iForest - Biogeosciences and Forestry


Spatial structure of the vertical layers in a subtropical secondary forest 57 years after clear-cutting

Yuanfa Li, Ji’an He, Sufang Yu, Deyi Zhu, Hongxiang Wang, Shaoming Ye   

iForest - Biogeosciences and Forestry, Volume 12, Issue 5, Pages 442-450 (2019)
doi: https://doi.org/10.3832/ifor2975-012
Published: Sep 16, 2019 - Copyright © 2019 SISEF

Research Articles

Stratification is an important phenomenon in natural forests. The mixed pine-oak forests along the Nanpan River in southwest China was clearly formed by two layers in the vertical direction. These forests developed in an area where the virgin forests suffered clear-cutting. After excluding habitat heterogeneity, we divided two plots into upper and lower layers according to the tree height, and then analyzed the spatial pattern, species distribution, and size differentiation using the pair correlation function g(r) and the mark correlation function (MCF) kmm (r), respectively. The following key results were obtained: (1) the upstory was slightly clustered, whereas the understory had an intensively clumped pattern. An uneven pattern of germplasm resources in the early stages of succession and seed dispersion limitation may have contributed to the aggregation of tree species. (2) The spatial correlation among the main populations in each layer, and between both layers, had a largely random association, suggesting that differences in tree growth and physiological characteristics play an important role in species association. (3) Species aggregation decreased as the observation scale increased; however, the aggregation intensity of the understory was significantly higher than that of the upstory, which may be related to negative density dependence and niche complementarity. (4) Size differentiation in the upstory was significant, with small trees gathered together. There were no obvious differences in tree size in the lower layer, indicating a higher competitive pressure among trees in the upper layer. In conclusion, the spatial structure of trees in terms of vertical stratification differs, which is of great significance for investigating the mechanisms by which species coexist.


Clear Cut, Secondary Forest, Size Differentiation, Spatial Pattern, Stratification

Authors’ address

Yuanfa Li
Ji’an He
Sufang Yu
Deyi Zhu
Hongxiang Wang 0000-0002-3224-5120
Shaoming Ye
Guangxi Key Laboratory of Forest Ecology and Conservation, College of Forestry, Guangxi University, Daxue East Road 100, Nanning, Guangxi, 530004 (China)

Corresponding author

Shaoming Ye


Li Y, He J, Yu S, Zhu D, Wang H, Ye S (2019). Spatial structure of the vertical layers in a subtropical secondary forest 57 years after clear-cutting. iForest 12: 442-450. - doi: 10.3832/ifor2975-012

Academic Editor

Francisco Lloret Maya

Paper history

Received: Oct 11, 2018
Accepted: Jun 30, 2019

First online: Sep 16, 2019
Publication Date: Oct 31, 2019
Publication Time: 2.60 months

Breakdown by View Type

(Waiting for server response...)

Article Usage

Total Article Views: 35325
(from publication date up to now)

Breakdown by View Type
HTML Page Views: 31311
Abstract Page Views: 1741
PDF Downloads: 1808
Citation/Reference Downloads: 4
XML Downloads: 461

Web Metrics
Days since publication: 1768
Overall contacts: 35325
Avg. contacts per week: 139.86

Article Citations

Article citations are based on data periodically collected from the Clarivate Web of Science web site
(last update: Feb 2023)

Total number of cites (since 2019): 7
Average cites per year: 1.40


Publication Metrics

by Dimensions ©

Articles citing this article

List of the papers citing this article based on CrossRef Cited-by.

Anonymous (1960)
Preliminary studies on regeneration and final felling of Pinus yunnanensis Franchet (Guangxi Zhuang Autonomous Region Forestry Research Institute, Guangxi Baise forestry experimental station, Lingle subbureau attached to Guangxi forest industry bureau, Guangxi Lingle forestry farm eds). Scientia Silvae Sinicae 1: 44-54.
Barbier S, Gosselin F, Balandier P (2008)
Influence of tree species on understory vegetation diversity and mechanisms involved. A critical review for temperate and boreal forests. Forest Ecology and Management 254: 1-15.
CrossRef | Gscholar
Beatty SW (1984)
Influence of microtopography and canopy species on spatial patterns of forest understory plants. Ecology 65 (5): 1406-1419.
CrossRef | Gscholar
Carrer M, Castagneri D, Popa I, Pividori M, Lingua E (2018)
Tree spatial patterns and stand attributes in temperate forests: the importance of plot size, sampling design, and null model. Forest Ecology and Management 407: 125-134.
CrossRef | Gscholar
Collet C, Manso R, Barbeito I (2017)
Coexistence, association and competitive ability of Quercus petraea and Quercus robur seedlings in naturally regenerated mixed stands. Forest Ecology and Management 390: 36-46.
CrossRef | Gscholar
Del Río M, Pretzsch H, Alberdi I, Bielak K, Bravo F, Brunner A, Condés S, Ducey MJ, Fonseca T, Von Lüpke N, Pach M, Peric S, Perot T, Souidi Z, Spathelf P, Sterba H, Tijardovic M, Tomé M, Vallet P, Bravo-Oviedo A (2016)
Characterization of the structure, dynamics, and productivity of mixed-species stands: review and perspectives. European Journal of Forest Research 135 (1): 23-49.
CrossRef | Gscholar
Ford ED (1975)
Competition and stand structure in some even-aged plant monocultures. Journal of Ecology 63: 311-333.
CrossRef | Gscholar
Gazol A, Ibáñez R (2009)
Different response to environmental factors and spatial variables of two attributes (cover and diversity) of the understorey layers. Forest Ecology and Management 258: 1267-1274.
CrossRef | Gscholar
Ghalandarayeshi S, Nord-Larsen T, Johannsen VK, Larsen JB (2017)
Spatial patterns of tree species in Suserup Skov - a semi-natural forest in Denmark. Forest Ecology and Management 406: 391-401.
CrossRef | Gscholar
Giesbrecht IJW, Saunders SC, MacKinnon A, Lertzman KP (2017)
Overstory structure drivers fine-scale coupling of understory light and vegetation in two temperature rainforest floodplains. Canadian Journal of Forest Research 47 (9): 1244-1256.
CrossRef | Gscholar
Hao Z, Zhang J, Song B, Ye J, Li B (2007)
Vertical structure and spatial associations of dominant tree species in an old-growth temperate forest. Forest Ecology and Management 252 (1): 1-11.
CrossRef | Gscholar
Hui G, Pommerening A (2014)
Analysing tree species and size diversity patterns in multi-species uneven-aged forests of Northern China. Forest Ecology and Management 316: 125-138.
CrossRef | Gscholar
Ishii H, Reynolds JH, Ford ED, Shaw DC (2000)
Height growth and vertical development of an old-growth Pseudotsuga-Tsuga forest in southwestern Washington State, USA. Canadian Journal of Forest Research 30: 17-24.
CrossRef | Gscholar
Janík D, Adam D, Hort L, Král K, Samonil P, Unar P, Vrška T (2014)
Tree spatial patterns of Abies alba and Fagus sylvatica in the Western Carpathians over 30 years. European Journal of Forest Research 133 (6): 1015-1028.
CrossRef | Gscholar
Kint V (2005)
Structural development in ageing temperate Scots pine stands. Forest Ecology and Management 214: 237-250.
CrossRef | Gscholar
Kuninaga T, Hirayama K, Sakimoto M (2015)
Negative canopy-understorey interaction shapes the sapling bank of Fagus crenata in a cool-temperate, conifer-hardwood mixed forest. Plant Ecology 16: 1191-1202.
CrossRef | Gscholar
Kuuluvainen T, Penttinen A, Leinonen K, Nygren M (1996)
Statistical opportunities for comparing stand structural heterogeneity in managed and primeval forests: an example from boreal spruce forests in Southern Finland. Silva Fennica 30: 315-328.
CrossRef | Gscholar
Li Z, Wang X (1981)
The distribution of Pinus yunnanensis var. tenuifolia in relation to the environmental conditions. Acta Phytoecologica et Geobotanica Sinica 5: 28-37.
Li L, Huang Z, Ye W, Cao H, Wei S, Wang Z, Lian J, Sun Y, Ma K, He F (2009)
Spatial distributions of tree species in a subtropical forest of China. Oikos 118: 495-502.
CrossRef | Gscholar
Li Y, Hui G, Zhao Z, Hu Y (2012)
The bivariate distribution characteristics of spatial structure in natural Korean pine broad-leaved forest. Journal of Vegetation Science 23: 1180-1190.
CrossRef | Gscholar
Li Y, Hui G, Yu S, Yao X, Ye S (2017)
Nearest neighbour relationships in Pinus yunnanensis var. tenuifolia forests along the Nanpan River, China. iForest 10: 746-753.
CrossRef | Gscholar
Martens SN, Breshears DD, Meyer CW (2000)
Spatial distributions of understory light along the grassland: forest continuum: effects of cover, height, and spatial pattern of tree canopies. Ecological Modelling 126 (1): 79-93.
CrossRef | Gscholar
Muvengwi J, Mbiba M, Chikumbindi J, Ndagurwa HGT, Mureva A (2018)
Population structure and spatial point-pattern analysis of a mono stand of Acacia polyacantha along a catena in a savanna ecosystem. Forest Ecology and Management 409: 499-508.
CrossRef | Gscholar
Nishimura N, Hara T, Miura M, Manabe T, Yamamoto S (2002)
Tree competition and species coexistence in a warm-temperate oldgrowth evergreen broad-leaved forest in Japan. Plant Ecology 164: 235-248.
CrossRef | Gscholar
Omelko A, Ukhvatkina O, Zhmerenetsky A, Sibirina L, Petrenko T, Bobrovsky M (2018)
From young to adult trees: how spatial patterns of plants with different life strategies change during age development in an old-growth Korean pine broadleaved forest. Forest Ecology and Management 411: 46-66.
CrossRef | Gscholar
Paluch JG (2005)
The influence of the spatial pattern of trees on forest floor vegetation and silver fir (Abies alba Mill.) regeneration in uneven-aged forests. Forest Ecology and Management 205 (1): 283-298.
CrossRef | Gscholar
Petritan IC, Marzano R, Petritan AM, Lingua E (2014)
Overstory succession in a mixed Quercus petraea-Fagus sylvatica old growth forest revealed through the spatial pattern of competition and mortality. Forest Ecology and Management 326 (326): 9-17.
CrossRef | Gscholar
Piao T, Comita LS, Jin G, Kim JH (2013)
Density dependence across multiple life stages in a temperate old-growth forest of northeast China. Oecologia 172: 207-217.
CrossRef | Gscholar
Picard N, Bar-Hen A, Mortier F, Chadoeuf J (2009)
Understanding the dynamics of an undisturbed tropical rain forest from the spatial pattern of trees. Journal of Ecology 97: 97-108.
CrossRef | Gscholar
Pommerening A, Uria-Diez J (2017)
Do large forest trees tend towards high species mingling? Ecological Informatics 42: 139-147.
CrossRef | Gscholar
Potvin C, Dutilleul P (2009)
Neighborhood effects and size asymmetric competition in a tree plantation varying in diversity. Ecology 90 (2): 321-327.
CrossRef | Gscholar
Salas C, LeMay V, Núñez P, Pacheco P, Espinosa A (2006)
Spatial patterns in an old-growth Nothofagus obliqua forest in south-central Chile. Forest Ecology and Management 231: 38-46.
CrossRef | Gscholar
Suzuki SN, Kachi N, Suzuki J-I (2008)
Development of local size hierarchy causes regular spacing of trees in an aven-aged Abies forest: analyses using spatial autocorrelation and the mark correlation function. Annual of Botany 102: 435-441.
CrossRef | Gscholar
Taylor AH, Huang J, Zhou S (2004)
Canopy tree development and undergrowth bamboo dynamics in old-growth Abies-Betula forests in southwestern China: a 12-year study. Forest Ecology and Management 200 (1-3): 347-360.
CrossRef | Gscholar
Torimaru T, Akada S, Ishida K, Matsuda S, Narita M (2013)
Spatial associations among major tree species in a cool-temperate forest community under heterogeneous topography and canopy conditions. Population Ecology 55 (2): 261-275.
CrossRef | Gscholar
Wang H, Wan P, Wang Q, Liu L, Zhang G, Hui G (2017)
Prevalence of inter-tree competition and its role in shaping the community structure of a natural Mongolian scots pine (Pinus sylvestris var. mongolica) forest. Forests 8 (84): 1-14.
CrossRef | Gscholar
Wang H, Peng H, Hui G, Hu Y, Zhao Z (2018)
Large trees are surrounded by more heterospecific neighboring trees in Korean pine broad-leaved natural forests. Scientific Reports 8: 9149.
CrossRef | Gscholar
Weiner J, Stoll P, Muller-Landau H, Jasentuliyana A (2001)
The effects of density, spatial pattern, and competitive symmetry on size variation in simulated plant populations. American Naturalist 158: 438-450.
CrossRef | Gscholar
Wiegand T, Moloney KA (2013)
Handbook of spatial point-pattern analysis in ecology. Chapman and Hall/CRC, New York, USA, pp. 538.
CrossRef | Gscholar
Xiang W, Liu S, Lei X, Frank SC, Tian D, Wang G, Deng X (2013)
Secondary forest floristic composition, structure, and spatial pattern in subtropical China. Journal of Forest Research 18 (1): 111-120.
CrossRef | Gscholar
Yu S, She G, Ye S, Zhou X, Yao X, Li Y (2018)
Characteristics of soil microbial biomass and community composition in Pinus yunnanensis var. tenuifolia secondary forests. Journal of Sustainable Forestry 37 (8): 753-770.
CrossRef | Gscholar
Yuan C, Liu W, Yang G (2015)
Diversity and spatial distribution of lianas in a mid-montane moist evergreen broad-leaved forest in the Ailao Mountains, SW China. Biodiversity Science 23 (3): 332-340.
CrossRef | Gscholar
Zhang J, Chen L, Guo Q, Nie D, Bai X, Jiang Y (1999)
Research on changes of dominant tree population distribution patterns during developmental processes of a climax forest community. Acta Phytoecology Sinica 23: 256-268.
Zhang J, Song B, Li B, Ye J, Wang X, Hao Z (2010)
Spatial patterns and associations of six congeneric species in an old-growth temperate forest. Acta Oecologica 36 (1): 1-10.
CrossRef | Gscholar

This website uses cookies to ensure you get the best experience on our website. More info