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Basic and Applied Ecology 11 (2010) 473–485

ERSPECTIVES

mplementing large-scale and long-term functional biodiversity research:he Biodiversity Exploratories

arkus Fischera,b,∗, Oliver Bossdorfa, Sonja Gockelc, Falk Hänselb, Andreas Hempb,ominik Hessenmöllerd, Gunnar Korteb, Jens Nieschulzed, Simone Pfeifferb, Daniel Pratia,wen Rennere, Ingo Schöningc, Uta Schumacherb, Konstans Wellse, Francois Buscotf,lisabeth K.V. Kalkoe, Karl Eduard Linsenmairg, Ernst-Detlef Schulzed, Wolfgang W. Weisserc

University of Bern, Institute of Plant Sciences, Botanical Garden and Oeschger Centre, Altenbergrain 21, CH-3013 Bern, SwitzerlandUniversity of Potsdam, Institute of Biochemistry and Biology, Maulbeerallee 1, D-14469 Potsdam, GermanyFriedrich-Schiller-University Jena, Institute of Ecology, Dornburger Str. 159, D-07743 Jena, GermanyMax-Planck Institute for Biogeochemistry, Hans-Knoell-Str. 10, D-07745 Jena, GermanyUniversity of Ulm, Institute of Experimental Ecology, Albert-Einstein Allee 11, D-89069 Ulm, GermanyHelmholtz Centre for Environmental Research-UFZ, Theodor-Lieser-Str. 4, D-04120 Halle, GermanyUniversity of Würzburg, Theodor-Boveri-Institute for Biosciences, Am Hubland, D-97074 Würzburg, Germany

eceived 13 July 2010; accepted 24 July 2010

bstract

Functional biodiversity research explores drivers and functional consequences of biodiversity changes. Land use change ismajor driver of changes of biodiversity and of biogeochemical and biological ecosystem processes and services. However,

and use effects on genetic and species diversity are well documented only for a few taxa and trophic networks. We hardlynow how different components of biodiversity and their responses to land use change are interrelated and very little abouthe simultaneous, and interacting, effects of land use on multiple ecosystem processes and services. Moreover, we do notnow to what extent land use effects on ecosystem processes and services are mediated by biodiversity change. Thus, overalloals are on the one hand to understand the effects of land use on biodiversity, and on the other to understand the modifyingole of biodiversity change for land-use effects on ecosystem processes, including biogeochemical cycles. To comprehensivelyddress these important questions, we recently established a new large-scale and long-term project for functional biodiversity,he Biodiversity Exploratories (www.biodiversity-exploratories.de). They comprise a hierarchical set of standardized field plots

n three different regions of Germany covering manifold management types and intensities in grasslands and forests. They serves a joint research platform for currently 40 projects involving over 300 people studying various aspects of the relationshipsetween land use, biodiversity and ecosystem processes through monitoring, comparative observation and experiments. Wentroduce guiding questions, concept and design of the Biodiversity Exploratories – including main aspects of selection and

e discuss the significance of this approach for further functional

mplementation of field plots and project structure – and w iodiversity research. This includes the crucial relevance of a common study design encompassing variation in both drivers

∗Corresponding author at: University of Bern, Institute of Plant Sciences and Botanical Garden, Altenbergrain 21, CH-3013 Bern, Switzerland.el.: +41 31 631 4943; fax: +41 31 631 4942.

E-mail addresses: markus.fischer@ips.unibe.ch, fischerm@uni-potsdam.de (M. Fischer).

439-1791/$ – see front matter © 2010 Gesellschaft für Ökologie. Published by Elsevier GmbH. All rights reserved.oi:10.1016/j.baae.2010.07.009

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74 M. Fischer et al. / Basic and Applied Ecology 11 (2010) 473–485

nd outcomes of biodiversity change and ecosystem processes, the interdisciplinary integration of biodiversity and ecosystemesearchers, the training of a new generation of integrative biodiversity researchers, and the stimulation of functional biodiversityesearch in real landscape contexts, in Germany and elsewhere.

2010 Gesellschaft für Ökologie. Published by Elsevier GmbH. All rights reserved.

usammenfassung

Funktionelle Biodiversitätsforschung erforscht die Ursachen und funktionellen Konsequenzen von Biodiversitätsverän-erungen. Landnutzung gehört zu den wichtigsten Ursachen von Änderungen von Biodiversität und biogeochemischen undiologischen Ökosystemprozessen und -leistungen. Allerdings sind Landnutzungsauswirkungen auf genetische und Artendi-ersität bisher nur für wenige Taxa und trophische Netzwerke gut dokumentiert. Zudem ist kaum bekannt, wie verschiedeneomponenten der Biodiversität und ihre Reaktion auf Landnutzungsänderungen zusammenhängen. Auch ist wenig überleichzeitige und miteinander wechselwirkende Effekte der Landnutzung auf mehrere verschiedene Ökosystemprozesse und -eistungen bekannt. So ist auch noch unklar, inwieweit Landnutzungseffekte auf Ökosystemprozesse und -leistungen durchiodiversitätsveränderungen vermittelt werden. Es gilt also einerseits Landnutzungseffekte auf Biodiversität zu verstehennd andererseits die modifizierende Rolle von Biodiversitätsveränderungen für Landnutzungseffekte auf Ökosystemprozesse,inschliesslich biogeochemischer Kreisläufe. Um diese wichtigen Fragen umfassend zu untersuchen, haben wir kürzlichin grosses und langfristiges Projekt zur funktionellen Biodiversitätsforschung gestartet, die Biodiversitäts-Exploratorienwww.biodiversity-exploratories.de). Diese umfassen einen Satz standardisierter Untersuchungsflächen in drei Regioneneutschlands, die jeweils vielfältige Typen und Intensitäten der Wald- und Grünlandnutzung umfassen. Sie dienen als gemein-

ame Forschungsplattform für gegenwärtig 40 Projekte mit über 300 Beteiligten, die verschiedenste Aspekte der Beziehungwischen Landnutzung, Biodiversität und Ökosystemprozessen durch Monitoring, vergleichende Beobachtung und Experimententersuchen. Wir stellen Leitfragen, Konzept und Design der Biodiversitätsexploratorien vor – einschliesslich der wesentlichenspekte der Auswahl und Einrichtung der Untersuchungsflächen und der Projektstruktur – und wir diskutieren die Bedeutunges Ansatzes für die weitere funktionelle Biodiversitätsforschung. Diese beinhaltet die zentrale Bedeutung des gemeinsamenorschungsdesigns, das sowohl Ursachen als auch Konsequenzen der Veränderungen von Biodiversität und Ökosystemprozessenmfasst, die interdisziplinäre Integration von Biodiversitäts- und Ökosystemforschenden, die Ausbildung einer neuen Generationon integrativen Biodiversitätsforschern und die Anregung funktioneller Biodiversitätsforschung im realen Landschaftskontext,n Deutschland und darüber hinaus.

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eywords: Biotic interactions; Ecological monitoring; Ecosystemichness

ntroduction

Functional biodiversity research explores drivers and func-ional consequences of biodiversity changes. Biodiversityomprises all scales of biological organization, spanningntra-specific genetic, morphological and demographic diver-ity, species and community diversity, the diversity ofiological interactions between organisms and the diversityf ecosystems in the landscape (Wilson 2001). Environmen-al effects on species diversity and genetic diversity are wellstablished only for few taxa, and no project in the worlds currently addressing all facets of biodiversity in a singleramework. Accordingly, we hardly know how different ele-ents of biodiversity are interrelated (Lamoreux et al. 2006;olters, Bengtsson, & Zaitsev 2006). This is exemplified

y the very scarcely studied relationship between speciesiversity and genetic diversity within species (Van Valen965; Vellend 2005; Vellend & Geber 2005), where currentnowledge is limited to a few case studies relating commu-

ity species diversity to molecular genetic diversity of singlepecies. Another example is soil biodiversity, where, despiteioneering large-scale programs such as the UK soil biodi-

ri(

es; Forest management; Grassland; Land use; Landscape; Species

ersity project (http://soilbio.nerc.ac.uk/), very little is stillnown about the relationship between plant diversity and theiversity of soil invertebrates, fungi and microbes (Wardle etl. 2004; van der Heijden, Bardgett, & van Straalen 2008).

he relevance of land use change in functionaliodiversity research

Among the major drivers of current changes of biodi-ersity and of ecosystem processes and services, land usehange is the most prominent (Sala et al. 2000) rangingrom local and regional (Hietel, Waldhardt, & Otte 2005)o national (Stöcklin, Bosshard, Klaus, Rudmann-Maurer,

Fischer 2007), international and global scales (Houghton994). In Germany, forests and grasslands are, along withroplands, among the most important ecosystems affectedy human requirements. The natural vegetation cover of Ger-

eplaced by production forests, e.g. spruce plantations, ands now restricted to rather small and highly fragmented areasEllenberg 1996; MCPFE 2007). Furthermore, nutrient-poor

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M. Fischer et al. / Basic and A

rasslands, formerly only used at low intensities by humansnd forming the most species-rich assemblages, are nowither threatened by abandonment or by land-use intensifi-ation (Poschlod, Bakker, & Kahmen 2005; Maskell, Smart,ullock, Thompson, & Stevens 2010). The Biodiversityxploratories described below focus on forest and grass-

and habitats ranging from unmanaged forest and hardly usedrassland to intensively used forest and grassland.

Currently, biodiversity experiences a general decline, inentral Europe especially at the local and regional scales

Settele et al. 2010). In terrestrial habitats, this is mainlyue to habitat loss and ongoing changes in land use. Inorests this is furthered by indirect effects of browsing rumi-ants, and enhanced by the displacement of native by alienpecies, by industrial emissions of nitrogen, organic com-ounds and heavy metals, and by climate change (Sala etl. 2000; Millenium Ecosystem Assessment 2005; Thuiller007). Although the mostly negative relationship betweeniodiversity and current land-use intensification is rather wellnown for some taxa, in particular vertebrates, vascular plantsnd some invertebrates such as grasshoppers or butterflies, its not known for many others. The most obvious ecological

echanisms contributing to declining grassland biodiver-ity after land use change are competitive exclusion due toncreased productivity (Grime 1973; Mittelbach et al. 2001)nd species loss due to increased disturbance (Grime 1973;iswas & Mallik 2010). However, as most case studies quiteoarsely compare fertilized with unfertilized or more withess disturbed sites, differentiated statements on the amountnd mechanism of biodiversity change after land use changere scarce. For instance, disturbance by grazing may enhancelant diversity on fertile grasslands and decrease it on less fer-ile grasslands (Proulx & Mazumder 1998). Last but not least,redicting the consequences of land use change for biodiver-ity change involves disentangling direct effects of land usen the diversity of higher trophic levels from indirect effectsediated by changes in plant diversity (Cardinale et al. 2006;race et al. 2007).In principle, similar considerations as for grasslands also

pply to forest trees and understory plants. Of course, effectsf forest management additionally depend on number anddentity of planted tree species (Hunter 1999). Moreover, theffect of age-class forests can only be studied by comparingeveral successional stages with unmanaged forests, whichre characterised by more continuous tree age distribution. Ofarticular additional importance in Central European forestss the pressure of large herbivores such as deer and wild boarMCPFE 2007; Ammer 2009).

cosystem processes and services

Society is interested in proper and continuous function-ng of ecosystem processes and services. These include theools and fluxes of water, carbon and nutrients, the mainte-ance of soil fertility, clean water and air, the provisioning of

isst

Ecology 11 (2010) 473–485 475

ood and construction material, pollination and pest controlDaily 1997). The biodiversity-ecosystem functioning rela-ionship is a rapidly expanding research area in ecology andvolution (Hooper et al. 2005). To date, a number of modelxperiments have addressed the functional consequences ofhanges of plant species composition and diversity (Hectort al. 1999; Tilman et al. 2001; Roscher et al. 2005). Whileverall effects of plant species composition, such as pres-nce or absence of legumes or grasses, on biogeochemicalycles and other ecosystem processes are widely acknowl-dged (Spehn et al. 2002), very little is known about the role ofndividual species. It has been shown repeatedly that ecosys-em processes and services depend not only on the speciesomposition of ecosystems but also, and mostly positively,n their species richness (e.g., Naeem, Thompson, Lawler,awton, & Woodfin 1994; Hooper et al. 2005; Balvanerat al. 2006). Moreover, higher plant species diversity mat-ers for the simultaneous maintenance of multiple ecosystemrocesses (Hector & Bagchi 2007).

In the real-world landscape context, positive correlationsetween biodiversity and ecosystem processes have beeneported for some taxa (Hooper et al. 2005; Balvanera etl. 2006). Moreover, restoration projects aiming at enhancediodiversity were reported to also enhance ecosystem ser-ices (Benayas, Newton, Diaz, & Bullock 2009). Firstxperiments manipulating plant species diversity at severalites indicate positive relationships of plant species diversitynd production (e.g. Stein, Auge, Fischer, Weisser, & Prati008). However, replicated experiments manipulating theiodiversity also of other taxa at many sites in real landscapesre missing completely. Overall, the relevance of positiveiodiversity-ecosystem functioning relationships establishedn model experiments for real-world ecosystems is far fromeing well established.

esearch approaches on drivers andonsequences of biodiversity change

Understanding the interactions between land use, bio-iversity, and ecosystem functioning requires long-termbservational, comparative and experimental studies at anppropriate spatial scale (Fischer et al. 2010). While short-erm laboratory and small field-experiments are useful toolso test particular hypotheses, such experiments are insuffi-ient to understand processes at the ecosystem level in natureCarpenter 1996; Symstad et al. 2003). Most experimentsn the biodiversity-ecosystem functioning relationship haveeen conducted in the laboratory or on small field plotsHooper et al. 2005). Thus, we need to conduct experiments inhe field, at a scale large enough to represent natural biogeo-hemical cycles. Moreover, because different regions differ

n their landscapes, resources, conditions, species pools, andocio-economic settings, studies need to be performed ineveral regions (Beck 2004). This is absolutely fundamen-al, because otherwise results cannot be generalised. Finally,

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76 M. Fischer et al. / Basic and A

o separate trends from fluctuations, to improve intra- andxtrapolation of available observations, to test ecologicalodels, and to allow parameterising models for predict-

ng change under future scenarios, functional biodiversityesearch needs to cover long periods of time.

Working in several real landscapes creates multiple chal-enges for data analysis and interpretation. The drivers ofiodiversity, such as land use and soil conditions, are rarelyompletely independent or perfectly balanced, as it is desir-ble in a planned experiment. Spatial auto-correlation in alot-based sampling design needs to be taken into account inome organisms, but not in others, depending on their mobil-ty. Moreover, research plots need to be large enough forxperiments not to influence each other, but still spatiallyomogeneous enough to allow the integration of differ-nt results from the same plot. Finally, the work load tossess biodiversity and to run experiments varies consider-bly between taxa and this requires some flexibility of theesign in terms of temporal and spatial replication. Thus, aundamental goal of any large-scale, interdisciplinary projects to integrate the diverging demands of individual disciplinesnd to foster their optimal cooperation.

he Biodiversity Exploratories

To comprehensively address important questions onhe feedback between land use, biodiversity and ecosys-em processes in real-world ecosystems we recentlystablished a new large-scale and long-term project forunctional biodiversity research, the Biodiversity Explorato-ies (www.biodiversity-exploratories.de). We use the termExploratories” to emphasize that the project is not onlyased on observation and between-plot comparison, whichould have been described by the term “observatories”, buthat the project very importantly also involves replicated fieldxperimentation in order to gain causal insights.

Here we illustrate the key requirements for successfulmplementation of a comprehensive land use-biodiversity-cosystem process project by describing the main features ofhe Biodiversity Exloratories. We emphasize the importance

f the common study design with many replicate plots fornterdisciplinary research, for addressing overarching ques-ions, and for quantitative synthesis of research results. Weonclude by highlighting the role and significance of the Bio-

vedi

able 1. Main geographic and environmental characteristics of the three

Schorfheide-Chorin Hain

ocation NE Germany Centize ∼1300 km2 ∼130eology Young glacial landscape Calcauman population density 23 km−1 116 kltitude a.s.l. 3–140 m 285–nnual mean temperature 8–8.5 ◦C 6.5–8nnual mean precipitation 500–600 mm 500–

Ecology 11 (2010) 473–485

iversity Exploratories in relation to other initiatives, and weraw conclusions for future research.

esign of the Biodiversity Exploratories

ain rationale of design

A key aspect of the Biodiversity Exploratories is the systemf standardized study plots, which are jointly used by allnvolved research groups. Such a common study design isivotal for statistical comparisons across taxa, land use types,r geographical regions, and a lack thereof has hamperedany cooperative projects in the past.To allow for the test of consistency and generality of

and-use effects across geographic regions, the Biodiversityxploratories were established in three different regions ofermany (see Table 1, Fig. 1). Within each region, we estab-

ished a network of field plots in grasslands and forests ofifferent land use types and intensities. All plots within oneegion are collectively termed an “exploratory”. Within eachxploratory, there are three hierarchical levels of study plots,n which research of different intensity and detail is con-ucted (Tables 2 and 3): (1) about 1000 grid plots (GPs),00 in grasslands and 500 in forests, which are mainly usedor large-scale analyses of biodiversity data and their rela-ionships to land use and other environmental factors, (2)00 experimental plots (EPs), 50 in grasslands and 50 inorests, which are a selected subset of the respective gridlots serving as a platform for more thorough biodiversityssessment and environmental monitoring, as well as foreveral manipulative experiments, and (3) 18 very inten-ive plots (VIPs), half in grasslands and half in forests,hich are a subset of the experimental plots used for study-

ng biodiversity or ecological processes in extreme detailr requiring very labour-intensive methods, for which these of the experimental plots is not feasible. In one of thexploratories (Hainich-Dün), where selection forests are anspecially important forest type, we established three further

ersity Exploratories comprise 57 very intensive plots, 300xperimental plots, and some 3000 grid plots. Below, weescribe the rationale and selection of each of these plot typesn more detail.

Biodiversity Exploratories.

ich-Dün Schwäbische Alb

ral Germany SW Germany0 km2 ∼422 km2

reous bedrock Calcareous bedrock with karst phenomenam−1 258 km−1

550 m 460–860 m◦C 6–7 ◦C

800 mm 700–1000 mm

M. Fischer et al. / Basic and Applied Ecology 11 (2010) 473–485 477

Fig. 1. Geographic location of the three Biodiversity Exploratories in Germany, a map of the exploratory Schwäbische Alb, and examplesof two ‘very intensive plots’ in grasslands and forests. In the plot charts, the subplots associated with various projects are shown in differentcolours.

Table 2. Frequency of land use types in grassland grid plots (GPs), experimental plots (EPs) and very intensive plots (VIPs) in the threeBiodiversity Exploratories in 2006/2007 as assessed during the plot selection process. One of the unfertilised very intensive plots in a pastureof the Hainich-Dün exploratory had received a very small amount of manure in 2006, but remained unfertilized ever since.

Schorfheide-Chorin Hainich-Dün Schwäbische Alb

GP EP VIP GP EP VIP GP EP VIP

MeadowsFertilized 91 7 3 69 7 3 155 18 3Unfertilized 178 11 – 12 – – 8 4 –

PasturesFertilized 25 – – 11 2 – 34 2Unfertilized 131 22 3 171 18 3 135 17 3

Mown pasturesFertilized 15 3 – 165 15 3 94 9 3Unfertilized 69 7 3 42 8 – 9 – –

Fallow 38 – – 14 – – – – –

478 M. Fischer et al. / Basic and Applied Ecology 11 (2010) 473–485

Table 3. Frequency of land use types in forest grid plots (GPs), experimental plots (EPs) and very intensive plots (VIPs) in the threeBiodiversity Exploratories in 2006/2007 as assessed during the plot selection process. In a few of the otherwise unmanaged forest plots in theSchwäbische Alb some treelets had been removed recently. As this intervention was very minor, we classified these plots as unmanaged. Forage class forests the dominant tree species is indicated in the table. The 22 pine age class forests in experimental plots in the Schorfheide-Chorinexploratory comprise 15 pure pine stands and 7 pine stands where beech has been added later.

Schorfheide-Chorin Hainich-Dün Schwäbische Alb

GP EP VIP GP EP VIP GP EP VIP

Unmanaged forest 65 5 3 81 13 3 33 5 3Age class forest

Beech 176 16 3 308 24 3 244 33 3Oak 58 7 – 6 – – 4 – –Spruce 10 – – 35 4 3 130 12 3Pine 251 22 3 – – – 10 –

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he three Exploratories

As a first step for establishing the Biodiversity Explorato-ies, we selected three study regions: (1) the UNESCOiosphere Reserve Schorfheide-Chorin, which is situated in

he lowlands of North-eastern Germany, a young glacial land-cape with many wetlands, (2) the National Park Hainichnd its surrounding areas, situated in the hilly lands ofentral Germany, and (3) the UNESCO Biosphere Reservechwäbische Alb (Swabian Jura), which is situated in the lowountain ranges of South-western Germany (Table 1, Fig. 1).ach of these areas represents most of the variation in landse typical for grasslands and forests in Germany, from hardlyanaged grasslands and unmanaged beech forests to highly

ertilized and intensively used meadows and pastures, andntensively managed forests (Tables 2 and 3).

election of grid plots

To select the grid plots within each exploratory, we firstsed several sources of information – aerial images, differ-nt types of maps, vegetation surveys and forest inventoriesto construct coarse distribution maps of habitats and land

se types in each region. After that, a virtual 100 × 100 mrid was placed over the entire area, and all grid plots (i.e.ells enclosed by grid lines) covering forests or grasslandsere identified. We generally discarded grid plots with inho-ogeneous land cover or partial overlap with settlements,

gricultural fields, water bodies and plots intersected byoads. From the remaining pool of candidate plots we selectedt least 500 grid plots in forests and 500 in grasslands in eachxploratory (Tables 2 and 3), which reflected the range of landse types and land use intensities in grasslands and forests,nd which were confirmed to be covered by the designated

and use type during field surveys. Since we aimed both atncluding plots of all major grassland and forest types and atepresenting the existing range of land use intensities, eachith a certain minimum number of plots, the grid plots are

tns(

– – 50 – –9 3 75 – –

ot a random sample of grassland and forest habitats withinach exploratory. Of course, when necessary, data from therid plots can be used together with data on the relative pro-ortion of land use types in the whole exploratory area torovide area-wide estimates.

For each of the selected grid plots, we collected data aboutey environmental variables such as slope, orientation, relief,nd altitude, and we conducted a soil and a land use inven-ory (see below) as well as a vegetation survey. In addition, tossess the potential suitability of plots for experimental stud-es, we examined the homogeneity of vegetation and land usend distance to other habitat types, and the chances for obtain-ng the necessary permission from the land owner. Later, allhis information was used for selecting the experimental plotsonsidering land use and soil type. In particular, we aimed atvoiding confounding of soil type with land use because soiluality very often determines how land is used.

oil inventory

To survey soil conditions in the grid plots, we con-ucted a standardized soil inventory. One soil sample wasaken in the centre of each of the 3000 grid plots. Min-ral soils were sampled with a motor-driven, 8 cm wideoil column cylinder (Eijkelkamp, Giesbeek, Netherlands)hat allowed us to take largely undisturbed samples. At for-st plots, the organic layer was removed with a 20 × 20 cmetal frame prior to sampling. In the Schorfheide-Chorin,

rganic soils could not be sampled with a motor-driven soilolumn cylinder because of the high groundwater tables.nstead, we used a split-tube sampler (Eijkelkamp, Gies-eek, Netherlands) to take undisturbed manual samples ofhe upper 30 cm of soil, and a peat sampler (Eijkelkamp,iesbeek, Netherlands) to sample the underlying horizons

o a depth of 100 cm. Additionally, we estimated the thick-ess of the organic layer. We described the soil type of alloil cores according to the German soil classification systemAd-Hoc-Arbeitsgruppe Boden 2005) and the World Refer-

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nce Base of Soil Resources (IUSS Working Group WRB006).

For more detailed soil physical and chemical analysis, weivided each soil core into several depth increments (0–10,0–30, 30–50, 50–70, 70–90, 90–110 cm), and analysed theseeparately for bulk density, pH, as well as carbon and nitrogenontents. These data were not used for initial plot selection,ut they will be available to analyse effects of land use oncosystem processes (e.g. Grüneberg, Schöning, Kalko, &eisser 2010) or soil effects on biodiversity.It is not surprising that, because of their different par-

nt materials, the three exploratories are characterised byifferent spectra of soil types (Table 4). In the Schorfheide-horin exploratory, the dominant geological substrate islacial till, often covered by glacio-fluvial or aeolianand. As a consequence, many forest soils have a tex-ure from sandy loam to pure sand. About 70% of theorest soils in Schorfheide-Chorin are Dystric Cambisols.ess frequent are Albeluvisols, Podzols, and Regosols.he grasslands in Schorfheide-Chorin are dominated by

ertile loamy soils and drained Histosols or Gleysols.n the Hainich-Dün exploratory, the dominant geologi-al substrate is loess over Triassic limestone. Thus, bothorest and grassland soils frequently have a loamy orlayey texture, and the main soil groups are Eutric Cam-isols, Luvisols, and Stagnosols. In the Schwäbische Albxploratory, soils developed mostly on Jurassic shell lime-tone, and they are therefore extremely rich in clay. Eutricambisols and Leptosols dominate in forests and grass-

ands.Soil types in the Biodiversity Exploratories are not only

etermined by geological substrate, but also by aeolianoess layers, relief and other environmental factors. In thechwäbische Alb, for instance, Leptosols are usually asso-iated with steep slopes, and 10% of the grassland soils arenthrosols, which indicates they have formerly been used as

rable fields.Inevitably, soil types are often partly correlated with

resent and past land use. For instance, slopes with shallowoils are usually used as sheep pastures in the Schwäbischelb because they do not support more intensive types of landse. Not all forest land use intensities from near-natural tontensively used are found on Cambisols in the Hainich-Dünxploratory, while they are found on Stagnosols. Extremelyandy soils are frequently associated with pine forests inchorfheide-Chorin. Since soil type and land use type usuallyre not entirely independent of each other, it is important tossess their correlation and to consider both factors togetherhen examining their effects on biodiversity and ecosystemrocesses.

Soil properties can vary at small scales, and one soil sam-le per plot is therefore not sufficient to fully capture the

atural heterogeneity of soils in the field. Therefore, charac-erising soil heterogeneity constitutes an important future taskn particular for the experimental plots where most researchs carried out. Ta

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and use inventory in grasslands

To enquire the land use of grassland grid plots, we devel-ped a questionnaire for all land owners and land users.his questionnaire covered all relevant information about

ype and intensity of annual land use, and further detailsbout plot-related agricultural practices, agri-environmentalchemes, socio-economic variables and plot history. For pro-ected areas, we also enquired conservation measures. The

ain land use inventory of all grassland grid plots wasone once, for the reference year 2006, and this informa-ion was used for the selection of experimental plots. Sinceand-use intensity may vary across years (particularly in thechorfheide-Chorin exploratory farmers frequently alternateetween mowing and grazing), we are repeating the land usenventory in all grassland experimental plots annually.

Grasslands fall into four main land use categories: mead-ws, pastures, mown pastures, and fallow land (Table 2).own pastures are grasslands that are both mown and grazedithin the same year. For meadows and mown pastures, theuestionnaire recorded the number of cuts per year and esti-ated the amount of hay or silage produced. For pastures

nd mown pastures, we recorded livestock species (cattle,orse, sheep, or goat), the type of grazing system (permanent,ationed or rotational pastures), and the estimated annualuration of grazing. In addition, to relate grazing data to live-tock units we collected data about the age and density of theivestock (cattle < 1 year: 0.3 livestock units (LU), cattle 2ears: 0.6 LU, cattle > 2 years: 1 LU, sheep and goat < 1 year:.05 LU, sheep and goat > 1 year: 0.1 LU, horse < 3 years:.7 LU, horse > 3 years: 1.1 LU). For cattle pastures, we alsoistinguished dairy, beef production, and breeding purposes.or all grasslands, we further recorded whether they wereertilized or not, the kind of fertilizer (mineral fertilizer orrganic fertilizer such as manure or dung), and the estimatednnual amount of added fertilizer.

In addition to these basic data about mowing, grazingnd fertilisation, we also enquired more special managementeasures such as drainage, levelling, harrowing, seed addi-

ion, hand weeding, or pesticide application. We also askedhether land use type had changed during the past 5 years,

nd whether plots had been used for agriculture during theast 20 years. Finally, for each involved farm, we enquiredarm size, organizational form (private farms or farm coop-ratives), number of employees, and the type of subsidieseceived.

On average across the three exploratories, 35% of therassland grid plots were meadows, 35% were pastures, 27%ere mown pastures, and 3% were fallow land (Table 2).round two-thirds of the meadows and mown pastures were

ertilized, but only 15% of the pastures. Mown pastures wereostly grazed by cattle, and less frequently by sheep. Pas-

ures, in contrast, were more equally grazed by cattle andheep. Both pastures and mown pastures were only occa-ionally grazed by horses. While sheep pastures were aboutqually frequent in the Hainich-Dün and Schwäbische Alb

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Ecology 11 (2010) 473–485

xploratories, almost all pastures in the Schorfheide-Chorinere grazed by cattle (see Appendix A: Table 1 for land useetails of the experimental plots in grassland).

and use inventory in forests

We assessed land use with a systematic forest inventory inll forest grid plots. This inventory was based on a circularampling area of 500 m2 in each plot, where we measuredhe diameter at breast height (DBH) and locations of all treesith DBH > 7 cm. We also recorded whether forests werenmanaged (mature forest not used (and mostly protected) fort least 60 years), age-class forests (harvested at 80–120 yearntervals) or uneven-aged selection forests (trees harvestedelectively). We also recorded the main tree species in eachlot, and the developmental stage (bare land, thicket stage,hicket, pole wood, young timber, old timber) for age-classorests. Further, we recorded stand structure, canopy density,requency of disturbances, and the amounts of coarse and fineoody debris as structural parameters.On average across the three exploratories, 13% of the

orests in grid plots were unmanaged forests, 83% wereanaged age-class forests, and 4% were selection forests.

n some of the otherwise unmanaged forest plots in thechwäbische Alb some treelets had been removed recently.s this intervention was a very minor one, we classified theselots as unmanaged. The selection forests mainly occurredn Hainich-Dün, which also harboured the largest propor-ion of unmanaged forests (28%). Across the exploratories,he most frequent dominant tree species was beech (78%f unmanaged and 46% of age-class forests), followed bypruce (10% of age-class forests), Scots pine (7% of age-lass forests) and oak (2% of age-class forests). Pine and oakorests occurred only in Schorfheide-Chorin, whereas spruceorests were restricted to the Hainich-Dün and Schwäbischelb exploratories. Across the exploratories, some 21% of the

ge-class forests were mixed forests, i.e. forests with less than0% cover of the dominant tree species. While many forestlots showed signs of management-related disturbances suchs logging trails, the proportion of disturbed plots did not dif-er much across the exploratories (see Appendix A: Table 2or land use details of the experimental plots in forest).

election of experimental plots

All environmental and land-use data described above weresed to select 100 locations in each exploratory – 50 inorests and 50 in grasslands – in which we establishedo-called experimental plots (EPs) as platforms for morehorough biodiversity assessment, environmental monitor-ng, and manipulative experiments. Experimental plots have

size of 100 × 100 m in forests and 50 × 50 m in grasslands.he selection was based on a stratified random sampling,ith strata representing land use and several other criteria.wo particularly important criteria were that the EPs should

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over the variation in land use intensities and in soil depthound in each exploratory, as illustrated in Fig. 2 for two vari-bles for grassland and forest, respectively. Moreover, landwners/users or regional authorities had to allow long-termtudies and experiments on the plots. Additional criteria wereonsistency and constancy of soil type, homogeneity of landse and vegetation composition within plots, and that theyhould have a slope of less than 20% and not be water-logged.n the partly steep Schwäbische Alb, however, eleven forestlots and seven grassland plots had to be selected with slopeslightly greater than 20%. Further criteria were a minimumistance of 200 m between the outer edges of two EPs, andhat forest EPs should be at least 100 m and grassland EPs0 m away from the nearest forest edge.

A particularly challenging aspect of experimental plotelection was soil type. Up to ten different soil types occurredn each exploratory, and soil type variation was partly con-ounded with land use type. To reduce this confounding and

ig. 2. The variability of study plots in the Biodiversity Explorato-ies, and the distribution of experimental plots and very intensivelots among grassland grid plots, illustrated by the variables soilepth and plant species richness (A), and among forest grid plots,llustrated by the variables standing biomass and stem density (B).ote that soil depth values are clustered at cut off at 100 cm, becauserecise values are not available beyond 100 cm for all of these plots.evertheless, for many of these plots, the soil depth is above 100 cm.

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Ecology 11 (2010) 473–485 481

he variation caused by soil type, we aimed to restrict exper-mental plots to the two most dominant and characteristicoil types in forests and grasslands, respectively, in eachxploratory (Table 4). As described above, these soil typesiffered between the three exploratories. In the Schwäbis-he Alb, Cambisols and Leptosols were dominant in bothorests and grasslands, and we therefore selected EPs onlyrom these soil types. As to Cambisols, EPs were restrictedo shallow Cambisols, excluding soils with chromic (“Terrausca”) horizons. In the Hainich-Dün exploratory, Luvisolsnd Stagnosols dominate in forests, and not all managementypes and spruce age-classes were available on Cambisols.ambisols and Stagnosols dominate in grasslands. Soils are

requently characterised by argic (forests) or vertic (grass-ands) horizons and stagnic properties. For the experimentallots in forests, we therefore selected Stagnic Luvisols anduvic Stagnisols, and for the experimental plots in grasslandse selected Cambisols, Vertic Stagnosols and a few Stagnicertisols. In the Schorfheide-Chorin, the characteristic soil

ype in forests was Cambisol, so most of the experimentallots were selected on this soil type. However, in order toully cover the land use gradient in this exploratory, we alsoncluded a few EPs on other soils (Table 4). The situationas similar in grasslands in Schorfheide-Chorin, where, in

ddition to the dominant Histosols and Cambisols, we alsoelected some EPs on Luvisols, Albeluvisols and Gleysols.e attempted to remove confounding of soil type with land

se within, but not necessarily across, each of the sets of plotsn grassland or forest in each exploratory. Matching betweenxploratories was not a main criterion as even the same soilypes are not necessarily comparable between exploratoriesue to the different bedrocks, which lead to differeneces e.g.n soil texture and pH. When selecting EPs, we generallyxcluded plots that were extreme outliers (>2 s.d.) in termsf solum thickness or stone content.

election of ‘very intensive plots’

Of the 100 experimental plots in each exploratory, we fur-her selected a subset of 18 plots – nine in forests and ninen grasslands, three per specific land use type and regionTables 2 and 3) – which we called very intensive plotsVIPs), for applying methods too labour-intensive to be donen all EPs. One example of such VIP research, with the goalo establish the method and to extend measurements in theuture to all experimental plots, is the detailed assessment ofoil microbial diversity with molecular methods. To cover thevailable range of land use intensities, we generally selectedhe VIPs equally from three land use types of increasing landse intensity. In grasslands we selected three VIPs from each,nfertilized pastures, strongly fertilized meadows, and mown

astures, which were unfertilized in Schorfheide-Chorin andertilized in the Schwäbische Alb and Hainich-Dün (Table 2).n forests we selected VIPs from natural forests, age-classorests, and conifer plantations (Table 3). In the Hainich-Dün

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xploratory, we additionally included three VIPs in selectionorests, because this forest management is very important inhis region.

lot implementation and instrumentation

We marked each experimental plot with subterranean metalies, and recorded their exact position in early 2008. To facili-ate orientation, we created detailed charts of each plot, whichlso contained the precise locations of forest tracks or anyther kind of disturbance on or close to the plots. The mapsere later digitized to create the interactive plot maps now

vailable in the database (see below). A key feature of thexploratories is that the research activities do not affect thengoing land use.

To measure climatic conditions in the Biodiversityxploratories, we set up 300 environmental monitoring units

n all experimental plots. The basic environmental sensorsnstalled in all EPs measure air temperature (nominal preci-ion: ±0.3 K) and relative air humidity (±1%) at 2 m aboveround, soil surface temperature (±0.1 K) at 10 cm aboveround, soil temperature (±0.1 K) at 5, 10, 20 and 50 cmelow ground, and soil moisture (±1% VWC) at 10 cm belowround. VIPs have a second soil moisture sensor which addi-ionally measures soil moisture at a depth of 20 cm. All

easurements are taken hourly, and all systems are workingith data loggers (ADL-MX, Meier-NT, Zwönitz, Germany).To also obtain information on variation in wind speed

nd direction, precipitation, solar radiation and baromet-ic pressure, we are currently equipping nine experimentallots per exploratory, geographically spread over the wholexploratory area, with additional sensors. In addition, were implementing one tower in closed beech forest in eachxploratory (and an additional one in Hainich-Dün), whichill measure climatic variables also above canopy level.

anagement structure

The Biodiversity Exploratories are a research platform forurrently over 40 projects involving more than 300 peopleequally divided between scientists, students and techni-

al staff – with a wide range of scientific backgrounds andpproaches. The success and synergistic effect of such a largend diverse project largely depends on an efficient manage-ent structure. The project is led by a steering committee

f six senior scientists, headed by a project speaker. Each ofhe three exploratories is managed by a local team consistingf a manager, a forester, an engineer, and two further tech-ical staff. These local management teams are responsibleor the set-up and maintenance of field plots and technicalacilities, they advise researchers about practical and admin-

strative issues in the exploratory, and they keep contactsith local stakeholders. All scientific activities are coordi-ated by the central project office (Biodiversity Exploratoryffice) as approved by the steering committee. This office

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Ecology 11 (2010) 473–485

lso assists with project administration, facilitates commu-ication among management teams and research groups,rganizes training courses for all PhD students, and it isesponsible for PR and outreach.

Although individual research projects in the Biodiversityxploratories typically last for 3 years, the platform has beenesigned as a long-term infrastructure. Therefore, to ensureong-term consistency and integrity of the project, a set ofommon rules for data and publication policy, as well asor conduct in the field have been established, which allroject members must acknowledge by signature to obtainhe necessary field permits.

atabase

To facilitate the compilation and dissemination of infor-ation and data across the different working groups, we set

p a central database, which serves both as central data stor-ge facility and as work-in-progress database actively usedy all researchers in the Biodiversity Exploratories includingn interactive platform for coordinating field work, manag-ng field plots, and distributing practical and administrativenformation among project participants.

To support field work, the database provides basicnformation such as photographs and directions for eachlot, a WebGIS (Dragicevic 2004; Mückschel, Schachtel,ieschulze, & Köhler 2004) for orientation in the field,etailed interactive maps of each plot (see Fig. 1 for exam-les), and an online booking system for field accommodationsnd facilities. All changes in the database are logged, so theatabase also functions as an archive.

The database sensu stricto, i.e. the part containing data andeta-data, employs XML technologies (W3C 2003) for data

armonization across the involved disciplines. XML allowsor a formalized yet flexible description of metadata, dataormats and syntaxes, which were determined prior to therst field season. The coverage of many different data for-ats and structures is essential for acceptance by the user

ommunity. Usage is further facilitated by web-service inter-aces (Heimann, Nieschulze, & König-Ries 2010) that can beead by the statistical package R (R Development Core Team008).

ore and contributing projects

The research in the Biodiversity Exploratories is organ-sed into core and contributing projects. Starting in 2006, sixore projects carried out the environmental, soil and land usenventories described above, selected experimental and veryntensive plots, and implemented the necessary technical anddministrative infrastructure for the project. Moreover, the

ore projects are ensuring consistent long-term monitoring ofhe species diversity of selected taxa of plants, fungi, birds,ats and other mammals, selected insect taxa and microor-anisms, and they assess genetic diversity of selected taxa

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f plants and microorganisms. The core projects addresselected ecosystem processes and services including poolsf biomass and carbon, pollination services, predation, seedispersal and community stability and are also carrying outxperimental studies. These include experiments manipulat-ng deadwood availability in the very intensive forest plots,encing off subplots of all forest experimental plots to excludearge herbivores, and increasing plant species diversity byeeding in subplots of the grassland experimental plots.

Currently, 34 contributing projects are addressing theenetic and species diversity of many taxa, including soilrganisms, ecosystem processes related to nutrients, foodebs and pollination webs, and modelling of species inter-

ctions and ecosystem processes in relation to land useAppendix A: Table 3). Where possible, all core and con-ributing projects are doing research on all experimental plots.nly those activities, for which this is not possible, are limited

o the very intensive plots. Moreover, many of the contribut-ng projects have experimental components replicated acrossll very intensive plots or all experimental plots. Working onhe same plots guarantees standardization of the study designnd enables quantitative synergy by data synthesis withinnd across projects. For research groups working on soilrganisms, common soil samples were taken in a coordinatedffort, during which soil or extracted DNA was distributed tohe involved groups, in order to minimize sampling hetero-eneity. More detailed information about the research in theiodiversity Exploratories is provided in Appendix A: Tableand on http://www.biodiversity-exploratories.de/Projects.

onclusions from the Biodiversityxploratories for functional biodiversityesearch

We are developing the Biodiversity Exploratories as anxemplary research platform for addressing feedback loopsetween land use, biodiversity change, and their conse-uences for ecosystem functioning and ecosystem services.ommon guiding questions (Lindenmayer & Likens 2009)nd the combination of monitoring of biodiversity andcosystem measures with manipulative experiments in aommon study design with replicated sites are essential forvercoming the disciplinary separation that is hamperingur comprehensive understanding of drivers and functionalonsequences of all facets of biodiversity. Working on com-on plots ensures that a common language is used by the

nvolved researchers and allows for quantitative data syn-hesis, two key elements in the integration of the strong butcattered research communities of the different biodiversitynd ecosystem research disciplines. At the same time, it is

n essential requirement for the training of a new generationf biodiversity explorers (Fischer, Schreier, & Larigauderie997). In addition to the spatial scale with many replicatedites in very different regions, the temporal scale with a

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Ecology 11 (2010) 473–485 483

lanned duration of more than 10 years is essential for cap-uring trends in the occurrence and abundance of differentaxa and temporal interactions between taxa and processes.

The concept of the Biodiversity Exploratories can bextended to other landscapes and habitat types, such as crop-ands, landscapes with freshwater bodies, urban systems,oasts or alpine landscapes. An interesting challenge is posedy integrating landscape features surrounding the study plots,uch as the type and areal proportion of habitat types, oregional topography and climate. Further challenges includehe quantification of land-use intensity, a concept which ishe more difficult to define and apply the more diverse theypes of land use are which are employed in a study area.

In Europe, a number of environmental monitoringrograms and initiatives are addressing parts of the land use-iodiversity-ecosystem process research agenda (Hammen etl. 2010). The long-term ecological research network LTER-urope combines a diverse array of ecological research sites,here biodiversity research is among several research foci

n some of the sites (Müller, Baessler, Schubert, & Klotz010). Further initiatives are addressing parts of the landse-biodiversity-ecosystem process research agenda outsideurope (North America (e.g. Keller, Schimel, Hargrove, &offman 2008), South America (e.g. Joly et al. 2010), Africa

e.g. Reyers & McGeoch 2007), Asia (e.g. Sodhi, Koh, Brook,Ng 2004), Australia (e.g. Abbott & Le Maitre 2010). In

ummary, there is the world-wide realization that long-termnd large-scale ecological research sites are needed to addressurrent ecological questions at the appropriate temporal andpatial scale.

The key messages of the Biodiversity Exploratories forny larger-scale efforts to monitor and understand the rela-ionships between drivers and functional consequences ofiodiversity change are:

1) consider all land uses and climatic conditions,2) consider all taxa, their genetic diversity, species diversity

and interaction diversity, and the relationships amongthese facets of biodiversity,

3) consider all relevant ecosystem processes and services,including means and variation of pools and fluxes inbiogeochemical cycles as well as biotic processes andservices including pollination, dispersal, predation,

4) reflect all these issues by involving the correspondingresearch disciplines in a common framework guided byoverarching questions and through a jointly used networkof replicated field plots.

In our opinion, an extended network of exploratory-likeesearch platforms across Europe and the world capturing theain types of ecosystems would allow for much more mecha-

istic understanding and prediction of changes in biodiversitynd ecosystem function than the manifold and very scattered

ctivities currently going on. Therefore we suggest integrat-ng such platforms into a future global network of biodiversitybservation systems, as envisaged by The Group on Earthbservations Biodiversity Observation Network (GEOBON,

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choles et al. 2008). We hope that the concept and examplef the Biodiversity Exploratories contributes to stimulatingunctional biodiversity research in realistic landscape con-exts in Germany and globally.

cknowledgments

We thank the Deutsche Forschungsgemeinschaft DFG forunding the DFG Priority Program 1374 “Infrastructure-iodiversity-Exploratories”, our institutions for administrat-

ng the exploratories, the participants of the exploratoryroject for their excellent and dedicated work, the authoritiesf the Biosphere Reserve Schorfheide-Chorin, the Nationalark Hainich and the Biosphere Area Schwäbische Alb forupport, responsible state environmental offices of Baden-

ürttemberg, Thüringen, and Brandenburg for field permitsaccording to § 7BbgNatSchG).

ppendix A. Supplementary data

Supplementary data associated with this article cane found, in the online version, at doi:10.1016/j.baae.010.07.009.

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