Abstract
Spatial and environmental processes are two ecological processes that have attracted considerable attention in plant community assembly, depending on sampling scale and life history. However, the processes that determine community assembly have not been studied in the karst region of southwest China. In this study, a 25-ha (500 m × 500 m) monitoring plot within the subtropical climax forest in the karst region was established and canonical correspondence analysis was used to reveal the effects of topography and soil on the spatial patterns of tree community assembly. Our study suggests that spatial processes dominate species composition and the combined effects of spatial and environmental processes play an important role. Overall interpretation rate increases with enlarging the sampling scale. However, the pattern of variation partitioning was similar in different life stages. Environmental variables significantly affected species composition at different sampling sizes and life histories and had a higher interpretation rate of species composition on larger sampling sizes. Topographic wetness index was the most important variable to explain species composition of the environmental variables. These results suggest that it is necessary to consider the relative importance of environmental and spatial factors on community assembly to better understand, conserve, and manage subtropical karst forests.
Similar content being viewed by others
References
Asefa M, Wen HD, Cao M, Hu YH (2020) Key community assembly processes switch between scales in shaping beta diversity in two primary forests, Southwest China. Forests 11(10):1106. https://doi.org/10.3390/f11101106
Bagchi R, Henrys PA, Brown PE, Burslem DFRP, Diggle PJ, Gunatilleke CVS, Gunatilleke IAUN, Kassim AR, Law R, Noor S, Valencia RL (2011) Spatial patterns reveal negative density dependence and habitat associations in tropical trees. Ecology 92(9):1723–1729. https://doi.org/10.1890/11-0335.1
Baldeck CA, Harms KE, Yavitt JB, John R, Turner BL, Valencia R, Navarrete H, Bunyavejchewin S, Kiratiprayoon S, Yaacob A, Supardi MNN, Davies SJ, Hubbell SP, Chuyong GB, Kenfack D, Thomas DW, Dalling JW (2013) Habitat filtering across tree life stages in tropical forest communities. P Roy Soc B-Biol Sci 280(1766):20130548. https://doi.org/10.1098/rspb.2013.0548
Bell G (2001) Neutral macroecology. Science 293(5539):2413–2418. https://doi.org/10.1126/science.293.5539.2413
Borcard D, Legendre P (2002) All-scale spatial analysis of ecological data by means of principal coordinates of neighbour matrices. Ecol Model 153(1–2):51–68. https://doi.org/10.1016/s0304-3800(01)00501-4
Cai H, Lyu L, Shrestha N, Tang Z, Su X, Xu X, Dimitrov D, Wang Z (2021) Geographical patterns in phylogenetic diversity of Chinese woody plants and its application for conservation planning. Divers Distrib 27(1):179–194. https://doi.org/10.1111/ddi.13180
Chang LW, Zeleny D, Li CF, Chiu ST, Hsieh CF (2013) Better environmental data may reverse conclusions about niche- and dispersal-based processes in community assembly. Ecology 94(10):2145–2151. https://doi.org/10.1890/12-2053.1
Chase JM, Myers JA (2011) Disentangling the importance of ecological niches from stochastic processes across scales. Philos T R Soc B 366(1576):2351–2363. https://doi.org/10.1098/rstb.2011.0063
Chen HJ, Peng WX, Du H, Song TQ, Zeng FP, Wang F (2020) Effect of different grain for green approaches on soil bacterial community in a karst region. Front Microbiol 11(30):577242. https://doi.org/10.3389/fmicb.2020.577242
Comita LS, Condit R, Hubbell SP (2007) Developmental changes in habitat associations of tropical trees. J Ecol 95(3):482–492. https://doi.org/10.1111/j.1365-2745.2007.01229.x
Comita LS, Thompson J, Uriarte M, Jonckheere I, Canham CD, Zimmerman JK (2010) Interactive effects of land use history and natural disturbance on seedling dynamics in a subtropical forest. Ecol Appl 20(5):1270–1284. https://doi.org/10.1890/09-1350.1
Condit R (1998) Tropical forest census plots: methods results from Barro Colorado Island, Panama and a comparison with other plots. Springer, Berlin, pp 3–7
Diniz-Filho JAF, Siqueira T, Padial AA, Rangel TF, Landeiro VL, Bini LM (2012) Spatial autocorrelation analysis allows disentangling the balance between neutral and niche processes in metacommunities. Oikos 121(2):201–210. https://doi.org/10.1111/j.1600-0706.2011.19563.x
Dixon P (2003) VEGAN, a package of R functions for community ecology. J Veg Sci 14(6):927–930. https://doi.org/10.1111/j.1654-1103.2003.tb02228.x
Dray S, Legendre P, Peres-Neto PR (2006) Spatial modelling: a comprehensive framework for principal coordinate analysis of neighbour matrices (PCNM). Ecol Model 196(3–4):483–493. https://doi.org/10.1016/j.ecolmodel.2006.02.015
Dray S, Pelissier R, Couteron P, Fortin MJ, Legendre P, Peres-Neto PR, Bellier E, Bivand R, Blanchet FG, De Caceres M, Dufour AB, Heegaard E, Jombart T, Munoz F, Oksanen J, Thioulouse J, Wagner HH (2012) Community ecology in the age of multivariate multiscale spatial analysis. Ecol Monogr 82(3):257–275. https://doi.org/10.1890/11-1183.1
Du H, Hu F, Zeng F, Wang K, Peng W, Zhang H, Zeng Z, Zhang F, Song T (2017) Spatial distribution of tree species in evergreen-deciduous broadleaf karst forests in southwest China. Sci Rep-UK 7(1):15664. https://doi.org/10.1038/s41598-017-15789-5
Du H, Zeng F, Song T, Liu K, Wang K, Liu M (2021) Water depletion of climax forests over humid karst terrain: Patterns, controlling factors and implications. Agr Water Manage 244:106541. https://doi.org/10.1016/j.agwat.2020.106541
Dungan JL, Perry JN, Dale MRT, Legendre P, Citron-Pousty S, Fortin MJ, Jakomulska A, Miriti M, Rosenberg MS (2002) A balanced view of scale in spatial statistical analysis. Ecography 25(5):626–640. https://doi.org/10.1034/j.1600-0587.2002.250510.x
Egler FE (1954) Vegetation science concepts I. Initial floristic composition, a factor in old-field vegetation development with 2 figs. Vegetation 4(6):412–417. https://doi.org/10.1007/BF00275587
Etienne RS, Alonso D (2007) Neutral community theory: How stochasticity and dispersal-limitation can explain species coexistence. J Stat Phys 128(1–2):485–510. https://doi.org/10.1007/s10955-006-9163-2
Francisco TM, Couto DR, Garbin ML, Misaki F, Ruiz-Miranda CR (2021) Role of spatial and environmental factors in structuring vascular epiphyte communities in two neotropical ecosystems. Perspect Plant Ecol 51:125621. https://doi.org/10.1016/j.ppees.2021.125621
Gaston KJ, Chown SL (2005) Neutrality and the niche. Funct Ecol 19(1):1–6. https://doi.org/10.1111/j.0269-8463.2005.00948.x
Gibert C, Shenbrot GI, Stanko M, Khokhlova IS, Krasnov BR (2021) Dispersal-based versus niche-based processes as drivers of flea species composition on small mammalian hosts: inferences from species occurrences at large and small scales. Oecologia 197(2):471–484. https://doi.org/10.1007/s00442-021-05027-1
Gravel D, Canham CD, Beaudet M, Messier C (2006) Reconciling niche and neutrality: the continuum hypothesis. Ecol Lett 9(4):399–409. https://doi.org/10.1111/j.1461-0248.2006.00884.x
Guerra-Castro EJ, Conde JE, Barcelo A, Cruz-Motta JJ (2021) Variation in fouling assemblages associated with prop roots of Rhizophora mangle L. in the Caribbean: The role of neutral and niche processes. Austral Ecol 46(6):991–1007. https://doi.org/10.1111/aec.13071
Guo Y, Wang B, Mallik AU, Huang F, Xiang W, Tao D, Wen S, Lu S, Li D, He Y (2017) Topographic species-habitat associations of tree species in a heterogeneous tropical karst seasonal rain forest, China. J Plant Ecol 10(3):450–460. https://doi.org/10.1093/jpe/rtw057
Gupta D, Garlaschi S, Suweis S, Azaele S, Maritan A (2021) Effective resource competition model for species coexistence. Phys Rev Lett 127(20):208101. https://doi.org/10.1103/PhysRevLett.127.208101
He MZ, Zheng JG, Li XR, Qian YL (2007) Environmental factors affecting vegetation composition in the Alxa Plateau, China. J Arid Environ 69(3):473–489. https://doi.org/10.1016/j.jaridenv.2006.10.005
Hu YH, Lan GY, Sha LQ, Cao M, Tang Y, Li YD, Xu DP (2012) Strong neutral spatial effects shape tree species distributions across life stages at multiple scales. PLoS ONE 7(5):e38247. https://doi.org/10.1371/journal.pone.0038247
Hubbell SP (2006) Neutral theory and the evolution of ecological equivalence. Ecology 87(6):1387–1398. https://doi.org/10.1890/0012-9658(2006)87[1387:ntateo]2.0.co;2
Jafari M, Chahouki MAZ, Tavili A, Azarnivand H, Amiri GZ (2004) Effective environmental factors in the distribution of vegetation types in Poshtkouh rangelands of Yazd Province (Iran). J Arid Environ 56(4):627–641. https://doi.org/10.1016/s0140-1963(03)00077-6
Jetz W, Rahbek C (2002) Geographic range size and determinants of avian species richness. Science 297(5586):1548–1551. https://doi.org/10.1126/science.1072779
Jiang Z, Lian Y, Qin X (2014) Rocky desertification in Southwest China: Impacts, causes, and restoration. Earth-Sci Rev 132:1–12. https://doi.org/10.1016/j.earscirev.2014.01.005
Jones MM, Tuomisto H, Borcard D, Legendre P, Clark DB, Olivas PC (2008) Explaining variation in tropical plant community composition: influence of environmental and spatial data quality. Oecologia 155(3):593–604. https://doi.org/10.1007/s00442-007-0923-8
Kraft NJB, Adler PB, Godoy O, James EC, Fuller S, Levine JM (2015) Community assembly, coexistence and the environmental filtering metaphor. Funct Ecol 29(5):592–599. https://doi.org/10.1111/1365-2435.12345
Lai JS, Mi XC, Ren HB, Ma KP (2009) Species-habitat associations change in a subtropical forest of China. J Veg Sci 20(3):415–423. https://doi.org/10.1111/j.1654-1103.2009.01065.x
Lai JS, Zou Y, Zhang JL, Peres-Neto PR (2022) Generalizing hierarchical and variation partitioning in multiple regression and canonical analyses using the rdacca.hp R package. Methods Ecol Evol 13:782–788. https://doi.org/10.1111/2041-210X.13800
Laliberte E, Zemunik G, Turner BL (2014) Environmental filtering explains variation in plant diversity along resource gradients. Science 345(6204):1602–1605. https://doi.org/10.1126/science.1256330
Legendre P, Borcard D, Roberts DW (2012) Variation partitioning involving orthogonal spatial eigenfunction submodels. Ecology 93(5):1234–1240. https://doi.org/10.1890/11-2028.1
Levine JM, Hillerislambers J (2009) The importance of niches for the maintenance of species diversity. Nature 461(7261):254-U130. https://doi.org/10.1038/nature08251
Liang S, Lin H, Bao H, Yao Y, Jiang Y, Li Y, Pan Y (2020) Distribution pattern of trait-based community assembly for Cyclobalanopsis Glauca in the Guilin Karst Mountainous Areas. China Trop Conserv Sci 13:1940082920980279. https://doi.org/10.1177/1940082920980279
Liu CC, Liu YG, Guo K, Wang SJ, Yang Y (2014) Concentrations and resorption patterns of 13 nutrients in different plant functional types in the karst region of south-western China. Ann Bot 113(5):873–885. https://doi.org/10.1093/aob/mcu005
Lu MZ, Du H, Song TQ, Peng WX, Su L, Zhang H, Zeng ZX, Wang KL, Tang WN, Wei L, Zeng FP (2021a) Drivers of tree survival in an evergreen-deciduous broadleaf karst forest in southwest China. For Ecol Manage 499:119598. https://doi.org/10.1016/j.foreco.2021.119598
Lu MZ, Du H, Song TQ, Peng WX, Su L, Zhang H, Zeng ZX, Wang KL, Zeng FP (2021b) Effects of density dependence in an evergreen-deciduous broadleaf karst forest in southwest China. For Ecol Manage 490:119142. https://doi.org/10.1016/j.foreco.2021.119142
McGill BJ, Maurer BA, Weiser MD (2006) Empirical evaluation of neutral theory. Ecology 87(6):1411–1423. https://doi.org/10.1890/0012-9658(2006)87[1411:eeont]2.0.co;2
McIntire EJB, Fajardo A (2009) Beyond description: the active and effective way to infer processes from spatial patterns. Ecology 90(1):46–56. https://doi.org/10.1890/07-2096.1
Peng WX, Zhu YF, Song M, Du H, Song TQ, Zeng FP, Zhang F, Wang KL, Luo YQ, Lan X, Zhang JY (2019) The spatial distribution and drivers of soil microbial richness and diversity in a karst broadleaf forest. For Ecol Manage 449:117241. https://doi.org/10.1016/j.foreco.2019.03.033
Peres-Neto PR, Legendre P (2010) Estimating and controlling for spatial structure in the study of ecological communities. Global Ecol Biogeogr 19(2):174–184. https://doi.org/10.1111/j.1466-8238.2009.00506.x
Peres-Neto PR, Legendre P, Dray S, Borcard D (2006) Variation partitioning of species data matrices: estimation and comparison of fractions. Ecology 87(10):2614–2625. https://doi.org/10.1890/0012-9658(2006)87[2614:vposdm]2.0.co;2
Perry GLW, Miller BP, Enright NJ (2006) A comparison of methods for the statistical analysis of spatial point patterns in plant ecology. Plant Ecol 187(1):59–82. https://doi.org/10.1007/s11258-006-9133-4
Punchi-Manage R, Getzin S, WiegT KR, Gunatilleke CVS, Gunatilleke IAUN, Wiegand K, Huth A (2013) Effects of topography on structuring local species assemblages in a Sri Lankan mixed dipterocarp forest. J Ecol 101(1):149–160. https://doi.org/10.1111/1365-2745.12017
Roberts CM, McClean CJ, Veron JEN, Hawkins JP, Allen GR, McAllister DE, Mittermeier CG, Schueler FW, Spalding M, Wells F, Vynne C, Werner TB (2002) Marine biodiversity hotspots and conservation priorities for tropical reefs. Science 295(5558):1280–1284. https://doi.org/10.1126/science.1067728
Rosindell J, Hubbell SP, Etienne RS (2011) The unified neutral theory of biodiversity and biogeography at age ten. Trends Ecol Evol 26(7):340–348. https://doi.org/10.1016/j.tree.2011.03.024
Seabloom EW, Batzer E, Chase JM, Harpole WS, Adler PB, Bagchi S, Bakker JD, Barrio IC, Biederman L, Boughton EH, Bugalho MN, Caldeira MC, Catford JA, Daleo P, Eisenhauer N, Eskelinen A, Haider S, Hallett LM, Jonsdottir IS, Kimmel K, Kuhlman M, MacDougall A, Molina CD, Moore JL, Morgan JW, Muthukrishnan R, Ohlert T, Risch AC, Roscher C, Schutz M, Sonnier G, Tognetti PM, Virtanen R, Wilfahrt PA, Borer ET (2021) Species loss due to nutrient addition increases with spatial scale in global grasslands. Ecol Lett 24(10):2100–2112. https://doi.org/10.1111/ele.13838
Silvertown J (2004) Plant coexistence and the niche. Trends Ecol Evol 19(11):605–611. https://doi.org/10.1016/j.tree.2004.09.003
Smith TW, Lundholm JT (2010) Variation partitioning as a tool to distinguish between niche and neutral processes. Ecography 33(4):648–655. https://doi.org/10.1111/j.1600-0587.2009.06105.x
Song XY, Yang J, Cao M, Lin LX, Sun ZH, Wen HD, Swenson NG (2021) Traits mediate a trade-off in seedling growth response to light and conspecific density in a diverse subtropical forest. J Ecol 109(2):703–713. https://doi.org/10.1111/1365-2745.13497
Stokes CJ, Archer SR (2010) Niche differentiation and neutral theory: an integrated perspective on shrub assemblages in a parkland savanna. Ecology 91(4):1152–1162. https://doi.org/10.1890/08-1105.1
Su L, Du H, Zeng F, Peng W, Rizwan M, Nunez-Delgado A, Zhou Y, Song T, Wang H (2019) Soil and fine roots ecological stoichiometry in different vegetation restoration stages in a karst area, southwest China. J Environ Manage 252:109694. https://doi.org/10.1016/j.jenvman.2019.109694
Tilman D (2004) Niche tradeoffs, neutrality, and community structure: a stochastic theory of resource competition, invasion, and community assembly. Proc Natl Acad Sci USA 101(30):10854–10861. https://doi.org/10.1073/pnas.0403458101
Trindade DPF, Sfair JC, de Paula AS, Barros MF, Tabarelli M (2020) Water availability mediates functional shifts across ontogenetic stages in a regenerating seasonally dry tropical forest. J Veg Sci 31(6):1090–1101. https://doi.org/10.1111/jvs.12896
Tuomisto H, Ruokolainen K, Yli-Halla M (2003) Dispersal, environment, and floristic variation of western Amazonian forests. Science 299(5604):241–244. https://doi.org/10.1126/science.1078037
Tuomisto H, Ruokolainen L, Ruokolainen K (2012) Modelling niche and neutral dynamics: on the ecological interpretation of variation partitioning results. Ecography 35(11):961–971. https://doi.org/10.1111/j.1600-0587.2012.07339.x
Vellend M (2010) Conceptual synthesis in community ecology. Q Rev Biol 85(2):183–206. https://doi.org/10.1086/652373
Volkov I, Banavar JR, Hubbell SP, Maritan A (2003) Neutral theory and relative species abundance in ecology. Nature 424(6952):1035–1037. https://doi.org/10.1038/nature01883
Wang MM, Chen HS, Zhang W, Wang KL (2019) Influencing factors on soil nutrients at different scales in a karst area. CATENA 175:411–420. https://doi.org/10.1016/j.catena.2018.12.040
Wei X, Deng X, Xiang W, Lei P, Ouyang S, Wen H, Chen L (2018) Calcium content and high calcium adaptation of plants in karst areas of southwestern Hunan. China Biogeosciences 15(9):2991–3002. https://doi.org/10.5194/bg-15-2991-2018
Wennekes PL, Rosindell J, Etienne RS (2012) The neutral-niche debate: a philosophical perspective. Acta Biotheor 60(3):257–271. https://doi.org/10.1007/s10441-012-9144-6
Yang J, El-Kassaby YA, Guan WB (2020) Multiple ecological drivers determining vegetation attributes across scales in a mountainous dry valley, southwest China. Forests 11(11):1140. https://doi.org/10.3390/f11111140
Yuan ZQ, Gazol A, Wang XG, Lin F, Ye J, Bai XJ, Li BH, Hao ZQ (2011) Scale specific determinants of tree diversity in an old growth temperate forest in China. Basic Appl Ecol 12(6):488–495. https://doi.org/10.1016/j.baae.2011.07.008
Zhang Z, Hu B, Hu G (2014) Spatial heterogeneity of soil chemical properties in a subtropical karst forest, Southwest China. Sciworld J 2014:473651. https://doi.org/10.1155/2014/473651
Zheng SL, Webber BL, Didham RK, Chen C, Yu MJ (2021) Disentangling biotic and abiotic drivers of intraspecific trait variation in woody plant seedlings at forest edges. Ecol Evol 11(14):9728–9740. https://doi.org/10.1002/ece3.7799
Acknowledgements
We are especially thanks to the Management Center for Guangxi Mulun National Nature Reserve and many field workers for their contributions to the establishment of the 25-ha Mulun plot.
Author information
Authors and Affiliations
Contributions
HD, FZ, WP, KW and TS conceived and designed the experiments. HD, LS and ML conducted the experiments. LS, HD, HW discussed the results. LS wrote the manuscript. All authors reviewed the manuscript.
Corresponding authors
Additional information
Publisher's Note
Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.
Project funding: This study was supported by the National Natural Science Foundation of China (42071073, 31971487), Youth Innovation Promotion Association of the Chinese Academy of Sciences (2021366), Guangxi Key Research and Development Program (AB17129009), and the Hechi Distinguished Expert Program to Fuping Zeng and the Guangxi Bagui Scholarship Program to Dejun Li.
The online version is available at http://www.springerlink.com
Corresponding editor: Yanbo Hu
Supplementary Information
Below is the link to the electronic supplementary material.
Rights and permissions
Springer Nature or its licensor (e.g. a society or other partner) holds exclusive rights to this article under a publishing agreement with the author(s) or other rightsholder(s); author self-archiving of the accepted manuscript version of this article is solely governed by the terms of such publishing agreement and applicable law.
About this article
Cite this article
Su, L., Du, H., Zeng, F. et al. Environmental and spatial contributions to tree community assembly across life stages and scales in evergreen-deciduous broadleaf karst forests, southwest China. J. For. Res. 34, 1323–1331 (2023). https://doi.org/10.1007/s11676-022-01587-x
Received:
Accepted:
Published:
Issue Date:
DOI: https://doi.org/10.1007/s11676-022-01587-x