Abstract
In arid and semi-arid ecosystems, the presence of woody neighbours affects the existence of several herbaceous species by modifying critical aspects of the environment (e.g., soil humidity, nutrient content or light availability) beneath their canopies. Herbaceous species growing in the understory of Pinus pinaster may be distinct from those in open areas due to litter fall, light interception and changes in nutrient availability. We suggest that the overall effect of woody neighbours on herbaceous layer diversity may vary with the scale focus of analysis. To examine this hypothesis, we collected data on the abundance of herbaceous species in open pineland forests of the central Iberian Peninsula (Spain) using sample quadrats of 0.5 m x 0.5 m distributed beneath, at the edge, and outside the canopy of pines in a landscape composed of dunes and plains. The results of CCA ordination revealed significant spatial segregation of herbaceous species reflecting the occurrence of pines and dunes in the landscape. Nested ANOVA disclosed markedly lower species richness beneath the pines, particularly in the dune sites. Species richness partitioning showed higher pine-induced heterogeneity than expected from the sample-based randomized model, leading to significantly increased species richness at the patch level. Hence, the outcome of pine-induced effects on the herbaceous plant diversity is scale-dependent, negative if we focus on separate communities, but positive if the scale focus is extended to whole patches comprising the sum of communities beneath, at the edge, and outside pine tree canopies. These results emphasize the necessity of using various scale perspectives to clarify the different ways in which pines and other woody nurse species affect structure of herbaceous communities in semi-arid Mediterranean ecosystems.
Article PDF
Similar content being viewed by others
Avoid common mistakes on your manuscript.
Abbreviations
- CCA:
-
Canonical Correspondence Analysis
References
Álvarez, A.M., C. Casquet, J.M. Fuster, L. Martín-Parra, J. Martínez-Salanova, and M. Pinado. 1987. Mapa geológico de España 1:50,000. Nava de la Asunción (hoja nº 456) Serv. Publ. Min. Industria y Energía, Madrid.
Álvarez, A.M., V. Cala and J. González. 1993. El factor edáfico en humedales de Cantalejo (provincia de Segovia). Ecología 7: 37–45.
Barbier, S., F. Gosselin and P. Balandier. 2008. Influence of tree species on understory vegetation diversity and mechanisms involved. A critical review for temperate and boreal forests. Forest Ecol. Manag. 254(1): 1–15.
Bertness, M.D. and R.M. Callaway. 1994. Positive interactions in communities. Trends Ecol. Evol. 9: 191–193.
Bertness, M.D. and S.D. Hacker. 1994. Physical stress and positive associations among plants. Am. Nat. 144: 363–372.
Birkett, M.A., K. Chamberlain, A.M. Hooper and J.A. Pickett. 2001. Does allelopathy offer real promise for practical weed management and for explaining rhizosphere interactions involving higher plants? Plant Soil 232: 31–39.
Brooker, R.W. and T.V. Callagham. 1998. The balance between positive and negative plant interactions and its relationship to environmental gradients: a model. Oikos 81: 196–207.
Callaway, R.M., N.M. Nadkarni and B.E. Mahall. 1991. Facilitation and interference of Quercus douglasii on understory productivity in central California. Ecology 72: 1484–1499.
Callaway, R.M. 1995. Positive interactions among plants. Bot. Rev. 61: 306–349.
Callaway, R.M. and L.R. Walker. 1997. Competition and facilitation: a synthetic approach to interactions in plant communities. Ecology 78: 1958–965.
Callaway, R.M. and F.I. Pugnaire. 1999. Facilitation in plant communities. In: Pugnaire, F.I. & F. Valladares (eds.), Handbook of Functional Plant Ecology. Marcel Dekker, New York. pp 272–313.
Callaway, R.M., Z. Kikvidze and D. Kikodze. 2000. Facilitation by unpalatable weeds may conserve plant diversity in overgrazed meadows in the Caucasus Mountains. Oikos 89: 275–282.
Callaway, R.M. 2007. Positive Interactions and Interdependence in Plant Communities. Springer, Dordrecht.
Calonge, G. 1987. El complejo ecológico y la organización de la explotación forestal en la Tierra de Pinares Segoviana. Dipu-tación Provincial de Segovia, Segovia.
Calvo, L., S. Santalla, L. Valbuena, E. Marcos, R. Tárrega and E. Luis-Calabuig. 2008. Post-fire natural regeneration of a Pinus pinaster forest in NW Spain. Plant Ecol. 197: 81–90.
Cavieres, L.A. and E.I. Badano. 2009. Do facilitative interactions increase species richness at the entire community level? J Ecol. 97: 1181–1191.
Cornell, H.V. 1999. Unsaturation and regional influences on species richness in ecological communties: a review of evidence. Eco-science 6: 1–13.
Cornell, H.V. and J.H. Lawton. 1992. Species interactions, local and regional processes, and limits to the richness of ecological communities: a theoretical perspective. J. Anim. Ecol. 61: 1–12.
Costa-Tenorio, M., C. Morla and H. Sáinz-Ollero. 1997. Los bosques ibéricos. Una interpretación Geobotánica. Planeta, Barcelona.
Crist, T.O., J.A. Veech, J.C. Gering and K.S. Summerville. 2003. Partitioning species diversity across landscapes and regions: a hierarchical analysis of α, β and γ diversity. Am. Nat. 162: 734–743.
Gering, J.C, T.O. Crist and J.A. Veech. 2003. Additive partitioning of species diversity across multiple spatial scales: implication for regional conservation of biodiversity. Conserv. Biol. 17: 488–499.
Gillian, F.S. and M.R. Roberts. 2003. Interactions between the herbaceous layer and overstory canopy of eastern forests: a mechanism for linkage. In: Gillian, F.S. and M.R. Roberts (eds.). The Herbaceous Layer in Forests of Eastern North America. Oxford University Press, New York. pp 198–223.
González-Alday, J., C. Martínez-Ruíz and F. Bravo. 2009. Evaluating different harvest intensities over understory diversity and pine seedlings, in a Pinus pinaster Aiton natural stand of Spain. Plant Ecol. 201:211–220.
Greenlee, J.P. and R.M. Callaway. 1996. Effects of abiotic stress on the relative importance of interference and facilitation. Am. Nat. 148: 386–396.
Gutiérrez J.R., P.L. Meserve, L.C. Contreras, H. Vásquez and F.M. Jaksic. 1993. Spatial distribution of soil nutrients and ephemeral plants underneath and outside the canopy of Porlieria chilensis shrubs (Zygophyllaceae) in arid coastal Chile. Oecologia 95: 347–352.
Hacker, S.D. and S.D. Gaines. 1997. Some implications of direct positive interactions for community species diversity. Ecology 78: 1990–2003.
Ha-Lin Zhao, Rui-Lian Zhou, Yong. Zong Su, Hua Zang, Li-Ya Zhao and S. Drake. 2007. Shrub facilitation of desert land restoration in the Horqin Sand Land of Inner Mongolia. Ecol. Eng. 31: 1–8.
Holzapfel, C.,K. Tielbörger, H.A. Parag, J. Kiegel and M. Sternberg. 2006. Annual plant-shrub interactions along an aridity gradient. Basic Appl. Ecol. 7: 268–279.
Huston, M.A. 1994. BiologicaL Diversity: The Coexistence of Species on Changing Landscapes. Cambridge University Press, Cambridge.
Huston, M.A. 1999. Local processes and regional patterns: appropriate scales for understanding variation in the diversity of plants and animals Oikos 86: 393–401.
Kawai, T. and M. Tokeshi. 2007. Testing the facilitation-competition paradigm under the stress-gradient hypothesis: decoupling multiple stress factors. Proc. Roy. Soc. B. 274: 2503–2508.
Koukoura, Z. and A. Kyriazopoulos. 2007. Adaptation of herbaceous plant species in the understory of Pinus brutia. Agroforest. Syst. 70: 11–16.
Lande, R. 1996. Statistics and partitioning of species diversity, and similarities among multiple communities. Oikos 76: 5–13.
Lepš, J. and P. Šmilauer. 2003. Multivariate Analysis of Ecological Data Using CANOCO. Cambridge University Press, Cambridge.
Liancourt, P., R.M. Callaway and R. Michalet. 2005. Stress tolerance and competitive – response ability determine the outcome of bi-otic interaction. Ecology 86: 1611–1618.
López, R.P., D.M. Larrea-Alzácar and T. Ortuño. 2009. Positive effects of shrubs on herbaceous species richness across several spatial scales: evidence from the semiarid Andean subtropics. J. Veg. Sci. 20:728–734.
López-Pintor, A., T. Espigares and J.M. Benayas. 2003. Spatial segregation of plant species caused by Retama sphaerocarpa influence in a Mediterranean pasture: a perspective from the soil seed bank. Plant Ecol. 167: 107–116.
López-Pintor, A., A. Gómez-Sal and J.M. Rey-Benayas. 2006. Shrubs as a source of spatial heterogeneity – the case of Retama sphaerocarpa in Mediterranean pastures of Central Spain. Acta Oecol. 29: 247–255.
Maestre, F.T., F. Valladares and J.F. Reynolds. 2005. Is the change of plant – plant interactions with abiotic stress predictable? A meta – analysis of field results in arid environments. J. Ecol. 93: 748–757.
Moir, W.H. 1966. Influence of Ponderosa Pine on herbaceous vegetation. Ecology 47: 1045–1048.
Moro, M.J., F.I. Pugnaire, P. Haase and J. Puigdefabregas. 1997. Mechanisms of interaction between Retama sphaerocarpa and its understory layer in a semi – arid environment. Ecography 20: 175–184.
Pausas, J.G., C. Bladè, A. Valdecantos, J.P. Seva, D. Fuentes, J.A. Alloza, A Milagrosa, S. Bautista, J. Cortina and R. Vallejo. 2004. Pines and oaks in the restoration of Mediterranean landscapes of Spain: new perspectives for an old practice – a review. Plant Ecol. 171: 209–220.
Pugnaire F.I., P. Haase, J. Puigdefábregas, M. Cueto, S.C. Clark and L.D. Incoll. 1996. Facilitation and succession under the leguminous shrub, Retama sphaerocarpa in a semi-arid environment in south-east Spain. Oikos 76: 455–464.
Pugnaire, F.I. and R. Lázaro. 2000. Seedbank and understorey species composition in a semiarid environment: the effect of shrub age and rainfall. Ann. Bot.-London 86: 807–813.
Pugnaire, F.I. and M.T. Luque. 2001. Changes in plant interactions along a gradient of environmental stress. Oikos 93: 42–49.
Pugnaire, F.I., C. Armas and R. Tirado. 2001. Capítulo 8: Balance de las interacciones entre las plantas en ambientes mediterráneos. In: Zamora, R. and F.I. Pugnaire (eds.). Ecosistemas Mediter-ráneos: Análisis Funcional. Consejo Superior de Investigacio-nes Científicas-Asociación Española de Ecología Terrestre, Granada. pp 213–235.
Salvador, L., R. Alía, D. Agúndez and L. Gil. 2000. Genetic variation and migration pathways of maritime pines (Pinus pinaster Ai-ton) in the Iberian Peninsula. Theor. Appl. Gent. 100: 89–95.
ter Braak, C.J.F. and P. Šmilauer. 2002. CANOCO 4.5 reference manual and CanoDraw for Windows. User’s guide to Canoco for Windows: software for canonical community ordination. Microcomputer Power, Ithaca.
Termiño, J., J.F. García-Hidalgo and M. Segura. 1997. Caracteri-zación y evolución geológica del sistema dunas – humedales de Cantalejo (Segovia). Estudios Geológicos 53: 135–143.
Tewksbury, J.J. and J.D. Lloyd. 2001. Positive interactions under nurse plant: spatial scale, stress gradients and benefactor size. Oecologia 127: 425–434.
Veech, J.A. and T.O. Crist. 2009a. Partition 3.0 user’s manual (unpublished manual) (https://doi.org/www.users.muohio.edu/cristto/partition.htm ).
Veech, J.A. and T.O. Crist. 2009b. PARTITION: software for hierarchical partitioning of species diversity, version 3.0. (https://doi.org/www.users.muohio.edu/cristto/partition.htm).
Wagner, H., O. Wildi and K.C. Ewald. 2000. Additive partitioning of plant species diversity in an agricultural mosaic landscape. Landscape Ecol. 15: 219–227.
Weedon J.T. and J.M. Facelli. 2008. Desert shrubs have negative or neutral effects on annuals at two levels of water availability in arid lands of South-Australia. J. Ecol. 96: 1230–1237.
Author information
Authors and Affiliations
Corresponding author
Rights and permissions
This article is distributed under the terms of the Creative Commons Attribution 4.0 International License (http://creativecommons.org/licenses/by/4.0/), which permits unrestricted use, distribution, and reproduction in any medium, provided you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons license, and indicate if changes were made.
About this article
Cite this article
Madrigal-González, J., García-Rodríguez, J.A., Puerto-Martín, A. et al. Scale-dependent effects of pines on the herbaceous layer diversity in a semi-arid mediterranean ecosystem. COMMUNITY ECOLOGY 11, 77–83 (2010). https://doi.org/10.1556/ComEc.11.2010.1.11
Received:
Revised:
Accepted:
Published:
Issue Date:
DOI: https://doi.org/10.1556/ComEc.11.2010.1.11