Topographic and environmental characteristics drive phylogenetic diversity and vegetation traits in Larix kaempferi plantations in South Korea

Abstract

Global diversification of the structure and composition of artificial forests and partial or complete conversion to natural forests is an important task for improving long-term biodiversity and counteracting climate change. Larix kaempferi is a tree species used widely in forests throughout northeast Asia that plays an important role in converting artificial forests to mixed forests. However, the phylogenetic diversity (PD) of the species remains unclear. We investigated L. kaempferi forests formed in Gayasan National Park, South Korea, categorized the community types, and quantified species composition, PD, and phylogenetic community structure depending on the vegetation type. Furthermore, we explored the factors regulating biodiversity in L. kaempferi forests to provide insights for promoting forests with high structural diversity. We observed unique vegetation characteristics and community formation mechanisms depending on the local environment, with vegetation types located in valleys and at the bottom of slopes having the highest PD. We revealed how the structural properties and local conditions of forests affect phylogenetic community structure for each vegetation type, leading to competitive interactions and competitive exclusion. For all vegetation types, PD showed a gradually increasing trend with older stand age, but piecewise structural equation modeling analysis showed that topographic environmental factors are the main factors regulating PD. Our findings highlight the need to introduce customized management approaches suited to the characteristics of each community rather than using the same method for all communities. This approach is crucial because species composition, ecological properties, rate of succession, and surrounding environmental conditions differ between vegetation types. In addition, by presenting management strategies to improve biodiversity depending on vegetation type, we expanded existing knowledge on the conversion of artificial forests to mixed forests. Our study provides important insights into establishing strategies for managing artificial coniferous forests and the mechanisms of community formation with changes in species composition after forest creation.

Keywords

Larix kaempferi / Phylogenetic diversity / Vegetation traits / Topographic characteristics / Artificial forests

References

[1] Ali A, Yan ER. Consequences of phylogenetic conservativeness and functional trait similarity on aboveground biomass vary across subtropical forest strata. For Ecol Manag, 2018, 429: 28-35

[2] Ali A, Lohbeck M, Yan ER. Forest strata-dependent functional evenness explains whole-community aboveground biomass through opposing mechanisms. For Ecol Manag, 2018, 424: 439-447

[3] Ammer C, Bickel E, Kölling C. Converting Norway spruce stands with beech: a review of arguments and techniques. Austrian J Forest Sci, 2008, 125(1): 3-26

[4] Barbour MG, Burk J, Pitts WD. Terrestrial Plant Ecology, 19872Menlo Park, California, The Benjamin/Cummings Publishing Company Inc

[5] Barsoum N, Henderson L. Converting planted non-native conifer to native woodlands: a review of the benefits, drawbacks and experience in Britain. For Commiss Res Note, 2016, 024: 1-10

[6] Beven KJ, Kirkby MJ. A physically based, variable contributing area model of basin hydrology/Un modèle à base physique de zone d’appel variable de l’hydrologie du bassin versant. Hydrol Sci Bull, 1979, 24(1): 43-69

[7] Bigelow SW, Canham CD. Community organization of tree species along soil gradients in a north-eastern USA forest. J Ecol, 2002, 90(1): 188-200

[8] Bolte A, Ammer C, Löf M, Madsen P, Nabuurs GJ, Schall P, Spathelf P, Rock J. Adaptive forest management in Central Europe: climate change impacts, strategies and integrative concept. Scand J for Res, 2009, 24(6): 473-482

[9] Braun-Blanquet J. Pflanzensoziologie: Grünzüge der Vegetationsgrunde, 19643Wien, Springer Vienna

[10] Cadotte MW, Tucker CM. Should environmental filtering be abandoned?. Trends Ecol Evol, 2017, 32(6): 429-437

[11] Cadotte MW, Cardinale BJ, Oakley TH. Evolutionary history and the effect of biodiversity on plant productivity. Proc Natl Acad Sci U S A, 2008, 105(44): 17012-17017

[12] Cahill JF, Kembel SW, Lamb EG, Keddy PA. Does phylogenetic relatedness influence the strength of competition among vascular plants?. Perspect Plant Ecol Evol Syst, 2008, 10(1): 41-50

[13] Canullo R, Allegrini MC, Campetella G. Reference field manual for vegetation surveys on the CONECOFOR LII network, Italy (National Programme of Forest Ecosystems Control—UNECE, ICP Forests). Braun-Blanquetia, 2012, 48: 5-65

[14] Cao JH, Qi R, Liu T, Li B, Gao BQ, Chen XL, Zhao Y, Zhao ZG. Patterns of species and phylogenetic diversity in Picea purpurea forests under different levels of disturbance on the northeastern Qinghai-Tibetan Plateau. Glob Ecol Conserv, 2021, 30 e01779

[15] Cavender-Bares J, Kozak KH, Fine PVA, Kembel SW. The merging of community ecology and phylogenetic biology. Ecol Lett, 2009, 12(7): 693-715

[16] Chavardès RD, Gennaretti F, Grondin P, Cavard X, Morin H, Bergeron Y. Role of mixed-species stands in attenuating the vulnerability of boreal forests to climate change and insect epidemics. Front Plant Sci, 2021, 12 658880

[17] Chen FQ, Song NN, Chen GH, Wang JZ. Effects of exotic species Larix kaempferi on diversity and activity of soil microorganisms in Dalaoling National Forest Park. Ecol Process, 2015, 4 10

[18] Chun JH, Lee CB. Temporal changes in species, phylogenetic, and functional diversity of temperate tree communities: insights from assembly patterns. Front Plant Sci, 2019, 10: 294

[19] Chun JH, Kim HH, Lim JH, Lee CB. Plant diversity patterns and the drivers on a long-term riparian forest research site in Mt. Gyebang, south Korea. J Agric Life Sci, 2019, 53(5): 13-23

[20] Coote L, Dietzsch AC, Wilson MW, Graham CT, Fuller L, Walsh AT, Irwin S, Kelly DL, Mitchell FJG, Kelly TC, O’Halloran J. Testing indicators of biodiversity for plantation forests. Ecol Indic, 2013, 32: 107-115

[21] Curtis JT, McIntosh RP. An upland forest continuum in the prairie-forest border region of Wisconsin. Ecology, 1951, 32(3476-496

[22] Dinnage R. Disturbance alters the phylogenetic composition and structure of plant communities in an old field system. PLoS ONE, 2009, 4(9 e7071

[23] Doyon F, Yamasaki S, Duchesneau R. The use of the natural range of variability for identifying biodiversity values at risk when implementing a forest management strategy. For Chron, 2008, 84(3): 316-329

[24] Duffy PJ, Jacobs DF, Kabrick JM, Thornton CD, Swaim JT, Jenkins MA. Response of understory vegetation to long-term occupancy by introduced Pinus species in temperate deciduous hardwood forests. For Ecol Manage, 2024, 572 122310

[25] FAO (2020) Global Forest Resources Assessment 2020: key findings. https://www.fao.org/interactive/forest-resources-assessment/2020/en/. Accessed 30 Jan 2025

[26] Finzi AC, Canham CD, Van Breemen N. Canopy tree–soil interactions within temperate forests: species effects on pH and cations. Ecol Appl, 1998, 8(2): 447-454

[27] Forest F, Grenyer R, Rouget M, Davies TJ, Cowling RM, Faith DP, Balmford A, Manning JC, Procheş S, van der Bank M, Reeves G, Hedderson TAJ, Savolainen V. Preserving the evolutionary potential of floras in biodiversity hotspots. Nature, 2007, 445(7129): 757-760

[28] Gilliam FS, Turrill NL, Adams MB. Herbaceous-layer and overstory species in clear-cut and mature central Appalachian hardwood forests. Ecol Appl, 1995, 5(4): 947-955

[29] Hamilton GJ (1980) Line thinning. Forestry Commission Archive: Leaflet 77.

[30] Harcombe PA, Marks PL. Tree diameter distributions and replacement processes in southeast Texas forests. For Sci, 1978, 24(2): 153-166

[31] Hartley MJ. Rationale and methods for conserving biodiversity in plantation forests. For Ecol Manag, 2002, 155(1–3): 81-95

[32] Heinrichs S, Schmidt W. Short-term effects of selection and clear cutting on the shrub and herb layer vegetation during the conversion of even-aged Norway spruce stands into mixed stands. For Ecol Manage, 2009, 258(5): 667-678

[33] Hill MO. TWINSPAN: A FORTRAN program for arranging multivariate data in an ordered two-way table by classification of the individuals and attributes, 1979, Ithaca, New York, Cornell University

[34] Hu Y, Li XY, Guo AX, Yue P, Guo XX, Lv P, Zhao SL, Zuo XA. Species diversity is a strong predictor of ecosystem multifunctionality under altered precipitation in desert steppes. Ecol Indic, 2022, 137 108762

[35] Jactel H, Moreira X, Castagneyrol B. Tree diversity and forest resistance to insect pests: patterns, mechanisms, and prospects. Annu Rev Entomol, 2021, 66: 277-296

[36] Jarzyna MA, Quintero I, Jetz W. Global functional and phylogenetic structure of avian assemblages across elevation and latitude. Ecol Lett, 2021, 24(2): 196-207

[37] Jin Y, Qian H. V.PhyloMaker2: an updated and enlarged R package that can generate very large phylogenies for vascular plants. Plant Divers, 2022, 44(4): 335-339

[38] Karger DN, Conrad O, Böhner J, Kawohl T, Kreft H, Soria-Auza RW, Zimmermann NE, Linder HP, Kessler M. Climatologies at high resolution for the Earth’s land surface areas. Sci Data, 2017, 4 170122

[39] Karizumi N. Illustrations of tree roots, 1979, Tokyo, Seibundo Shinkosha Publishing

[40] Kembel SW, Cowan PD, Helmus MR, Cornwell WK, Morlon H, Ackerly DD, Blomberg SP, Webb CO. Picante: R tools for integrating phylogenies and ecology. Bioinformatics, 2010, 26(11): 1463-1464

[41] Kirby KJ. Changes in the ground flora under plantations on ancient woodland sites. Forestry, 1988, 61(4): 317-338

[42] Landeweert R, Hoffland E, Finlay RD, Kuyper TW, van Breemen N. Linking plants to rocks: ectomycorrhizal fungi mobilize nutrients from minerals. Trends Ecol Evol, 2001, 16(5): 248-254

[43] Lawley V, Parrott L, Lewis M, Sinclair R, Ostendorf B. Self-organization and complex dynamics of regenerating vegetation in an arid ecosystem: 82 years of recovery after grazing. J Arid Environ, 2013, 88: 156-164

[44] Lee IK, Monsi M. Ecological studies on Pinus densiflora forest. 1. Effect of plant substances on the floristic composition of the undergrowth. Bot Mag Tokyo, 1963, 76 400

[45] Lee HI, Seo YO, Kim H, Ali A, Lee CB, Chung Y. Species evenness declines but specific functional strategy enhances aboveground biomass across strata in subtropical–warm-temperate forests of South Korea. For Ecol Manag, 2022, 512 120179

[46] Lee SJ, Lee AR, Byeon JG, Oh SH. Pre-drought effects on northern temperate trees and vine invasion in forest gaps hindering regeneration. Sci Total Environ, 2024, 951 175707

[47] Lefcheck JS. piecewiseSEM: Piecewise structural equation modelling in r for ecology, evolution, and systematics. Meth Ecol Evol, 2016, 7(5): 573-579

[48] Legendre P, Legendre L. Numerical ecology, 20123Amsterdam, Elsevier

[49] Li JY, Wu GX, Guo QX, Korpelainen H, Li CY. Fast-growing Larix kaempferi suffers under nutrient imbalance caused by phosphorus fertilization in larch plantation soil. For Ecol Manag, 2018, 417: 49-62

[50] Liu MX, Che YD, Jiao J, Li LR, Jiang XX. Exploring the community phylogenetic structure along the slope aspect of subalpine meadows in the eastern Qinghai–Tibetan Plateau, China. Ecol Evol, 2019, 9(9): 5270-5280

[51] Losos JB, Leal M, Glor RE, De Queiroz K, Hertz PE, Rodríguez Schettino L, Lara AC, Jackman TR, Larson A. Niche lability in the evolution of a Caribbean lizard community. Nature, 2003, 424(6948): 542-545

[52] Lu YH, Coops NC, Wang TL, Wang GY, Lu YH, Coops NC, Wang TL, Wang GY. A process-based approach to estimate Chinese fir (Cunninghamia lanceolata) distribution and productivity in Southern China under climate change. Forests, 2015, 6(2): 360-379

[53] Maherali H, Klironomos JN. Influence of phylogeny on fungal community assembly and ecosystem functioning. Science, 2007, 316(5832): 1746-1748

[54] Mamet SD, Brown CD, Trant AJ, Laroque CP. Shifting global Larix distributions: northern expansion and southern retraction as species respond to changing climate. J Biogeogr, 2019, 46(1): 30-44

[55] Mastrogianni A, Kallimanis AS, Chytrý M, Tsiripidis I. Phylogenetic diversity patterns in forests of a putative refugial area in Greece: a community level analysis. For Ecol Manag, 2019, 446: 226-237

[56] Nagaike T. A review of ecological studies on plant species diversity in plantation ecosystems. J Jpn for Soc, 2000, 82(4): 407-416

[57] Nagaike T, Hayashi A, Abe M, Arai N. Differences in plant species diversity in Larix kaempferi plantations of different ages in central Japan. For Ecol Manag, 2003, 183(1–3): 177-193

[58] Nakamura Y, Krestov PV. Anderson F. Coniferous forests of the temperate zone of Asia. Ecosystems of the world. 6: Coniferous forests, 2005, Amsterdam, Elsevier163220

[59] Oh JH, Hwang JS, Cha DS. Analysis of slope stability effect of arbors’ roots-on tensile strength of the roots. Korean J Environ Ecol, 2011, 25(1): 31-36

[60] Oh SH, Kim JS, Cho JH, Cho HJ. Vegetation types and ecological characteristics of Larix kaempferi plantations in baekdudaegan protected area, South Korea. J Korean Soc for Sci, 2021, 110(4): 530-542

[61] Park YS, Fowler DP. A provenance test of Japanese larch in eastern Canada, including comparative data on European larch and tamarack. Silvae Genet, 1983, 32: 96-101

[62] Pastore AI, Scherer BP. Changes in community phylogenetic structure in a North American forest chronosequence. Ecosphere, 2016, 7(12 e01592

[63] Pielou EC. An introduction to mathematical ecology, 1969, Hoboken, Wiley

[64] Piwczyński M, Puchałka R, Ulrich W. Influence of tree plantations on the phylogenetic structure of understorey plant communities. For Ecol Manag, 2016, 376: 231-237

[65] Poorter L, van Der Sande MT, Amissah L, Bongers F, Hordijk I, Kok J, Lohbeck M (2024) A comprehensive framework for vegetation succession. Ecosphere 15(4):e4794

[66] Puettmann KJ, Ares A, Burton JI, Dodson EK. Forest restoration using variable density thinning: lessons from Douglas-fir stands in western Oregon. Forests, 2016, 7(12): 310

[67] Qin H, Dong G, Zhang YB, Zhang F, Wang MB. Patterns of species and phylogenetic diversity of Pinus tabuliformis forests in the eastern Loess Plateau, China. For Ecol Manag, 2017, 394: 42-51

[68] Rice EL. Alleopathy, 1984, London, Academic Press

[69] Seliger A, Puffpaff S, Manthey M, Kreyling J. Management options for the conversion of allochthonous coniferous forest patches towards more natural species composition in the Vorpommersche Boddenlandschaft National Park, NE Germany. Balt for, 2021

[70] Seliger A, Ammer C, Kreft H, Zerbe S. Diversification of coniferous monocultures in the last 30 years and implications for forest restoration: a case study from temperate lower montane forests in Central Europe. Eur J for Res, 2023, 142(61353-1368

[71] Seliger A, Ammer C, Kreft H, Zerbe S. Changes of vegetation in coniferous monocultures in the context of conversion to mixed forests in 30 years: implications for biodiversity restoration. J Environ Manag, 2023, 343 118199

[72] Shannon CE. A mathematical theory of communication. Bell Syst Tech J, 1948, 27(3): 379-423

[73] Shao XZ. Study on the ecological survey of Larix kaempferi. Shandong for Sci Technol, 1985, 1: 9-18(in Chinese)

[74] Shipley B. The AIC model selection method applied to path analytic models compared using a d-separation test. Ecology, 2013, 94(3): 560-564

[75] Socolar JB, Gilroy JJ, Kunin WE, Edwards DP. How should beta-diversity inform biodiversity conservation?. Trends Ecol Evol, 2016, 31(1): 67-80

[76] Spracklen BD, Lane JV, Spracklen DV, Williams N, Kunin WE. Regeneration of native broadleaved species on clearfelled conifer plantations in upland Britain. For Ecol Manag, 2013, 310: 204-212

[77] Srivastava DS, Cadotte MW, MacDonald AAM, Marushia RG, Mirotchnick N. Phylogenetic diversity and the functioning of ecosystems. Ecol Lett, 2012, 15(7): 637-648

[78] Sykes JM, Lowe VPW, Briggs DR. Some effects of afforestation on the flora and fauna of an upland sheepwalk during 12 years after planting. J Appl Ecol, 1989, 26(1): 299-320

[79] Tucker CM, Davies TJ, Cadotte MW, Pearse WD. On the relationship between phylogenetic diversity and trait diversity. Ecology, 2018, 99(6): 1473-1479

[80] Turley NE, Brudvig LA. Agricultural land-use history causes persistent loss of plant phylogenetic diversity. Ecology, 2016, 97(9): 2240-2247

[81] Ujházy K, Hederová L, Máliš F, Ujházyová M, Bosela M, Čiliak M. Overstorey dynamics controls plant diversity in age-class temperate forests. For Ecol Manag, 2017, 391: 96-105

[82] Wagner S, Fischer H, Huth F. Canopy effects on vegetation caused by harvesting and regeneration treatments. Eur J Forest Res, 2011, 130(1): 17-40

[83] Wang BZ, Yang XH, Zhu YJ, Tu Y, Wang DY, Indree T. Plant diversity mediates the response of ecosystem multifunctionality to climate factor in Eastern Eurasian Steppe. Glob Ecol Conserv, 2024, 50 e02827

[84] Webb CO, Ackerly DD, McPeek MA, Donoghue MJ. Phylogenies and community ecology. Annu Rev Ecol Evol Syst, 2002, 33: 475-505

[85] Winter M, Devictor V, Schweiger O. Phylogenetic diversity and nature conservation: where are we?. Trends Ecol Evol, 2013, 28(4): 199-204

[86] Wu CY, Chen DS, Shen JP, Sun XM, Zhang SG. Estimating the distribution and productivity characters of Larix kaempferi in response to climate change. J Environ Manag, 2021, 280 111633

[87] Xiao YM, Yang LC, Nie XQ, Li CB, Xiong F, Wang LL, Zhou GY. Examining differences in phylogenetic composition enhances understanding of the phylogenetic structure of the shrub community in the northeastern Qinghai-Tibetan Plateau. Ecol Evol, 2020, 10(13): 6723-6731

[88] Xie J, Fang S, Yan QL. Effects of thinning on functional and phylogenetic diversity of regenerated woody plants in plantations over time. Appl Veg Sci, 2023, 26(3 e12737

[89] Yamamoto SI. Forest gap dynamics and tree regeneration. J for Res, 2000, 5(4): 223-229

[90] Yang J, Zhang GC, Ci XQ, Swenson NG, Cao M, Sha LQ, Li J, Baskin CC, Ferry Slik JW, Lin LX. Functional and phylogenetic assembly in a Chinese tropical tree community across size classes, spatial scales and habitats. Funct Ecol, 2014, 28(2): 520-529

[91] Yang J, Li EH, Yang C, Xia Y, Zhou R. Effects of south-to-north water diversion project cascade dams on riparian vegetation along the middle and lower reaches of the Hanjiang River, China. Front Plant Sci, 2022, 13 849010

[92] Zhang XP, Wang MB, Liang XM. Quantitative classification and carbon density of the forest vegetation in Lüliang Mountains of China. Plant Ecol, 2009, 201(1): 1-9

[93] Zhang M, Peng PH, Shi SL, Wang GY. The environmental interpretation of Pinus yunnanensis community differentiation after the invasion of Ageratina adenophora in Panxi region, China. Community Ecol, 2024, 25(1): 45-64

[94] Zhao Y, Li M, Wang XH, Deng JY, Zhang ZZ, Wang BT. Influence of habitat on the phylogenetic structure of Robinia pseudoacacia forests in the eastern Loess Plateau, China. Glob Ecol Conserv, 2020, 24 e01199

Funding

Kyungpook National University

RIGHTS & PERMISSIONS

The Author(s)