Abstract
In peri-urban regions, the establishment of new forest patches can be affected by specific community assembly rules driving species extinction debts and colonization credits. This might result in species surpluses or deficits and shifts in species composition along forest establishment, whose association with human-driven landscape transformation remains largely unknown. We have compared plant assemblages in peri-urban Mediterranean forests of contrasting history (recent vs. preexisting; post- vs. pre-1956) according to species’ habitat requirements (forest, scrub-grassland, synanthropic or other) and dispersal mode (vertebrate- vs. non-vertebrate), while considering site and landscape characteristics. Species richness and cover were mostly associated to site characteristics, but also to forest connectivity and history. Connectivity increased total species cover, forest species richness and both richness and cover of vertebrate-dispersed species, while reducing non-forest species richness and synanthropic species cover. Regarding forest history, recent stands showed higher total and non-forest species richness, higher scrub-grassland species cover and lower cover of vertebrate-dispersed species than preexisting forests. Thus, recent stands exhibit a species surplus due to the extinction debt of scrub-grassland species and the transient abundance of synanthropic species, despite the observed colonization credit of vertebrate-dispersed species. These results emphasize the importance of forest connectivity when managing plant species assemblage in peri-urban forest patches but also highlight the need to consider recent history of habitat patches to manage time lags in biodiversity change in human-transformed landscapes.
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References
Bagaria G, Helm A, Rodà F, Pino J (2015) Assessing coexisting plant extinction debt and colonization credit in a grassland–forest change gradient. Oecologia 179:823–834. https://doi.org/10.1007/s00442-015-3377-4
Bailey S (2007) Increasing connectivity in fragmented landscapes: an investigation of evidence for biodiversity gain in woodlands. For Ecol Manage 238:7–23. https://doi.org/10.1016/j.foreco.2006.09.049
Bascompte J, Rodriguez MA (2001) Habitat patchiness and plant species richness. Ecol Lett 4:417–420. https://doi.org/10.1046/j.1461-0248.2001.00242.x
Basnou C, Álvarez E, Bagaria G, Guardiola M, Isern R, Vicente P, Pino J (2013) Spatial patterns of land use changes across a mediterranean metropolitan landscape: implications for biodiversity management. Environ Manage 52:971–980. https://doi.org/10.1007/s00267-013-0150-5
Basnou C, Iguzquiza J, Pino J (2015) Examining the role of landscape structure and dynamics in alien plant invasion from urban mediterranean coastal habitats. Landsc Urban Plan 136:156–164. https://doi.org/10.1016/j.landurbplan.2014.12.001
Basnou C, Vicente P, Espelta JM, Pino J (2016) Of niche differentiation, dispersal ability and historical legacies: what drives woody community assembly in recent mediterranean forests? Oikos 125:107–116. https://doi.org/10.1111/oik.02534
Bellemare J, Motzkin G, Foster DR (2002) Legacies of the agricultural past in the forested present: an assessment of historical land-use effects on rich mesic forests. J Biogeogr 29:1401–1420. https://doi.org/10.1046/j.1365-2699.2002.00762.x
Bolòs O, Vigo J, Masalles RM, Ninot JM (2005) Flora manual dels països catalans. Pòrtic, Barcelona
Brothers TS, Spingarn A (1992) Forest fragmentation and alien plant invasion of central indiana old-growth forests. Conserv Biol 6:91–100. https://doi.org/10.1046/j.1523-1739.1992.610091.x
Clotet M, Basnou C, Bagaria G, Pino J (2016) Contrasting historical and current land-use correlation with diverse components of current alien plant invasions in mediterranean habitats. Biol Invasions 18:2897–2909. https://doi.org/10.1007/s10530-016-1181-7
Connell JH (1978) Diversity in tropical rain forests and coral reefs. Science 199:1302–1310. https://doi.org/10.1126/science.199.4335.1302
Cousins SAO, Vanhoenacker D (2011) Detection of extinction debt depends on scale and specialisation. Biol Cons 144:782–787. https://doi.org/10.1016/j.biocon.2010.11.009
Cristofoli S, Piqueray J, Dufrêne M, Bizoux JP, Mahy G (2010) Colonization credit in restored wet heathlands. Restor Ecol 18:645–655. https://doi.org/10.1111/j.1526-100X.2008.00495.x
Dambrine E, Dupouey JL, Laüt L, Humbert L, Thinon M, Beaufils T, Richard H (2007) Present forest biodiversity patterns in France related to former Roman agriculture. Ecology 88:1430–1439. https://doi.org/10.1890/05-1314
Dale MRT, Fortin MJ (2014) Spatial analysis. A guide for ecologists. Cambridge University Press, Cambridge
Deconchat M, Balent G (2001) Effects of soil disturbance and increased light following logging on vegetation at a fine scale. Ann For Sci 58:315–328. https://doi.org/10.1051/forest:2001129
Díaz S, Cabido M, Casanoves F (1998) Plant functional traits and environmental filters at a regional scale. J Veg Sci 9:113–122. https://doi.org/10.2307/3237229
Dobbs C, Kendal D, Nitschke C (2013) The effects of land tenure and land use on the urban forest structure and composition of melbourne. Urban For Urban Gree 12:417–425. https://doi.org/10.1016/j.ufug.2013.06.006
Dormann CM, McPherson JB, Araujo M, Bivand R, Bolliger J, Carl GG, Davies R, Hirzel A, Jetz W, Kissling D, Kuhn I, Ohlemuller RR, Peres-Neto P, Reineking B, Schroder BM, Schurr F, Wilson R (2007) Methods to account for spatial autocorrelation in the analysis of species distributional data a review. Ecography 30:609–628. https://doi.org/10.1111/j.2007.0906-7590.05171.x
Fahrig L (2003) Effects of habitat fragmentation on biodiversity. Annu Rev Ecol Evol Syst 34:487–515. https://doi.org/10.1146/annurev.ecolsys.34.011802.132419
Flinn KM, Vellend M (2005) Recovery of forest plant communities in post-agricultural landscapes. Front Ecol Environ 3:243–250. https://doi.org/10.1890/1540-9295(2005)003[0243:ROFPCI]2.0.CO;2
Godefroid S, Koedam N (2003) How important are large vs. small forest remnants for the conservation of the woodland flora in an urban context? Glob Ecol Biogeogr 12:287–298. https://doi.org/10.1046/j.1466-822X.2003.00035.x
González-Moreno P, Pino J, Gassó N, Vilà M (2013) Landscape context modulates plant invasion in mediterranean forest edges. Biol Invasions 15:547–557. https://doi.org/10.1007/s10530-012-0306-x
Grove AT, Rackham O (2001) The nature of mediterranean Europe: an historical ecology. Yale University Press, New Haven, CT
Guirado M, Pino J, Rodà F (2006) Understorey plant species richness and composition in metropolitan forest archipelagos: effects of forest size, adjacent land use and distance to the edge. Glob Ecol Biogeogr 15:50–62. https://doi.org/10.1111/j.1466-822X.2006.00197.x
Guirado M, Pino J, Rodà F (2007) Comparing the role of site disturbance and landscape properties on understory species richness in fragmented periurban mediterranean forests. Landscape Ecol 22:117–129. https://doi.org/10.1007/s10980-006-9009-y
Hermy M, Verheyen K (2007) Legacies of the past in the present-day forest biodiversity: a review of past land-use effects on forest plant species composition and diversity. Ecol Restor 22:361–371. https://doi.org/10.1007/978-4-431-73238-9_1
Herrera J (1986) Flowering and fruiting phenology in the coastal shrublands of Doñana, south Spain. Vegetatio 68:91–98. https://doi.org/10.1007/BF00045059
Hobbs RJ (2000) Land-use changes and invasions. In: Mooney HA, Hobbs RJ (eds) Invasive species in a changing world. Island Press, Washington, pp 55–64
Hobbs RJ, Arico S, Aronson J, Baron JS, Bridgewater P, Cramer VA, Epstein PR, Ewel JJ, Klink CA, Lugo AE, Norton D, Ojima D, Richardson DM, Sanderson EW, Valladares F, Vilà M, Zamora R, Zobel M (2006) Novel ecosystems: theoretical and management aspects of the new ecological world order. Glob Ecol Biogeogr 15:1–7. https://doi.org/10.1111/j.1466-822X.2006.00212.x
Holt RD, Robinson GR, Gaines MS (1995) Vegetation dynamics in an experimentally fragmented landscape. Ecology 76:1610–1624. https://doi.org/10.2307/1938162
Honnay O, Endels P, Vereecken H, Hermy M (1999) The role of patch area and habitat diversity in explaining native plant species richness in disturbed suburban forest patches in northern Belgium. Divers Distrib 5:129–141. https://doi.org/10.1046/j.1472-4642.1999.00047.x
Hylander K, Ehrlén J (2013) The mechanisms causing extinction debts. Trends Ecol Evol 28:341–346. https://doi.org/10.1016/j.tree.2013.01.010
Jackson ST, Sax DF (2010) Balancing biodiversity in a changing environment: extinction debt, immigration credit and species turnover. Trends Ecol Evol 25:153–160. https://doi.org/10.1016/j.tree.2009.10.001
Jacquemyn H, Butaye J, Hermy M (2003) Influence of environmental and spatial variables on regional distribution of forest plant species in a fragmented and changing landscape. Ecography 26:768–776. https://doi.org/10.1111/j.0906-7590.2003.03620.x
Kowarik I (2011) Novel urban ecosystems, biodiversity, and conservation. Environ Pollut 159:1974–1983. https://doi.org/10.1016/j.envpol.2011.02.022
Kuussaari M, Bommarco R, Heikkinen RK, Helm A, Krauss J, Lindborg R, Öckinger E, Pärtel M, Pino J, Rodà F, Stefanescu C, Teder T, Zobel M, Steffan-Dewenter I (2009) Extinction debt: a challenge for biodiversity conservation. Trends Ecol Evol 24:564–571. https://doi.org/10.1016/j.tree.2009.04.011
Lundholm JT (2009) Plant species diversity and environmental heterogeneity: spatial scale and competing hypotheses. J Veg Sci 20:377–391. https://doi.org/10.1111/j.1654-1103.2009.05577.x
Martín-Queller E, Gil-Tena A, Saura S (2011) Species richness of woody plants in the landscapes of central Spain: the role of management disturbances, environment and non-stationarity. J Veg Sci 22:238–250. https://doi.org/10.1111/j.1654-1103.2010.01242.x
McKinney ML (2006) Urbanization as a major cause of biotic homogenization. Biol Cons 127:247–260. https://doi.org/10.1016/j.biocon.2005.09.005
Montoya D, Zavala MA, Rodriguez MA, Purves D (2008) Animal versus wind dispersal and the robustness of tree species to deforestation. Science 320:1502–1504. https://doi.org/10.1126/science.1158404
Morse NB, Pellissier PA, Cianciola EN, Brereton RL, Sullivan MM, Shonka NK, Wheeler TB, McDowell WH (2014) Novel ecosystems in the anthropocene: a revision of the novel ecosystem concept for pragmatic applications. Ecol Soc 19:12. https://doi.org/10.5751/ES-06192-190212
Murcia C (1995) Edge effects in fragmented forests: implications for conservation. Trends Ecol Evol 10:58–62. https://doi.org/10.1016/S0169-5347(00)88977-6
Naaf T, Kolk J (2015) Colonization credit of post-agricultural forest patches in NE Germany remains 130–230 years after reforestation. Biol Cons 182:155–163. https://doi.org/10.1016/j.biocon.2014.12.002
Navarro-González I, Pérez-Luque AJ, Bonet FJ, Zamora R (2013) The weight of the past: land-use legacies and recolonization of pine plantations by oak trees. Ecol Appl 23:1267–1276. https://doi.org/10.1890/12-0459.1
Piessens K, Honnay O, Nackaerts K, Hermy M (2004) Plant species richness and composition of heathland relics in north-western Belgium: evidence for a rescue-effect? J Biogeogr 31:1683–1692. https://doi.org/10.1111/j.1365-2699.2004.01056.x
Pino J, Arnan X, Rodrigo A, Retana J (2013) Post-fire invasion and subsequent extinction of Conyza spp. in mediterranean forests is mostly explained by local factors. Weed Res 53:470–478. https://doi.org/10.1111/wre.12040
Puerta-Piñero C, Espelta JM, Sánchez-Humanes B, Rodrigo A, Coll L, Brotons L (2012) History matters: previous land use changes determine post-fire vegetation recovery in forested Mediterranean landscapes. For Ecol Manage 279:121–127. https://doi.org/10.1016/j.foreco.2012.05.020
Ramage BS, Roman LA, Dukes JS (2013) Relationships between urban tree communities and the biomes in which they reside. Appl Veg Sci 16:8–20. https://doi.org/10.1111/j.1654-109X.2012.01205.x
Ruiz-Carbayo H, Bonal R, Pino J, Espelta JM (2018) Zero-sum landscape effects on acorn predation associated with shifts in granivore insect community in new holm oak (Quercus ilex) forests. Divers Distrib 24:521–534. https://doi.org/10.1111/ddi.12701
Selkimäki M, González-Olabarria JR, Pukkala T (2012) Site and stand characteristics related to surface erosion occurrence in forests of catalonia (Spain). Eur J For Res 131:727–738. https://doi.org/10.1007/s10342-011-0545-x
Svenning JC, Baktoft KH, Balslev H (2009) Land-use history affects understorey plant species distributions in a large temperate-forest complex, Denmark. Plant Ecol 201:221–234. https://doi.org/10.1007/s11258-008-9557-0
Tilman D, May RM, Lehman CL, Nowak MA (1994) Habitat destruction and the extinction debt. Nature 371:65–66. https://doi.org/10.1038/371065a0
Tomaz C, Alegria C, Monteiro J, Teixeira MC (2013) Land cover change and afforestation of marginal and abandoned agricultural land: a 10-year analysis in a mediterranean region. For Ecol Manage 308:40–49. https://doi.org/10.1016/j.foreco.2013.07.044
Vellend M (2003) Habitat loss inhibits recovery of plant diversity as forests regrow. Ecology 84:1158–1164. https://doi.org/10.1890/0012-9658
Vellend M, Verheyen K, Jacquemyn H, Kolb A, Van Calster H, Peterken G, Hermy M (2006) Extinction debt of forest plants persists for more than a century following habitat fragmentation. Ecology 87:542–548. https://doi.org/10.1890/05-1182
Vellend M, Verheyen K, Flinn KM, Jacquemyn H, Kolb A, Van Calster H, Graae BJ, Bellemare J, Honnay O, Wulf M, Gerhardt F, Hermy M (2007) Homogenization of forest plant communities and weakening of species-environment relationships via agricultural land use. J Ecol 95:565–573. https://doi.org/10.1111/j.1365-2745.2007.01233.x
Verheyen K, Guntenspergen G, Hermy M (2003) An integrated analysis of the effects of past land-use on forest plant species colonization at the landscape scale. J Ecol 91:731–742. https://doi.org/10.1111/j.1365-2745.2007.01233.x
Willson MF, Whelan CJ (1990) Variation in postdispersal survival of vertebrate-dispersed seeds: effects of density, habitat, location, season, and species. Oikos 57:191–198. https://doi.org/10.2307/3565939
Acknowledgements
The study was partially funded by the FORASSEMBLY (CGL2015-70558-P) and NEWFORLAND (RTI2018-099397-B-C22 MCIU/AEI/ERDF, EU) projects of the Spanish Ministry of Economy and Competitiveness, the project SPONFOREST (EU Biodiversa, APCIN-2016-0174), the SGR research group (Generalitat de Catalunya SGR-1006) and the Fondo de Incentivos of the Consejo Nacional para Investigaciones Científicas y Tecnológicas (CONICIT), Ministerio de Ciencia, Tecnología y Telecomunicaciones (MICITT) of Costa Rica.
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The study exclusively received public funding from: The FORASSEMBLY (CGL2015-70558-P) and NEWFORLAND (RTI2018-099397-B-C22) projects of the Spanish Ministry of Economy and Competitiveness. The project SPONFOREST (EU Biodiversa, APCIN-2016-0174), The SGR research group (Generalitat de Catalunya SGR-1006). The Fondo de Incentivos of the Consejo Nacional para Investigaciones Científicas y Tecnológicas (CONICIT), Ministerio de Ciencia, Tecnología y Telecomunicaciones (MICITT) of Costa Rica.
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Tello-García, E., Gamboa-Badilla, N., Álvarez, E. et al. Plant species surplus in recent peri-urban forests: the role of forest connectivity, species’ habitat requirements and dispersal types. Biodivers Conserv 30, 365–384 (2021). https://doi.org/10.1007/s10531-020-02094-3
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DOI: https://doi.org/10.1007/s10531-020-02094-3