Research article

Faunal diversity of Fagus sylvatica forests: A regional and European perspective based on three indicator groups

H. Walentowski , S. Müller-Kroehling, E. Bergmeier, M. Bernhardt-Römermann, M.M. Gossner, A. Reif, E.-D. Schulze, H. Bußler, C. Strätz, W. Adelmann

H. Walentowski
Bavarian State Institute of Forestry, Hans-Carl-von-Carlowitz-Platz 1, 85354 Freising. Email: Helge.Walentowski@lwf.bayern.de
S. Müller-Kroehling
Bavarian State Institute of Forestry, Hans-Carl-von-Carlowitz-Platz 1, 85354 Freising
E. Bergmeier
Department of Vegetation & Phytodiversity Analysis, University of Göttingen, Germany
M. Bernhardt-Römermann
Institute of Ecology, Friedrich Schiller University Jena, Germany
M.M. Gossner
Terrestrial Ecology Research Group, Department of Ecology and Ecosystem Management, Center for Food and Life Sciences Weihenstephan, Munich University of Technology, Germany
A. Reif
Department of Vegetation Science and Site Classification, University of Freiburg, Germany
E.-D. Schulze
Max-Planck-Institute for Biogeochemistry, Jena, Germany
H. Bußler
Bavarian State Institute of Forestry, Freising, Germany
C. Strätz
Office for Ecological Studies, Bayreuth, Germany
W. Adelmann
Bavarian Academy for Nature Conservation and Landscape Management, Germany

Online First: May 08, 2014
Walentowski, H., Müller-Kroehling, S., Bergmeier, E., Bernhardt-Römermann, M., Gossner, M., Reif, A., Schulze, E., Bußler, H., Strätz, C., Adelmann, W. 2014. Faunal diversity of Fagus sylvatica forests: A regional and European perspective based on three indicator groups. Annals of Forest Research DOI:10.15287/afr.2014.172


While the postglacial history of European beech (Fagus sylvatica) and the plant species composition of beech forests in  Central Europe are fairly well understood, the faunal biodiversity has been less well investigated. We studied three groups of  mostly sedentary organisms in beech forest at regional and European scales by combining field studies with a compilation of existing literature and expert knowledge. Specifically, we examined the relationship between host tree genera and saproxylic  beetles, and the diversity and composition of forest ground-dwelling molluscs and ground beetles in relation to the abundance  of beech. At a west central European scale (Germany), where beech has a “young” ecological and biogeographical history,  we found 48 primeval forest relict species of saproxylic beetles associated with beech, 124 ground beetles and 91 molluscs  inhabiting beech forest, yet none exclusive of west central European beech forests. High levels of faunal similarity between beech and other woodland trees suggested that many of the beech forest dwelling species are euryoecious and likely to  originate from mid-Holocene mixed broadleaf forests. Beech forests of the mountain ranges in southern and east central  Europe, which are ecologically and biogeographically “old”, were found to harbour distinct species assemblages, including  beech forest specialists (such as 10 carabid species in the Carpathians) and narrow-range endemics of broadleaf forest. The  observed biodiversity patterns suggest differentiated conservation priorities in “young” and “old” European beech forest  regions.


Andersson G.K.S., Birkhofer K., Rundlöf M., Smith H.G., 2013. Landscape heterogeneity and farming practice alter the species composition and taxonomic breadth of pollinator communities. Basic Appl Ecol 14 (7):540-546. DOI: 10.1016/j.baae.2013.08.003

Birks H.J.B., 1980. British trees and insects: a test of the time hypothesis over the last 13,000 years. Am Nat 115 (4): 600-605. DOI: 10.1086/283584

Bohn U., Neuhäusl R., Gollub G., Hettwer C., Neuhäuslová Z., Schlüter H., Weber H., 2000 / 2003. Karte der natürlichen Vegetation Europas (Map of the Natural Vegetation of Europe). Maßstab (Scale) 1 : 2 500 000. Teil 1: Erläuterungstext mit CD-ROM; Teil 2: Legende; Teil 3: Karten. Münster.

Brandmayr, P., 1983. The main axes of the coenoclinal continuum from macroptery to brachyptery in carabid communities of the temperate zone. Report 4th Symp Carabidol 1981, pp 147-169.

Brandmayr P., Pizzolotto R., Scalercio S., Alfieri M.C., Zetto T., 2003. Diversity patterns of carabids in the Alps and theApennines. In: Nagy L., Grabherr G., Körner C., Thompson D.B.A. (eds.), Alpine Biodiversity inEurope. Ecol Stud 167: 307-317.

Brunet J., Fritz Ö., Richnau G., 2010. Biodiversity in European beech forests – a review with recommendations for sustainable forest management. Ecological Bulletins 53: 77-94.

Buse, J., 2012. "Ghosts of the past": flightless saproxylic weevils (Coleoptera: Curculionidae) are relict species in ancient woodlands. J Insect Conserv 16(1):93-102. DOI: 10.1007/s10841-011-9396-5

Cameron, R.A.D., Pokrysko B.M., Horsák M., SirbuI., Gheoca V., 2011. Forest snail faunas from Transylvania (Romania) and their relationship to the faunas of Central and Northern Europe. Biol J Linn Soc 104 (2): 471-479. DOI: 10.1111/j.1095-8312.2011.01730.x

Čiliak M., Šteffek J., 2011. Preliminary malacological survey of the Site of Community Importance Kamenná Baba (Branisko-Bachureň). Folia faun Slov 16 (2): 85-89.

Comandini F., Vigna Taglianti A., 1990. Ground beetle communities in a Mediterranean area (TolfaMountains,Central Italy). In: Stork, N.E. (ed.), The role of ground beetles in ecological and environmental studies. Intercept Ltd,Andover, pp 171-179.

Delagrange S., Montpied P., Dreyer E., Messier Ch., Sinoquet H., 2006. Does shade improve light interception efficiency? A comparison among seedlings from shade-tolerant and -intolerant temperate deciduous tree species. New Phytologist 172: 293-304. DOI: 10.1111/j.1469-8137.2006.01814.x

Diekmann M., Eilertsen O., Fremstad E., Lawesson J.E., Aude E., 1999. Beech forest communities in the Nordic countries – a multivariate analysis. Plant Ecology 140: 203-220. DOI: 10.1023/A:1009768225798

Dierschke H., 1990. Species-rich beech woods in mesic habitats in central and western Europe: a regional classification into suballiances. Vegetatio 87: 1-10. DOI: 10.1007/ BF00045649

Dierschke H.,BohnU., 2004. Eutraphente Rotbuchenwälder in Europa (Eutrophic beech forests inEurope). Tuexenia 24: 19-56.

Dierßen K., 1996. Vegetation Nordeuropas (Vegetation of Northern Europe). Ulmer,Stuttgart.

Ellenberg H., Leuschner C., 2010. Vegetation Mitteleuropas mit den Alpen (Vegetation Ecology of Central Europe), 6th ed. Ulmer,Stuttgart.

Fang J., Lechowicz M.J., 2006. Climatic limits for the present distribution of beech (Fagus L.) species in the world. J. Biogeogr. 33: 1804-1819. DOI: 10.1111/j.1365-2699.2006.01533.x

Gedda B., 2001. Environmental and climatic aspects of the early to mid Holocene calcareous tufa and land mollusc fauna in southernSweden. Dissertation,Lund University,Sweden.

Gedda B. 2006. Terrestrial mollusc succession and stratigraphy of a Holocene calcareous tufa deposit from the Fyledalen valley, southern Sweden. The Holocene 16 (1): 137-147. DOI: 10.1191/0959683606hl914rr

Giesecke T., Hickler T., Kunkel T., Sykes M.T., Bradshaw R.H.W., 2007. Towards an understanding of the Holocene distribution of Fagus sylvatica L. J Biogeogr 34: 118-131. DOI: 10.1111/j.1365-2699.2006.01580.x

Gutiérrez D., Rosa Menéndez R., Méndez M., 2003. Habitat-based conservation priorities for carabid beetles within the Picos de Europa National Park, northern Spain. Biol Cons 115: 379-393. DOI: 10.1016/S0006-3207(03)00154-X

Hässlein, L., 1960. Weichtierfauna der Landschaften an der Pegnitz - Ein Beitrag zur Ökologie und Soziologie niederer Tiere (Molluscs from the Pegnitz Region - A contribution to the ecology and sociology of lower animals). Abh Naturhist Ges Nürnb 29: 1-148.

Hennig C., 2010. fpc: Flexible procedures for clustering. R package version 2.0-3.

HorvatI., Glavac V., Ellenberg H., 1974. Vegetation Südosteuropas (Vegetation of Southeast-Europe). Fischer,Stuttgart.

Jacob M., Viedenz K., Polle A., Thomas F.M., 2010. Leaf litter decomposition in temperate deciduous forest stands with a decreasing fraction of beech (Fagus sylvatica). Oecologia 164(4): 1083-1094. DOI: 10.1007/s00442-010-1699-9

Jansson N., 2009. Habitat requirements and preservation of the beetle assemblages associated with hollow oaks. Dissertation, Linköping University, Sweden.

Jonsson BG, Kruys N, Ranius T (2005) Ecology of species living on dead wood: lessons for dead wood management. Silva Fennica 39(2):289–309 DOI: 10.14214/sf.390

Lang G., 1994. Quartäre Vegetationsgeschichte Europas (Quaternary vegetation history ofEurope). Fischer,Stuttgart.

Lovei G.L., SunderlandK.D., 1996. Ecology and behavior of Ground Beetles (Coleoptera: Carabidae). Annu Rev Entomol 41: 231-256. DOI: 10.1146/ annurev.en.41.010196.001311

Ložek V., 1982a. Faunengeschichtliche Grundlinien zur spät- und nacheiszeitlichen Entwicklung der Molluskenbestände in Mitteleuropa (Faunal historical baselines of the late- and postglacial development of mollusk populations inCentral Europe). Rozpr Českoslov Akad Věd 92(4): 1-106.

Ložek V., 1982b. Nacheiszeitliche Entwicklung der mitteleuropäischen Vertebraten- und Molluskenfauna in gegenseitiger Korrelation (Postglacial evolution of central European vertebrates and mollusks in mutualistic relationships). Z Geol Wiss 10 (7): 1019-1027.

Ložek V., 1982c. Contribution of malacology to the chronological subdivision of the Central European Holocene. In: Mangerud J., Birks H.J.B., Jager K.D. (eds.), Chronostratigraphic subdivision of the Holocene. Striae 16: 84-87.

Magri D., 2008. Patterns of post-glacial spread and the extent of glacial refugia of European beech (Fagus sylvatica). J Biogeogr 35: 450-463.
DOI: 10.1111/j.1365-2699.2007.01803.x

Magri D., Vendramin G.G., Comps B., Dupanloup I., Geburek T., Gömöry D., Latalowa M., Litt T., Paule L., Roure J.M., Tantau I., van der Knaap W.O., Petit R.J., de Beaulieu, J.L., 2006. Anew scenario for the Quaternary history of European beech populations: palaeobotanical evidence and genetic consequences. New Phytol 171 (1): 199-221. DOI: 10.1111/j.1469-8137.2006.01740.x

Meyer N., Meierott L., Schuwerk H., Angerer, O., 2005. Beiträge zur Gattung Sorbus in Bayern (Systematics of the genus Sorbus inBavaria). Ber Bayer Bot Ges (Sonderband): 1-216.

Mitrović B., 2007. Pleistocene malacofauna of the Požarevac Danube Area (NE Serbia). Ann Géol Pénins Balk 68: 81-89. DOI: 10.2298/GABP0701081M

Meyrick R.A., 2003. Holocene molluscan faunal history and environmental change at Kloster Mühle, Rheinland-Pfalz, western Germany. J Quart Sci 18 (2): 121-132. DOI: 10.1002/jqs.728

Müller J., Brunet J., Brin A., Bouget C., Brustel H., Bussler H., Förster B., Isacsson, G., Köhler F., Lachat T., Gossner M.M., 2012. Implications from large-scale spatial diversity patterns of saproxylic beetles for the conservation of European Beech forests. Insect Conservation and Diversity. DOI: 10.1111/j.1752-4598.2012.00200.x

Müller J., Bußler H., Bense U., Brustel H., Flechtner G., Fowles A., Kahlen M., Möller G., Mühle H., Schmidl J., Zabransky P., 2005. Urwald relict species - Saproxylic beetles indicating structural qualities and habitat tradition. Waldökol online 2: 106-113.

Müller-Kroehling S., Jantsch, M.C., Fischer, H.S., Fischer, A., 2014. Modelling the effects of global warming on the ground beetle fauna of beech forests in Bavaria, Germany. Eur. J. Entomol. 111(1): 35-49. DOI: 10.14411/eje.2014.005

Müller-Kroehling, S., Zehetmair T., 2014: Laufkäfer in den Kronen europäischer Buchenwälder (Ground beetles in the canopy of European beech forests). Angew Carabidologie 10: 101-107.

Nieto A., Alexander K.N.A., 2010. European Red List of Saproxylic Beetles. http://ec.europa.eu/environment/nature/conservation/species/redlist/downloads/European_saproxylic_beetles.pdf. Accessed 20 November 2012.

Nilsson G., Baranowski R., 1993. Skoghistorikens betydelse för artsammsättning av vedskalbaggar i urskogsartad blandskog. (Species composition of wood beetles in an unmanaged, mixed forest in relation to forest history). Ent Tidskr 114 (4): 133-146.

Nilsson G., Baranowski R., 1994. Indikatorer pa jätteträdskontinuitet – svenska förekomster av knäppare son är beroende av grova, levande träd. (Indicators of megatree continuity – Swedish distribution of click beetles dependent on hollow trees). Ent Tidskr 115 (3): 81-97.

Nitzu E., Nae A.,PopaI., 2008. The fauna of soil beetles (edaphic Coleoptera) as a sensitive indicator of evolution and conservation of ecosystems. A study on the altitudinal gradient in the Rodnei Mountains Biosphere Reserve (The Carpathians). In: Makarov, S.E., Dimitrijevic R.N. (eds.), Advances in Arachnology and Developmental Biology, Vienna-Belgrade-Sofia Monogr 12: 405-417

Økland B., Bakke A., Hagvar S., Kvamme T., 1996. What factors influence the diversity of saproxylic beetles? A multiscaled study from a spruce forest in southern Norway. Biodiver Conserv 5(1):75–100. DOI: 10.1007/BF00056293

Oksanen J., Guillaume Blanchet F., Kindt R., Legendre P., Minchin PR., O'Hara R.B., Simpson G.L., Solymos P., Stevens M.H.H., Wagner H., 2011. vegan: Community Ecology Package. R package version 2.0-1.

Pott, R., 1989. Die Formierung von Buchenwaldgesellschaften im Umfeld der Mittelgebirge Nordwestdeutschlands unter dem Einfluss des Menschen. Ber Geobot Inst Univ Hannover 1: 30-44.

Pott, R., 1997. Invasion of beech and establishment of beech forests inEurope. Annali di Botanica 55: 27-57.

Pott, R. 2000. Palaeoclimate and vegetation - long-term vegetation dynamics in central Europewith particular reference to beech. Phytocoenologia 30 (3-4): 285-333. DOI: 10.1127/ phyto/30/2000/285

R Development Core Team, 2011. R: A language and environment for statistical computing. R Foundation for Statistical Computing.Vienna,Austria. ISBN 3-900051-07-0.

Riley K.N., Browne R.A., 2011. Changes in ground beetle diversity and community composition in age structured forests (Coleoptera, Carabidae). Zookeys 147: 601–621. DOI: 10.3897/ zookeys.147.2102

Rizun V., Chumak V., 2003. Carabid Beetle Communities (Coleoptera, Carabidae) in virgin beech forests of the Ukrainian Carpathians. Vestnik zoologii. Suppl N 16: 114-120.

Rosenzweig M.L., 1995. Species Diversity in Space and Time. University Press, Cambridge. DOI: 10.1017/CBO9780511623387

Russo D., Cistrone L., Garonna A.P., 2010. Habitat selection by the highly endagered long-horned beetle Rosalia alpina in Southern Europe: a multiple spatial scale assessment. J Insect Conserv 15: 685-693. DOI: 10.1007/s10841-010-9366-3

Schmidl J., Bußler, H., 2008. Xylobiontic beetle guild composition and diversity driven by forest canopy structure and management. In: Floren A., Schmidl, J. (eds.), Canopy arthropod research in Europe, Bioform entomology, Nuremberg, pp 299-323.

Southwood T.R.E., 1961. The number of species of insect associated with various trees. J Animal Ecol 30: 1-8. DOI: 10.2307/2109

Speight M.C.D., 1989. Saproxylic invertebrates and their conservation. Nat Environ 42: 1-79.

Sverdrup-Thygeson A., Skarpaas O., Odegaard F., 2010. Hollow oaks and beetle conservation: the significance of the surroundings. Biodiver Conserv 19(3): 837-852. DOI: 10.1007/s10531-009-9739-7

Sverdrup-Thygeson A., Gustafsson L., Kouki J., 2014. Spatial and temporal scales relevant for conservation of dead-wood associated species: current status and perspectives. Biodiver Conserv 23 (3): 513-535. DOI: 10.1007/s10531-014-0628-3

Thoms C., Gattinger A., Jacob M., Thomas F.M., Gleixner G., 2010. Direct and indirect effects of tree diversity drive soil microbial diversity in temperate deciduous forest. Soil Biol Biochem 42 (9): 1558-1565. DOI: 10.1016/ j.soilbio.2010.05.030

Tinner W., Lotter A.F., 2006. Holocene expansions of Fagus silvatica and Abies alba in Central Europe: where are we after eight decades of debate? Quarter Sci Rev 25: 526-549. DOI: 10.1016/j.quascirev.2005.03.017

Valladares F., Niinemets Ü., 2008. Shade tolerance, a key plant feature of complex nature and consequences. Annu Rev Ecol Evol Syst 39: 237-257. DOI: 10.1146/annurev.ecolsys.39. 110707.173506

Vigna Taglianti A., De Felici S., 1994. Ground beetle communities in Central Appeninesbeech woods. In: Desender K., Dufrêne M., Loreau M., Luff M.L., Maelfait J.P. (eds.), Carabid beetles: Ecology and Evolution. Kluwer Acad Publ Dordrecht, pp 71-78. DOI: 10.1007/978-94-017-0968-2_11

Walentowski H., Bußler H., Bergmeier E., Blaschke M., Finkeldey R., Gossner M.M., Litt T., Müller-Kroehling S., Philippi G., Popp V.V., Reif A., Schulze E.D., Strätz C., Wirth V., 2010.Sinddie deutschen Waldnaturschutzgebiete adäquat für die Erhaltung der buchenwaldtypischen Flora und Fauna? Eine kritische Bewertung basierend auf der Herkunft der Waldarten des mitteleuropäischen Tief- und Hügellandes (Are the German conceptions suitable for the conservation of the specific flora and fauna of native beech forests? - A critical assessment based on the origin of forest species of central European lowland and submontane regions). Forstarchiv 71: 95-117. doi: 10.2376/0300-4112-81-195

Walentowski H., Ewald J., Fischer A., Kölling C., Türk W., 2013. Handbuch der natürlichen Waldgesellschaften Bayerns (Forest Vegetation Classification Manual forBavaria), 3rd ed. Geobotanica, Freising

Walentowski H., Gulder H.J., Kölling C., Ewald J., Türk W., 2001. Die regionale natürliche Waldzusammensetzung Bayerns (The regionally natural forest tree species composition ofBavaria). LWF Wissen 32: 1-99.

Walentowski H., Winter S., 2013. Phytodiversitätsmuster in mitteldeutschen Buchenwäldern (Patterns of plant diversity in beech forests ofCentral Germany). Waldökol Landschaftsforsch Natursch 13: 33-55.

Walentowski H., Zehm A., 2010. Reliktische und endemische Gefäßpflanzen im Waldland Bayern - eine vegetationsgeschichtliche Analyse zur Schwerpunktsetzung im botanischen Artenschutz (Occurrences of relict species and endemics in the temperate deciduous forest region ofBavaria- A geobotanical analysis as a contribution for setting priorities in botanical species conservation). Tuexenia 30: 59-81.

Warren M.S., Key R.S., 1991. Woodlands: past, present and potential for insects. In: CollinsN.M., Thomas J.A. (eds.), The conservation of insects and their habitats. Academic Press, London, pp 155-211. DOI: 10.1016/B978-0-12-181370-3.50014-2

Willner W., 2002. Syntaxonomische Revision der südmitteleuropäischen Buchenwälder. Phytocoenologia 32: 337-453. DOI: 10.1127/0340-269X/2002/0032-0337

Willner W., Di Pietro R., Bergmeier E., 2009. Phytogeographical evidence for post-glacial dispersal limitation of European beech forest species. Ecography 32 (6): 1011-1018. DOI: 10.1111/j.1600-0587.2009.05957.x


Supporting Information I-III
| DOWNLOAD 130KB
No metrics available for this article.