾期;B:散粉前期;C:散粉初期;D:散粉中期;E:散粉末期;F:末花期。
Fig. 8 Flowering dynamic observation of Gymnospermium kiangnanense A:Bud stage;B:Pre-dehiscence;C:Initial dehiscence;D:Full dehiscence;E:Terminal dehiscence;F:Fade stage.
图10 江南牡丹草柱头可授性变化 A:散粉前期;B:散粉初期;C:散粉中期;D:散粉末期。
Fig. 10 Stigma reproducibility of Gymnospermium kiangnanense A:Pre-dehiscence;B:Initial dehiscence;C:Full dehiscence;D:Terminal dehiscence.
图12 江南牡丹草访花昆虫 A:中华蜜蜂;B:隧蜂属;C:黑带食蚜蝇;D:斜斑鼓额食蚜蝇。
Fig. 12 The species of flower-visiting insect of Gymnospermium kiangnanense A:Apis cerana;B:Halictussp.;C:Episyrphus balteatus;D:Scaeva pyrastri.
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