Comparing woody plant seedling composition and distribution patterns under different sampling intensities in the 24 ha Gutianshan forest dynamics plot
(2.State Key Laboratory of Vegetation and Environmental Change, Institute of Botany, Chinese Academy of Sciences, Beijing 100093)
(3.Gutianshan National Nature Reserve Administrative Bureau, Kaihua, Zhejiang 324300)
【Abstract】As the seedling stage is the bottleneck in forest tree regeneration, knowledge of the composition and structure of seedlings is crucial to understanding the mechanisms of community assemblage and diversity maintenance. However, based on the limited sampling intensity which is common in previous studies, seedling census datasets are representative of common species, but are ineffective for monitoring rare species. In this study, we established a new seedling monitoring network in the 24 ha GTS (GTS) forest dynamics plot (FDP) consisting of 285 seedling quadrats (5 m × 5 m) in 2012. In the seedling quadrats, all woody plants (DBH < 1 cm and height ≥ 10 cm) were tagged, mapped and measured every two years. We used the first census data to investigate seedling composition, species diversity patterns, seedling, and non-seedling species habitat association. Our results are as follows. (1) There were 138 species, consisting of a total of 20,581 individuals in 285 seedling quadrats of 5 m × 5 m. The Shannon-Wiener diversity index and Rarefied species abundance were higher than those of the 1 m × 1 m seedling quadrats and 24 ha FDP census (DBH ≥ 1 cm). Similar results were found when using rarefaction approach. Compared with 1 m × 1 m seedling quadrats, the number of species showed a significant increase and the species-area curve became more asymptotic. (2) The relationship between seedling abundance and tree species exhibited negative allometry, suggesting that the mortality of plants at the seedling stage was relatively high compared with the rate found at the non-seedling stage due to the negative density dependence. (3) Sampling intensity and selection of DBH cutoffs influence the definition of rare species. Some rare species defined by trees with DBH ≥ 1 cm were not actually rare when individuals with DBH < 1 cm were considered (i.e., Ardisia crenata, Litsea cubeba, Lespedeza thunbergii subsp. formosa). (4) The indicator species of the 5 m × 5 m seedling quadrats were different from those found in the 24 ha FDP and the 1 m × 1 m seedling census. A total of 13 species were significantly correlated with single habitat type in these 5 m × 5 m seedling quadrats, which shared only three species that identified as indicator species for 1 m × 1 m seedling quadrats and two species for non-seedling quadrats. In conclusion, sampling intensity influenced patterns of tree seedling composition and community diversity, and increasing sampling intensity can provide deeper insights into the processes of diversity maintenance.
【Keywords】 Gutianshan (GTS) ; woody plant seedling; sampling intensity; rare species; mechanisms of maintenance of species diversity; Gutianshan (GTS) ; tree seedling; sampling intensity; rare species; mechanisms of maintenance of species diversity;
Antos JA, Guest HJ, Parish R (2005) The tree seedling bank in an ancient montane forest: stress tolerators in a productive habitat. Journal of Ecology, 93, 536–543.
Bachelot B, Kobe RK, Vesk P (2013) Rare species advantage? Richness of damage types due to natural enemies increases with species abundance in a wet tropical forest. Journal of Ecology, 101, 846–856.
Bai XJ, Queenborough SA, Wang XG, Zhang J, Li BH, Yuan ZQ, Xing DL, Lin F, Ye J, Hao ZQ (2012) Effect of local biotic neighbors and habitat heterogeneity on tree and shrub seedling survival in an old-growth temperate forest. Oecologia, 170, 755–765.
Bin Y, Lin GJ, Li BH, Wu LF, Shen Y, Ye WH (2012) Seedling recruitment patterns in a 20 ha subtropical forest plot: hints for niche-based processes and negative density dependence. European Journal of Forest Research, 131, 453–461.
Bunyavejchewi S, Baker P, La Frankie J, Ashton P (2004) Structure, history, and rarity in a seasonal evergreen forest in western Thailand. In: Forest Diversity and Dynamism: Findings from a Network of Large-scale Tropical Forest Plots (eds Losos EC, Leigh EG), pp. 145–158. University of Chicago Press, Chicago.
Cáceres MD, Legendre P, Moretti M (2010) Improving indicator species analysis by combining groups of sites. Oikos, 119, 1674–1684.
Carson WP, Jill TA, Egbert GL, Schnitzer SA (2008) Challenges associated with testing and falsifying the Janzen–Connell hypothesis: a review and critique. In: Tropical Forest Community Ecology (eds Carson WP, Schnitzer SA), pp. 210–241. Wiley–Blackwell Publishing, Chichester.
Chen GK, Kery M, Plattner M, Ma KP, Gardner b (2013) Imperfect detection is the rule rather than the exception in plant distribution studies. Journal of Ecology, 101, 183–191.
Chen L, Mi XC, Comita LS, Zhang LW, Ren HB, Ma KP (2010) Community-level consequences of density dependence and habitat association in a subtropical broadleaved forest. Ecology Letters, 13, 695–704.
Chen ZH, Zhang DM, Lin FP (1996) Floristic and ecological studies of natural seedlings in the lower subtropical forest in Heishiding, Guangdong Province. Acta Phytoecologica Sinica, 20, 568–579 (in Chinese with English abstract).
Comita LS, Aguilar S, Pérez R, Lao S, Hubbell SP (2007) Patterns of woody plant species abundance and diversity in the seedling layer of a tropical forest. Journal of Vegetation Science, 18, 163–174.
Condit R (1998) Tropical Forest Census Plots: Methods and Results from Barro Colorado Island, Panama and a Comparison With Other Plots. Springer Science and Business Media, Berlin.
Connell JH (1971) On the role of natural enemies in preventing competitive exclusion in some marine animals and in rain forest trees. In: Dynamics of Number in Populations (eds Boer PJ, Gradwell GR), pp. 298–310. Centre for Agricultural Publishing and Documentation, Wageningen, The Netherlands.
de Cáceres M, Legendre P (2009) Associations between species and groups of sites: indices and statistical inference. Ecology, 90, 3566–3574.
de Cáceres M, Legendre P, Moretti M (2010) Improving indicator species analysis by combining groups of sites. Oikos, 119, 1674–1684.
Dufrêne M, Legendre P (1997) Species assemblages and indicator species: the need for a flexible asymmetrical approach. Ecological Monographs, 67, 345–366.
Dungan JL, Perry JN, Dale MRT, Legendre P, Citraon-Pousty S, Fortin MJ, Jakomulska A, Miriti M, Rosenberg MS (2002) A balanced view of scale in spatial statistical analysis. Ecography, 25, 626–640.
Goldsmith GR, Comita LS, Morefield LL, Condit R, Hubbell SP (2006) Long-term research impacts on seedling community structure and composition in a permanent forest plot. Forest Ecology and Management, 234, 34–39.
Green PT, Harms KE, Connell JH (2014) Nonrandom, diversifying processes are disproportionately strong in the smallest size classes of a tropical forest. Proceedings of the National Academy of Sciences, USA, 111, 18649–18654.
Harms KE, Wright SJ, Calderón O, Hernández A, Herre EA (2000) Pervasive density-dependent recruitment enhances seedling diversity in a tropical forest. Nature, 404, 493–495.
Heck KL Jr, van Belle G, Simberloff D (1975) Explicit calculation of the rarefaction diversity measurement and the determination of sufficient sample size. Ecology, 56, 1459–1461.
Hu ZH, Yu MJ, Ding BY, Fang T, Qian HY, Chen QC (2003) Types of evergreen broadleaved forests and their species diversity in GTS National Nature Reserve. Chinese Journal of Applied Environmental Biology, 9, 341–345 (in Chinese with English abstract).
Hubbell SP, Foster RB (1986) Commonness and rarity in a neotropical forest: implications for tropical tree conservation. In: Conservation Biology: the Science of Scarcity and Diversity (ed. Soule ME), pp. 205–231. Sinauer Associates, Sunderland, Massachusetts.
Hubbell SP, Dallmeier F, Comiskey J (1998) The maintenance of diversity in a neotropical tree community: conceptual issues, current evidence, and challenges ahead. In: Forest Biodiversity Research, Monitoring and Modeling (eds Dallmeier F, Comiskey JA), pp. 17–44. UNESCO, Paris.
Hurlbert SH (1971) The nonconcept of species diversity: a critique and alternative parameters. Ecology, 52, 577–586.
Jansen PA, Visser MD, Joseph WS, Rutten G, Muller-Landau HC (2014) Negative density dependence of seed dispersal and seedling recruitment in a neotropical palm. Ecology Letters, 17, 11–20.
Janzen DH (1970) Herbivores and the number of tree species in tropical forests. The American Naturalist, 104, 501–528.
Jin Y, Chen JH, Mi XC, Ren HB, Ma KP, Yu MJ (2015) Impacts of the 2008 ice storm on structure and composition of an evergreen broadleaved forest community in eastern China. Biodiversity Science, 23, 610–618 (in Chinese with English abstract).
Kallimanis AS, Mazaris AD, Tzanopoulos J, Halley JM, Pantis JD, Sgardelis SP (2008) How does habitat diversity influence the species–area relationship? Global Ecology and Biogeography, 17, 532–538.
Lai JS (2008) Species Habitat Associations and Species Coexistence on Evergreen Broadleaved Forest in GTS, Zhejiang. Ph D dissertation, Institute of Botany, Chinese Academy of Sciences, Beijing (in Chinese with English abstract).
Lambers JHR, Clark JS, Beckage b (2002) Density-dependent mortality and the latitudinal gradient in species diversity. Nature, 417, 732–735.
Lebrija-Trejos E, Pérez-García EA, Meave JA, Bongers F, Poorter L (2010) Functional traits and environmental filtering drive community assembly in a species-rich tropical system. Ecology, 91, 386–398.
Legendre P, Mi XC, Ren HB, Ma KP, Yu MJ, Sun YF, He FL (2009) Partitioning beta diversity in a subtropical broadleaved forest of China. Ecology, 90, 663–674.
Li L, Chen JH, Ren HB, Mi XC, Yu MJ, Yang b (2010) Spatial patterns of Castanopsis eyrei and Schima superba in mid-subtropical broadleaved evergreen forest in GTS National Reserve, China. Chinese Journal of PlantEcology, 34, 241–252 (in Chinese with English abstract).
Lin LX, Comita LS, Zheng Z, Cao M (2012) Seasonal differentiation in density-dependent seedling survival in a tropical rain forest. Journal of Ecology, 100, 905–914.
Lou LH, Jin SH (2000) Spermatophyta flora of GTS Nature Reserve in Zhejiang. Journal of Beijing Forestry University, 22 (5), 33–39 (in Chinese with English abstract).
Lu ZJ, Bao DC, Guo YL, Lu JM, Wang QG, He D, Zhang KH, Xu YZ, Liu HB, Meng HJ, Huang HD, Wei XZ, Liao JX, Qiao XJ, Jiang MX, Gu ZR, Liao CL (2013) Community composition and structure of Badagongshan FDP in a mid-subtropical mountain evergreen and deciduous broadleaved mixed forest, central China. Chinese Journal of Plant Ecology, 31, 336–344 (in Chinese with English abstract).
Mc Gill BJ, Enquist BJ, Weiher E, Westoby M (2006) Rebuilding community ecology from functional traits. Trends in Ecology and Evolution, 21, 178–185.
Mi XC, Swenson NG, Valencia R, Kress WJ, Erickson DL, Perez AJ, Ren HB, Su SH, Gunatilleke N, Gunatilleke S, Hao ZQ, Ye WH, Cao M, Suresh HS, Dattaraja HS, Sukumar R, Ma KP (2012) The contribution of rare species to community phylogenetic diversity across a global network of forest plots. The American Naturalist, 180, 17–30.
Muscarella R, Uriarte M, Forero-Montaña J, Comita LS, Swenson NG, Thompson J, Nytch CJ, Jonckheere I, Zimmerman JK, Zuidema P (2013) Life-history trade-offs during the seed-to-seedling transition in a subtropical wet forest community. Journal of Ecology, 101, 171–182.
Oksanen J, Blanchet FG, Kindt R, Legendre PR, O’Hara R, Simpson GL, Solymos P, Stevens MH, Wagner H (2013) Vegan: community ecology package. R package version2.0-10. http://CRAN. R-project.org/package=vegan. (accessed on 2015–10–15)
Paine C, Norden N, Chave J, Forget PM, Fortunel C, Dexter KG, Baraloto C (2012) Phylogenetic density dependence and environmental filtering predict seedling mortality in a tropical forest. Ecology Letters, 15, 34–41.
Paine CT, Harms KE (2009) Quantifying the effects of seed arrival and environmental conditions on tropical seedling community structure. Oecologia, 160, 139–150.
Parent S, Simard MJ, Morin H, Messier C (2003) Establishment and dynamics of the balsam fir seedling bank in oldforests of northeastern Quebec. Canadian Journal of Forest Research, 33, 597–603.
Pielou EC (1975) Ecological Diversity. Wiley, New York.
Shannon CE (1948) A mathematical theory of communication. Bell System Technical Journal, 27, 623–656.
Song YC, Yan ER, Song K (2015) Synthetic comparison of eight dynamics plots in evergreen broadleaved forests, China. Biodiversity Science, 23, 139–148 (in Chinese with English abstract).
Song YC, Chen XY, Wang XH (2005) Studies on evergreen broadleaved froests of China: aretrospect and prospect. Journal of East China Normal University (Natural Science), (1), 1–8 (in Chinese with English abstract).
Svenning JC, Wright SJ (2005) Seed limitation in a Panamanian forest. Journal of Ecology, 93, 853–862.
Szwagrzyk J, Szewczyk J, Bodziarczyk J (2001) Dynamics of seedling banks in beech forest: results of a 10-year study on germination, growth and survival. Forest Ecology and Management, 141, 237–250.
Tian K, Chen L, Mi XC, Ma KP, Chen JH (2013) The effect of habitat filtering on tree seedling distribution in a subtropical evergreen broadleaved forest in China. Chinese Science Bulletin, 58, 3561–3569 (in Chinese with English abstract).
Ugland KI, Ellingsen KE (2003) The species–accumulation curve of estimation of species abundance. Journal of Animal Ecology, 72, 888–897.
Wang XG, Wiegand T, Wolf A, Howe R, Davies SJ, Hao ZQ (2011) Spatial patterns of tree species abundance in two temperate forests. Journal of Ecology, 99, 1382–1393.
Webb CO, Gilbert GS, Donoghue MJ (2006) Phylodiversity-dependent seedling mortality, size structure, and disease in a Bornean rain forest. Ecology, 87, 123–131.
Wu ZY (1980) 中国植被. Science Press, Beijing (in Chinese).
Yang QS, Ma ZP, Xie YB, Zhang ZG, Wang ZH, Liu HM, Li P, Zhang N, Wang DL, Yang HB, Fang XF, Yan ER, Wang XH (2011) Community structure and species composition of an evergreen broadleaved forest in Tiantong’s 20 ha dynamic plot, Zhejiang Province, eastern China. Biodiversity Science, 19, 215–223 (in Chinese with English abstract).
Ye WH, Cao HL, Huang ZL, Lian JY, Wang ZG, Li L, Wei SG, Wang ZM (2008) Community structure of a 20 ha lower subtropical evergreen broadleaved forest plot in Dinghushan, China. Journal of Plant Ecology (Chinese Version), 32, 274–286 (in Chinese with English abstract).
Zhu Y, Zhao GF, Zhang LW, Shen GC, Mi XC, Ren HB, YuMJ, Chen JH, Chen SW, Fang T, Ma KP (2008) Community composition and structure of GTS FDP in a mid-subtropical evergreen broadleaved forest, East China. Journal of Plant Ecology (Chinese Version), 32, 262–273 (in Chinese with English abstract).