Interspecific and intraspecific variation in functional traits of subtropical evergreen and deciduous broad-leaved mixed forests

Tang Qingqing 1 Huang Yongtao 1 Ding Yi 1,2,3 Zang Runguo 1,2,3

(1.Key Laboratory of Forest Ecology and Environment of State Forestry Administration, Institute of Forest Ecology, Environment and Protection, Chinese Academy of Forestry, Beijing 100091)
(2.Co-Innovation Center for Sustainable Forestry in Sounthern China, Nanjing Forestry University, Nanjing 100091)
(3.Co-Innovation Center for Sustainable Forestry in Sounthern China, Nanjing Forestry University, Nanjing 210037)

【Abstract】Interspecific variation in plant functional traits is the basis of species coexistence in natural ecosystems. However, intraspecific variation is also extremely important for community assemblage and distribution. Here, we sampled 28 dominant tree species with two different leaf forms (14 evergreen species and 14 deciduous species, respectively, obtained by species abundance ranking) in a subtropical evergreen and deciduous broad-leaved mixed forest in Hubei Xingdoushan National Nature Reserve. Differences in interspecific and intraspecific variations of four functional traits were explored, including specific leaf area (SLA), leaf dry matter content (LDMC), leaf area (LA), and stem specific density (SSD). Results demonstrated that: (1) There were significant differences in the four functional traits between evergreen and deciduous trees. The SLA and LA of evergreen species were significantly lower than those of deciduous species, but LDMC and SSD showed the opposite pattern; (2) Leaf habit was the main source (57.49%) for variation of SLA. Interspecific variability (66.80%) played an important role in LA than intraspecific variability (27.52%). LDMC variation was contributed relatively evenly by interspecific (38.12%) and intraspecific (33.88%) variability. On the contrary, the variation in SSD was explained more by intraspecific (51.50%) than interspecific (32.52%) variability; (3) Correlations of different functional traits on intraspecific levels of both evergreen and deciduous trees were higher than those found for interspecific levels. Our results showed that the variation of functional traits between communities could be influenced significantly by intraspecific variability. However, there were differences in the degree of variation of different functional traits.

【Keywords】 functional trait; coefficient of variation; interspecific variability; intraspecific variability; evergreen and deciduous plants; leaf habit;


【Funds】 Special Program for Science and Technology Basic Work (2013FY111600-2) Special Scientific Research Fund of Public Welfare Profession (201304308)

Download this article


    Ackerly DD, Cornwell WK(2007)A trait-based approach to community assembly:partitioning of species trait values into within-and among-community components. Ecology Letters, 10, 135-145.

    Albert CH, Thuiller W, Yoccoz NG, Douzet R, Aubert S, Lavorel S(2010a)A multi-trait approach reveals the structure and the relative importance of intra-vs. interspecific variability in plant traits. Functional Ecology, 24, 1192-1201.

    Albert CH, Thuiller W, Yoccoz NG, Soudant A, Boucher F, Saccone P, Lavorel S(2010b)Intraspecific functional variability:extent, structure and sources of variation. Journal of Ecology, 98, 604-613.

    Auger S, Shipley B(2013)Interspecific and intraspecific trait variation along short environmental gradients in an old-growth temperate forest. Journal of Vegetation Science, 24, 419-428.

    Bao L, Liu YH(2009)Comparison of leaf functional traits in different forest communities in Mountains Dongling of Beijing. Acta Ecologica Sinica, 29, 3692-3703. (in Chinese with English abstract)

    Baraloto C, Paine CET, Patino S, Bonal D, Herault B, Chave J(2010)Functional trait variation and sampling strategies in species-rich plant communities. Functional Ecology, 24, 208-216.

    Chapin FSI, Zavaleta ES, Eviner VT, Naylor RL, Vitousek PM, Reynolds HL, Hooper DU, Lavorel S, Sala OE, Hobbie SE, Mack MC, Díaz S(2000)Consequences of changing biodiversity. Nature, 405, 234-242.

    Condit R(1998)Ecological implications of changes in drought patterns:shifts in forest composition in Panama. Climatic Change, 39, 413-427.

    Cornelissen JHC, Lavorel S, Garnier E, Díaz S, Buchmann N, Gurvich DE, Reich PB, Steege HT, Morgan HD, Heijden MGAV, Pausas JG, Poorter H(2003)A handbook of protocols for standardised and easy measurement of plant functional traits worldwide. Australian Journal of Botany, 51, 335.

    Cornwell WK, Ackerly DD(2009)Community assembly and shifts in plant trait distributions across an environmental gradient in coastal California. Ecological Monographs, 79, 109-126.

    Díaz S, Asri Y, Band SR, Basconcelo S, Castro-Díez P, Hamzehee GFB, Khoshnevi M, Pérez-Harguindeguy N, Pérez-RontoméMC, Shirvany A, Hodgson JG, Vendramini F, Yazdani S, Abbas-Azimi R, Bogaard A, Boustani S, Charles M, Dehghan M, de Torres-Espuny L, Falczuk V, Guerrero-Campo J, Thompson K, Hynd A, Jones G, Kowsary E, Kazemi-Saeed F, Maestro-Martínez M, Romo-Díez A, Shaw S, Siavash B, Villar-Salvador P, Zak MR, Cabido M, Cornelissen JHC, Jalili A, Montserrat-MartíG, Grime JP, Zarrinkamar F(2004)The plant traits that drive ecosystems:evidence from three continents. Journal of Vegetation Science, 15, 295-304.

    Ding Y, Zang RG, Letcher SG, Liu SR, He FL(2012)Disturbance regime changes the trait distribution, phylogenetic structure and community assembly of tropical rain forests. Oikos, 121, 1263-1270.

    Editorial Board of Forest in China(2000)Forest in China, Vol3, Broad-leaved Forest. China Forestry Publishing House, Beijing(in Chinese).

    Fortunel C, Paine CET, Fine PVA, Kraft NJB, Baraloto C(2014)Environmental factors predict community functional composition in Amazonian forests. Journal of Ecology, 102, 145-155.

    Garnier E, Cortez J, Billes G, Navas ML, Roumet C, Debussche M, Laurent G, Blanchard A, Aubry D, Bellmann A, Neill C, Toussaint JP(2004)Plant functional markers capture ecosystem properties during secondary succession. Ecology, 85, 2630-2637.

    Gratani L, Meneghini M, Pesoli P, Crescente MF(2003)Structural and functional plasticity of Quercus ilex seedlings of different provenances in Italy. Trees-Structure and Function, 17, 515-521.

    Grime JP(1977)Evidence for existence of three primary strategies in plants and its relevance to ecological and evolutionary theory. The American Naturalist, 111, 1169-1194.

    Huang YT, Yao L, Ai XR, LüSA, Ding Y(2015)Quantitative classification of the subtropical evergreen-deciduous broadleaved mixed forest and the deciduous and evergreen species composition structure across two national nature reserves in the southwest of Hubei, China. Chinese Journal of Plant Ecology, 39, 990-1002. (in Chinese with English abstract)

    Jackson BG, Peltzer DA, Wardle DA(2013)The withinspecies leaf economic spectrum does not predict leaf litter decomposability at either the within-species or whole community levels. Journal of Ecology, 101, 1409-1419.

    Jung V, Albert CH, Violle C, Kunstler G, Loucougaray G, Spiegelberger T(2014)Intraspecific trait variability mediates the response of subalpine grassland communities to extreme drought events. Journal of Ecology, 102, 45-53.

    Jung V, Violle C, Mondy C, Hoffmann L, Muller S(2010)Intraspecific variability and trait-based community assembly. Journal of Ecology, 98, 1134-1140.

    Kang M, Chang SX, Yan E, Wang X(2014)Trait variability differs between leaf and wood tissues across ecological scales in subtropical forests. Journal of Vegetation Science, 25, 703-714.

    Kitajima K(1994)Relative importance of photosynthetic traits and allocation patterns as correlates of seedling shade tolerance of 13 tropical trees. Oecologia, 98, 419-428.

    Kraft NJB, Valencia R, Ackerly DD(2008)Functional traits and niche-based tree community assembly in an Amazonian forest. Science, 322, 580-582.

    Laforest-Lapointe I, Martínez-Vilalta J, Retana J(2014)Intraspecific variability in functional traits matters:case study of Scots pine. Oecologia, 175, 1337-1348.

    Laughlin DC(2014)Applying trait-based models to achieve functional targets for theory-driven ecological restoration. Ecology Letters, 17, 771-784.

    Lavorel S, Garnier E(2002)Predicting changes in community composition and ecosystem functioning from plant traits:revisiting the Holy Grail. Functional Ecology, 16, 545-556.

    Liu XJ, Ma KP(2015)Plant functional traits-concepts, applications and future directions. Science China:Life Science, 45, 325-339. (in Chinese with English abstract)

    Messier J, Mcgill BJ, Lechowicz MJ(2010)How do traits vary across ecological scales?A case for trait-based ecology. Ecology Letters, 13, 838-848.

    Mouillot D, Graham N, Villeger S, Mason N, Bellwood DR(2013)A functional approach reveals community responses to disturbances. Trends in Ecology&Evolution, 28, 167-177.

    Perez-Harguindeguy N, Díaz 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, Quetier F, Hodgson JG, Thompson K, Morgan HD, ter Steege H, van der Heijden MGA, 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.

    Plourde BT, Boukili VK, Chazdon RL, Anten N(2015)Radial changes in wood specific gravity of tropical trees:inter‐and intraspecific variation during secondary succession. Functional Ecology, 29, 111-120.

    Poorter L, Kitajima K(2007)Carbohydrate storage and light requirements of tropical moist and dry forest tree species. Ecology, 88, 1000-1011.

    Quigley MF, Platt WJ(2003)Composition and structure of seasonally deciduous forests in the Americas. Ecological Monographs, 73, 87-106.

    Read QD, Moorhead LC, Swenson NG, Bailey JK, Sanders NJ(2014)Convergent effects of elevation on functional leaf traits within and among species. Functional Ecology, 28, 37-45.

    Reich PB, Walters MB, Ellsworth DS(1992)Leaf life-span in relation to leaf, plant, and stand characteristics among diverse ecosystems. Ecological Monographs, 62, 365-392.

    Siefert A, Violle C, Chalmandrier L, Albert CH, Taudiere A, Fajardo A, Aarssen LW, Baraloto C, Carlucci MB, Cianciaruso MV, Dantas LV, de Bello F, Duarte LDS, Fonseca CR, Freschet GT, Gaucherand S, Gross N, Hikosaka K, Jackson B, Jung V, Kamiyama C, Katabuchi M, Kembel SW, Kichenin E, Kraft NJB, Lagerström A, Bagousse-Pinguet YL, Li Y, Mason N, Messier J, Nakashizuka T, Overton JM, Peltzer DA, Pérez-Ramos IM, Pillar VD, Prentice HC, Richardson S, Sasaki T, Schamp BS, Schöb C, Shipley B, Sundqvist M, Sykes MT, 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.

    Siso S, Camarero JJ, Gil-Pelegrin E(2001)Relationship between hydraulic resistance and leaf morphology in broadleaf Quercus species:a new interpretation of leaf lobation. TreesStructure and Function, 15, 341-345.

    Swenson NG, Enquist BJ(2009)Opposing assembly mechanisms in a Neotropical dry forest:implications for phylogenetic and functional community ecology. Ecology, 90, 2161-2170.

    Tomlinson KW, Poorter L, Bongers F, Borghetti F, Jacobs L, van Langevelde F(2014)Relative growth rate variation of evergreen and deciduous savanna tree species is driven by different traits. Annals of Botany, 114, 315-324.

    Valverde BOJ, Smemo KA, Feinstein LM, Kershner MW, Blackwood CB, Guo D(2013)The distribution of below-ground traits is explained by intrinsic species differences and intraspecific plasticity in response to root neighbours. Journal of Ecology, 101, 933-942.

    Violle C, Enquist BJ, Mcgill BJ, Jiang L, Albert CH, Hulshof C, Jung V, Messier J(2012)The return of the variance:intraspecific variability in community ecology. Trends in Ecology&Evolution, 27, 244-252.

    Violle C, Navas M, Vile D, Kazakou E, Fortunel C, Hummel I, Garnier E(2007)Let the concept of trait be functional. Oikos, 116, 882-892.

    Webb CT, Hoeting JA, Ames GM, Pyne MI, Poff NL(2010)Astructured and dynamic framework to advance traits-based theory and prediction in ecology. Ecology Letters, 13, 267-283.

    Weiher E, Freund D, BuntonT, Lee T(2011)Advances, challenges and a developing synthesis of ecological community assembly theory. Philosophical Transactions of the Royal Society B:Biological Sciences, 366, 2403-2413

    Wright IJ, Reich PB, Westoby M, Ackerly DD, Baruch Z, Bongers F, Cavender-Bares J, Chapin T, Cornelissen J, 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.

    Yao L, Ai XR, LüSA, Feng G, Liu JC, Huang YT(2015)Characteristics of community types and structures, and species diversity of natural secondary forests in Xingdou Mountain of Hubei Province. Scientia Silvae Sinicae, 51(11), 1-7. (in Chinese with English abstract)

This Article



Vol 24, No. 03, Pages 262-270

March 2016


Article Outline



  • 1 Materials and methods
  • 2 Results
  • 3 Discussion
  • References