Abstract
In temperate mountains, fens have been reported as relict habitats subject to geographical fragmentation and broad climatic gradients, but few studies have analyzed the influence of these factors on plant diversity. Here we investigate the effect of isolation on the vegetation diversity of rich fens (Caricion davallianae) in the mountains of the Iberian Peninsula, the distribution limit of these habitats in south-western Europe. We used plot-based vegetation data from the Pyrenees and the Cantabrian mountain range to evaluate their regional species-pool, occurrence of specialists, beta-diversity and the effect of geo-climatic variables on their species-richness and species-composition. We found a lower ratio of rare specialists in the Pyrenees than in the Cantabrian range, but similar estimates in the species pools, total species-richness per plot and beta-diversity. The isolation of the two mountain regions resulted in different species assemblages best predicted by summer precipitation and bedrock types, showing region-based differences in the response of vegetation and plant specialists to the environment. The tighter correlation between local climate and diversity estimates in the Cantabrian range suggests relict character of rich fens in that region, where climatic conditions have restricted local distribution of formerly more widely distributed specialists. Although there is no relevant evidence of vegetation impoverishment in that region, historical isolation has probably resulted in the existence of fragmentary plant communities. We conclude that fen vegetation may experience long-time persistence in climatically sub-optimal mountain refugia, but related plant specialists may be sensitive to climatic changes and subject to the extinction of local populations.
Similar content being viewed by others
References
Amon JP, Thompson CA, Carpenter QJ, Miner J (2002) Temperate zone fens of the glaciated Midwestern USA. Wetlands 22:301–317
Anderson MJ, Ellingsen KE, McArdle BH (2006) Multivariate dispersion as a measure of beta diversity. Ecol Lett 9:683–693
Bedford B, Godwin K (2003) Fens of the United States: Distribution, characteristics, and scientific connection versus legal isolation. Wetlands 23:608–629
Benito Alonso JL (2003) Las comunidades con Carex bicolor All. del Pirineo. Actas del VI Coloquio de Botánica Pirenaico-Cantábrica. Acta Bot Barcinon 49:229–243
Bergamini A, Peintinger M, Fakheran S, Moradi H, Schmid B, Joshi J (2009) Loss of habitat specialists despite conservation management in fen remnants 1995–2006. Perspect Pl Ecol Evol Syst 11:65–79
Braun-Blanquet J (1932) Plant sociology, the study of plant communities. McGraw-Hill, New York, London (translated and edited by Fuller GD, Conarn HS)
Braun-Blanquet J (1948) La végétation alpine des Pyrénées Orientales. Monografías de la Estación de Estudios Pirenaicos y del Instituto Español de Edafología, Ecología y Fisiología Vegetal, Barcelona
Casanovas L (1991) Estudis sobre l’estructura i l’ecologia de les molleres pirinenques. Thesis doctoral, Facultad de Biología, Universidad de Barcelona, Barcelona
Casas C, Brugués M, Cros RM, Sérgio C (2006) Handbook of mosses of the Iberian Peninsula and the Balearic Islands. Institut d’Estudis Catalans, Barcelona
Castroviejo S (1986–2011) Flora Ibérica, plantas vasculares de la Península Ibérica e Islas Baleares. Real Jardín Botánico, CSIC, Madrid
Chytrý M (2001) Phytosociological data give biased estimates of species richness. J Veg Sci 12:439–444
Cires E, Samain M-S, Goetghebeur P, Fernández Prieto JA (2011) Genetic structure in peripheral Western European populations of the endangered species Cochlearia pyrenaica (Brassicaceae). Pl Syst Evol 297:75–85
Colwell R, Mao CX, Chang J (2004) Interpolating, extrapolating, and comparing incidence-based species accumulation curves. Ecology 85:2717–2727
Csergö A-M, Schönswetter P, Mara G, Deák T, Boşcaiu N and Höhn M. (2009) Genetic structure of peripheral, island-like populations: a case study of Saponaria bellidifolia Sm. (Caryophyllaceae) from the Southeastern Carpathians. Pl Syst Evol 278:33–41
Dierssen K, Dierssen B (1985) Corresponding Caricion bicolori-atrofuscae communities in western Greenland, northern Europe and the central European mountains. Vegetatio 59:151–157
Ellenberg H (1988) Vegetation ecology of central Europe. Cambridge University Press, Cambridge
Felton A (2009) Climate change, conservation and management: an assessment of the 2009 peer-reviewed scientific journal literature. Biodivers & Conservation 18:2243–2253
Fillat F (1984) Estacionalidad de las precipitaciones en España: Clasificación de zonas homogéneas. In Blanco de Pablos A (ed) Avances sobre la investigación en Bioclimatología. Universidad de Salamanca, Salamanca
Flinn KM, Lechowicz MJ, Waterway MJ (2008) Plant species diversity and composition of wetlands within an upland forest. Amer J Bot 95:1216–1224
Font X, Rodríguez-Rojo MP, Acedo C, Biurrun I, Fernández-González F, Lence C, Loidi J, Ninot JM (2010) SIVIM: an on-line database of Iberian and Macaronesian vegetation. Waldökol Landschaftsf Naturschutz 9:15–22
Gilbrich WH (2000) International hydrogeological map of Europe. Waterway 19:1–11
Grootjans AP, Adema EB, Bleuten W, Joosten H, Madaras M, Janáková M (2006) Hydrological landscape settings of base-rich fen mires and fen meadows: an overview. Appl Veg Sci 9:175–184
Habel JC, Assmann T, Schmitt T (2009) Relict species: from past to future. In Habel JC, Assmann T (eds) Relict species, phylogeography and conservation biology. Springer Verlag, Heidelberg, pp 1–8
Hájek M, Horsák M, Hájková P, Dítě D (2006) Habitat diversity of central European fens in relation to environmental gradients and an effort to standardise fen terminology in ecological studies. Perspect Pl Ecol Evol Syst 8:97–114
Hájek M, Hájková P, Sopotlieva D, Apostolova I, Velev N (2008) The Balkan wet grassland vegetation: a prerequisite to better understanding of European habitat diversity. Pl Ecol 195:197–213
Hájek M, Hájková P, Apostolova I, Horsák M, Plášek V, Shaw B, Lazarova M (2009). Disjunct occurrences of plant species in the refugial mires in Bulgaria. Folia Geobot 44:365–386
Hájek M, Horsák M, Tichý L, Hájková P, Dítě D, Jamrichová E (2011) Testing a relict distributional pattern of fen plant and terrestrial snail species at the Holocene scale: a null model approach. J Biogeogr 38:742–755
Hájková P, Hájek M, Apostolova I (2006) Diversity of wetland vegetation in the Bulgarian high mountains, main gradients and context-dependence of the pH role. Pl Ecol 184:111–130
Hájková P, Hájek M, Apostolova I, Zelený D, Dítě D (2008) Shifts in the ecological behaviour of plant species between two distant regions: evidence from the base richness gradient in mires. J Biogeogr 35:282–294
Hijmans RJ, Cameron SE, Parra JL, Jones PG, Jarvis A (2005) Very high resolution interpolated climate surfaces for global land areas. Int J Climatol 25:1965–1978
Hooftman DA, Billeter RC, Schmid B, Diemer M (2004) Genetic effects of habitat fragmentation on common species of swiss fen meadows. Conservation Biol 18:1043–1051
Horsák M, Hájek M, Dítě D, Tichý L (2007) Modern distribution patterns of snails and plants in the Western Carpathian spring fens: is it a result of historical development? J Molluscan Stud 73:53–60
Jiménez-Alfaro B, Díaz González TE, Fernández Pascual E (2011) Grupos de vegetación y hábitats de turberas neutro-basófilas pirenaico-cantábricas. Acta Bot Barcinon 53:47–60
Johnson JB, Steingraeber DA (2003) The vegetation and ecological gradients of calcareous mires in the South Park valley, Colorado. Canad J Bot 81:201–219
Kramp K, Huck S, Niketić M, Tomović G, Schmitt T (2009) Multiple glacial refugia and complex postglacial range shifts of the obligatory woodland plant species Polygonatum verticillatum (Convallariaceae). Pl Biol 11:392–404
Lájer K (2007) Statistical tests as inappropriate tools for data analysis performed on non-random samples of plants community. Folia Geobot 42:115–122
Lepš J, Šmilauer P (2003) Multivariate analysis of ecological data using CANOCO. Cambridge University Press, Cambridge
Lepš J, Šmilauer P (2007) Subjectively sampled vegetation data: Don’t throw out the baby with the bath water. Folia Geobot 42:169–178
Lobo J, Castro I, Moreno JC (2001) Spatial and environmental determinants of vascular plant species richness distribution in the Iberian Peninsula and Balearic Islands. Biol J Linn Soc 73:233–253
Lomolino MV, Riddle BR, Brown JH (2006) Biogeography. Sinauer, Sunderland, MA
McClellan MH, Brock T, Baichtal JF (2003) Calcareous fens in Southeast Alaska. Research Note PNW-RN-536. US Department of Agriculture, Forest Service, Pacific Northwest Research Station. Portland, OR
Michalcová D, Lvončík S, Chytrý M, Hájek O (2011) Bias in vegetation databases? A comparison of stratified-random and preferential sampling. J Veg Sci 22:1654–1103
Nagy L, Grabherr G, Körner Ch, Thompson DBA (2003) Alpine biodiversity in Europe. Ecological Studies, Springer Verlag, Berlin
Nava HS, Fernández Casado MA (2004). Salix hastata subsp. picoeuropeana. In Bañares A, Blanca G, Moreno JC, Ortiz S (eds) Atlas y libro rojo de la flora vascular amenazada de España. Dirección General para la Biodiversidad, Ministerio de Medio Ambiente, Madrid, p 907
Nekola JC (1999) Palaorefugia and neorefugia: The influence of colonization history on community pattern and process. Ecology 80:2459–2473
Nekola JC (2004) Vascular plant compositional gradients within and between Iowa fens. J Veg Sci 15:771–780
Økland RH (1990) A phytoecological study of the mire Northern Kisselbergmosen, SE Norway. III. Diversity and habitat niche relationships. Nordic J Bot 10:191–220
Otýpková Z, Chytrý M (2006) Effects of plot size on the ordination of vegetation samples. J Veg Sci 17:465–472
Pärtel M, Moora M, Zobel M (2001) Variation in species richness within and between calcareous (alvar) grassland stands: the role of core and satellite species. Pl Ecol 157:205–213
Peintinger M, Bergamini A, Schmid B (2003) Species-area relationships and nestedness of four taxonomic groups in fragmented wetlands. Basic Appl Ecol 4:385–394
Peredo EL, Revilla MA, Jiménez-Alfaro B, Bueno A, Fernández Prieto JA, Abbott R (2009) Historical biogeography of a disjuntly distributed, Spanish alpine plant, Senecio boisseri (Asteraceae). Taxon 58:883–892
Prentice IC (1992) A global biome model based on plant physiology and dominance, soil properties and climate. J Biogeogr 19:117–134
Rivas-Martínez S (2007) Mapa de series, geoseries y geopermaseries de vegetación de España (Memoria del mapa de vegetación potencial de España, parte I). Itinera Geobot 17:1–436
Rivas-Martínez S, Díaz TE, Fernández Prieto JA, Loidi J, Penas A (1984) La vegetación de la alta montaña cantábrica. Los Picos de Europa. Ediciones Leonesas, León
Rivas-Martínez S, Díaz González TE, Fernández González F, Izco J, Loidi J, Lousa P, Penas A (2002) Vascular plant communities of Spain and Portugal. Itinera Geobot 15:5–922
Robinson S, Bueno Sánchez A, Jiménez-Alfaro B (2008) Juncus balticus subsp. cantabricus. In Bañares A, Blanca G, Moreno JC, Ortiz S (eds) Atlas y libro rojo de la flora vascular amenazada de España. Dirección General para la Biodiversidad-Sociedad Española de Biología de la Conservación de Plantas, Madrid, pp 72–73
Roekaerts M (2002) The biogeographical regions map of Europe. European Environment Agency, Copenhagen
Roleček J, Chytrý M, Hájek M, Lvončík S, Tichý L (2007) Sampling design in large-scale vegetation studies: do not sacrifice ecological thinking to statistical purism! Folia Geobot 42:199–208
Sekulová L, Hájek M, Hájková P, Mikulášková E, Rozbrojová Z (2011) Alpine wetlands in the West Carpathians: vegetation survey and vegetation-environment relationships. Preslia 83:1–24
Tichý L (2002) JUICE, software for vegetation classification. J Veg Sci 13:451–453
Tichý L, Chytrý M (2006) Statistical determination of diagnostic species for site groups of unequal size. J Veg Sci 17:809–818
Tiner RW (2003) Geographically isolated wetlands of the United States. Wetlands 23:494–516
Topić J, Stančić Z (2006) Extinction of Fen and Bog Plants and their Habitats in Croatia. Biodivers & Conservation 15:3371–3381
van Diggelen R, Middleton BA, Bakker JP, Grootjans A, Wassen M (2006) Fens and floodplains of the temperate zone: Present status, threats, conservation and restoration. Appl Veg Sci 9:157–162
Vargas P (2003) Molecular evidence for multiple diversification patterns of alpine plants in Mediterranean Europe. Taxon 52:463–476
Virolainen M, Suomi, T, Suhonen J, Kuitunen M (1998) Conservation of vascular plants in single large and several small mires: species richness, rarity and taxonomic diversity. J Appl Ecol 35:700–707
Wheeler BD, Proctor MCF (2000) Ecological gradients, subdivisions and terminology of North-West European Mires. J Ecol 88:187–203
Zimmermann M, Vischer-Leopold M, Ellwanger G, Ssymank S, Schröder E (2009) The EC habitats directive and the German Natura 2000 Network of Protected Areas as tool for implementing the conservation of relict species. In Habel JC, Assmann T (eds) Relict species, phylogeography and conservation biology. Springer, Heidelberg, pp 323–340
Acknowledgments
We thank D. Gómez and G. Corriol for providing unpublished data and complete literature about Pyrenean rich fens. We also thank the reviewers and associate editor for the comments and suggestions that permitted improvements to the manuscript. BJA was grant by the European Social Fund through the Spanish Ministry of Science (PTA2007-0726-I). EFP had a PhD grant from the Government of the Principality of Asturias (Plan de Ciencia, Tecnología e Innovación del Principado de Asturias).
Author information
Authors and Affiliations
Corresponding author
Rights and permissions
About this article
Cite this article
Jiménez-Alfaro, B., Fernández-Pascual, E., Díaz González, T.E. et al. Diversity of Rich Fen Vegetation and Related Plant Specialists in Mountain Refugia of the Iberian Peninsula. Folia Geobot 47, 403–419 (2012). https://doi.org/10.1007/s12224-012-9125-1
Received:
Revised:
Accepted:
Published:
Issue Date:
DOI: https://doi.org/10.1007/s12224-012-9125-1