基于功能性状及系统发育的亚高寒草甸群落构建

doi:10.17521/cjpe.2017.0211

摘要/Abstract

摘要：

为了解甘南藏族自治州亚高寒草甸植物群落在坡向梯度上的构建机制, 该文选取5个坡向的样地, 构建了植物群落系统发育树, 测定了各坡向土壤环境因子和植物叶片的功能性状, 检验了叶片功能性状的系统发育信号。结果表明: 坡向变化对土壤含水量、土壤养分含量影响显著。大部分植物的叶片特征在不同坡向的差异显著, 叶片干物质含量在南坡、西南坡较高, 比叶面积和叶片氮、磷含量在北坡和西北坡较高。叶片的磷含量具有微弱的系统发育信号, 而叶片干物质含量、比叶面积、叶片的氮含量均没表现出显著的系统发育信号。从南坡到北坡, 群落的系统发育结构由发散到聚集。生境过滤作用是南坡、西南坡群落构建的驱动因素, 种间竞争是北坡和西北坡群落构建的主要驱动力。西坡系统发育指数相反, 其构建机制比较复杂, 可能是几种机制共同作用的结果。

关键词: 群落构建, 系统发育, 功能性状, 坡向, 亚高寒草甸

Abstract:

Aims The community assembly has been a prominent issue in community ecology. This work was intended to explore the mechanisms of the species coexistence and biodiversity in communities. Our objective was to explore the mechanisms of community assembly in subalpine meadow plant communities along slope gradients in Gannan Tibetan Autonomous Prefecture, Gansu Province, Northwest China.
Methods We selected five slope-oriented plots to construct a super-tree representing the species pool. We surveyed the leaf functional traits and soil environmental factors in different slopes. Then we tested the phylogenetic signal of leaf dry matter content (LDMC), specific leaf area (SLA), leaf nitrogen content (LNC) and leaf phosphorus content (LPC).
Important findings The changes of slope aspect had significant influence on soil water content (SWC) and soil nutrient content. Most of the plants leaf functional traits had significant difference along different slope aspects. The LDMC was higher in south and southwest slope than north slope, while SLA, LNC and LPC were relatively high in north and northwest slope. The LPC showed feeble phylogenetic signal, while LDMC, SLA, LNC did not have a significant phylogenetic signal. With changes in the slope aspect from south to north, community phylogenetic structure shifted from over-dispersion to clustered dispersion. In south and southwest slope, habitat filtering was the driving force for community assembly. Interspecific competition was the main driving factor for community assembly in north and northwest slope aspects. But in west slope, two indices showed contrary consequence. This means the process of community assembly in west slope was more complicated and its phylogenetic index may be the result of several mechanisms working together.

Key words: community assembly, phylogenetic, functional traits, slope aspect, subalpine meadow

车应弟, 刘旻霞, 李俐蓉, 焦骄, 肖卫. 基于功能性状及系统发育的亚高寒草甸群落构建. 植物生态学报, 2017, 41(11): 1157-1167. DOI: 10.17521/cjpe.2017.0211

Ying-Di CHE, Min-Xia LIU, Li-Rong LI, Jiao JIAO, Wei XIAO. Exploring the community assembly of subalpine meadow communities based on functional traits and community phylogeny. Chinese Journal of Plant Ecology, 2017, 41(11): 1157-1167. DOI: 10.17521/cjpe.2017.0211

导出引用管理器 EndNote|Ris|BibTeX

链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2017.0211

https://www.plant-ecology.com/CN/Y2017/V41/I11/1157

图/表 7

表1 不同生态性状和群落构建过程结合的群落期望系统发育结构(引自Webb et al., 2002; Kraft et al., 2007)

Table 1 The excepted distribution of community phylogenetic, given various community assembly processes and different evolutionary characteristic of ecological traits after( Webb et al., 2002; Kraft et al., 2007)

群落构建过程 Community assembly processes		生态性状的进化特征 Evolutionary characteristic of ecological trait
群落构建过程 Community assembly processes		性状保守 Traits conserved	性状趋同 Traits convergent
生态位理论 Neutral theory	生境过滤 Habitat filtering	系统发育聚集 Cluster dispersion	系统发育发散 Over dispersion
生态位理论 Neutral theory	极限相似 Limiting similarity	系统发育发散 Over dispersion	系统发育聚集或随机 Clustered or random dispersion
中性理论 Neutral theory	中性作用 Neutral assembly	系统发育随机 Random dispersion	系统发育随机 Random dispersion

表2 研究区样地概况

Table 2 The information of plots in study area

坡向 Aspect	海拔 Altitude (m)	纬度 Latitude (° N)	经度 Longitude (° E)	坡度 Gradient (°)	坡向定位 Aspect orientation (°)
南坡 South slope	3 034	34.65	102.53	32	44
西南坡 Southwest slope	3 032	34.65	102.53	27	88
西坡 West slope	3 031	34.65	102.53	24	132
西北坡 Northwest slope	3 038	34.65	102.53	20	176
北坡 North slope	3 042	34.65	102.53	22	220

图1 不同坡向土壤环境因子的变化(平均值±标准偏差)。不同小写字母表示坡向间差异显著(p < 0.05)。

Fig. 1 Variation on soil environmental factors in different slope aspect (mean ± SD). Different lowercase letters indicate significant difference among different slope aspects (p < 0.05).

图2 植物功能性状沿坡向变化(平均值±标准偏差)。不同小写字母表示坡向间差异显著(p < 0.05)。

Fig. 2 Variation of plant functional traits in different slope aspects (mean ± SD). Different lowercase letters indicate significant difference among different slope aspects (p < 0.05).

表3 甘南藏族自治州亚高寒草甸植物功能性状的系统发育信号

Table 3 The phylogenetic signal of plants functional traits in subalpine meadow communities in Gannan Tibetan Autonomous Prefecture

功能性状 Functional traits	物种数 Species No.	K值 K value	p
叶干物质含量 Leaf dry matter content (mg·g^-1)	60	0.378	0.259
比叶面积 Specific leaf area (cm²·g^-1)	60	0.352	0.167
叶片氮含量 Leaf nitrogen content (mg·g^-1)	60	0.412	0.135
叶片磷含量 Leaf phosphorus content (mg·g^-1)	60	0.235	0.040

图3 甘南藏族自治州亚高寒草甸基于坡向样地60个物种构建的群落系统发育树。

Fig. 3 A super-tree representing the 60 species based on slope-oriented plots in subalpine meadow communities in Gannan Tibetan Autonomous Prefecture.

图4 不同坡向生境内的物种数、谱系多样性指数及系统发育指数(平均值±标准偏差)。不同小写字母表示坡向间差异显著 (p < 0.05)。

Fig. 4 Variation of species, phylogenetic diversity and community phylogenetic relatedness along slope aspect (mean ± SD). Different lowercase letters indicate significant difference among different slope aspects (p < 0.05).

参考文献 40

[1]	Blomberg SP, Garland T, Ives AR (2003). Testing for phylogenetic signal in comparative data: Behavioral traits are more labile.Evolution, International Journal of Organic Evolution, 57, 717-745. DOI URL PMID
[2]	Cao K, Rao MD, Yu JZ, Liu XJ, Mi XC, Chen JH (2013). The phylogenetic signal of functional traits and their effects on community structure in an evergreen broad-leaved forest.Biodiversity Science, 21, 564-571. (in Chinese with English abstract)[曹科, 饶米德, 余建中, 刘晓娟, 米湘成, 陈建华 (2013). 古田山木本植物功能性状的系统发育信号及其对群落结构的影响. 生物多样性, 21, 564-571.] DOI URL
[3]	Chai YF, Yue M (2016). Research advances in plant community assembly mechanisms.Acta Ecologica Sinica, 36, 4557-4572. (in Chinese with English abstract)[柴永福, 岳明 (2016). 植物群落构建机制研究进展. 生态学报, 36, 4557-4572.] DOI URL
[4]	Ding Y, Zang R, Letcher SG, Liu S, He F (2012). Disturbance regime changes the trait distribution, phylogenetic structure and community assembly of tropical rain forests.Oikos, 121, 1263-1270. DOI URL
[5]	Dong SK, Tang L, Zhang XF, Liu SL, Liu QR, Su XK, Zhang Y, Wu XY, Zhao ZZ, Li Y, Sha W (2017). Relationship between plant species diversity and functional diversity in alpine grasslands.Acta Ecologica Sinica, 37, 1472-1483. (in Chinese with English abstract)[董世魁, 汤琳, 张相锋, 刘世梁, 刘全儒, 苏旭坤, 张勇, 武晓宇, 赵珍珍, 李钰, 沙威 (2017). 高寒草地植物物种多样性与功能多样性的关系. 生态学报, 37, 1472-1483.] DOI URL
[6]	Fang S, Yuan ZQ, Lin F, Ye J, Hao ZQ, Wang XG (2014). Functional and phylogenetic structures of woody plants in broad-leaved Korean pine mixed forest in Changbai Mountains,Chinese Science Bulletin, 59, 2342-2348. (in Chinese with English abstract)[房帅, 原作强, 蔺非, 叶吉, 郝占庆, 王绪高 (2014). 长白山阔叶红松林木本植物系统发育与功能性状结构. 科学通报, 59, 2342-2348.]
[7]	Faith DP (1992). Conservation evaluation and phylogenetic diversity.Biological Conservation, 61, 1-10. DOI URL
[8]	Felsenstein J (1985). Phylogenies and the comparative method.The American Naturalist, 125, 1-15.
[9]	Gerhold P, Price JN, Pussa K, Kalamess R, Aher K, Kaasik A (2013). Functional and phylogenetic community assembly linked to changes in species diversity in a long-term resource manipulation experiment.Journal of Vegetation Science, 24, 843-852. DOI URL
[10]	Gong X, Brueck H, Giese KM, Zhang L, Sattelmacher B, Lin S (2008). Slope aspect has effects on productivity and species composition of hilly grassland in the Xilin River Basin, Inner Mongolia, China.Journal of Arid Environments, 72, 483-493. DOI URL
[11]	Gong X, Wang G (2015). Based on trait and community phylogenetic community assembly along a slope aspect gradient in an alpine meadow. (in Chinese with English abstract)[宫骁, 王刚 (2015). 沿坡向梯度亚高寒草甸基于功能性状和群落系统发育结构的群落构建机制. .]
[12]	Gravel D, Canham CD, Beaudet M, Messier C (2006). Reconciling niche and neutrality: The continuum hypothesis.Ecology Letters, 9, 399-409.
[13]	Honorio Coronado EN, Dexter KG, Pennington RT, Chave J, Lewis SL, Alexiades MN (2015). Phylogenetic diversity of Amazonian tree communities.Diversity and Distributions, 21, 1295-1307. DOI URL
[14]	Hubbell SP (2001). The Unified Neutral Theory of Biodiversity and Biogeography. Princeton University Press, Princeton, USA.
[15]	Kraft NJ, Cornwell WK, Webb CO, Ackerly DD (2007). Trait evolution, community assembly, and the phylogenetic structure of ecological communities. The American Naturalist, 170, 271-283. DOI URL PMID
[16]	Li HQ, Mao SJ, Zhu JB, Yang YS, He HD, LI YN (2017). Effects of grazing intensity on the ecological stoichiometry characteristics of alpine meadow.Pratacultural Science, 34, 449-455. (in Chinese with English abstract)[李红琴, 毛绍娟, 祝景彬, 杨永胜, 贺慧丹, 李英年 (2017). 放牧强度对高寒草甸群落碳氮磷化学计量特征的影响. 草业学报, 34, 449-455.] DOI URL
[17]	Liu JJ, Zhang X, Song F, Zhou S, Cadotte MM, Bradshaw CJ (2015). Explaining maximum variation in productivity requires phylogenetic diversity and single functional traits.Ecology, 96, 176-183. DOI URL PMID
[18]	Liu MX, Liu JJ, Du XG, Zhang XG (2010). Fitting different models to species abundance distribution patterns in three plant communities in sub-alpine meadow.Acta Ecologica Sinica, 30, 6935-6942. (in Chinese with English abstract)[刘梦雪, 刘佳佳, 杜晓光, 郑小刚 (2010). 亚高寒草甸不同生境植物群落物种多度分布格局的拟合. 生态学报, 30, 6935-6942.]
[19]	Liu MX, Ma JZ (2012). Responses of plant functional traits and soil factors to slope aspect in alpine meadow of South Gansu, Northwest China.Chinese of Applied Ecology, 23, 3295-3300. (in Chinese with English abstract)[刘旻霞, 马建祖 (2012). 甘南高寒草甸植物功能性状和土壤因子对坡向的响应. 应用生态学报, 23, 3295-3300.]
[20]	Liu MX, Wang G, Sheng HM (2013). Environmental factors characteristic of sunny and shady slope gradients in relation to above-ground biomass and species richness in an alpine meadow. Journal of Lanzhou University (Natural Science), 49, 76-81. (in Chinese with English abstract)[刘旻霞, 王刚, 盛红梅 (2013). 高寒草甸阳坡-阴坡梯度上环境因子特征及其与地上生物量和物种丰富度的关系. 兰州大学学报(自然科学版), 49, 76-81.]
[21]	Liu XJ, Ma KP (2015). Plant functional traits—Concepts, applications and future directions.Scientia Sinica Vitae, 45, 325-339. (in Chinese with English abstract)[刘晓娟, 马克平 (2015). 植物功能性状研究进展. 中国科学: 生命科学, 45, 325-339.]
[22]	Lu MM, Huang XC, Ci XQ, Yang GP, Li J (2014). Phylogenetic community structure of subtropical forests along elevational gradients in Ailao Mountains of Southwest China.Biodiversity Science, 22, 438-448. (in Chinese with English abstract)[卢孟孟, 黄小翠, 慈秀芹, 杨国平, 李捷 (2014). 沿海拔梯度变化的哀牢山亚热带森林群落系统发育结构. 生物多样性, 22, 438-449.]
[23]	Lü MQ, Zhu ZH, Li YN, Yao TH, Pan SY, Kong BB (2014). Assembly patterns of plant functional traits in alpine meadow under disturbances by mowing and fertilization.Chinese Journal of Plant Ecology, 38, 916-928. (in Chinese with English abstract)[吕美强, 朱志红, 李英年, 姚天华, 潘石玉, 孔彬彬 (2014). 刈割与施肥干扰下高寒草甸植物功能性状的构建模式. 植物生态学报, 38, 916-928.] DOI URL
[24]	Meng TT, Ni J, Wang GH (2007). Plant functional traits, environments and ecosystem functioning.Journal of Plant Ecology (Chinese Version), 31, 150-165. (in Chinese with English abstract)[孟婷婷, 倪健, 王国宏 (2007). 植物功能性状与环境和生态系统. 植物生态学报, 31, 150-165.] DOI URL
[25]	Nie YY, Li XE, Wang G (2010). Variation mode of α diversity and β diversity of plant community of different habitat gradients from south-facing slope to north-facing slope and its relation with different environmental factors.Journal of Lanzhou University (Natural Science), 46, 73-79. (in Chinese with English abstract)[聂莹莹, 李新娥, 王刚 (2010). 阳坡—阴坡生境梯度上植物群落α多样性与β多样性的变化模式与环境因子的关系. 兰州大学学报: 自然科学版, 46, 73-79.]
[26]	Niu HY, Wang ZF, Lian JY, Ye WH, Shen H (2011). New progress in community assembly: Community phylogenetic structure combining evolution and ecology.Biodiversity Science, 19, 275-283. (in Chinese with English abstract)[牛红玉, 王峥峰, 练琚愉, 叶万辉, 沈浩 (2011). 群落构建研究的新进展: 进化和生态相结合的群落谱系结构研究. 生物多样性, 19, 275-283.] DOI URL
[27]	Niu KC, Liu YN, Shen ZH, He FL, Fang JY (2009). Community assemble: The relative important of neutral theory and niche theory.Biodiversity Science, 17, 579-593. (in Chinese with English abstract)[牛克昌, 刘怿宁, 沈泽昊, 何芳良, 方精云 (2009). 群落构建的中性理论和生态位理论. 生物多样性, 17, 579-593.] DOI URL
[28]	Rosindell J, Hubbell SP, Etienne RS (2011). The unified neutral theory of biodiversity and biogeography, at age ten.Trends in Ecology & Ecology, 26, 340-348. DOI URL
[29]	Simberloff DS (1970). Taxonomic diversity of island biotas.Evolution, 24, 23-47. DOI URL PMID
[30]	Swenson NG (2013). The assembly of tropical tree communities—The advances and shortcomings of phylogenetic and functional trait analyses.Ecography, 36, 264-276.
[31]	Wang HD, Zhang LL, Zhu ZH (2013). Effects of clipping and fertilizing on the relationships between species diversity and ecosystem functioning and mechanisms of community stability in alpine meadow.Chinese Journal of Plant Ecology, 37, 279-295. (in Chinese with English abstract)[王海东, 张璐璐, 朱志红 (2013). 刈割、施肥对高寒草甸物种多样性与生态系统功能关系的影响及群落稳定性机制. 植物生态学报, 37, 279-295.] DOI URL
[32]	Webb CO (2000). Exploring the phylogenetic structure of ecological communities: An example for rain forest trees.The American Naturalist, 156, 145-155.
[33]	Webb CO, Ackerly DD, Mcpeek MA, Donoghue MJ (2002). Phylogenies and community ecology.Annual Review of Ecology & Systematics, 33, 475-505.
[34]	Webb CO, Donoghue MJ (2005). Phylomatic: Tree assembly for applied phylogenetics.Molecular Ecology Notes, 5, 181-183.
[35]	Webb CO, Ackerly DD, Kembel SW (2008). Phylocom: Software for the analysis of phylogenetic community structure and trait evolution.Bioinformatics, 24, 2098-2100. DOI URL PMID
[36]	Xiao XS, Wang G (2015). Based on trait and community phylogeny community assembly along a slope aspect gradient in an alpine meadow. (in Chinese with English abstract)[肖欣爽, 王刚 (2015). 基于功能性状的亚高寒草甸阳坡-阴坡生境梯度上群落构建机制的研究. .]
[37]	Yang J, Lu MM, Cao M, Li J, Lin LX (2014). Phylogenetic and functional alpha and beta diversity in mid-mountain humid evergreen broad-leaved forest.Chinese Science Bulletin, 59, 2349-2358. (in Chinese with English abstract)[杨洁, 卢孟孟, 曹敏, 李捷, 林露湘 (2014). 中山湿性常绿阔叶林系统发育和功能性状的α及β多样性. 科学通报, 59, 2349-2358.] DOI URL
[38]	Zhang H, John R, Peng ZC, Yuan JL, Chu CG, Du GZ, Zhou SR (2012). The relationship between species richness and evenness in plant communities along a successional gradient: A study from sub-alpine meadows of the eastern Qinghai-Tibetan Plateau, China.PLOS ONE, 7, e49024. doi: 10.1371/journal.pone.0049024. DOI URL PMID
[39]	Zhang H, Qi W, John R, Wang WB, Song FF, Zhou SR (2015). Using functional trait diversity to evaluate the contribution of multiple ecological processes to community assembly during succession.Ecography, 38, 1176-1186. DOI URL
[40]	Zhu GL, Li J, Wei XH, He NP (2017). Longitudinal patterns of productivity and plant diversity in Tibetan alpine grasslands.Journal of Natural Resources, 32, 210-222. (in Chinese with English abstract)[朱桂丽, 李杰, 魏学红, 何念鹏 (2017). 青藏高寒草地植被生产力与生物多样性的经度格局. 自然资源学报, 32, 210-222.]