Phylogenetic structure and diversity of herbaceous communities in the conifer forests along an elevational gradient in Luya Mountain, Shanxi, China

doi:10.17521/cjpe.2016.0247

Abstract

Abstract:

Aims Incorporating phylogenetic data in the studies of species diversity patterns along elevational gradients can bridge the gap between ecological and evolutionary processes, and thus shed light on the issues related to community assemblage. We aim is to explore the elevational patterns of phylogenetic relatedness and phylodiversity in the herbaceous angiosperm assemblages of alpine conifer forest within mountain ecosystem, and to quantify the relationship between the patterns and habitat factors. Methods We sampled 17 plots (20 m × 30 m) in the coniferous forest communities at ca. 50 m altitudinal intervals along the elevation gradient. In each plot, we documented all species encountered and the environmental conditions. We used the Net Relatedness Index (NRI) and Nearest Taxon Index (NTI) to quantify the phylogenetic structures of each herbaceous assemblage, and used the Mean Pairwise Distance and Mean Nearest Taxon Distance to quantify phylogenetic beta diversity. Ordinary least square regression and multiple regression on distance matrices were employed respectively to explore the elevational trends of phylogenetic structure and phylobetadiversity. We analyzed the taxonomic composition of the herbaceous assemblages within forests, and demonstrated the relationship between the clustering of clades and the significant habitat descriptors using principal coordinates of phylogenetic structure (PCPS). Important findings The result showed that the herbaceous communities tended to be more phylogenetically overdispersed at lower elevations, suggesting that intraspecific competition potentially influences the local assemblages. In contrast, species occurring at high-elevation sites tended to be more closely related, implying that these communities are structured primarily by environmental filtering. However, we found that all of the NRI (or NTI) were confined within 95% confidence intervals, suggesting strong contributions of stochastic processes on species assembly. Phylogenetic beta diversity significantly increased with the elevational distance between community pairs, also suggesting that habitat filtering probably played an important role on structuring the herbaceous communities. The first two axes of PCPS contained 55.9% of total variation in phylogeny-weighted species composition, and were both significantly related with elevation and arboreal basal area. We found that species of large families, including Asteraceae and Poaceae, were phylogenetically clustered at high elevations, whereas the others were phylogenetically overdispersed at low elevation region. Our findings suggest that exploring the patterns of phylogenetic structures across elevational gradients is important, which can provide insights into the underlying mechanisms shaping community composition within montane ecosystems.

Key words: diversity elevational pattern, phylogenetic alpha diversity, phylogenetic beta diversity, principal coordinates of phylogenetic structure, herbaceous plants

Ming-Fei ZHAO, Feng XUE, Yu-Hang WANG, Guo-Yi WANG, Kai-Xiong XING, Mu-Yi KANG, Jing-Lan WANG. Phylogenetic structure and diversity of herbaceous communities in the conifer forests along an elevational gradient in Luya Mountain, Shanxi, China[J]. Chin J Plan Ecolo, 2017, 41(7): 707-715.

Add to citation manager EndNote|Ris|BibTeX

URL: https://www.plant-ecology.com/EN/10.17521/cjpe.2016.0247

https://www.plant-ecology.com/EN/Y2017/V41/I7/707

Figures/Tables 4

Fig. 1 Elevational patterns of the phylogenetic structures of herbaceous assemblages in the coniferous forest plots in Luya Mountain.

Fig. 2 The relationships of phylogenetic beta diversity with elevational distance for the herbaceous plants assemblages in the coniferous forest plots in Luya Mountain.

Fig. 3 Scatter diagram between the first two axes of the principal coordinates of phylogenetic structure (PCPS) for herbaceous plants occurring in the coniferous forest plots in Luya Mountain. Color points represent large families (>5 species) grouped in monocots and dicotyledon clades. Ele, elevation; BA, total basal area of breast height.

Table 1 Correlations between the first two constrained ordination axes (PCPS 1 and PCPS 2) and environmental factors

环境因子 Environmental variables	第1排序轴 PCPS 1	第2排序轴 PCPS 2	R²	p
海拔 Elevation	-0.495	0.869	0.653 8	0.001
坡向 Aspect	-0.620	0.784	0.203 9	0.224
坡度 Slope	0.305	-0.952	0.290 0	0.109
冠层高 Delta height	0.296	-0.955	0.145 6	0.349
胸高断面积和 Total basal area of breast height	-0.701	0.713	0.359 0	0.049
立木密度 Stem density	0.300	0.954	0.156 9	0.325
土壤深度 Soil depth	-0.436	-0.900	0.175 0	0.289
凋落物厚度 Litter thickness	-0.762	0.648	0.010 6	0.928
土壤有机碳 Soil organic carbon	-0.914	-0.406	0.183 4	0.267
土壤总氮 Soil total nitrogen	-0.384	-0.923	0.134 8	0.389
土壤总磷 Soil total phosphorus	0.165	0.986	0.045 5	0.733

References 54

[1]	APG III (2009). An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG III.Botanical Journal of the Linnean Society, 161, 105-121.
[2]	Bryant JA, Lamanna C, Morlon H, Kerkhoff AJ, Enquist BJ, Green JL (2008). Microbes on mountainsides: Contrasting elevational patterns of bacterial and plant diversity.Proceedings of the National Academy of Sciences of the United States of American, 105, 11505-11511.
[3]	Cavender-Bares J, Kozak KH, Fine PV, Kembel SW (2009). The merging of community ecology and phylogenetic biology.Ecology Letters, 12, 693-715.
[4]	Cook JE (2015). Structural effects on understory attributes in second-growth forests of northern Wisconsin, USA.Forest Ecology and Management, 347, 188-199.
[5]	Donoghue MJ (2008). Phylogenetic perspective on the distribution of plant diversity.Proceedings of the National Academy of Sciences of the United States of American, 105, 11549-11555.
[6]	Duarte LDS (2011). Phylogenetic habitat filtering influences forest nucleation in grasslands.Oikos, 120, 208-215.
[7]	Elton C (1946). Competition and the structure of ecological communities.Journal of Animal Ecology, 15, 54-68.
[8]	Fine PVA, Kembel SW (2011). Phylogenetic community structure and phylogenetic turnover across space and edaphic gradients in western Amazonian tree communities.Ecography, 34, 552-565.
[9]	Gaston KJ (2000). Global patterns in biodiversity.Nature, 405, 220-227.
[10]	Gilbert B, Lechowicz MJ (2004). Neutrality, niches, and dispersal in a temperate forest understory.Proceedings of the National Academy of Sciences of the United States of America, 101, 7651-7656.
[11]	Gilliam FS (2007). The ecological significance of the herbaceous layer in temperate forest ecosystems.BioScience, 57, 845-858.
[12]	Graham CH, Fine PV (2008). Phylogenetic beta diversity: Linking ecological and evolutionary processes across space in time. Ecology Letters, 11, 1265-1277.
[13]	Graham CH, Mcguire JA (2009). Phylogenetic structure in tropical hummingbird communities.Proceedings of the National Academy of Sciences of the United States of American, 106, 19673-19678.
[14]	Hardy OJ, Senterre B (2007). Characterizing the phylogenetic structure of communities by an additive partitioning of phylogenetic diversity.Journal of Ecology, 95, 493-506.
[15]	Hawkins BA, Rueda M, Rangel TF, Field R, Diniz-Filho JAF (2014). Community phylogenetics at the biogeographical scale: Cold tolerance, niche conservatism and the structure of North American forests.Journal of Biogeography, 41, 23-38.
[16]	Helmus M, Savage K, Diebel MJ, Ives A (2007). Separating the determinants of phylogenetic community structure.Ecology Letters, 10, 917-925.
[17]	Kembel SW, Hubbell SP (2006). The phylogenetic structure of a neotropical forest tree community.Ecology, 87, 86-99.
[18]	Kluge J, Kessler M (2011). Phylogenetic diversity, trait diversity and niches: Species assembly of ferns along a tropical elevational gradient.Journal of Biogeography, 38, 394-405.
[19]	Körner C (2007). The use of “altitude” in ecological research.Trends in Ecology & Evolution, 22, 569-574.
[20]	Latham RE, Ricklefs RE (1993). Global patterns of tree species richness in moist forests: Energy-diversity theory does not account for variation in species richness.Oikos, 67, 325-333.
[21]	Li XH, Zhu XX, Niu Y, Sun H (2014). Phylogenetic clustering and overdispersion for alpine plants along elevational gradient in the Hengduan Mountains Region, southwest China.Journal of Systematics and Evolution, 52, 280-288.
[22]	Li YJ, Wang SY, Niu JJ, Fang KY, Li XL, Li Y, Bu WL, Li YH (2016). Climate-adial growth relationship of Larix principis-rupprechtii at different altitudes on Luya Mountain. Acta Ecologica Sinica, 36, 1608-1618. (in Chinese with English abstract)[李颖俊, 王尚义, 牛俊杰, 方克艳, 李晓岚, 栗燕, 布文丽, 李玉晗 (2016). 芦芽山华北落叶松(Larix principis-rupprechtii)树轮宽度年表对气候因子的响应. 生态学报, 36, 1608-1618.]
[23]	Lichstein JW (2007). Multiple regression on distance matrices: A multivariate spatial analysis tool.Plant Ecology, 188, 117-131.
[24]	Lomolino MV (2001). Elevation gradients of species-density: Historical and prospective views.Global Ecology Biogeography, 10, 3-13.
[25]	Losos JB (2008). Phylogenetic niche conservatism, phylogenetic signal and the relationship between phylogenetic relatedness and ecological similarity among species.Ecology Letters, 11, 995-1003.
[26]	Lozupone CA, Hamady M, Kelley ST, Rob K (2007). Quantitative and qualitative beta diversity measures lead to different insights into factors that structure microbial communities.Applied & Environmental Microbiology, 73, 1576-1585.
[27]	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谱系结构. 生物多样性, 22, 438-448.]
[28]	Machac A, Janda M, Dunn RR, Sanders NJ (2011). Elevational gradients in phylogenetic structure of ant communities reveal the interplay of biotic and abiotic constraints on diversity.Ecography, 34, 364-371.
[29]	Márialigeti S, Tinya F, Bidló A, Ódor P (2016). Environmental drivers of the composition and diversity of the herb layer in mixed temperate forests in Hungary.Plant Ecology, 217, 549-563.
[30]	McCain CM (2009). Global analysis of bird elevational diversity.Global Ecology and Biogeography, 18, 346-360.
[31]	Oommen MA, Shanker K (2005). Elevational species richness patterns emerge from multiple local mechanisms in Himalayan woody plants.Ecology, 86, 3039-3047.
[32]	Pillar VD, Duarte LDS (2010). A framework for metacommunity analysis of phylogenetic structure.Ecology Letters, 13, 587-596.
[33]	Qian H, Hao Z, Zhang J (2014). Phylogenetic structure and phylogenetic diversity of angiosperm assemblages in forests along an elevational gradient in Changbaishan, China.Journal of Plant Ecology, 7, 154-165.
[34]	Rahbek C (2005). The role of spatial scale and the perception of large-scale species-richness patterns.Ecology Letters, 8, 224-239.
[35]	Ricklefs RE (2004). A comprehensive framework for global patterns in biodiversity.Ecology Letters, 7, 1-15.
[36]	Roberts DW, Cooper SV (1989). Concepts and techniques of vegetation mapping. In: Ferguson D, Morgan P, Johnson FD eds. Land Classifications Based on Vegetation: Applications for Resource Management, General Technical Report INF-257. Department of Agriculture, Forest Service, Intermountain Research Station, Odgen, USA. 90-96.
[37]	Sheldon KS, Yang S, Tewksbury JJ (2011). Climate change and community disassembly: Impacts of warming on tropical and temperate montane community structure.Ecology Letters, 14, 1191-1200.
[38]	Soininen J, McDonald R, Hillebrand H (2007). The distance decay of similarity in ecological communities.Ecography, 30, 3-12.
[39]	Stevens GC (1992). The elevational gradient in altitudinal range: An extension of Rapoport’s latitudinal rule to altitude.The American Naturalist, 140, 893-911.
[40]	Swenson NG (2011). Phylogenetic beta diversity metrics, trait evolution and inferring the functional beta diversity of communities.PLOS ONE, 6, e21264. doi: 10.1371/journal. pone.0021264.
[41]	Swenson NG, Anglada-Cordero P, Barone JA (2011). Deterministic tropical tree community turnover: Evidence from patterns of functional beta diversity along an elevational gradient.Proceedings of the Royal Society of London B: Biological Sciences, 278, 877-884.
[42]	Tang ZY, Fang JY, Chi XL, Feng JM, Liu YN, Shen ZH,Wang XP, Wang ZH, Wu XP, Zheng CY (2012a). Patterns of plant beta-diversity along elevational and latitudinal gradients in mountain forests of China.Ecography, 35, 1083-1091.
[43]	Tang ZY, Fang JY, Chi XL, Yang YY, Ma WH, Mohhamot A, Guo ZD, Liu YN, Gaston KJ (2012b). Geography, environment, and spatial turnover of species in China’s grasslands.Ecography, 35, 1103-1109.
[44]	Tello JS, Myers JA, Macía MJ, Fuentes AF, Cayola L, Arellano G, Loza MI, Torrez V, Cornejo M, Miranda TB (2015). Elevational gradients in β-diversity reflect variation in the strength of local community assembly mechanisms across spatial scales.PLOS ONE, 10, e0121458. doi: 10.1371/ journal.pone.0121458.
[45]	Vamosi SM, Heard SB, Vamosi JC, Webb CO (2009). Emerging patterns in the comparative analysis of phylogenetic community structure.Molecular Ecology, 18, 572-592.
[46]	Webb CO (2000). Exploring the phylogenetic structure of ecological communities: An example for rain forest trees.The American Naturalist, 156, 145-155.
[47]	Webb CO, Ackerly DD, Kembel SW (2008). Phylocom: Software for the analysis of phylogenetic community structure and trait evolution.Bioinformatics, 24, 2098-2100.
[48]	Webb CO, Ackerly DD, McPeek MA, Donoghue MJ (2002). Phylogenies and community ecology.Annual Review of Ecology and Systematics, 33, 475-505.
[49]	Webb CO, Donoghue MJ (2005). Phylomatic: Tree assembly for applied phylogenetics.Molecular Ecology Notes, 5, 181-183.
[50]	Wiens JJ, Donoghue MJ (2004). Historical biogeography, ecology and species richness.Trends in Ecology & Evolution, 19, 639-644.
[51]	Wiens JJ, Parra-Olea G, García-París M, Wake DB (2007). Phylogenetic history underlies elevational biodiversity patterns in tropical salamanders.Proceedings of the Royal Society of London B: Biological Sciences, 274, 919-928.
[52]	Zanne AE, Tank DC, Cornwell WK, Eastman JM, Smith SA, FitzJohn RG, McGlinn DJ, O’Meara BC, Moles AT, Reich PB (2014). Three keys to the radiation of angiosperms into freezing environments.Nature, 506, 89-92.
[53]	Zhang JT (1989). Vertical zone of vegetation in Luya Mountain in Shanxi Province.Scientia Geographica Sinica, 9, 346-353. (in Chinese with English abstract)[张金屯 (1989). 山西芦芽山植被垂直带的划分. 地理科学, 9, 346-353.]
[54]	Zhang WT, Jiang Y, Wang MC, Zhang LN, Dong MY (2015). Responses of radial growth in Larix principis-rupprechtii to climate change along an elevation gradient on the southern slope of Luya Mountain.Acta Ecologica Sinica, 35, 1-10. (in Chinese with English abstract)[张文涛, 江源, 王明昌, 张凌楠, 董满宇 (2015). 芦芽山阳坡不同海拔华北落叶松径向生长对气候变化的响应. 生态学报, 35, 1-10.]