Abstract
In recent decades, the European populations of wild boar have grown substantially, as has the impact of this species, owing above all to its rooting activity. Our aim was to investigate the relationships between vascular plant understorey and wild boar rooting intensity. The questions we addressed are: does rooting intensity influence understorey species composition and diversity? Which functional traits are associated with different levels of rooting? We performed a comparative analysis of plant communities in areas with contrasting levels of rooting intensity within a Mediterranean deciduous lowland forest in central Italy. Besides comparing species composition and diversity, we tested the association between species traits and rooting levels through fourth-corner analysis. We found that contrasting levels of rooting were associated to different understorey species composition and evenness, while we observed no significant difference in species richness. In contrast with our expectations, sites with lower rooting returned i) lower evenness values and ii) a higher proportion of species characterized by traits related to resistance or response to herbivory, i.e., spinescence, clonality, endozoochory, underground storage organs, and low height values. Our findings suggest that current vegetation patterns partly depend on the legacy effect of past rooting disturbance, since the areas currently subjected to low rooting intensity were likely to be intensely rooted in the past. These areas may have developed a marked dominance of clonal thorny species that, in turn, inhibited further feeding activities by wild boar.
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Abbate, G., S. Bonacquisti, S. Burrascano, E. Giovi, A. Giuliani, F. Pretto and E. Scassellati. 2015. Woody flora as a predictor of vascular plant richness: An insight in Italy. Plant Biosyst. 149: 565–573.
Anderson, M.J. 2001. A new method for non-parametric multivariate analysis of variance. Austral. Ecol. 26: 32–46.
Avery, E.T. and E.H. Burkhart. 2002. Forest Measurements. 5th ed. McGraw-Hill Higher Education, New York, N Y.
Ballari, S.A. and M.N. Barrios-Garcia. 2014. A review of wild boar Sus scrofa diet and factors affecting food selection in native and introduced ranges. Mammal. Rev. 44: 124–134.
Barrios-Garcia, M.N. and S.A. Ballari. 2012. Impact of wild boar (Sus scrofa) in its introduced and native range: a review. Biol. Invasions 14: 2283–2300.
Bieber, C. and T. Ruf. 2005. Population dynamics in wild boar Sus scrofa: ecology, elasticity of growth rate and implications for the management of pulsed resource consumers. J. Appl. Ecol. 42: 1203–1213.
Blasi, C., G. Capotorti, R. Copiz, D. Guida, B. Mollo, D. Smiraglia, and L. Zavattero. 2014. Classification and mapping of the ecore-gions of Italy. Plant Biosyst. 148: 1255–1345.
Blasi, C., L. Filesi, A. Stanisci, R. Frondoni, R. Di Pietro, and M. L. Carranza. 2002. Excursion to the Circeo National Park. Fitosociologia 39: 91–130.
Boitani, L., L. Mattei, D. Nonis and F. Corsi. 1994. Spatial and activity patterns of wild boars in Tuscany, Italy. J. Mammal. 75: 600–612.
Bratton, S.P. 1975. The Effect of the European Wild Boar, Sus scrofa, on Gray Beech Forest in the Great Smoky Mountains. Ecology 56: 1356–366.
Brooker, R.W., F.T. Maestre, R.M. Callaway, C.L. Lortie, L.A. Cavieres, G. Kunstler, P. Liancourt, K. Tielboerger, J.M.J. Travis, F. Anthelme, C. Armas, L. Coll, E. Corcket, S. Delzon, E. Forey, Z. Kikvidze, J. Olofsson, F.I. Pugnaire, C.L. Quiroz, P. Saccone, K. Schiffers, M. Seifan, B. Touzard and R. Michalet. 2008. Facilitation in plant communities: the past, the present, and the future. J. Ecol. 96: 18–34.
Burrascano, S., F.M. Sabatini and C. Blasi. 2011. Testing indicators of sustainable forest management on understorey composition and diversity in southern Italy through variation partitioning. Plant Ecol. 212: 829–841.
Burrascano, S., E. Giarrizzo, S. Bonacquisti, R. Copiz, E. Del Vico, S. Fagiani, A. Mortelliti and C. Blasi. 2015. Quantifying Sus scrofa rooting effects on the understorey of the deciduous broadleaf forests in Castelporziano Estate (Italy). Rendiconti Lincei. 26, Suppl. 3: 317–324.
Catorci, A., R. Gatti and S. Cesaretti. 2012. Effect of sheep and horse grazing on species and functional composition of sub-Mediterranean grasslands. Appl. Veg. Sci. 15: 459–469.
Catorci, A., S. Cesaretti and F.M. Tardella. 2014. Effect of tall-grass invasion on the flowering-related functional pattern of submedi-terranean hay-meadows. Plant Biosyst. 148: 1127–1137.
Champagnon, J., J. Elmberg, M. Guillemain, M. Gauthier-Clerc and J.D. Lebreton. 2012. Conspecifics can be aliens too: A review of effects of restocking practices in vertebrates. J. Nat. Conserv. 20: 231–241.
Conti, F., G. Abbate, A. Alessandrini and C. Blasi. 2005. An annotated checklist of Italian vascular flora, 1st edn. Palombi Editore, Roma.
Cornelissen, J.H.C., S. Lavorel, E. Garnier, S. Diaz, N. Buchmann, D.E. Gurvich, P.B. Reich, H. ter Steege, H.D. Morgan, M.G.A. van der Heijden, J.G. Pausas and H. Poorter. 2003. A handbook of protocols for standardised and easy measurement of plant functional traits worldwide. Aust. J. Bot. 51: 335–380.
Cuevas, M.F., A. Novillo, C. Campos, M.A. Dacar and R.A. Ojeda. 2010. Food habits and impact of rooting behaviour of the invasive wild boar, Sus scrofa, in a protected area of the Monte Desert, Argentina. J. Arid Environ. 74: 1582–1585.
Cushman, J.H., T.A. Tierney and J.M. Hinds. 2004. Variable effects of feral pig disturbances on native and exotic plants in a California grassland. Ecol. Appl. 14: 1746–1756.
Debussche, M., G. Debussche and J. Lepart. 2001. Changes in the vegetation of Quercus pubescens woodland after cessation of coppicing and grazing. J. Veg. Sci. 12: 81–92.
Dowgiallo, G. and D. Bottini. 1998. Aspetti pedologici del Parco Nazionale del Circeo. In: A. Stanisci and S. Zerunian (eds), Flora e Vegetazine del Parco Nazionale del Circeo. Ministero per le Politiche Agricole, gestione ex A.S.F.D., Sabaudia. pp. 33–46.
Dray, S. and A.B. Dufour. 2007. The ade4 package: implementing the duality diagram for ecologists. J. Stat. Softw. 22: 1–20.
Dray, S. and P. Legendre. 2008. Testing the species traits-environment relationships: the fourth-corner problem revisited. Ecology 89: 3400–3412.
Dufrêne, M. and P. Legendre. 1997. Species assemblages and indicator species: the need for a flexible asymmetrical approach. Ecol. Monogr. 61: 345–366.
Fagiani, S., D. Fipaldini, L. Santarelli, S. Burrascano, E. Del Vico, E. Giarrizzo, M. Mei, A.V. Taglianti, L. Boitani and A. Mortelliti. 2014. Monitoring protocols for the evaluation of the impact of wild boar (Sus scrofa) rooting on plants and animals in forest ecosystems. Hystrix 25: 31–38.
Gilliam, F.S. 2007. The ecological significance of the herbaceous layer in temperate forest ecosystems. Bioscience 57: 845–858.
Gomez, J.M. and J.A. Hodar. 2008. Wild boars (Sus scrofa) affect the recruitment rate and spatial distribution of holm oak (Quercus ilex). For. Ecol. Manag. 256: 1384–1389.
Haaverstad, O., O. Hjeljord and H.K. Wam. 2014. Wild boar rooting in a northern coniferous forest - minor silviculture impact. Scand. J. Forest Res. 29: 90–95.
Heinken, T., M. Schmidt, G. von Oheimb, W.U. Kriebitzsch and H. Ellenberg. 2006. Soil seed banks near rubbing trees indicate dispersal of plant species into forests by wild boar. Basic Appl. Ecol. 7: 31–44.
Hone, J. 2002. Feral pigs in Namadgi National Park, Australia: dynamics, impacts and management. Biol. Conserv. 105: 231–242.
Hunter, M.L. 1990. Wildlife, Forests and Forestry: Principles of Managing Forests for Biological Diversity. Prentice Hall, Englewood Cliffs.
Ickes, K., C.J. Paciorek and S.C. Thomas. 2005. Impacts of nest construction by native pigs (Sus scrofa) on lowland Malaysian rain forest saplings. Ecology 86: 1540–1547.
Kattge, J., S. Diaz, S. Lavorel, C. Prentice, P. Leadley et al. 2011. TRY - a global database of plant traits. Global Change Biol. 17: 2905–2935.
Keddy, P.A. 1992. Assembly and response rules - 2 goals for predictive community ecology. J. Veg. Sci. 3: 157–164.
Kleyer, M., R.M. Bekker, I.C. Knevel, J.P. Bakker, K. Thompson, M. Sonnenschein, P. Poschlod, J.M. van Groenendael, L. Klimes, J. Klimesova, S. Klotz, G.M. Rusch, M. Hermy, D. Adriaens, G. Boedeltje, B. Bossuyt, A. Dannemann, P. Endels, L. Gotzenberger, J.G. Hodgson, A.K. Jackel, I. Kuhn, D. Kunzmann, W.A. Ozinga, C. Romermann, M. Stadler, J. Schlegelmilch, H. J. Steendam, O. Tackenberg, B. Wilmann, J.H.C. Cornelissen, O. Eriksson, E. Garnier and B. Peco. 2008. The LEDA Traitbase: a database of life-history traits of the Northwest European flora. J. Ecol. 96: 1266–1274.
Klimesova, J. and F. de Bello. 2009. CLO-PLA: the database of clonal and bud bank traits of Central European flora. J. Veg. Sci. 20: 511–516.
Lavorel, S., S. Díaz, J.H.C. Cornelissen, E. Garnier, S.P. Harrison, S. McIntyre, J.G. Pausas, N. Pérez-Harguindeguy, C. Roumet and C. Urcelay. 2007. Plant Functional Types: Are We Getting Any Closer to the Holy Grail? In: J. Canadell, D. Pataki and L. Pitelka (eds), Terrestrial Ecosystems in a Changing World. Springer-Verlag, Berlin Heidelberg. pp. 149–164.
Lavorel, S. and E. Garnier. 2002. Predicting changes in community composition and ecosystem functioning from plant traits: revisiting the Holy Grail. Funct. Ecol. 16: 545–556.
Li, L.L., J.B. Shi, J. Wang, Y.M. Gao, L.B. Wang, J.Q. Wang and X. Ying. 2013. Factors influencing wild boar damage in Taohongling National Nature Reserve in China: a model approach. Eur. J. Wildl. Res. 59: 179–184.
Ma, M., X. Zhou and G. Du. 2013. Effects of disturbance intensity on seasonal dynamics of alpine meadow soil seed banks on the Tibetan Plateau. Plant Soil 369: 283–295.
Martinez-Palle, E. and G. Aronne. 1999. Flower development and reproductive continuity in Mediterranean Ruscus aculeatus L. (Liliaceae). Protoplasma 208: 58–64.
Massei, G. and P. Genov. 2004. The environmental impact of wild boar. Galemys 16: 135–145.
Nuttle, T., T.E. Ristau and A.A. Royo. 2014. Long-term biological legacies of herbivore density in a landscape-scale experiment: forest understoreys reflect past deer density treatments for at least 20 years. J. Ecol. 102: 221–228.
Ohashi, H., M. Saito, R. Horie, H. Tsunoda, H. Noba, H. Ishii, T. Kuwabara, Y. Hiroshige, S. Koike, Y. Hoshino, H. Toda and K. Kaji. 2013. Differences in the activity pattern of the wild boar Sus scrofa related to human disturbance. Eur. J. Wildl. Res. 59: 167–177.
Onaindia, M., I. Dominguez, I. Albizu, C. Garbisu and I. Amezaga. 2004. Vegetation diversity and vertical structure as indicators of forest disturbance. For. Ecol. Manag. 195: 341–354.
Paušič, A. and A. Čarni. 2013. Records of past land use are best stored in soil properties. Plant Biosyst. 147: 654–663.
Perea, R. and L. Gil. 2014. Tree regeneration under high levels of wild ungulates: The use of chemically vs. physically-defended shrubs. For. Ecol. Manag. 312: 47–54.
Pignatti, S. 1982. Flora d’Italia. Edagricole, Bologna.
Pinna, W., G. Nieddu, G. Moniello and M.G. Cappai. 2007. Vegetable and animal food sorts found in the gastric content of Sardinian Wild Boar (Sus scrofa meridionalis). J. Anim. Physiol. Anim. Nutr. 91: 252–255.
Royo, A.A. and W.P. Carson. 2006. On the formation of dense under-story layers in forests worldwide: consequences and implications for forest dynamics, biodiversity, and succession. Can. J. For. Res. 36: 1345–1362.
Sabatini, F.M., S. Burrascano, H. Tuomisto and C. Blasi. 2014a. Ground layer plant species turnover and beta diversity in Southern-European old-growth forests. PLoS ONE 9: e95244.
Sabatini, F.M., J.I. Burton, R.M. Scheller, K.L. Amatangelo and D.J. Mladenoff. 2014b. Functional diversity of ground-layer plant communities in old-growth and managed northern hardwood forests. Appl. Veg. Sci. 17: 398–407.
Saezroyuela, C. and J.L. Telleria. 1986. The increased population of the wild boar (Sus scrofa L.) in Europe. Mammal. Rev. 16: 97– 101.
Sandom, C.J., J. Hughes and D.W. Macdonald. 2013. Rewilding the Scottish Highlands: Do wild boar, Sus scrofa, use a suitable foraging strategy to be effective ecosystem engineers? Restor. Ecol. 21: 336–343.
Schley, L., M. Dufrene, A. Krier and A.C. Frantz. 2008. Patterns of crop damage by wild boar (Sus scrofa) in Luxembourg over a 10-year period. Eur. J. Wildl. Res. 54: 589–599.
Schmidt, M., K. Sommer, W.U. Kriebitzsch, H. Ellenberg and G. von Oheimb. 2004. Dispersal of vascular plants by game in northern Germany. Part I: Roe deer (Capreolus capreolus) and wild boar (Sus scrofa). Eur. J. Forest Res. 123: 167–176.
Siemann, E., J.A. Carrillo, C.A. Gabler, R. Zipp and W.E. Rogers. 2009. Experimental test of the impacts of feral hogs on forest dynamics and processes in the southeastern US. For. Ecol. Manag. 258: 546–553.
Sims, N.K., E.A. John and A.J.A. Stewart. 2014. Short-term response and recovery of bluebells (Hyacinthoides non-scripta) after rooting by wild boar (Sus scrofa). Plant Ecol. 215: 1409–1416.
Valenzuela, A.E.J., C.B. Anderson, L. Fasola and J.L. Cabello. 2014. Linking invasive exotic vertebrates and their ecosystem impacts in Tierra del Fuego to test theory and determine action. Acta Oecol. 54: 110–118.
Van Reeuwijk, L.P. (ed.). 2002. Procedures for Soil Analysis, 6th edition. Tech. Pap. 9, ISRIC. Wageningen.
Višnjić, Ć., S. Solaković, F. Mekić, B. Balić, S. Vojniković, M. Dautbašić, S. Gurda, F. Ioras, J. Ratnasingam and I.V. Abrudan. 2013. Comparison of structure, regeneration and dead wood in virgin forest remnant and managed forest on Grmeč Mountain in Western Bosnia. Plant Biosyst. 147: 913–922.
Welander, J. 2000. Spatial and temporal dynamics of wild boar (Sus scrofa) rooting in a mosaic landscape. J. Zool. 252: 263–271.
Wirthner, S., M. Schutz, D.S. Page-Dumroese, M.D. Busse, J.W. Kirchner and A.C. Risch. 2012. Do changes in soil properties after rooting by wild boars (Sus scrofa) affect understory vegetation in Swiss hardwood forests? Can. J. For. Res. 42: 585–592.
Wright, I.J., P.B. Reich, M. Westoby, D.D. Ackerly, Z. Baruch, F. Bongers, J. Cavender-Bares, T. Chapin, J.H.C. Cornelissen, M. Diemer, J. Flexas, E. Garnier, P.K. Groom, J. Gulias, K. Hikosaka, B.B. Lamont, T. Lee, W. Lee, C. Lusk, J.J. Midgley, M.L. Navas, U. Niinemets, J. Oleksyn, N. Osada, H. Poorter, P. Poot, L. Prior, V.I. Pyankov, C. Roumet, S.C. Thomas, M.G. Tjoelker, E.J. Veneklaas and R. Villar. 2004. The worldwide leaf economics spectrum. Nature 428: 821–827.
Zerunian S. 2005. Studio di fattibilità per la reintroduzione di specie significative e gestione sostenibile del cinghiale nel Parco Nazionale del Circeo, Contributo di Ricerca con atto del 02 Febbraio 2005. Parco Nazionale del Circeo.
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Burrascano, S., Copiz, R., Del Vico, E. et al. Wild boar rooting intensity determines shifts in understorey composition and functional traits. COMMUNITY ECOLOGY 16, 244–253 (2015). https://doi.org/10.1556/168.2015.16.2.12
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DOI: https://doi.org/10.1556/168.2015.16.2.12
Keywords
- Circeo National Park
- Deciduous oak forest
- Fourth-corner analysis
- Indicator Species Analysis
- Ruscus aculeatus
- Sus scrofa