ACTA PHYTOGEOGRAPHICA SUECICA 88 EDIDIT SVENSKA VAXTGEOGRAFISKA SALLSKAPET Structure and dynamics •• of alvar vegetation on Oland and some related dry grasslands - Dedicated to Ejvind Rosen on his 65th birthday - Edited by Eddy van der Maarel UPPSALA 2007 ACTA PHYTOGEOGRAPHICA SUECICA 88 EDIDIT SVENSKA VAXTGEOGRAFISKA SALLSKAPET Structure and dynamics •• of alvar vegetation on Oland and some related dry grasslands - Dedicated to EjvindRosen on his 65th birthday - Edited by Eddy van der Maarel UPPSALA 2007 ISBN 978-91-7210-088-6 (paperback) ISBN 978-91-7210-488-4 (cloth) ISSN 0084-5914 Editor: Bengt Carlsson Guest editor: Eddy van der Maarel Technical editor: Marijke van der Maarel-Versluys © Respective author Edidit: Svenska Vaxtgeografiska Sallskapet Villavagen 14 SE-752 36 Uppsala Lay-out: Opulus Press AB Printed by Veiters in Riga, Latvia Acta Phytogeogr. Suec. 88 Contents Ejvind Rosen 65 years 5 E. van der Maarel & J.P. Bakker Structure and dynamics of alvar vegetation on Gland and some related drygrasslands: Introduction 7 E. van der Maarel Drygrassland communities on southern Gland: phytosociology, ecology, and diversity 13 S. Label & J. Dengler Plant diversity and species characteristics of alvar grasslands in Estonia and Sweden 33 A. Helm, P. Urbas & M. Parte/ Plant species turnover in an alvar grassland under different environmental conditions and 43 at different scales R. Huber & E. van der Maarel Species mobility and frequency dynamics in a Swiss limestone grassland 55 K. Studer-Ehrensberger & D.M. Newbery Re-establishment of alvar plant species in abandoned arable fields on Gland 73 J.P. Bakker,E. Rosen & K. Steg Fragmented grasslands on the Baltic island of Gland: Plant community composition and land-use history 83 H. C. Prentice, B.O. Jonsson, M.T. Sykes, M. Ihse & M. Kindstrom Acta Phytogeogr. Suec. 88 4 E. van der Ma arel & J.P Bakker Left above: Eje Rosen, 'promotion day' Uppsala university, June 1982, with laurel-wreath; after obtaining his Ph.D. degree at Uppsala University, June 1982. Left below: Eje enjoying coffee on Gettlinge alvar, June 1989 (photo: Renate Huber); Right above: Lunch pause on the alvar, June 1994; but Eje (left) busy with an identification;further (left to right) Graciela Rusch, Camilla Wessberg, Dai Xiao-bing; Cecilia Dupre, Martin Sykes, Mikael Lonn & Eddy van der Maarel; Right middle: Eje Rosen taking photographs of 1-m2 permanent plots on the alvar, June 1994; Right below: Eje Rosen in typical position during a course for students from Groningen University and Universitat Oldenburg, May 2006 (photo: Roos Ve eneklaas). Acta Phytogeogr. Suec. 88 Ejvind Rosen 65 years * Eddy van der Maare11 & Jan P. Bakker1•2 1Communityand Conservat ion Ecology Group, Groningen University,P.O . Box 14, NL-9750 AA Haren, The Net herlands; 2E-mail [email protected] *Corresponding aut hor; E-mail [email protected] Ejvind Gosta Rosen, Eje for his many friends, young and species known for their irregular occurrence on the alvar, old, was born in Gavle, the capital town of the County e.g. Ranunculus illyricus. of Gavleborgs Ian, north of Uppsala, an old coastal town Eje used to make long seasons of field work on the along the Baltic. Eje studied Biology at Uppsala University island; he lived at the Ecological Research Station of and became a Ph.D. student at the Department of Plant Uppsala University (which has practically ceased to exist Ecology, then called Department of Ecological Botany as such, despite the many efforts of Eje and others). He (Vaxtbiologiska institutionen). His doctoral thesis dealt usually arrived in April and stayed usually until late au­ with the dynamics of the limestone vegetation of the tumn, interruptedby short visits to Uppsala and excursions Great Alvar, especially in relation to sheep grazing. His elsewhere. His family spent summer holidays and shorter supervisor was docent Dr. Erik Sjogren, who knew the periods on bland, but Eje was separated from them for island from his childhood and whose father, a well-known most of the very long season. painter, had portrayed the scenery of the Great Alvar in The willingness of Eje to help colleagues and students many paintings and etches. Sjogren, who continued a much with their projects was proverbial. He used to visit their longer tradition of ecological research on bland - which research sites and give advice on the setup of descriptions started with the lichen research of G. Einar Du Rietz, had and experiments. He was similarly helpful to other guests studiedthe deciduous woodlands on bland and stimulated at the Station and he often gave slide shows on the alvar Rosen to study the open landscape. vegetation and its rarities. After his PhD studies Eje continued with researchon During the 1990s two new developments occurred bland, for which he obtained the 'docent' degree in 1988, in the career of Eje Rosen. First the Department of Plant comparable to the German habilitation degree, a qualifica­ Ecology extended the existing ecological fieldcourse to an tion for a professorship. international event where the second author (who spent a Inspired by Eje 's work, the firstauthor who had joined sabbatical leave at Uppsala University) participated with the Uppsala department in the early 1980s, decided to students from the University of Groningen, The Neth­ build a research group on the fascinating dynamics of erlands, and later also students from the University of the limestone grasslands; the group also included Martin Marburg, Germany, under Prof. Peter Poschlod, and from Sykes, then post-doe, now professor in Lund, who is one Lund under Prof. HorrorPrentice joined the course. Honor of the authors to this volume. When our colleague Robert had started her Swedish career as a postdoc in Uppsala K. Peet from Chapel Hill, NC, US suggested cooperation where she developed genecology with particular interest in in a transcontinental study of species-rich grasslands in typical alvar species; after she became a professor in Lund relation to productivity and dynamics, we decided to extend she continued research and teaching and she contributes our project and from 1984 onwards Eje was instrumental with a paper in this volume. in selecting suitable study sites and maintaining them, in Following our interest in the historical development of the identifying the species and in putting our findings in the landscape in relation to ups and downs in the agricultural broader perspective of the historical development of the management of the alvar, and possibilities for restora­ Alvar landscape in relation to livestock grazing. tion after bush encroachment, the Groningen department Parallel to our common project on fine-scaledynamics introduced seed bank analysis in the course, and also in Eje continued and newly established permanent plot new research projects. A paper in this volume is one of research on scrub encroachment (notably Juniperus the outcomes. communis) under reduction or temporary absence of The ecology course was a highlight for many years grazing, and on the population dynamics of rare vascular because the students (often 25 - 30) combined intensive listening to lectures with field work and close contacts with the teachers, including Eje. Acta Phytogeogr. Suec. 88: 5-6. ISBN 978 91-7210-088-6. A second development was the shift of emphasis from Acta Phytogeogr. Suec. 88 6 E. van der Maarel & J.P. Bakker ecology of the alvar to management. This was stimulated classification and ecology of dry grassland communities on ( by a EU funded LIFE project (1996-1999), which exam­ Hands Stora Alvar. - Nord. J. Bot. 6: 797-809. ined the effects ofgrazing and scrub clearance. Valuable Prentice, H.C., Rosen, E., Moberg, R. & Sjogren, E. 1988. The b informationwas obtained for the large-scale restoration and dry alvar grasslands of land: ecological amplitudes of plant species in relation to vegetation composition. -Acta management progr e supported by the European Union amm Phytogeogr. Suec. 76: 21-46. agri-environment schemes. Eje with his wide knowledge Rejmanek, M. & Rosen, E. 1988. The effects of colonizing shrubs of grazing and its impact on the landscape was the obvi­ (Juniperus communis and Potentilla fruticosa) on species ous person to get involved in research and management richness in the grasslands of Stora Alvaret, (Hand. - Acta plans. Many visitors and students were shown around at Phytogeogr. Suec. 76: 67-72. new experimental sites. Rosen, E. 1988. Shrub expansion in alvar grasslands on bland. One of the major outcomes of this new development, - Acta Phytogeogr. Suec. 76: 87-100. to which Eje has greatly contributed, was the recognition Rosen, E. 198 8. Development and seedling establishment within of the agricultural landscape of southernb land as a World a Juniperus communis stand on bland, Sweden. - ActaBot. Heritage. The description and justification presented by Need. 37: 193-201. Rosen, E. & Sjogren, E. Plant cover in alvar junipers on UNESCO includes: "Human beings have lived here for 1988. bland. Distribution features correlated to shrub size and some fivethousand years and adapted their way of life to shape. -Acta Phytogeogr. Suec. 76: 101-112. the physical constraints of the island. As a consequence, Zobel, K., Zobel, M. & Rosen, E. 1994. An experimental test the landscape is unique, with abundant evidence of con­ of diversity maintenance mechanisms, by a species removal tinuous human settlement from prehistoric times to the experiment in a species-rich wooded meadow. -Folia Geobot. present day." "The landscape of Southern bland takes its Phytota:x. 29: 433-437. contemporary form from its long cultural history, adapting Rosen, E. 1995. Periodic droughts and long-term dynamics to the physical constraints of the geology and topography. of alvar grassland vegetation on bland, Sweden. - Folia Sodra bland is an outstanding example of human settle­ Geobot. Phytotax. 30: 13 1-140. ment, making the optimum use of diverse landscape types Bakker, J.P., Bakker, E.S., Rosen, E., Bekker, R.M.