What is the optimal number of leaves when measuring leaf area of tree species in a forest community?

doi:10.17521/cjpe.2018.0087

Abstract

Abstract:

Aims Leaf size, as one of the easier measured plant morphological traits, reflects response and adaptation of plants to environment and indicates functions and processes of ecosystem. When measuring leaf size (the leaf area, LA) on the field, the common accepted practice considers that the number of leaves picked off is often 10-20. However, what is the optimal number of leaves remains unknown. In this study, we attempt to determine how many leaves should be investigated when the leaf size of a tree is measured.

Methods This study selected two dominant tree species (Schima superba, Ss and Liquidambar formosana, Lf) from a broadleaved evergreen and deciduous mixed forest in Jinhua Mountain of Zhejiang Province, eastern China. On the basis of sampling (>2 500 leaves for each tree) in five classes of the diameter of breast height (DBH) of tree species and at six directions for each individual, variations of LA in the two tree species are statistically analyzed. The optimal number of leaves, which can mostly represent the common leaf size feature, is further investigated.

Important findings Mean LAof the evergreen tree Ss was smaller than that of the deciduous tree Lf. The former was (41.60 ± 10.88) cm ² (16.74-100.80 cm ²) and the latter was (57.65 ± 19.35) cm ² (11.31-129.51 cm ²). LA of Lf was significantly related to the DBH, but LAof Ss was not. LA of both trees in the middle DBH class (15-20 cm) was not significantly different from their means. LA of two trees have no significant correlations with the sampling directions, but LA at the east, west and bottom had no significant relationships with their means. Considering the representativeness and practicality in the field sampling, the priority of selecting leaves can target the bottom direction of middle diameter mature trees. Random sampling analysis indicated that, the optimal number of leaves for tree LA measurement is species specific. The optimal number of leaves for Ss is 40 and for Lf is at least 170, respectively. Therefore, when measuring leaf area in a forest community, the optimal sampling number of leaves should not be limited to 10-20 leaves. Under sufficient labor, material and time, more leaves should be measured.

http://jtp.cnki.net/bilingual/detail/html/ZWSB201809003

Key words: plant functional traits, leaf area, sampling number, sampling direction, tree age, evergreen and deciduous broadleaved mixed forest

GAO Si-Han, GE Yu-Xi, ZHOU Li-Yi, ZHU Bao-Lin, GE Xing-Yu, LI Kai, NI Jian. What is the optimal number of leaves when measuring leaf area of tree species in a forest community?[J]. Chin J Plant Ecol, 2018, 42(9): 917-925.

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URL: https://www.plant-ecology.com/EN/10.17521/cjpe.2018.0087

https://www.plant-ecology.com/EN/Y2018/V42/I9/917

Figures/Tables 5

Fig. 1 Relationship between leaf area and diameter at breast height of two tree species. A, B, Quartile map; C, D, Scatter plot; E, F, Confidence interval. Ss, Lf, mean values of all samples in all classes of diameter at breast height for Schima superba and Liquidambar formosana, respectively. D2-D6, D2-D6 classes of diameter at breast height. *, p < 0.05; **, p < 0.01.

Fig. 2 Relationship between leaf area and sampling direction of two tree species. A, B, Quartile map; C, D, Scatter plot; E, F, Confidence interval. *, p < 0.05; **, p < 0.01.

Fig. 3 Deviation of leaf area to diameter and sampling direction of two tree species. A, Diameter of Ss; B, Sampling direction of Ss; C, Diameter of Lf; D, Sampling direction of Lf. *, p < 0.05; **, p < 0.01. The vertical line indicates the mean value returning to zero.

Fig. 4 Deviation of mature and non-mature leaf areas to diameter at breast height (DBH) of two tree species. A, Schima superba (Ss). B, Liquidambar formosana (Lf). The horizontal line indicates the mean value returning to zero.

Fig. 5 Mean of random sampling, 95% confidence interval and significant test of leaf areas of two tree species. A, C, All leaf samples. B, D, Leaf samples of medium diameter class and bottom sampling direction. The black line of the leaf area plot indicates the mean value of random sampling, and the red line indicates the 95% confidence interval; the light blue block is the area of significant difference, indicating that the sample size of the right position of the block is the optimal sampling number of leaves. Ss, Schima superba; Lf, Liquidambar formosana.

References 25

[1]	Aerts R ( 1995). The advantages of being evergreen. Trends in Ecology and Evolution, 10, 402-407. DOI URL PMID
[2]	Albert CH, Thuiller W, Yoccoz NG, Soudant A, Boucher F, Saccone P, Lavorel S ( 2010). Intraspecific functional variability: Extent, structure and sources of variation. Journal of Ecology, 98, 604-613. DOI URL
[3]	Auger S, Shipley B ( 2013). Inter-specific and intra-specific trait variation along short environmental gradients in an old-growth temperate forest. Journal of Vegetation Science, 24, 419-428. DOI URL
[4]	Bolnick DI, Amarasekare P, Araojo MS, Burger R, Levine JM, Novak M, Rudolf VHW, Schreiber SJ, Urban MC, Vasseur DA ( 2011). Why intraspecific trait variation matters in community ecology. Trends in Ecology and Evolution, 26, 183-192. DOI URL PMID
[5]	Bu WS, Schmid B, Liu XJ, Li Y, Hardtle W, von Oheimb G, Liang Y, Sun ZK, Huang YY, Bruelheide H, Ma KP ( 2017). Interspecific and intraspecific variation in specific root length drives aboveground biodiversity effects in young experimental forest stands. Journal of Plant Ecology, 10, 158-169. DOI URL
[6]	Chen YT, Xu ZZ ( 2014). Review on research of leaf economics spectrum. Chinese Journal of Plant Ecology, 38, 1135-1153. DOI URL
	[ 陈莹婷, 许振柱 ( 2014). 植物叶经济谱的研究进展. 植物生态学报, 38, 1135-1153.] DOI URL
[7]	Cornelissen JHC, Lavorel S, Garnier E, Diaz S, Buchmann N, Gurvich DE, Reich PB, Ter Steege H, Morgan HD, van der Heijden MGA, Pausas JG, Poorter H ( 2003). A handbook of protocols for standardized and easy measurement of plant functional traits worldwide. Australian Journal of Botany, 51, 335-380. DOI URL
[8]	Díaz S, Kattge J, Cornelissen JHC, Wright IJ, Lavorel S, Dray S, Reu B, Kleyer M, Wirth C, Prentice IC, Garnier E, Boenisch G, Westoby M, Poorter H, Reich PB, Moles AT, Dickie J, Gillison AN, Zanne AE, Chave J, Wright SJ, Sheremet’ev S, Jactel H, Baraloto C, Cerabolini B, Pierce S, Shipley B, Kirkup D, Casanoves F, Joswig J, Günther A, Falczuk V, Rüger N, Mahecha MD, Gorné LD ( 2016). The global spectrum of plant form and function. Nature, 529, 167-171. DOI URL PMID
[9]	Donovan LA, Maherali H, Caruso CM, Huber H, de Kroon H ( 2011). The evolution of the worldwide leaf economics spectrum. Trends in Ecology and Evolution, 26, 88-95. DOI URL PMID
[10]	Fang F, Guo SL ( 2004). DCA ordination on woody plant communities in Beishan Mountain of Jinhua, Zhejiang Province. Journal of Zhejiang Normal University (Natural Science), 27(1), 55-61. DOI URL
	[ 方芳, 郭水良 ( 2004). 浙江金华北山木本植物群落的DCA排序. 浙江师范大学学报(自然科学版), 27(1), 55-61.] DOI URL
[11]	Givnish TJ ( 2002). Adaptive significance of evergreen vs. deciduous leaves: Solving the triple paradox. Silva Fennica, 36, 703-743.
[12]	Guo SL, Liu P, Chen G, Lu X ( 1993). Observations on the floristic and vegetation in Beishan Mountain of Jinhua in Zhejiang Province. Journal of Zhejiang Normal University (Natural Science), 16(2), 59-67.
	[ 郭水良, 刘鹏, 陈刚, 卢晓 ( 1993). 浙江金华北山植物区系及植被. 浙江师范大学学报(自然科学版), 16(2), 59-67.]
[13]	Li MC, Zhu JJ, Sun YR ( 2009). Responses of specific leaf area of dominant tree species in Northeast China secondary forests to light intensity. Chinese Journal of Ecology, 28, 1437-1442.
	[ 李明财, 朱教君, 孙一荣 ( 2009). 东北次生林主要树种比叶面积对光照强度的响应. 生态学杂志, 28, 1437-1442.]
[14]	Liu XJ, Ma KP ( 2015). Plant functional traits—Concepts, applications and future directions. Scientia Sinica Vitae, 45, 325-339. DOI URL
	[ 刘晓娟, 马克平 ( 2015). 植物功能性状研究进展. 中国科学: 生命科学, 45, 325-339.] DOI URL
[15]	McDonald PG, Fonseca CR, Overton J, Westoby M ( 2003). Leaf-size divergence along rainfall and soil-nutrient gradients: Is the method of size reduction common among clades? Functional Ecology, 17, 50-57. DOI URL
[16]	Meng TT, Ni J, Wang GH ( 2007). Plant functional traits, environments and ecosystem functioning. Journal of Plant Ecology (Chinese Version), 31, 150-165. DOI URL
	[ 孟婷婷, 倪健, 王国宏 ( 2007). 植物功能性状与环境和生态系统功能. 植物生态学报, 31, 150-165.] DOI URL
[17]	Messier J, Lechowicz MJ, McGill BJ, Violle C, Enquist BJ ( 2017). Interspecific integration of trait dimensions at local scales: The plant phenotype as an integrated network. Journal of Ecology, 105, 1775-1790. DOI URL
[18]	Peppe DJ, Royer DL, Cariglino B, Oliver SY, Newman S, Leight E, Enikolopov G, Fernandez-Burgos M, Herrera F, Adams JM, Correa E, Currano ED, Erickson JM, Hinojosa LF, Hoganson JW, Iglesias A, Jaramillo CA, Johnson KR, Jordan GJ, Kraft NJ, Lovelock EC, Lusk CH, Niinemets U, Pe?uelas J, Rapson G, Wing SL, Wright IJ ( 2011). Sensitivity of leaf size and shape to climate: Global patterns and paleoclimatic applications. New Phytologist, 190, 724-739. DOI URL PMID
[19]	Pérez-Harguindeguy N, Diaz S, Garnier E, Lavorel S, Poorter H, Jaureguiberry P, Bret-Harte MS, Cornwell WK, Craine JM, Gurvich DE, Urcelay C, Veneklaas EJ, Reich PB, Poorter L, Wright IJ, Ray P, Enrico L, Pausas JG, de Vos AC, Buchmann N, Funes G, Quétier F, Hodgson JG, Thompson K, Morgan HD, ter Steege H, van der Heijden MG A, Sack L, Blonder B, Poschlod P, Vaieretti MV, Conti G, Staver AC, Aquino S, Cornelissen JHC ( 2013). New handbook for standardised measurement of plant functional traits worldwide. Australian Journal of Botany, 61, 167-234. DOI URL
[20]	Royer DL, McElwain JC, Adams JM, Wilf P ( 2008). Sensitivity of leaf size and shape to climate within Acer rubrum and Quercus kelloggii. New Phytologist, 179, 808-817.
[21]	Siefert A, Violle C, Chalmandrier L, Albert CH, Taudiere A, Fajardo A, Aarssen LW, Baraloto C, Carlucci MB, Cianciaruso MV, Dantas VD, DeBello F, Duarte LDS, Fonseca CR, Freschet GT, Gaucherand S, Gross N, Hikosaka K, Jackson B, Jung V, Kamiyama C, Katabuchui M, Kembel SW, Kichenin E, Kraft NJB, Lagerstrom A, Le Bagousse-Pinguer Y, Li YZ, Mason N, Messier J, Nakashizuka T, Overton JM, Peltzer DA, Perez-Ramos IM, Pillar VD, Prentice HC, Richardson S, Sasaki T, Schamp BS, Vandewalle M, Wardle DA ( 2015). A global meta-analysis of the relative extent of intraspecific trait variation in plant communities. Ecology Letters, 18, 1406-1419. DOI URL PMID
[22]	Violle C, Enquist BJ, McGill BJ, Jiang L, Albert CH, Huishof C, Jung V, Messier J ( 2012). The return of the variance: Intraspecific variability in community ecology. Trends in Ecology and Evolution, 27, 244-252. DOI URL
[23]	Wang CS, Wang SP ( 2015). A review of research on responses of leaf traits to climate change. Chinese Journal of Plant Ecology, 39, 206-216. DOI URL
	[ 王常顺, 汪诗平 ( 2015). 植物叶片性状对气候变化的响应研究进展. 植物生态学报, 39, 206-216.] DOI URL
[24]	Westoby M, Falster DS, Moles AT, Vesk PA, Wright IJ ( 2002). Plant ecological strategies: Some leading dimensions of variation between species. Annual Review of Ecology and Systematics, 33, 125-159. DOI URL
[25]	Wright IJ, Reich PB, Westoby M, Ackerly DD, Baruch Z, Bongers F, Cavender-Bares J, Chapin T, Cornelissen JHC, Diemer M, Flexas J, Garnier E, Groom PK, Gulias J, Hikosaka K, Lamont BB, Lee T, Lee W, Lusk C, Midgley JJ, Navas ML, Niinemets U, Oleksyn J, Osada N, Poorter H, Poot P, Prior L, Pyankov VI, Roumet C, Thomas SC, Tjoelker MG, Veneklaas EJ, Villar R ( 2004). The worldwide leaf economics spectrum. Nature, 428, 821-827. DOI URL