Challenges for the conservation of calcareous grasslands in ...

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Challenges for the conservation of calcareous grasslands in ...

Abstract

Editorial

Challenges for the conservation of calcareous grasslands in

northwestern Europe: integrating the requirements of flora and

fauna

The conservation of biological diversity requires an integrated approach covering the ecological demands of a multitude of species.

Integration may be achieved by focusing on a careful selection of target species, which is rare in practice. Calcareous grasslands

offer a case in point. Although they harbour a high diversity of both plant and insect species, in management the emphasis is placed

on the flora. This results in an underestimation of, notably, the importance of structural heterogeneity in the vegetation. As an

apparent consequence, conservation management in the Netherlands has been much more successful for the flora than for butterflies.

In contrast, Germany shows promising efforts to integrate both plant and animal species in conservation management and

landscape planning. The main constraints for a successfully integrated conservation management presently consist of a limited

availability and exchange of information, and an insufficient organisation of research and management at an international level.

# 2002 Elsevier Science Ltd. All rights reserved.

Keywords: Conservation; Management; Calcareous grassland; Target species; Invertebrates; Butterflies; Plants

1. Introduction

The conservation of biological diversity typically follows

one of three main approaches: a species, a systems

or a process-oriented approach (Sturm, 1993; Meffe and

Carroll, 1994). The species approach focuses on particular

species for a variety of reasons (Noss, 1990). Species

may be targeted for conservation because of their

appeal as flagship species to the public (especially rare

birds and large mammals), or because of a biological

function as keystone species which determine community

structure (Paine, 1969; WallisDeVries, 1998),

umbrella species whose ecological requirements encompass

those of other members of the ecosystem, or indicator

species which simply indicate the occurrence of

rare communities or biodiversity hotspots (Noss, 1990).

The systems approach concentrates on ecosystems, in a

narrower perspective on plant communities, or on biodiversity

hotspots. In Europe, the process approach

focuses on the fact that natural or even anthropogenic

processes are not included within the current conservation

management (Sturm, 1993). Processes may consist

of more or less regular environmental catastrophes

(floods, fires, wind throws; Plachter, 1998) but also succession

(Beinlich et al., 1995). Besides these biological

arguments, an area approach may be chosen because

areas, especially when large, are easy to define and

handle from a practical point of view of management

Biological Conservation 104 (2002) 265–273

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feasibility and the cooperation of owners and stakeholders.

Thus, conservation issues either focus on a low level

of integration (the species) or a high level of integration

(system and processes). This gives rise to problems of

scale when developing recommendations for conservation

management. The species approach may fail to

cover the needs for all threatened species by focusing

on one or just a few taxa, whereas the systems or the

process approach may fail by only addressing general

issues of ecosystem functioning where more detailed

measures are required for particular species (Henle et

al., 1999). This is well illustrated by recent developments

in metapopulation ecology, where populations of single

species may collapse because of increasing habitat fragmentation,

even though the quality of individual habitat

patches remains unchanged (Hanski, 1999). The spatial

scale at which conservation takes place has different

consequences for different taxonomic groups, varying in

dispersal capacity, home range size and life history

(Reck et al., 1996; WallisDeVries, 1999). This should be

taken into consideration when selecting target species or

species groups for conservation. One obvious example is

dispersal capacity. Whereas animals are able to move,

plants stand still. Despite that, conservation management

of fragmented habitats includes only the creation

of static corridors which may be successful to link animal

but not plant populations which rather need


266 Editorial / Biological Conservation 104 (2002) 265–273

‘‘moving’’ corridors such as grazing animals (Poschlod

et al., 1996; Willerding and Poschlod, 2002).

The challenge for the future clearly is to arrive at

an integration of conservation at the level of species,

level of communities and level of processes. Just how

this integration should be achieved remains a problematic

issue. The option pursued in the papers from this

special issue is to take a closer look at a particular

community, the calcareous grasslands in northwestern

Europe, by examining the conservation issues from

various angles for different species groups, plants and

butterflies in particular.

This special issue on the conservation of calcareous

grasslands is the product of an international workshop,

with participants from seven northwest European

countries, hosted by Dutch Butterfly Conservation in

Wageningen, 22–23 November 1999. It is an attempt to

work towards a more integrated approach for research

and conservation management. At the same time it

offers an overview of current topics in conservation,

which can be taken as a starting point for future challenges.

The three questions we want to address in this

introduction are: first, what are the common concerns

for the preservation of the different species groups, second,

how can we develop a strategy for an integrated

conservation, and, third, what are the main gaps in our

knowledge to arrive at such a strategy?

2. The case of calcareous grasslands

The calcareous grasslands of northwestern Europe

comprise the dry grasslands on limestone and chalk as

well those on calcareous loess. They are predominantly

semi-natural communities originating after man felled

the primeval forests in prehistoric times, ca. 7000 years

B.P. Plant species from natural open habitats assembled

and formed species-rich grasslands especially on dry

calcareous slopes and plateaus in northwest Europe

which were also called ‘‘Steppenheide’’ according to

Gradmann (1950; see also Willems, 1980; Ellenberg,

1996; Wilmanns, 1997; Pott, 1998; Prins, 1998; Poschlod

and WallisDeVries, 2002). It should be noted that the

role of wild herbivores in maintaining grassland vegetation

before human impact is still under debate (Gerken

and Meyer, 1996; Vera, 2000). Since the Bronze Age,

grazing by livestock and, since the Middle Ages, occasional

mowing, prevented these grasslands from spontaneous

afforestation. However, from the Roman era

onwards and especially since the Middle Ages, calcareous

grasslands have developed by different types of

land use (Poschlod and WallisDeVries, 2002). It should

also be clear that many species of calcareous grasslands

are not indigenous but have established only since the

Roman era, e.g. herbs such as Hippocrepis comosa and

Salvia pratensis, or even since modern times, e.g. grasses

such as Bromus erectus and Koeleria pyramidata

(Poschlod and WallisDeVries, 2002). Finally, specific

types of land use such as grazing and mowing had not

only a tremendous effect on species composition but

also in evolving ‘‘seasonal ecotypes’’ in calcareous

grassland plants (Poschlod et al., 2000).

Calcareous grasslands provide a particularly suitable

example to illustrate the problem of developing an integrated

approach for conservation. First, their biological

diversity is high and includes a variety of rare species

from different taxonomic groups (Van Helsdingen et al.,

1996; Ssymank et al., 1998). Among plant communities,

chalk grasslands rank as one of the richest in species at

a small spatial scale (


have strongly invaded the calcareous grasslands (Hagen,

1996).

Finally, the conservation interest of calcareous grasslands

has long been recognised in policy by the Council

of Europe (Wolkinger and Plank, 1981). Rare species

from different taxonomic groups are listed under the

Habitat Directives of the European Union (Van Helsdingen

et al., 1996; Ssymank et al., 1998). This is

important if any measures are to be implemented.

3. Integration issues: managing for plants or animals

In managing grasslands for conservation, the emphasis

is usually placed on plants (Van Wieren and Bakker,

1998; Crofts and Jefferson, 1999), especially orchids in

calcareous grasslands. Despite the extensive work by

M.G. Morris (Morris, 1990), animals are often given

minor attention, except for birds, despite the abovementioned

importance of calcareous grasslands especially

for invertebrates. A more balanced approach was

presented in an earlier symposium on calcareous grasslands

(Hillier et al., 1990). Yet, as noted by Jones-

Walters (1990) the management practice typically shows

an apparent lack of consideration for invertebrates. One

of the factors hampering the integration across taxonomic

groups is a mere lack of information on the

occurrence of invertebrates, their ecology and their

management in comparison with plants (Jones-Walters,

1990; Crofts and Jefferson, 1999). The bias is similar in

the field of scientific research: out of 101 papers from

1996 and later on European calcareous grasslands in

ecological journals 67 were devoted to studies on plants

and vegetation, only 11 on invertebrates and just four

treated both plants and invertebrates (the other 11

papers concerned abiotic processes, fungi and soil

microbiota). This not only indicates a bias in interest

but also an excessive specialization instead of integration.

Although current developments indicate that an

integrated approach is gaining importance, as shown

below for Germany, there is still a long way to go to

achieve a proper balance.

The need for integration is stressed by management

conflicts that result from different effects across different

taxonomic groups. In a long-term experiment on the

management of grasslands in Baden-Wu¨ rttemberg

(started in 1975) which included calcareous grasslands

(Schiefer, 1981; Brauckmann et al., 1997), various

groups of invertebrates were recorded as well. The

results have clearly shown that every selected taxonomic

group reacts in a different way on the respective management

treatment (Table 1). In managing for plant

diversity, low nutrient levels are stressed, which can be

arrived at by grazing, frequent mowing, mowing in

summer or even mulching (Bobbink and Willems, 1987;

Willems et al., 1993; Kahmen et al., 2001). Heterogeneity

in vegetation structure is emphasized for the

invertebrates, at least some groups of which require a

less intensive management by grazing at low stocking

rates or by rotational mowing (e.g. Duffey et al., 1974;

BUTT, 1986; Morris, 1990; Gibson et al., 1992; Settele

et al., 1995). Botanically centred management may thus

lead to an impoverished invertebrate fauna. This can

only be avoided by an integrated approach that considers

both taxonomic groups.

4. The example of the Netherlands: lack of integration

In the Netherlands the threat to calcareous grasslands,

in this case chalk grasslands, is extreme. Only ca.

20 fragments remain in the chalk district of South-

Limburg, varying in size between 0.05 and 5 ha, and

representing only a few percent of the original area

(Bobbink and Willems, 1998). The introduction of chemical

fertilisers meant the end of the traditional rural

landscape in which the flocks of sheep were a characteristic

feature. Most of the traditional chalk grasslands

lost their significance for wildlife, as illustrated by

plants and butterflies (Fig. 1). Only a limited number of

sites have been set aside as nature reserves, due to their

extraordinary richness in plant and animal species.

Among these species a number reach their northwestern

outpost from continental Europe in the chalk

Table 1

Species number (vascular plants, selected groups of invertebrates) and density of indivduals at the study site St. Johann (Baden-Wu¨ rttemberg,

Germany) after 20 to 25 years applying three different management treatments: low-intensity sheep grazing in early and late summer, mulching twice

a year and abandonment (from Brauckmann et al., 1997). Vascular plants—number of species per 25 m 2 permanent plot; earthworms—number of

species (in brackets individuals per m 2 ); arachnids—number of species (in brackets number of individuals per five traps and day); ground beetles (in

brackets number of individuals per five traps and day); bugs—number of species (in brackets number of individuals caught)

Management treatments Vascular plants Invertebrates

Editorial / Biological Conservation 104 (2002) 265–273 267

Earthworms (Lumbricidae) Arachnids (Arachnida) Ground beetles (Carabidae) Bugs (Hemiptera)

Grazing 32 4 (ca. 400/m 2 ) 35 (3.9) 17 (3.3) 20 (122)

Mulching 36 6 (ca. 400/m 2 ) 62 (4.3) 20 (2.9) 16 (38)

Abandonment 28 5 (ca. 250/m 2 ) 71 (3.9) 18 (2.2) 33 (170)


268 Editorial / Biological Conservation 104 (2002) 265–273

Fig. 1. Changes in species distribution of plants and butterflies in the

chalk district in the Netherlands since 1950 on a 5 5 km scale. Plants

represented by 46 target species for calcareous grassland after Bal et

al. (1995) with distribution data from Mennema et al. (1980, 1985),

Van der Meijden et al. (1989) and Blink (1997). Butterflies comprise all

65 observed indigenous species with distribution data from Dutch

Butterfly Conservation.

district of South-Limburg. Undoubtedly, the south–

north flowing river Meuse, originating in Central

France, fulfilled the role of a migration route for many

plant and animal species (De Leeuw, 1935). The reserves

are managed either by mowing late in the year or by

seasonal grazing, especially by flocks of sheep in short

rotation. This type of management is largely aimed at

the preservation and restoration of botanical diversity,

prompted by the results of experimental investigations

on the management of plant communities (Bobbink and

Willems, 1987; Willems et al., 1993; Bobbink and

Willems, 1998; Willems, 2001). Unfortunately, parallel

research on the management of invertebrates has been

lacking.

In comparing the loss of diversity, it can be noted that

the distribution of butterflies has declined more severely

than plants (Fig. 1; G-test P


hand, a management that does not value the importance

of structural heterogeneity and, on the other hand, a

continued decline of both rare and common grassland

butterflies. A link between the two can be suggested but

not substantiated as the necessary data are lacking. This

highlights the importance of integrating efforts for

research, monitoring and management across taxonomical

groups.

5. The example of Germany: promising developments

Compared to The Netherlands and the United Kingdom,

Germany covers a broad range of calcareous

grassland types affected by either suboceanic, continental

or even submediterranean climate. Calcareous

grasslands are one of, if not the most, species-rich

habitats in Germany (Bundesamt fu¨ r Naturschutz,

1999). The decrease of calcareous grasslands locally has

been extremely high (e.g. Fig. 3). The change in agricultural

practices in the 1960s contributed most to this

decrease due to abandonment and afforestation but also

the conversion of calcareous grasslands into fertile pastures

or meadows or arable fields (Mattern et al., 1980

and 1992). Therefore, they belong to the most endangered

habitats. The actual area which is still covered by

calcareous grasslands is ca. 60,000 ha (Table 2; Schumacher

et al., 1995) with the highest proportion in the

south German countries Baden-Wuerttemberg and

Bavaria due to the Jurassic mountain belt crossing both

countries (Swabian Alb, Franconian Alb) which is,

however, only between 0.1 and 0.2% of the total area of

Germany. As in the Netherlands, recent management

has mainly consisted of shrub clearance followed by

mowing with little consideration of the dominant traditional

land use by grazing. Locally, grazing could be

maintained, mostly by private initiative of sheep farmers

which, under the influence of subsidies, has changed the

Fig. 3. Loss of calcareous grasslands (‘‘Wacholderheiden’’) in the

administrative division Stuttgart of Baden-Württemberg (Germany)

between 1900 and 1990 (after Mattern et al., 1980, 1992).

Editorial / Biological Conservation 104 (2002) 265–273 269

emphasis from farming to landscape management.

More recently, triggered by the increasing amount of

money necessary to manage mechanically those areas

that cannot be longer maintained (Petermann, 1995),

there is a strong emphasis in an integrated approach of

conservation and socioeconomic issues. In south Germany,

two projects on large calcareous grassland areas

included such an approach (Swabian Alb—Beinlich and

Plachter, 1995, Beinlich, 1997; Garchinger Haide—Pfadenhauer

et al., 2000). Hampicke and Tampe (1995)

have shown that subsidizing farmers is much cheaper in

the long term than artificial management by mowing.

However, conflicts may arise locally when reintroducing

grazing since species as orchids, which have established

during artificial conservation management, might go

extinct (Petermann, 1995).

There is no comparative analysis of the loss of diversity

for different taxonomic groups. From >400 plant

species more or less restricted to this habitat about 1%

is extinct, 42% are threatened and 1% is extremely rare

(Korneck et al., 1998; Bundesamt fu¨ r Naturschutz,

1999). Calcareous grasslands host the highest number of

endangered plant species (205; Korneck et al., 1998).

Similar data have not been summarised for animals but

data are available for selected groups and at a regional

level. Whereas in birds only a few species are restricted

to calcareous grasslands (indicating large areas and

complex habitat situations at a landscape level), the

opposite is true of many invertebrate groups such as

butterflies, grasshoppers, wild bees, owl flies (Ascalaphidae),

beetles, cicadas, bugs, spiders and land snails

(Bra¨ u in Quinger et al., 1994). Calcareous grasslands

host the largest proportion of butterflies (Ebert, 1991)

and grasshoppers (Detzel, 1998). Butterflies restricted to

calcareous grasslands are extinct only at a regional level.

The same is true for grasshopper species except Arcyptera

microptera which has its main distribution in the

pontic Steppe region (Bra¨ u in Quinger et al., 1994;

Detzel, 1998).

Integrating information of plants and animals is

becoming ever more frequent in Germany. A recent

extensive field study on the Swabian Alb in southwest

Germany, which included a review of the literature on

the occurrence of four different groups of invertebrates

along a successional sere after the abandonment of calcareous

grasslands, has shown that the species number

in recently abandoned grasslands is highest. However,

the highest number of threatened species was found in

grazed calcareous grasslands (Table 3). More recently,

animals have become strongly embedded in conservation

management and landscape planning (Henle et al.,

1999). Many endangered invertebrates are now

acknowledged as strong indicators of biodiversity and,

therefore, are increasingly taken into account to determine

conservation value (Bra¨ u in Quinger et al., 1994;

Reck et al., 1996; Henle et al., 1999).


270 Editorial / Biological Conservation 104 (2002) 265–273

Table 2

Size of calcareous grassland areas in the respective countries of

Germany (from Schumacher et al., 1995)

Country Area (km 2 ) Dominant historical land use

Lower Saxony 3.75 Grazing

Northrhine-Westfalia ca. 22 Grazing

Hessen 25.30 Grazing

Thuringia 95.50 Grazing, mowing

Saxony-Anhalt 38.37 Grazing

Rheinland-Pfalz 7.52 Mowing, grazing

Saarland 7.54 Mowing

Baden-Württemberg 254.56 Grazing, mowing

Bavaria 134.6 Grazing, mowing

6. Scope of this special issue

The papers in the issue offer some answers to the three

questions raised in the introduction.

As to the common concerns, the papers show a great

similarity in conservation themes for plants and invertebrates.

Habitat fragmentation and limitations by dispersal

are apparent for both groups and perhaps

constitute the most immediate threat for biodiversity.

Management continues to be an important issue for

both groups as well. In general, again both in plants and

animals, different species show contrasting responses to

different management treatments. Therefore, spatial

heterogeneity is a prerequisite for the maintenance of

species diversity. The scale at which this heterogeneity is

best achieved remains to be established. In small areas it

should perhaps be sought in a different management

between areas. In large areas extensive grazing may

provide the best option. However, where areas are both

small and isolated, such as in the Netherlands, a wise

approach to prevent a further loss of species seems to

optimize small-scale heterogeneity by a careful rotational

grazing or mowing regimes. In time the creation

of ‘corridors’ and ‘stepping stones’ should reduce the

threats of fragmentation.

As to the gaps in our knowledge, various observations

can be made. First, the lack of baseline distribution data

is emphasized for butterflies by Van Swaay (2001), but it

reflects a general need for a comprehensive European

approach that also deserves more attention for plants.

The sheer amount of time and resources required for

such an endeavour prevents its fulfilment. However, a

careful selection of a comparatively small set of target

species (Bal et al., 1995), may overcome this problem.

Second, as shown by Bourn and Thomas (2002), climate

change can have profound effects for species at the edge

of their range, as is the case for many species of calcareous

grasslands in northwestern Europe. These effects

increase with habitat fragmentation. In the case of

a general warming, Bourn and Thomas (2002) suggest a

positive impact on thermophilous species. However, if

climate change is accompanied by the expected increase

of winter rainfall (Schuurmans, 1995; Van Boxel and

Cammeraat, 1999), the balance might become negative

for species with high thermal requirements in spring.

More detailed information on the reaction of individual

species to various aspects of climate change within distinct

seasonal periods is therefore necessary. Third,

despite the attempt at integration, the papers still reflect

the specialization of botanists versus entomologists. On

both sides attempts are made to arrive at an increased

understanding of community dynamics by focusing on

functional groups, such as annual plants or pollinating

insects. The integration in terms of plant-herbivore and

plant-pollinator interactions in a spatially heterogeneous

environment still requires considerable further

study. Restoration experiments, such as reported by

Mortimer et al. (2002), show that joint efforts of plant

and animal ecologists are not yet sufficient to reveal

success stories in managing for both plants and animals.

It is encouraging, however, to show in this issue that

such integrated studies are undertaken.

As to the strategy for an integrated conservation, the

main constraints appear present at three levels: availability

of information, dissemination of information

and organization at an international level. These levels

are evidently interrelated. There is an urgent need to

organize conservation efforts at a European level, such

that information on the distribution of species and

experience with management practices can be exchanged

more readily. In the field of management practice

the development of the Downland Practitioners Network

by English Nature seems a good initiative. However,

in the field of scientific expertise on management

Table 3

Number of species (in brackets number of Red List species in the country Baden-Württemberg) of four invertebrate groups along a successional sere

in calcareous grasslands (from Beinlich, 1995)

Successional stage Intensive

grazing

Extensive

grazing or

mowing

Abandoned,

young fallows

Abandoned,

older fallows

Abandoned,

overgrown

fallows (shrubs)

Butterflies (day flying Lepidoptera) 35 (22) 48 (28) 48 (22) 34 (13) 27 (8) 9 (4)

Grasshoppers (Saltatoria) 7 (3) 6 (4) 11 (2) 2 (0) 4 (1) 4 (0)

Ground beetles (Carabidae) 23 (10) 15 (4) 21 (2) 16 (1) 11 (1) 5 (1)

Ants (Formicidae) 10 (6) 15 (6) 15 (4) 10 (3) 12 (3) 4 (2)

Abandoned,

overgrown fallows

(shrubs and trees)


and the establishment of international databases on

species distribution and trends, the degree of fragmentation

is still enormous. This precludes a meaningful

setting of international conservation priorities. We

hope that this special issue may strengthen this international

perspective on the conservation of calcareous

grasslands.

Acknowledgements

The first author wants to thank the Expert Centre for

Nature Management from the Dutch Ministry of Agriculture,

Nature Conservation and Fisheries for financial

support to organize the workshop. Dennis Slotboom

offered valuable assistance in analysing the data for the

Netherlands.

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Editorial / Biological Conservation 104 (2002) 265–273 273

Michiel F. WallisDeVries

Dutch Butterfly Conservation (De Vlinderstichting)

PO Box 506

6700 AM Wageningen

The Netherlands

email address: michiel.wallisdevries@vlinderstichting.nl

Peter Poschlod

University of Marburg/Faculty of Biology

Department of Nature Conservation

D-35032, Marburg

Germany

email address: peter.poschlod@biologie.uniregensburg.de

Jo H.Willems

Department of Plant Ecology and Evolutionary Biology

Utrecht University

PO Box 800.84

2508 TB Utrecht

The Netherlands

email address: j.h.willems@bio.uu.nl

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