
生物多样性 ›› 2024, Vol. 32 ›› Issue (4): 23352. DOI: 10.17520/biods.2023352 cstr: 32101.14.biods.2023352
李雪萌1,2, 蒋际宝1,2(
), 张曾鲁1,2, 刘晓静3(
), 王亚利4, 吴宜钊1,2, 李银生1,2(
), 邱江平1,2(
), 赵琦1,2,*(
)(
)
收稿日期:2023-09-20
接受日期:2024-03-17
出版日期:2024-04-20
发布日期:2024-04-26
通讯作者:
* E-mail: 基金资助:
Xuemeng Li1,2, Jibao Jiang1,2(
), Zenglu Zhang1,2, Xiaojing Liu3(
), Yali Wang4, Yizhao Wu1,2, Yinsheng Li1,2(
), Jiangping Qiu1,2(
), Qi Zhao1,2,*(
)(
)
Received:2023-09-20
Accepted:2024-03-17
Online:2024-04-20
Published:2024-04-26
Contact:
* E-mail: 摘要:
宝天曼国家级自然保护区独特的地理位置和气候条件使其成为生物多样性研究热点地区之一, 但关于该地区蚯蚓物种多样性的研究却十分匮乏。本研究在系统调查宝天曼国家级自然保护区蚯蚓物种的基础上, 联合5个线粒体基因(COI、COII、ND1、12S、16S)构建蚯蚓系统发育树, 并结合海拔、土壤、凋落物质量等环境因素建立结构方程模型, 初步探讨了宝天曼蚯蚓多样性形成的主要影响因素。研究结果表明: (1)宝天曼国家级自然保护区共记录到蚯蚓3科5属14种, 多为广布种, 其中巨蚓科为优势科; (2)蚯蚓物种数随海拔升高而增加, 但500-1,000 m海拔段的蚯蚓α多样性更高, 分布也更均匀, 符合中间高度膨胀格局; (3)在局域尺度上, 海拔高度对蚯蚓物种多样性和系统发育多样性的影响较大, 其次为土壤性质和凋落物质量。宝天曼国家级自然保护区蚯蚓物种组成及多样性成因的研究丰富了我国蚯蚓物种库和基因库。
李雪萌, 蒋际宝, 张曾鲁, 刘晓静, 王亚利, 吴宜钊, 李银生, 邱江平, 赵琦 (2024) 宝天曼国家级自然保护区蚯蚓物种多样性及其影响因素. 生物多样性, 32, 23352. DOI: 10.17520/biods.2023352.
Xuemeng Li, Jibao Jiang, Zenglu Zhang, Xiaojing Liu, Yali Wang, Yizhao Wu, Yinsheng Li, Jiangping Qiu, Qi Zhao (2024) Earthworm biodiversity and its influencing factors in Baotianman National Nature Reserve. Biodiversity Science, 32, 23352. DOI: 10.17520/biods.2023352.
| 样方 Quadrat | 海拔 Altitude (m) | 地理坐标 Geographical coordinates | 植被 Vegetation | 土壤类型 Soil types | 凋落物厚度 Litter thickness |
|---|---|---|---|---|---|
| 1 | 1,352 | 111.93° E, 33.52° N | 锐齿槲栎和华山松混交林 Mixed forest of Quercus aliena var. acutiserrata and Pinus armandii | 潮湿黑壤 Moist black soil | 约5 cm厚 About 5 cm |
| 2 | 1,334 | 111.93° E, 33.51° N | 锐齿槲栎和华山松混交林 Mixed forest of Quercus aliena var. acutiserrata and Pinus armandii | 干硬黄棕壤 Dry hard yellow brown soil | 约5 cm厚 About 5 cm |
| 3 | 1,389 | 111.93° E, 33.50° N | 锐齿槲栎、短柄枹混交林 Mixed forest of Quercus aliena var. acutiserrata and Q. glandulifera var. brevipetiolata | 黄棕壤 Yellow brown soil | 约5 cm厚 About 5 cm |
| 4 | 1,401 | 111.93° E, 33.50° N | 锐齿槲栎林 Quercus aliena var. acutiserrata forest | 黄棕壤 Yellow brown soil | 约5 cm厚 About 5 cm |
| 5 | 1,360 | 111.74° E, 33.51° N | 短柄枹林 Quercus glandulifera var. brevipetiolata forest | 黄棕壤 Yellow brown soil | 约5 cm厚 About 5 cm |
| 6 | 1,074 | 111.92° E, 33.50° N | 锐齿槲栎、短柄枹混交林, 小溪旁 Mixed forest of Quercus aliena var. acutiserrata and Q. glandulifera var. brevipetiolata, beside the stream | 潮湿黄棕壤, 土层薄 Wet yellow brown soil, thin soil layer | 约20 cm厚, 下层凋落物腐烂 About 20 cm, the litter in the lower layer is rotten |
| 7 | 338 | 112.10° E, 33.57° N | 杨树林 Poplar forest | 黄棕壤 Yellow brown soil | 无凋落物覆盖 No litter cover |
| 8 | 557 | 112.05° E, 33.55° N | 野核桃树、草本植物, 小溪旁 Wild walnut trees, herbs, beside stream | 潮湿黑壤 Wet black soil | 无凋落物覆盖 No litter cover |
| 9 | 613 | 112.05° E, 33.55° N | 香樟和闽楠 Cinnamomum camphora and Phoebe bournei | 黄棕壤 Yellow brown soil | 少量凋落物覆盖 Small amount of litter cover |
表1 河南宝天曼国家级自然保护区蚯蚓、土壤和凋落物采样信息
Table 1 Sampling information of earthworm, soil and litter in Baotianman National Nature Reserve, Henan Province
| 样方 Quadrat | 海拔 Altitude (m) | 地理坐标 Geographical coordinates | 植被 Vegetation | 土壤类型 Soil types | 凋落物厚度 Litter thickness |
|---|---|---|---|---|---|
| 1 | 1,352 | 111.93° E, 33.52° N | 锐齿槲栎和华山松混交林 Mixed forest of Quercus aliena var. acutiserrata and Pinus armandii | 潮湿黑壤 Moist black soil | 约5 cm厚 About 5 cm |
| 2 | 1,334 | 111.93° E, 33.51° N | 锐齿槲栎和华山松混交林 Mixed forest of Quercus aliena var. acutiserrata and Pinus armandii | 干硬黄棕壤 Dry hard yellow brown soil | 约5 cm厚 About 5 cm |
| 3 | 1,389 | 111.93° E, 33.50° N | 锐齿槲栎、短柄枹混交林 Mixed forest of Quercus aliena var. acutiserrata and Q. glandulifera var. brevipetiolata | 黄棕壤 Yellow brown soil | 约5 cm厚 About 5 cm |
| 4 | 1,401 | 111.93° E, 33.50° N | 锐齿槲栎林 Quercus aliena var. acutiserrata forest | 黄棕壤 Yellow brown soil | 约5 cm厚 About 5 cm |
| 5 | 1,360 | 111.74° E, 33.51° N | 短柄枹林 Quercus glandulifera var. brevipetiolata forest | 黄棕壤 Yellow brown soil | 约5 cm厚 About 5 cm |
| 6 | 1,074 | 111.92° E, 33.50° N | 锐齿槲栎、短柄枹混交林, 小溪旁 Mixed forest of Quercus aliena var. acutiserrata and Q. glandulifera var. brevipetiolata, beside the stream | 潮湿黄棕壤, 土层薄 Wet yellow brown soil, thin soil layer | 约20 cm厚, 下层凋落物腐烂 About 20 cm, the litter in the lower layer is rotten |
| 7 | 338 | 112.10° E, 33.57° N | 杨树林 Poplar forest | 黄棕壤 Yellow brown soil | 无凋落物覆盖 No litter cover |
| 8 | 557 | 112.05° E, 33.55° N | 野核桃树、草本植物, 小溪旁 Wild walnut trees, herbs, beside stream | 潮湿黑壤 Wet black soil | 无凋落物覆盖 No litter cover |
| 9 | 613 | 112.05° E, 33.55° N | 香樟和闽楠 Cinnamomum camphora and Phoebe bournei | 黄棕壤 Yellow brown soil | 少量凋落物覆盖 Small amount of litter cover |
| 基因 Gene | 引物名称 Primer | 引物序列 Primer sequence | 参考文献 Reference |
|---|---|---|---|
| COI | LCO1490 | 5°-GGTCAACAAATCATAAAGATATTGG-3° | Folmer et al, |
| COI-E | 5°-TATACTTCTGGGTGTCCGAAGAATCA-3° | Bely & Wray, | |
| COII | COII-F | 5°-GGCACCTATTTGTTAATTAGG-3° | Pérez-Losada et al, |
| COII-R | 5°-GTGAGGCATAGAAATACACC-3° | ||
| ND1 | ND1-F | 5°-GAATAGTGCCACAGGTTTAAAC-3° | Pérez-Losada et al, |
| ND1-R | 5°-TTAACGTCATCAGAGTTATC-3° | ||
| 12S | 12S-F | 5°-CTTAAAGATTTTGGCGGTGTC-3° | Pérez-Losada et al, |
| 12S-R | 5°-CCTTTGCACGGTTAGGATAC-3° | ||
| 16S | 16SarL | 5°-CCGGTCTGAACTCAGATCACGT-3° | Moritz & Hillis, |
表2 线粒体5个基因引物序列
Table 2 The primers of 5 mitochondrial genes
| 基因 Gene | 引物名称 Primer | 引物序列 Primer sequence | 参考文献 Reference |
|---|---|---|---|
| COI | LCO1490 | 5°-GGTCAACAAATCATAAAGATATTGG-3° | Folmer et al, |
| COI-E | 5°-TATACTTCTGGGTGTCCGAAGAATCA-3° | Bely & Wray, | |
| COII | COII-F | 5°-GGCACCTATTTGTTAATTAGG-3° | Pérez-Losada et al, |
| COII-R | 5°-GTGAGGCATAGAAATACACC-3° | ||
| ND1 | ND1-F | 5°-GAATAGTGCCACAGGTTTAAAC-3° | Pérez-Losada et al, |
| ND1-R | 5°-TTAACGTCATCAGAGTTATC-3° | ||
| 12S | 12S-F | 5°-CTTAAAGATTTTGGCGGTGTC-3° | Pérez-Losada et al, |
| 12S-R | 5°-CCTTTGCACGGTTAGGATAC-3° | ||
| 16S | 16SarL | 5°-CCGGTCTGAACTCAGATCACGT-3° | Moritz & Hillis, |
| 物种名 Catalogue | 海拔 Altitude (m) | 数量 Number | |
|---|---|---|---|
| 巨蚓科 Megascolecidae | |||
| 远盲蚓属 Amynthas | |||
| 枯萎远盲蚓 A. emarcidus | 338 | 7 | |
| 无管远盲蚓 A. exsiphonus | 557、613、1,334、1,360、 1,389、1,401 | 44 | |
| 齿盲远盲蚓 A. filicinus | 557 | 5 | |
| 腔蚓属 Metaphire | |||
| Metaphire_W20519-01* | 1,334、1,360、1,389、1,401 | 22 | |
| Metaphire_W21111-05* | 557、613 | 6 | |
| 直隶腔蚓指名亚种 M. tschiliensis tschiliensis | 557、1,334 | 4 | |
| 通俗腔蚓 M. vulgaris | 338 | 1 | |
| 正蚓科 Lumbricidae | |||
| 丛林蚓属 Dendrodrilus | |||
| 红枝蚓 D. rubidus | 1,074、1,352 | 4 | |
| 爱胜蚓属 Eisenia | |||
| Eisenia_W20518-02* | 1,074、1,334、1,352、1,360、 1,389、1,401 | 105 | |
| 链胃蚓科 Moniligastridae | |||
| 杜拉蚓属 Drawida | |||
| 朝鲜杜拉蚓 D. koreana | 1,389 | 1 | |
| Drawida_W20518-01* | 1,334、1,352、1,360 | 13 | |
| Drawida_W20523-03* | 1,074 | 2 | |
| Drawida_W21110-03* | 338、557 | 12 | |
| Drawida_W21112-04* | 613 | 4 | |
表3 河南宝天曼国家级自然保护区蚯蚓物种名录(截至2022年10月)
Table 3 Checklist of earthworm species in Baotianman National Nature Reserve (until Oct. 2022)
| 物种名 Catalogue | 海拔 Altitude (m) | 数量 Number | |
|---|---|---|---|
| 巨蚓科 Megascolecidae | |||
| 远盲蚓属 Amynthas | |||
| 枯萎远盲蚓 A. emarcidus | 338 | 7 | |
| 无管远盲蚓 A. exsiphonus | 557、613、1,334、1,360、 1,389、1,401 | 44 | |
| 齿盲远盲蚓 A. filicinus | 557 | 5 | |
| 腔蚓属 Metaphire | |||
| Metaphire_W20519-01* | 1,334、1,360、1,389、1,401 | 22 | |
| Metaphire_W21111-05* | 557、613 | 6 | |
| 直隶腔蚓指名亚种 M. tschiliensis tschiliensis | 557、1,334 | 4 | |
| 通俗腔蚓 M. vulgaris | 338 | 1 | |
| 正蚓科 Lumbricidae | |||
| 丛林蚓属 Dendrodrilus | |||
| 红枝蚓 D. rubidus | 1,074、1,352 | 4 | |
| 爱胜蚓属 Eisenia | |||
| Eisenia_W20518-02* | 1,074、1,334、1,352、1,360、 1,389、1,401 | 105 | |
| 链胃蚓科 Moniligastridae | |||
| 杜拉蚓属 Drawida | |||
| 朝鲜杜拉蚓 D. koreana | 1,389 | 1 | |
| Drawida_W20518-01* | 1,334、1,352、1,360 | 13 | |
| Drawida_W20523-03* | 1,074 | 2 | |
| Drawida_W21110-03* | 338、557 | 12 | |
| Drawida_W21112-04* | 613 | 4 | |
| 海拔 Altitude (m) | Chao1 多样性指数 Chao1 diversity index | Shannon-Wiener 指数 Shannon- Wiener index | Simpson 多样性指数 Simpson diversity index | Pielou均匀度指数 Pielou evenness index |
|---|---|---|---|---|
| 0-500 | 3.00 ± 0.01 | 0.90 ± 0.02 | 0.56 ± 0.01 | 0.82 ± 0.01 |
| 500-1,000 | 4.00 ± 1.00 | 1.25 ± 0.17 | 0.69 ± 0.04 | 0.93 ± 0.05 |
| 1,000-1,500 | 3.83 ± 1.07 | 0.91 ± 0.11 | 0.50 ± 0.08 | 0.72 ± 0.07 |
表4 河南宝天曼国家级自然保护区不同海拔蚯蚓α多样性指数(平均值 ± 标准差)
Table 4 The α diversity index of earthworms at different altitudes in Baotianman National Nature Reserve (mean ± SD)
| 海拔 Altitude (m) | Chao1 多样性指数 Chao1 diversity index | Shannon-Wiener 指数 Shannon- Wiener index | Simpson 多样性指数 Simpson diversity index | Pielou均匀度指数 Pielou evenness index |
|---|---|---|---|---|
| 0-500 | 3.00 ± 0.01 | 0.90 ± 0.02 | 0.56 ± 0.01 | 0.82 ± 0.01 |
| 500-1,000 | 4.00 ± 1.00 | 1.25 ± 0.17 | 0.69 ± 0.04 | 0.93 ± 0.05 |
| 1,000-1,500 | 3.83 ± 1.07 | 0.91 ± 0.11 | 0.50 ± 0.08 | 0.72 ± 0.07 |
图1 基于5个基因联合构建的蚯蚓系统发育树。分支结点处的数字表示贝叶斯树的支持率。W加四位数字为样本编号。
Fig. 1 Phylogenetic tree of earthworms based on the combination of five genes. The number at the branch node represents the support percent of the Bayesian tree. W plus four digits is the sample number.
| 海拔 Altitude (m) | 系统发育多样性 标准化效应指数 Standardized effect size-phylogenetic diversity (SES-PD) | 净种间亲缘关系指数 Net relatedness index (NRI) | 净最近种间亲缘关系指数 Net nearest taxa index (NTI) | |
|---|---|---|---|---|
| 0-500 | -0.65 ± 0.00 | -0.65 ± 0.00 | -0.61 ± 0.00 | |
| 500-1,000 | -0.94 ± 0.32 | -0.99 ± 0.34 | -0.51 ± 0.14 | |
| 1,000-1,500 | 0.76 ± 0.70 | 0.73 ± 0.66 | 0.96 ± 0.61 | |
表5 宝天曼国家级自然保护区不同海拔段蚯蚓系统发育多样性指数(SES-PD)与系统发育结构(NRI和NTI)
Table 5 Phylogenetic diversity index and phylogenetic structure of earthworms at different altitudes in Baotianman National Nature Reserve
| 海拔 Altitude (m) | 系统发育多样性 标准化效应指数 Standardized effect size-phylogenetic diversity (SES-PD) | 净种间亲缘关系指数 Net relatedness index (NRI) | 净最近种间亲缘关系指数 Net nearest taxa index (NTI) | |
|---|---|---|---|---|
| 0-500 | -0.65 ± 0.00 | -0.65 ± 0.00 | -0.61 ± 0.00 | |
| 500-1,000 | -0.94 ± 0.32 | -0.99 ± 0.34 | -0.51 ± 0.14 | |
| 1,000-1,500 | 0.76 ± 0.70 | 0.73 ± 0.66 | 0.96 ± 0.61 | |
图2 土壤、海拔、凋落物对蚯蚓物种多样性(a)和系统发育多样性(b)影响的结构方程模型。“/”表示与对应变量的对应关系较弱。数字表示各变量间的路径系数。
Fig. 2 Structural equation model depicting the influences of soil, elevation and litter on species diversity (a) and phylogenetic diversity (b) of earthworms in Baotianman National Nature Reserve. “/” indicates a weak correlation with the corresponding variable. The number represents the path coefficient between the variables.
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