iForest - Biogeosciences and Forestry


Towards a functional phytosociology: the functional ecology of woody diagnostic species and their vegetation classes in Northern Italy

Magda Zanzottera (1-2), Michele Dalle Fratte (1)   , Marco Caccianiga (2), Simon Pierce (3), Bruno Enrico Leone Cerabolini (1)

iForest - Biogeosciences and Forestry, Volume 14, Issue 6, Pages 522-530 (2021)
doi: https://doi.org/10.3832/ifor3730-014
Published: Nov 22, 2021 - Copyright © 2021 SISEF

Research Articles

Vegetation is often classified through phytosociology, which defines floristically and ecologically coherent units identified by diagnostic species. Since species- and community-environment relations are regulated by plant functional traits, it is likely that phytosociology has a strong functional underpinning, although the past and current phytosociology does not explicitly tackle this issue. Here we provide an analysis of functional traits of 221 woody species from Northern Italy, diagnostic of 21 European woody vegetation classes (including alien dominated ones). We assessed whether the functional space occupied by selected species corresponds to the physiognomy and ecology of the vegetation they represent, and whether this could help to evaluate major threats, such as invasion by alien species. For each species we collected from Authors’ datasets leaf trait data (leaf area, specific leaf area, leaf dry matter content, carbon to nitrogen ratio, leaf nitrogen content) and whole plant (plant height, seed mass) traits, and calculated Competitor, Stress-tolerant, Ruderal (CSR) scores. We identified the multidimensional functional trait space of diagnostic species and clustered classes according to their scores in the functional space, to check whether their physiognomy was coherently represented and mirrored in the CSR space. Lastly, we tested for differences between native and neophyte species and their overlap with classes. Diagnostic species mirrored the global spectrum of plant form and function, and classes showed a functional pattern coherent with their physiognomy and ecology. Evergreen dominated classes showed a similar convergence toward conservative characteristics and the stress-tolerant strategy, as opposed to deciduous forest classes that showed a tendency toward the competitive strategy. None of the classes showed a marked ruderal strategy, thus abiotic stress and biotic competition are the main ecological drivers affecting woody vegetation. Neophyte woody species exhibited relatively more competitive strategies compared to natives, and their invasion could be facilitated in resource-limited or mildly disturbed environments, should climate warming or increased nutrient availability occur. We demonstrated that plant traits and CSR strategies of woody diagnostic species reliably indicate the structure and functions of the phytosociological classes they represent, opening the way to the development of a “functional phytosociology”.


CSR Adaptive Strategies, Forests, Shrublands, Global Spectrum, Neophytes, Plant Functional Traits, Structure and Functions, Typical Species

Authors’ address

Magda Zanzottera
Michele Dalle Fratte 0000-0002-7907-1586
Bruno Enrico Leone Cerabolini 0000-0002-3793-0733
Department of Biotechnology and Life Science - DBSV, University of Insubria, v. Dunant 3, 21100 Varese (Italy)
Magda Zanzottera
Marco Caccianiga 0000-0001-9715-1830
Department of Biosciences - DBS, University of Milano, v. Celoria 26, 20133 Milan (Italy)
Simon Pierce 0000-0003-1182-987X
Department of Agricultural and Environmental Sciences - DiSAA, University of Milan, v. Celoria 2, 20133 Milan (Italy)

Corresponding author

Michele Dalle Fratte


Zanzottera M, Dalle Fratte M, Caccianiga M, Pierce S, Cerabolini BEL (2021). Towards a functional phytosociology: the functional ecology of woody diagnostic species and their vegetation classes in Northern Italy. iForest 14: 522-530. - doi: 10.3832/ifor3730-014

Academic Editor

Paola Mairota

Paper history

Received: Dec 23, 2020
Accepted: Sep 23, 2021

First online: Nov 22, 2021
Publication Date: Dec 31, 2021
Publication Time: 2.00 months

Breakdown by View Type

(Waiting for server response...)

Article Usage

Total Article Views: 23669
(from publication date up to now)

Breakdown by View Type
HTML Page Views: 20736
Abstract Page Views: 1158
PDF Downloads: 1496
Citation/Reference Downloads: 3
XML Downloads: 276

Web Metrics
Days since publication: 970
Overall contacts: 23669
Avg. contacts per week: 170.81

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 2021): 2
Average cites per year: 0.67


Publication Metrics

by Dimensions ©

Articles citing this article

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

Barbati A, Marchetti M, Chirici G, Corona P (2014)
European forest types and forest Europe SFM indicators: tools for monitoring progress on forest biodiversity conservation. Forest Ecology and Management 321: 145-157.
CrossRef | Gscholar
Becking RW (1957)
The Zürich-Montpellier school of phytosociology. The Botanical Review 23 (7): 411-488.
CrossRef | Gscholar
Blasi C, Burrascano S (2013)
The role of plant sociology in the study and management of European forest ecosystems. iForest - Biogeosciences and Forestry 6 (2): 55-58.
CrossRef | Gscholar
Bonari G, Fantinato E, Lazzaro L, Sperandii MG, Acosta ATR, Allegrezza M, Assini S, Caccianiga M, Di Cecco V, Frattaroli A, Gigante D, Rivieccio G, Tesei G, Valle B, Viciani D, Albani Rocchetti G, Angiolini C, Badalamenti E, Barberis D, Barcella M, Bazan G, Bertacchi A, Bolpagni R, Bonini F, Bricca A, Buffa G, Calbi M, Cannucci S, Cao Pinna L, Caria MC, Carli E, Cascone S, Casti M, Cerabolini BEL, Copiz R, Cutini M, De Simone L, De Toma A, Dalle Fratte M, Di Martino L, Di Pietro R, Filesi L, Foggi B, Fortini P, Gennaio R, Gheza G, Lonati M, Mainetti A, Malavasi M, Marcenò C, Micheli C, Minuzzo C, Mugnai M, Musarella CM, Napoleone F, Nota G, Piga G, Pittarello M, Pozzi I, Praleskouskaya S, Rota F, Santini G, Sarmati S, Selvaggi A, Spampinato G, Stinca A, Tozzi FP, Venanzoni R, Villani M, Zanatta K, Zanzottera M, Bagella S (2021)
Shedding light on typical species: implications for habitat monitoring. Plant Sociology 58 (1): 157-166.
CrossRef | Gscholar
Braun-Blanquet J (1932)
Plant sociology, the study of plant communities (1st edn). McGraw-Hill Book Co., New York, USA, pp. 439.
Bruelheide H, Dengler J, Purschke O, Lenoir J, Jiménez-Alfaro B, Hennekens SM, Botta-Dukát Z, Chytry M, Field R, Jansen F, Kattge J, Pillar VD, Schrodt F, Mahecha MD, Peet RK, Sandel B, Van Bodegom P, Altman J, Alvarez-Dávila E, Arfin Khan MAS, Attorre F, Aubin I, Baraloto C, Barroso JG, Bauters M, Bergmeier E, Biurrun I, Bjorkman AD, Blonder B, Cayuela L, Cornelissen JHC, Craven D, Dainese M, Derroire G, De Sanctis M, Díaz S, Doleal J, Farfan-Rios W, Feldpausch TR, Fenton NJ, Garnier E, Guerin GR, Gutiérrez AG, Haider S, Hattab T, Henry G, Hérault B, Higuchi P, Hölzel N, Homeier J, Jentsch A, Jürgens N, Karger DN, Kessler M, Kleyer M, Knollová I, Korolyuk AY, Kühn I, Laughlin DC, Lens F, Loos J, Louault F, Lyubenova MI, Malhi Y, Marcen A, Mencuccini M, Müller JV, Munzinger J, Myers-Smith IH, Neill DA, Niinemets U, Orwin KH, Ozinga WA, Penuelas J, Pérez-Haase A, Pet A, Phillips OL, Pärtel M, Reich PB, Römermann C, Rodrigues AV, Sabatini FM, Sardans J, Schmidt M, Seidler G, Espejo JES, Silveira M, Smyth A, Sporbert M, Svenning JC, Tang Z, Thomas R, Tsiripidis I, Vassilev K, Violle C, Virtanen R, Weiher E, Welk E, Wesche K, Winter M, Wirth C, Jandt U (2018)
Global trait-environment relationships of plant communities. Nature Ecology and Evolution 2 (12): 1906-1917.
CrossRef | Gscholar
Brzeziecki B, Kienast F (1994)
Classifying the life-history strategies of trees on the basis of the Grimian model. Forest Ecology and Management 69 (1-3): 167-187.
CrossRef | Gscholar
Chytry M, Tichy L, Holt J, Botta-Dukát Z (2002)
Determination of diagnostic species with statistical fidelity measures. Journal of Vegetation Science 13 (1): 79-90.
CrossRef | Gscholar
Costa-Saura JM, Trabucco A, Spano D, Mereu S (2019)
A height-wood-seed axis which is preserved across climatic regions explains tree dominance in European forest communities. Plant Ecology 220 (4-5): 467-480.
CrossRef | Gscholar
Dalle Fratte M, Bolpagni R, Brusa G, Caccianiga M, Pierce S, Zanzottera M, Cerabolini BEL (2019a)
Alien plant species invade by occupying similar functional spaces to native species. Flora 257 (2): 151419.
CrossRef | Gscholar
Dalle Fratte M, Brusa G, Cerabolini BEL (2019b)
A low-cost and repeatable procedure for modelling the regional distribution of Natura 2000 terrestrial habitats. Journal of Maps 15 (2): 79-88.
CrossRef | Gscholar
Dalle Fratte M, Pierce S, Zanzottera M, Cerabolini BEL (2021)
The association of leaf sulfur content with the leaf economics spectrum and plant adaptive strategies. Functional Plant Biology 48 (9): 924-935.
CrossRef | Gscholar
Davis MA, Grime JP, Thompson K (2000)
Fluctuating resources in plant communities: a general theory of invasibility. Journal of Ecology 88 (3): 528-534.
CrossRef | Gscholar
De Bello F, Thuiller W, Lepš J, Choler P, Clément JC, Macek P, Sebastià MT, Lavorel S (2009)
Partitioning of functional diversity reveals the scale and extent of trait convergence and divergence. Journal of Vegetation Science 20 (3): 475-486.
CrossRef | Gscholar
De La Riva EG, Tosto A, Pérez-Ramos IM, Navarro-Fernández CM, Olmo M, Anten NPR, Marañón T, Villar R (2016)
A plant economics spectrum in Mediterranean forests along environmental gradients: is there coordination among leaf, stem and root traits? Journal of Vegetation Science 27 (1): 187-199.
CrossRef | Gscholar
Díaz S, Cabido M (2001)
Vive la différence: plant functional diversity matters to ecosystem processes. Trends in Ecology and Evolution 16: 646-655.
CrossRef | Gscholar
Díaz S, Kattge J, Cornelissen JHC, Wright IJ, Lavorel S, Dray S, Reu B, Kleyer M, Wirth C, Prentice IC, Garnier E, Bönisch G, Westoby M, Poorter H, Reich PB, Moles AT, Dickie J, Gillison AN, Zanne AE, Chave J, Wright SJ, Sheremetev SN, Jactel H, Baraloto C, Cerabolini BEL, Pierce S, Shipley S, Kirkup D, Casanoves F, Joswig JS, Günther A, Falczuk V, Rüger N, Mahecha MD, Gorné LD (2016)
The global spectrum of plant form and function. Nature 529 (7585): 167-171.
CrossRef | Gscholar
Dick JTA, Laverty C, Lennon JJ, Barrios-ON, Mensink PJ, Britton JR, Médoc V, Boets P, Alexander ME, Taylor NG, Dunn AM, Hatcher MJ, Rosewarne PJ, Crookes S, MacIsaac HJ, Xu M, Ricciardi A, Wasserman RJ, Ellender BR, Weyl OLF, Lucy FE, Banks PB, Dodd JA, MacNeil C, Penk MR, Aldridge DC, Caffrey JM (2017)
Invader Relative Impact Potential: a new metric to understand and predict the ecological impacts of existing, emerging and future invasive alien species. Journal of Applied Ecology 54 (4): 1259-1267.
CrossRef | Gscholar
EEA (2006)
European forest types. Categories and types for sustainable forest management and reporting. Report no. 9/2006, European Environment Agency - EEA, Copenhagen, Denmark, pp. 114.
Online | Gscholar
Freschet GT, Cornelissen JH, Van Logtestijn RS, Aerts R (2010)
Evidence of the “plant economics spectrum” in a subarctic flora. Journal of Ecology 98 (2): 362-373.
CrossRef | Gscholar
Fyllas NM, Michelaki C, Galanidis A, Evangelou E, Zaragoza-Castells J, Dimitrakopoulos PG, Tsadilas C, Arianoutsou M, Lloyd J (2020)
Functional trait variation among and within species and plant functional types in mountainous Mediterranean forests. Frontiers in Plant Science 11: 25.
CrossRef | Gscholar
Garnier E, Navas ML, Grigulis K (2016)
Plant functional diversity: organism traits, community structure, and ecosystem properties. Oxford University Press, Oxford, UK, pp. 231.
CrossRef | Gscholar
Giannetti F, Barbati A, Mancini LD, Travaglini D, Bastrup-Birk A, Canullo R, Nocentini S, Chirici G (2018)
European forest types: toward an automated classification. Annals of Forest Science 75 (1): 1-14.
CrossRef | Gscholar
Grime JP (2001)
Plant strategies, vegetation processes, and ecosystem properties (2nd edn). John Wiley and Sons Ltd, Chichester, UK, pp. 456.
Grime JP (2006)
Trait convergence and trait divergence in herbaceous plant communities: mechanisms and consequences. Journal of Vegetation Science 17 (2): 255-260.
CrossRef | Gscholar
Grime JP, Pierce S (2012)
The evolutionary strategies that shape ecosystems. John Wiley and Sons Ltd, Chichester, UK, pp. 264.
Online | Gscholar
Guo WY, Van Kleunen M, Winter M, Weigelt P, Stein A, Pierce S, Pergl J, Moser D, Maurel N, Lenzner B, Kreft H, Essl F, Dawson W, Pyšek P (2018)
The role of adaptive strategies in plant naturalization. Ecology Letters 21 (9): 1380-1389.
CrossRef | Gscholar
Kattge J, Bönisch G, Díaz S, Lavorel S, Prentice IC, Leadley P, Tautenhahn S, Werner GDA, et al. (2020)
TRY plant trait database-enhanced coverage and open access. Global Change Biology 26 (1): 119-188.
CrossRef | Gscholar
Kleinbauer I, Dullinger S, Peterseil J, Essl F (2010)
Climate change might drive the invasive tree Robinia pseudacacia into nature reserves and endangered habitats. Biological Conservation 143 (2): 382-390.
CrossRef | Gscholar
Kunstler G, Falster D, Coomes DA, Hui F, Kooyman RM, Laughlin DC, Poorter L, Vanderwel M, Vieilledent G, Wright SJ, Aiba M, Baraloto C, Caspersen J, Cornelissen JHC, Gourlet-Fleury S, Hanewinkel M, Herault B, Kattge J, Kurokaw H, Onoda Y, Peñuelas J, Poorter H, Uriarte M, Richardson S, Ruiz-Benito P, Sun IF, Ståhl G, Swenson NG, Thompson J, Westerlund B, Wirth C, Zavala MA, Zeng H, Zimmerman JK, Zimmermann NE, Westoby M (2016)
Plant functional traits have globally consistent effects on competition. Nature 529 (7585): 204-207.
CrossRef | Gscholar
Leuschner C, Ellenberg H (2017)
Ecology of central European forests: vegetation ecology of Central Europe. Springer, Berlin, Germany, vol. 1, pp. 971.
Leuschner C, Meier IC (2018)
The ecology of Central European tree species: trait spectra, functional trade-offs, and ecological classification of adult trees. Perspectives in Plant Ecology, Evolution and Systematics 33: 89-103.
CrossRef | Gscholar
Macedo JA, Alves P, Lomba Vicente J, Henriques R, Granja H, Honrado J (2010)
On the interest of plant functional classifications to study community level effects of increased disturbance on coastal dune vegetation. Acta Botanica Gallica 157 (2): 305-315.
CrossRef | Gscholar
Mucina L, Bültmann H, Dieren K, Theurillat JP, Raus T, Carni A, Sumberová KI, Willner W, Dengler J, Gavilán García R, Chytry M, Hájek M, Di Pietro R, Iakushenko D, Pallas J, Daniëls FJA, Bergmeier E, Santos Guerra A, Ermakov N, Valachovič M, Schaminee JHJ, Lysenko T, Didukh YP, Pignatti S, Rodwell JS, Capelo J, Weber HE, Solomeshch A, Dimopoulos P, Aguiar C, Hennekens SM, Tichy L (2016)
Vegetation of Europe: hierarchical floristic classification system of vascular plant, bryophyte, lichen, and algal communities. Applied Vegetation Science 19 (1): 3-264.
CrossRef | Gscholar
Pierce S, Brusa G, Vagge I, Cerabolini BEL (2013)
Allocating CSR plant functional types: the use of leaf economics and size traits to classify woody and herbaceous vascular plants. Functional Ecology 27 (4): 1002-1010.
CrossRef | Gscholar
Pierce S, Negreiros D, Cerabolini BEL, Kattge J, Díaz S, Kleyer M, Shipley B, Wright SJ, Soudzilovskaia NA, Onipchenko VG, Van Bodegom PM, Frenette-Dussault C, Weiher E, Pinho BX, Cornelissen JHC, Grime JP, Thompson K, Hunt R, Wilson PJ, Buffa G, Nyakunga OC, Reich PB, Caccianiga M, Mangili F, Ceriani RM, Luzzaro A, Brusa G, Siefert A, Barbosa NPU, Chapin III FS, Cornwell WK, Fang J, Fernandes JW, Garnier E, Le Stradic S, Peñuelas J, Melo FPL, Slaviero A, Tabarelli M, Tampucci D (2017)
A global method for calculating plant CSR ecological strategies applied across biomes world-wide. Functional Ecology 31 (2): 444-457.
CrossRef | Gscholar
Pignatti S, Oberdorfer E, Schaminée JHJ, Westhoff V (1995)
On the concept of vegetation class in phytosociology. Journal of Vegetation Science 6 (1): 143-152.
CrossRef | Gscholar
Pignatti S, Guarino R, La Rosa M (2017)
Flora d’italia [Italian Flora] (2nd edn). Edagricole, Bologna, Italy, pp. 1178. [in Italian]
Poldini L, Sburlino G, Buffa G, Vidali M (2011)
Correlations among biodiversity, biomass and other plant community parameters using the phytosociological approach: a case study from the south-eastern Alps. Plant Biosystems 145 (1): 131-140.
CrossRef | Gscholar
Pyšek P, Pergl J, Essl F, Lenzner B, Dawson W, Kreft H, Weigelt P, Winter M, Kartesz J, Nishino M, Antonova LA, Barcelona JF, Cabezas FJ, Cárdenas D, Cárdenas-Toro J, CastaA, Chacón E, Chatelain C, Dullinger S, Ebel AL, Figueiredo E, Fuentes N, Genovesi P, Groom QJ, Henderson L, Inderjit Kupriyanov A, Masciadri S, Maurel N, Meerman J, Morozov O, Moser D, Nickrent D, Nowak PM, Pagad S, Patzelt A, Pelser PB, Seebens H, Shu W, Thomas J, Velayos M, Weber E, Wieringa JJ, Baptiste MP, Van Kleunen M (2017)
Naturalized alien flora of the world. Preslia 89 (3): 203-274.
CrossRef | Gscholar
Pyšek P, Sádlo J, Mandák B, Jarošík V (2003)
Czech alien flora and the historical pattern of its formation: what came first to Central Europe? Oecologia 135 (1): 122-130.
CrossRef | Gscholar
R Core Team (2020)
R: a language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria.
Online | Gscholar
Ricotta C, Acosta AT, Caccianiga M, Cerabolini BEL, Godefroid S, Carboni M (2020)
From abundance-based to functional-based indicator species. Ecological Indicators 118: 106761.
CrossRef | Gscholar
Rodwell JS, Evans D, Schaminée JH (2018)
Phytosociological relationships in European Union policy-related habitat classifications. Rendiconti Lincei - Scienze Fisiche e Naturali 29 (2): 237-249.
CrossRef | Gscholar
Rueda M, Godoy O, Hawkins BA (2018)
Trait syndromes among North American trees are evolutionarily conserved and show adaptive value over broad geographic scales. Ecography 41 (3): 540-550.
CrossRef | Gscholar
Silva MAM, Pinto ADVF, Do Nascimento LM, Lins-e-Silva ACB, De Lima ALA, Rodal MJN (2015)
Does the plant economics spectrum change with secondary succession in the forest? Trees 29 (5): 1521-1531.
CrossRef | Gscholar
Smith TM, Shugart HH, Woodward FI, Burton PJ (1992)
Plant functional types. In: “Vegetation Dynamics and Global Change” (Solomon AM, Shugart HH eds). Chapman and Hall, New York, USA, pp. 272-292.
CrossRef | Gscholar
Thonicke K, Billing M, Von Bloh W, Sakschewski B, Niinemets U, Peñuelas J, Cornelissen JHC, Onoda Y, Van Bodegom P, Schaepman ME, Schneider FD, Walz A (2020)
Simulating functional diversity of European natural forests along climatic gradients. Journal of Biogeography 47 (5): 1069-1085.
CrossRef | Gscholar
Van Der Maarel E, Franklin J (2012)
Vegetation ecology (2nd edn). John Wiley and Sons Ltd, Chichester, UK, pp. 556.
Online | Gscholar
Viciani D, Vidali M, Gigante D, Bolpagni R, Villani M, Acosta ATR, Adorni M, Aleffi M, Allegrezza M, Angiolini C, Assini S, Bagella S, Bonari G, Bovio M, Bracco F, Brundu G, Buffa G, Caccianiga M, Carnevali L, Ceschin S, Ciaschetti G, Cogoni A, Di Cecco V, Foggi B, Frattaroli AR, Genovesi P, Gentili R, Lazzaro L, Lonati M, Lucchese F, Mainetti A, Mariotti M, Minissale P, Paura B, Pellizzari M, Perrino EV, Pirone G, Poggio L, Poldini L, Poponessi S, Prisco I, Prosser F, Puglisi M, Rosati L, Selvaggi A, Sottovia L, Spampinato G, Stanisci A, Stinca A, Venanzoni R, Lastrucci L (2020)
A first checklist of the alien-dominated vegetation in Italy. Plant Sociology 57 (1): 29-54.
CrossRef | Gscholar
Westoby M, Leishman MR, Lord JM (1995)
On misinterpreting the phylogenetic correction. Journal of Ecology 83 (3): 531-534.
CrossRef | Gscholar
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 MJ, Villar R (2004)
The worldwide leaf economics spectrum. Nature 428 (6985): 821-827.
CrossRef | Gscholar
Zanzottera M, Dalle Fratte M, Caccianiga M, Pierce S, Cerabolini BEL (2020)
Community-level variation in plant functional traits and ecological strategies shapes habitat structure along succession gradients in alpine environment. Community Ecology 21: 55-65.
CrossRef | Gscholar

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