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Inland Sand Ecosystems: Dynamics and restitution as a consequence of the use of different grazing systems

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Abstract

In Germany, sand ecosystems are among the endangered habitats, particularly the open and the ecotone-rich forms including open oak and pine woodland. A practicable nature conservation concept has to take into account the often anthropo-zoogenically caused dynamics of sand ecosystems. The conservation of these ecosystems is problematic, as they form dynamic systems which lose relevance for species and habitat conservation if they are not used or if they are intensively used or fertilized. Therefore, apart from existing sandy regions, restitution areas are studied, among them an inland dune complex the morphology of which was altered in the context of a trial and development project. The areas studied are located in the north-German lowland plain (“Hase valley”, “Ems valley” in the Emsland region) and in the northern Upper Rhine valley (sandy regions near Darmstadt). Differentiated grazing systems with cattle, sheep, goats, horses, Mangalitza pigs and donkeys are employed. The central aim of the project is to analyse the effects of different grazing systems on the vegetation, on nutrient dynamics, on selected animal groups and on socio-economics. The article presents conclusions for nature conservation and socio-economic aspects.
lnland Sand Ecosystems: Dynamics and
restitution as a consequence of the use of
different grazing systems
Angelika Schwabe', Dominique Remy', Thorsten Assmann', Anselm Kratochwil',
Albrecht Mährleinn, Michael Nobis', Christian Storm', Andreas Zehm', Harald
Schlemmert, Robert Seußt, Sabine Bergmann', Carsten Eichbergr, Uwe Menzelt,
Markus Persigehl', Kai Zimmermann', Mareike Weinert2
rDepartment of Biology, Geobotany/Vegetation Ecology, Darmstadt University of
Technology, Germany
'Dep. of Biology/Chemistry, Division of Ecology, University of Osnabrück,
Germany
'Institute for Ecology and Environmental Chemistry, University of Lüneburg,
Germany
nGreven, Germany
'Institute of Geodesy, Darmstadt University of Technology, Germany
Abstract
In Germany, sand ecosystems zue irmong the endangered habitats, particularly the open and
the ecotone-rich forms including open oak and pine woodland. A practicable nature
conservation concept has to take into account the often anthropo-zoogenically caused
dynamics of sand ecosystems. The conservation of these ecosystems is problematic, as they
form dynamic systems which lose relevance for species and habitat conservation if they are
not used or if they are intensively used or fertilized.
Therefore, apart from existing sandy regions, restitution areas are studied, among them
an inland dune complex the morphology of which was altered in the context of a trial and
development project. The areas studied are located in the north-German lowland plain
("Hase valley", "Ems valley" in the Emsland region) and in the northem Upper Rhine
valley (sandy regions near Darmstadt). Differentiated grazing systems with cattle, sheep,
goats, horses, Mangalitza pigs and donkeys are employed.
The central aim of the project is to analyse the effects of different grazing systems on
the vegetation, on nutrient dynamics, on selected animal groups and on socio-economics.
The article presents conclusions for nature conservation and socio-economic aspects.
1 Background
Most open habitats developed as a result of human activities, especially as a
consequence of the many and diverse forms of land use practised in historical
times. Consequently, there was a wide variety of such habitats in Central Europe;
many of them, however, have now disappeared.
Pasture Landscapes and Nature Conservation
B. Redecker, P. Finck, W Härdtle, U. Riecken, E. Schröder (Eds.)
@ Springer-Verlag Berlin Heidelberg
240 Angelika Schwabe et al.
In Germany, sand ecosystems are among these endangered habitats, particularly
the open types and those rich in ecotones including open oak and pine woodland.
A practicable nature conservation concept has to take into account the often
anthropo-zoogenically caused dynamics of sand ecosystems. The conservation of
these ecosystems is problematic, as they form dynamic systems which are no
longer of relevance for species and habitat conservation if they are not used or if
they are intensively used or fertilized. Mosaic complexes need certain disturbance
factors to also retain pioneer stages (BAKKER et al. 1983, Or-rn et al. 1999, WHrTE
and JeNtsctt 2001, KRATocHwTL and ScHwesE 2001). Many poorly competitive
plant species as well as xerophilous and thermophilous arthropods depend on such
early successional stages.
Tabelle 1. The concepts of redynamization and restitution in sand ecosystems (schematic)
status quo management oractices
changes of abiotic
conditions ("abiotic
redvnqmizlfinn"I
transfer of diaspores
(vascular plants,
crvntosnms)
grazing impacl
(hypothesis)
o
d
N
r!
o
!
mono-
dominance of
species (e.g.
Poa
angustifulia,
Calamagrostis
epigejos,
Cynodon
dactylon)
rncrease ln specles
of early
successional
stages (indirect
effects through
the initiation of
gaps)
selective grazing
on nutrient-rich
species, increase
in some avoided
endangered
species (e.g.
Corynephorus
canescens,R:
Koeleria glauca,
Helichrysum)
specific effects:
flower phenology
(e.g. induction of
second flowering
after srazins)
€€E
= b0'=
E€ E
O!:9E
E9
5!
arable field,
fallow land
(e.g. former
Zea mays
fields)
manipulation of the
upper soil layer (0-60
cm) which is rich in N
and P: transfer of the
upper layer or
substitution by the lower
layer poor in N and P,
modelling of dune
structures. hollows
the study sites are
treated with diaspores of
higher plant species and
cryptogams by transfer
of hay, litter or sods of
sand-specific
vegetation, seed bank
and seed rain are studied
selective grazing on
nutrient-rich,
especially ruderal
species, therefore
increase in some
avoided endangered
species which are
poor in nutrients,
grazing of
endangered species
will be analysed
Inland Sand Ecocsystems: Dynamics and restitution 241
Table I (cont.)
target
structures
(ahiotio)
target species/communities which still occur in intact sand
ecosystems (at the top) and target species/communities for the
restitution ereäs (af the hollnml /erennlcs)
vesetation. nlant snecies animal snecies
N
C!
gaps, dune
structures Corynephoretalia and Armerion
elongatae (e.g. Corynephorus
cane sc ens, Te e sda lia nud.ic au I i s,
Dianthus deltoides,
R: Phleum arenarium, Silene
conica, Veronica praecox, V.
verna, Arme ria* e longata)
R: Koelerion glaucae (e.9. Jurinea
cyanoide s, Koe le ria glauca,
Alys sum* gme linii, Medicago
minima, Bassia laniflora, Poa
badensis)
R: Oenanthe oenanthe (Aves),
Oedipoda caerulescens
(Caelif era), H ar pal us ne g le c tu s
(Carabidae)
R, H: My rme leote ttix mac ulatus
(Caelifera), C i c i nde Ia hyb r i da
s.l. (Cicindelidae), Poecilus
le pidus, Masore us we r I e r halli i
(Carabidae)
R: Eresus cinnaberinus s.l.
(Araneida), Bembix rostrata
(S phecidae), N o m i o i d e s
minutissimus, Andrena
carbonaria s.1., Halictus
sexcinctus
R, H: Andrena barbilabris
(Apoidea)
€€E
.E€ E
9€
EO
5L
redynamization
of dune
structures
H:
fluviodynamic
structures
in the floodplain
dry habitats: Corynephoretalia
/Armerion elongatae
R: Koelerion glaucae, elements of
Allio-Stipetum
H (moisUwet habitats): e.g. Elatine
species, different Nanocyperion
species
dry habitats: Poecilus lepidus,
Masoreus wetterhallii
(Carabidae) (question of
dispersal),
translocation experiments with
Harpalus neglectus (H)
(wingless species),
H = moist/wet habitats:
Agonum viridicupreum
(Carabidae)
R = Upper Rhine valley near Darmstadt; H = Ems/Hase region (NW Germany)
* = abbr. for species name
A management form aiming to maintain dynamic systems but also considering
economic aspects has to be devised. It should be applicable to both remnants of
these habitats and to restitution areas, i.e. areas where the re-establishment of
such ecosystems is attempted. Especially the latter aspect is of great importance in
regions where remnants of sand ecosystems are so small and fragmented that a
gene exchange has become almost impossible. Therefore, apart from existing
sandy regions, restitution areas are studied, among them an inland dune complex
upmodelled in the framework of a trial and development project. "projects that
combine protection and utilization aspects are of special significance" (FEDERAL
AcsNcy FoR NATURE CoNsenvlrroN 2000).
Angelika Schwabe et al.
The areas studied are representative oftheir respective regions; they are located
in the north-German lowland plain ("Hase valley", "Ems valley,'in the Emsland
region) and in the northern upper Rhine valley (sandy areas near Darmstadt).
Differentiated grazing systems with cattle, sheep, goats, horses, Mangalitza
pigs and donkeys are employed.
The Darmstadt University of Technology, the University of Osnabrück, the
University of Lüneburg and the university of Göttingen take part in this research
project.
2 Hypothesis, questions and the concept of restitution
and redynamization
Our hypothesis is that grazing systems with extensive management (< 0.7 live-
stock units/ha) and hardy breeds are suitable to promote dynamic processes:
- to redynamize sand ecosystems which have become consolidated and contain
mono-dominant species such as Poa angustifulia, Calamagrostis epigejos,
Cynodon dactylon and
- to develop restitution areas e.g. by selective grazing of nutrient-rich, especially
ruderal species; therefore populations of endangered plant species - often
avoided by grazing stock - may increase. (Possible negative effects will be
taken into consideration, see "Outlook".)
The following questions need to be addressed:
- which dynamic effects are caused by grazing in existing and restituted sand
ecosystems?
- which effects can be identified concerning abiotic and biotic resource
availability and preservation?
- How are the dispersal strategies in redynamised and restituted sand ecosystems
for selected plant and animal species?
- which effects are important for nature conservation management and for socio-
economics?
To summarize: which dynamic responses of abiotic and biotic structures can be
identified, given the impact of the grazing regime employed, and what
consequences for nature conservation and socio-economics can be deducted from
this?
Table I summarizes the concepts of redynamization and restitution including
the proposed hypothesis and relevant questions.
Inland Sand Ecocsystems: Dynamics and restitution 243
3 Some information about study sites and sampling
design
The sandy sites in the Upper Rhine valley near Darmstadt (R) and in the
Ems/Hase valley (H) still contain a number of endangered species and vegetation
types (e.g. R: Bromo tectori-Phleetum arenarii, Jurineo-Koelerietum, Armerio-
Festucetum trachyphyllae, H: Spergulo morisonii-Corynephoretum, Diantho
deltoidis-Armerietum), as well as larger populations of endangered plant and
animal species (e.g. R: Jurinea cyanoides, Bassia lanifTora, Nomioides
minutissimus, H: Harpalus neglectus); see Table L In the Ems/Hase region we
have endangered mosaic complexes consisting of inland dunes and temporarily
flooded hollows. r'*es
Fig. l. Example of a fenced 14 x 14 m exclosure and the grazed plots outside the fence
(colour infrared aerial photograph, May 2000, pixel size: 7 cm; "Ehemaliger August-Euler-
Flugplatz von Darmstadt" in the Upper Rhine valley). The grazed area is dominated by
flowering Cereutium arvense (white) and open structures which are very rich in
therophytes, such as Silene conica; the fenced area (since March 1999) already has a
different structure. The pathways for the study of flower phenology (exclosure: bottom left)
are clearly discernible
244 Angelika Schwabe et al.
E Bnün-Bhquct plot Elth plotl fd' phtn0toglcC sildrr (dotfcd)
I ptotr for nndüt rcolosr
[l vcücrt vcgrttlü cmcürr
n hor{zcüel YcAatalo lructlrr
a loggqß(tcmprnhc)
..... fdcc ddoroc
IIIIII
f--:T]
l---1o o
f:T1
l--1a a
Fig. 2. Diagram of the experimental design shown in Fig. L In our project about 35 fenced
exclosures will be studied
In open habitat sites, monodominant stands of Poa angustifulia, Calamagrostis
epigejos and, in the recent past, an increase in the Ca grass Cynodon dactylon have
been observed (R), especially when the land was not utilized.
The study areas in the Emsland region are grazed extensively with cattle and/or
horses. In the Darmstadt region mainly sheep grazing regimes with different sheep
breeds can be compared (hardy indigenous sheep breeds such as Rhoen sheep,
Skudde, Moorschnucke); Zatr,t et al. (2002). No supplementary feeding is
provided during the pasture period (for Rhoen sheep: May - December).
In addition to classic approaches (e.g. vegetation mapping) we have been able
to study replicated fenced and grazed plots in different vegetation complexes (e.g.
typical and ruderalised Koelerion glaucae complex, Armerio-Festucetum
trachyphyllae complex, Spergulo-Corynephoretum complex, Diantho-Armerietum
compfex and others) (see Figs. I and2).In both regions two replicated exclosures
were established in each vegetation complex. In a further step the data will be
scaled up in order to obtain results for the landscape level (see below).
Inland Sand Ecocsystems: Dynamics and restitution 245
4 Structure of the proiect
The investigations focus on the impact of the different grazing animals on
vegetation structure, nutrient dynamics and selected arthropod groups, as well as
on agricultural yield, applied economics and socio-economics. Moreover the
colonization dynamics of arthropod species in "habitat islands" (original situation
of restitution sites) is studied as an important aspect in nature conservation
management. The data are processed, analysed in a four-dimensional Geographic
Information System (GIS) and scaled up for larger areas.
4.1 Modute: Vegetation - structure and dynamics
The effects of grazing (mainly with cattle, sheep, goats, horses and Mangalitza
pigs) on the vegetation and structure of the stratified sampling plots are studied
and compared to reference plots. Existing sand ecosystems and restitution areas
are studied with respect to their horizontal and vertical vegetation structure, their
flower and fruit phenology and the diaspore dynamics. Relationships between
vegetation structure and animals like grasshoppers (Ensifera/Caelifera) and bees
(Hymenoptera Apoidea) are also studied. Two examples of grazing preferences
are given in Fig. 3.
Fig. 3. Grazing preferences demonstrated using wicker baskets (examples from the nature
reserve "Ehemaliger August-Euler-Flugplatz von Darmstadt"). Vegetation undemeath a
wicker basket of I x 2 m (right) and after two weeks of Rhoen sheep grazing (left).
Example I (left): Bromo-Phleetum-/Koelerion glaucae-complex with 6 endangered species
(e.g. Koeleria glauca, Medicago minima, Silen^e conica, Veronica verna, Veronica
praecox); standing crop (above stubbles): 87 glm'dry weight, grazed percentage of dry
weight26%o, nearly no endangered species grazed.
Example 2 (right): Armerio elongatae-Festucetum rich in the leguminous plant species
Medicago falcata with one endangered species (Medicago minima); standing crop (above
stubbles): 585 g/m2dry weight, grazed percentage ofdry weight6lVo
248 Angelika Schwabe et al.
communis). River engineering like the building of dikes hydrodynamically
separated the riparian grassland from the river. The frequency and duration of
floods was thus reduced and the ground-water level lowered. The region was then
intensively used agriculturally, leading to a quantitative and qualitative reduction
of the semi-open pasture landscapes and their biocoenoses.
We assume that it is possible to restore a levelled, heavily fertilized agricultural
area to an extensively used complex of inland dunes and temporarily flooded
hollows and to develop this complex as a dynamic system in the long term. Under
an extensive grazing regime site-typical plant and animal species with their
communities should develop from residual populations, from persistent seed
banks, from transferred diaspores (hay, sods; see Table l) or immigrate from the
vicinity.
Fig. 4. Concept for the restitution (including abiotic redynamization) within the "Hammer
Schleife" (Ems region, north-westem Germany)
4a Situation around 1900 (survey: Königlich Preußische Landesaufnahme)
4b Present situation (from topographic map I : 25,000)
4c Future relief of the restitution area adapted from the historic situation and considering
the local deposits of sandy material (relief modelling using the Surfer Software). (Height
given in m)
Inland Sand Ecocsystems: Dynamics and restitution 249
The biotic targets for the ecological restoration (BAKKER et al. 2000) have been
defined according to typical vegetation types (BerreR and BERENDSE 1999) and
animal communities of nutrient-poor sand ecosystems (target
species/communities, see Table l). For this reason the impact of grazing on intact
vegetation complexes of the Ems and Hase valley has also been studied. Grazed
and ungrazed (fenced) plots were set up in different vegetation units in these study
areas (different types of Spergulo-Corynephoretum, Diantho deltoidis-
Armerietum) with the experimental design shown in Fig. 2.
The restitution project has been divided into three phases: 1. Planning and
preliminary investigations; 2. Restoration and initial phase; 3. Phase of dynamic
development.
In the first phase, completed at the end of the year 2000, models of the
landscape including the abiotic target structures (Table l) were developed. A plan
for altering the dune morphology were drawn up (Fig. 2) on the basis of historical
data (maps, aerial photographs) and the models as well as the available soil masses
were assessed with regard to quantity and quality (soil type, nutrient content).
Ground modelling was completed in a short second phase in the winter of 2001 :
an artificial relief is to be created and simultaneously natural hydrodynamics are to
be restored. The upper soil layer (0-60 cm) which is rich in N and P was
transferred to the deeper part of the dune structures and was substituted by layers
poor in nutrients. Immediately after having moved the soil, site typical grassland
species are to be sown in the main area to prevent ruderalization. The reshaped
dune areas (Fig. a) are to be "inoculated" through the transfer ofhay or sods from
sand-specific vegetation. Sporadically, site-typical shrubs will be planted (e.g.
Prunus spinosa). Eight permanent plots will be established, four of which will be
ungrazed fenced areas. Measurement devices for hydrogeological / geochemical
investigations (WetNrnr et al. 2000) was placed in the soil (soil moisture samplers
with nylon cups, water-level indicators, lysimeters).
In the third phase of the development of natural morphological structures and
biocoenoses under the influence of abiotic processes and extensive grazing, these
systems will be studied carefully (monitoring). "Monitoring is a conditio sine qua
non for the evaluation of the effects of ecological restoration" (Bnrxen et al.
2000).
Other restoration projects in smaller areas (l-2 ha) were initiated in the
Darmstadt-Dieburg region. One project has already been running since 1998 and
demonstrates the effects of grazing by sheep preferring nutrient-rich plant species
and neglecting e.g. Corynephorus canescens, Koeleria glauca and other
endangered species.
7 Outlook
Having completely analysed and summarized the data, recommendations for
nature conservation measures can be formulated, which should be applicable to
250 Angelika Schwabe et al.
other sand ecosystems and restoration projects. The measures are to be judged as
to their efficiency, also considering economic aspects.
Following the first year of study there are some initial results;
- an increase in plant species biodiversity in mono-dominant stands of
competitively aggressive grasses, such as Calamagrostis epigejos, after grazing
(Zzuvt et a1.,2002)
- an increasing number of gaps and therefore of therophytes in the grazed plots
(e.g. Silene conica)
- strong structural changes in both grazed and ungrazed plots. Results concerning
connections e.g. between structural changes, plant and animal populations will
be outlined.
Possible negative effects of local overgrazing and grazing of endangered
species as reported by RosEN and vnN DER MAAREL (2000) for the Alvar
vegetation (Öland) will be taken into consideration. In our extensive grazing
systems local overgrazing probably does not play a role. Grazing of endangered
species will be analysed, such as Armeria*elongata and Dianthus deltoides.In the
case of Armeria a second flowering induction phase after grazing can be observed.
We are confident that our project can contribute empirical and synoptic data to
answer questions on the importance of dynamic processes in landscapes grazedby
hardy breeds (Rncr et al. 1998).
8 German summary
In der Landschaft sowohl der nördlichen Oberrheinebene als auch des Emslandes sollten
Sandstandorte Beispiele von Ökosystemen darstellen, die z.T. noch großflächigeren
dynamischen Prozessen unterliegen. Die Sandstandorte des nördlichen Teils der
Obenheinebene weisen eine Reihe extrem gefährdeter Vegetationstypen, Pflanzen- und
Tierarten auf (s. Tab. l). Im Zuge von Ruderalisierungen nehmen in den letzten
Jahrzehnten unduldsame Gräser wie z.B. Schmalblättriges Rispengras (Poa angustifulia)
und Sandrohr (Calamagrostis epigejos) überhand. Die flußnahen Sand-
Vegetationskomplexe im Ems-/Hase-Gebiet bergen mit Sand-Pionierfluren und -Rasen in
ihrer Verzahnung mit Flutmulden ebenfalls eine Fülle gefährdeter Vegetationstypen,
Pflanzen- und Tierarten (Tab. l).
.. Das typische Vegetationsmosaik der gefährdeten Pflanzengesellschaften von Sand-
Okosystemen bildet sich in der Regel nur bei vorhandenen Störungen aus, die die
Sukzessionsprozesse immer wieder initiieren; so kommen konkunenzschwache Pionier-
Pflanzenarten zur Entwicklung und xero-/thermobionte Arthropoden finden entsprechende
offene Lebensräume.
Neben einer möglichen Redynamisierung gewachsener Ökosysteme spielt für den
Naturschutz die Frage der Restitution eine bedeutende Rolle. Unsere Hypothese ist, daß
Beweidung ein tragftihiges Konzept darstellt, um konsolidierte Sandstandorte zu
redynamisieren und um Restitutionsflächen im Sinne des Naturschutzes zu entwickeln.
In einem BMBF-Projekt werden in der nördlichen Obenheinebene und im Emsland
neben bestehenden Sandgebieten auch Restitutionsflächen untersucht, darunter eine in dem
E+E-Vorhaben ,,Hasetal" aufmodellierte Binnendünenfläche. Differenzierte Weidesysteme
Inland Sand Ecocsystems: Dynamics and restitution 25r
(u.a. mit Rindem, Schafen, Ziegen) werden eingesetzt. Wesentliches Ziel des Projektes ist
die Untersuchung der Auswirkungen unterschiedlicher Weidesysteme auf die Vegetations-
entwicklung, die Nährstoffdynamik und ausgewählte Tiergruppen. Ertrags- und
betriebswirtschaftliche sowie sozioökonomische Untersuchungen ermöglichen eine
Bewertung der ökonomischen Gesichtspunkte der verschiedenen Beweidungsvarianten.
Unter Berücksichtigung der naturschutzfachlichen und sozioökonomischen Aspekte wird
eine Szenarienbildung mit Hilfe eines vierdimensionalen GIS (Raum und Zeit) erarbeitet.
In allen Gebieten wurden neben größerflächigen Untersuchungsansätzen (Luftbilder,
Vegetationskartierungen) nicht beweidete Vergleichsfl ächen eingerichtet.
Nach vollständiger Analyse der Daten sollen Empfehlungen für den Naturschutz
gegeben werden, die auch auf andere Sandökosysteme und Restitutionsprojekte übertragbar
sind. Hierbei wird auch dieEffrzienz der möglichen Maßnahmen betrachtet.
Acknowledgements and corresponding authors
We are grateful to the German Federal Ministry for Education and Research
(Bundesministerium für Bildung und Forschung, BMBF; funded 2000-2003, No.
01LN0003) and the Federal Agency for Nature Conservation (Bundesamt für Naturschutz,
BfN) for supporting our projects. The studies have been made possible by the cooperation
of the competent authorities; in some instances cooperation contracts exist with these
authorities. We would especially like to thank the Weser-Ems regional administration, the
Darmstadt regional administration and the Darmstadt-Dieburg district authority. We are
also grateful to the livestock owners in both study areas for their cooperation.
More information about the project can be found at:
http://www.tu-darmstadt.de/fblbio/bot/geobot/BMB FSTAR.htm.
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... Aufgrund der Zunahme von Ruderalisierungs-und Sukzessionsprozessen in den wertvollen primär basenreichen Sand- Ökosystemen der nördlichen Oberrheinebene haben wir zu Beginn des BMBF- Projektes " Sand-Ökosysteme im Binnenland: Dynamik und Restitution " die Hypothese formuliert, dass Beweidung zur Redynamisierung der Standortskomplexe führt. Das Ziel hierbei ist, mit Hilfe von Beweidung konsolidierende Prozesse umzukehren und die Lebensräume zu " dynamisieren " (SCHWABE et al. 2002). In unseren Hauptuntersuchungsfl ächen im Bereich des Offenlandes ( " Ehemaliger-Euler-Flugplatz von Darmstadt " und " Griesheimer Düne und Eichwäldchen " ) ging es vor allem um die Zurückdrängung von Ruderalisierungs-und fortgeschritteneren Sukzessionsstadien im Zusammenhang mit einer Beweidungs-induzierten Dynamik. ...
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... These studies also aim to indicate management measures for the conservation of both lichens and vascular plants in these important Corynephorus grasslands. Possible management measures to restore and manage this particular habitat could include mechanical disturbance (Jentsch 2004;Ketner-Oostra et al. 2012), dispersal of lichen fragments (Heinken 1999;Jeschke 2012) and grazing by sheep (Schwabe et al. 2002). Grasslands are generally considered of high conservation concern because they are rich in species, and thus host a high proportion of Europe's biodiversity (Pärtel et al. 2007;Janišová et al. 2011;Blasi et al. 2012). ...
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In the course of a four-year project the effects of extensive grazing in sand ecosystems were studied. The research work focused on two sandy areas: (1) sand ecosystems as well as restoration areas situated in the upper Rhine valley which were newly grazed with mostly landrace breeds of sheep (partly also with donkeys and goats); (2) target areas characterized by semi-open cattle-grazed pastures as well as a restitution area of 37 ha situated in the Ems region. Exclosures and grazed areas in the upper Rhine valley and in the restitution areas, and since 2000 also fenced areas exhibiting spontaneous successional processes on the permanent pastures situated in the Emsland region (target area), were the basis for a field experiment. Moreover, vegetation dynamics were studied on the basis of a regular grid. As part of a pilot study concerning restitution measures and grazing impact, studies have been realized for five vegetation periods comparing plots inoculated with mown, raked material or sods and non-inoculated plots. The grazing impact on newly grazed areas depended to a high degree on the respective grazing systems. In the case of a grazing regime with relatively small paddocks of 1 ha exposed to temporary intensive grazing (only a few days within one year), the following effects, which are important for nature conservation, could be demonstrated: an increase in plant species numbers and especially an advancement of Red Data species. This correlated e.g. with changes in the vertical vegetation structure (thinning). The soil temperature of grazed areas rose by 0.8 °C on average enabling above all the advancement of thermobiotic animal species. Regression effects became obvious in the case of, among others, Calamagrostis epigejos. Trails of sheep play an overall role concerning regressive dynamics. The successional processes observed in the formerly grazed, exclosures in the Emsland region are characterized by extended lag-phases. Some effects have been evident after a four year period; these include for example a lack of plant populations dynamics, such as in the case ofthe acrocarpic moss species Ceratodon purpureus, which is favoured by grazing. To date, the restoration experiment studied over five vegetation periods has been successful. Many Red Data Book species have been transferred to the restitution area and tall species (often ruderal plants) have been repressed. The repression of ruderal plant species became especially obvious where successive multispecies grazing systems with ruminants (here: sheep) and non-ruminants (here: donkeys) were used. The most successful inoculation method used in the upper Rhine valley - "inoculation with mown/raked material" (material taken from donor plots in target areas) - was subsequently also used in the Emsland study areas.
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In the Emsland region the impact of cattle grazing on the vegetation of semi-open pasture landscapes was studied by exclosure experiments and a raster-based analysis (50-m spacing) of grazing pressure. The exclosure experiment refers to inland sand dune complexes, whereas the raster-based analysis comprises the whole vegetation mosaic of the open pasture landscape (grazed sand dunes, fresh and moist pasture habitats). Both approaches were undertaken in target and newly restored areas. For the exclosure experiment a set of grazed plots was fenced and the cattle excluded in the target area in the year 2000. There were only slight changes in the fl oristic composition of the studied sand vegetation (Spergulo-Corynephoretum and Diantho-Armerietum) in the 4-year period. Especially the moss Ceratodon purpureus and the grass Festuca fi liformis show higher population dynamics in the grazed plots. We expect this “lag period” to be followed by stronger effects of fl oristic change in the coming years. According to the results of the raster-based analysis (which used phytosociological relevés carried out yearly and a special grazing-pressure evaluation: three times a year) cattle generally prefer plant species of moist to fresh or not too dry grasslands in the studied Vegetation mosaic (e.g. species of Lolio-Cynosuretum, but also of Diantho-Armerietum and Agrostis capillaris-community). Preferred species were: Agrostis capillaris, A. stolonifera, Alopecurus geniculatus, A. pratensis, Bidens frondosa, Cerastium arvense, Dactylis glomerata, Elymus repens, Holcus lanatus, Juncus effusus, Poa pratensis and Rorippa palustris. Target species of the Spergulo-Corynephoretum are not preferred but the open dune structures had a function as “wellness” places (wind infl uence causes lower densities of Diptera). This basal preference list is modified mainly by three aspects: 1. stocking density, 2. ratio between moist, fresh and dry grassland in the vegetation mosaic, 3. specific weather conditions during a given year. Although all investigation areas were extensively grazed by cattle, there were differences in the stocking densities (counted for May to October): target area with 0.7-0.8 livestock units/ha, restoration areas “Hammer Schleife” with 0.6 livestock units/ha and “Wester Schleife” with 0.9 livestock units/ha. In the case of the target area all vegetation types were grazed, including the sand dune complexes. In the case of the restoration areas (which currently have more fresh and moist grassland than the target area) the sand ecosystems were sufficiently grazed in the “Wester Schleife”, with 0.9 livestock units/ha, but not in the area “Hammer Schleife”. Therefore in the case of dominance of fresh/moist grassland the stocking density should be increased to 0.9 in the area “Hammer Schleife”. In future it is expected that both restoration areas may change their vegetation mosaic more and more by accumulation of fluviatile sand (as already happened in winter 2002/03); the stocking density has then to be adapted to the value of the target area. The faeces are more or less homogeneously distributed in the area and do not correlate with the sites of high grazing pressure, which guarantees the dispersal of endozoochorous species throughout vegetation mosaic.
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In a sand-ecosystem (Allio-Stipetum complex) situated in the nature reserve “Griesheimer Düne und Eichwäldchen” (Darmstadt) influences of sheep grazing on wild-bee coenosis and on their food resources were studied. A three-year study (following initial grazing) comparing vegetation on grazed and ungrazed areas shows a significant increase of some plant species used by wild bees as well as an increase of Calamagrostis epigejos in ungrazed areas. Even after three years, grazed and ungrazed areas hardly show any difference concerning their flowering phenology. Concerning their species and abundance, there are also no significant differences between wild-bee species on grazed and ungrazed areas. However, according to a more detailed analysis, grazed older sand dune complexes show signifi cantly higher numbers of wild-bee species than younger ones, the latter having previously been exposed to severe anthropogenic impact. According to a Detrended and Canonical Correspondence Analysis, the species composition of wild bees on the specific grid areas depends on the pollen and nectar resources available, whereas, in the present vegetation stage, grazing and non-grazing factors do not play any role. Some species (e.g. Centaurea stoebe) are characterised by a correlation between increasing resources and increasing numbers of wild-bee species. Patchy sheep grazing on small areas for a limited time starting at the beginning of July has not caused any impact on the wild-bee community. On the contrary, grazing produces nesting sites of bare ground for endogeic wild-bee species, promotes important pollen and nectar resources on the one hand and prevents the further succession of mono-dominant grass species (e.g. Calamagrostis epigejos) on the other.
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The main subject of the following study is the question of synecological interactions between flowers and insects on the complex level of plant communities and communities of flower-visiting insects. The studied area represents a fallow limestone grassland complex (Mesobrometum) in the Kaiserstuhl. In the two years (1979 and 1980) 3600 flower-visiting insects (Hymenoptera, Lepidoptera, Diptera and Coleoptera) could be observed visiting 71 different plant species. The high species diversity is shown by the fact that 102 bee species and 56 butterfly species were recorded in an area of 4000 m². Botanical, zoological and flower-ecological results and their discussion form the three main parts of this paper. Special attention is payed to the phenology of flowers and insects (the "co-phenology") and to questions of population dynamics (flight activities of insects, flowering_ times of entomophilous plants) especially under consideration of area geography. It is shown that the history of the flora and fauna is important in the Interpretation of the flower-visitor communities beside the different competitive exclusion mechanisms (e.g., staggering of flowering times by plants, different lengths of mouth parts by insects, food niche contraction or niche expansion). The limestone grasslands ( Mesobrometum ) in southwestern Germany are characterized by a high percentage of submediterranean and subcontinental plant and animal species. A phenological analysis (flowering times of entomophilous plants, flight activities of Hymenoptera Apoidea and Lepidoptera) yields four seasonal periods named by the area geographical centre of their species: eurosibirian period (March, April), submedirerranean period (May, June), eurosibirian period (July), eurosiberian period wirh subcontinental and submediterranean elements (August, September). Vegetation, Apido- and Lepidofauna of the same area type correspond in their phenology (co-phenology). The flower-visiting insects of the studied area prefer plants of the same geoelement. It is argued that the euryanthic behaviour of Hymenoptera Apoidea is a highly developed feature representing the precondition ("preadaptive plateau") of longer time of flower-visiting during the year, of bivoltinism and socioevolution. Stenanthy can be interpreted as an original feature and, in middle European bees, can only be understood with regard to the history of flora and fauna of the postglacial period.
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Studies of disturbance have a long tradition in vegetation ecology (Cooper 1926; Raup 1941; White 1979) and have increased lly during the last 30 years (Dayton 1971; Heinselman 1973; Levin and Paine 1974; Borman and Likens 1979; Sousa 1979a,b, 1984; Pickett 1980; Pickett and White 1985; Van der Maarel 1993; Bornette and Amoros 1996; Paine et al. 1998; Freiich and Reich 1999; White et al. 1999). We have learned a tremendous amount about the significance of disturbance as an ecological factor in various habitats and communities (Knapp 1974; Grubb 1977; Miles 1979; Oliver 1981; Pickett and White 1985; Goldberg 1988; Frelich and Lorimer 1991; Milton et al. 1997), about disturbance regimes (Romme 1982; Turner et al. 1993; White et al. 1999), about functional adaptations of plants (Garcia-Mora et al. 1999; Walker et al. 1999), about responses of ecosystems (Bornette and Amoros 1996; Johnson et al. 1998; Engelmark et al. 1999) and about restoring disturbance as an ecosystem process (White and Walker 1997; Covington et al. 1999). During this period, a few theories and synthetic concepts have been proposed, but we do not yet have an inclusive general paradigm for this important body of work.
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An initially uniform Holcus lanatus-dominated sward came partly under hay-making and partly under sheep-grazing. Preferential grazing by sheep resulted in grazing at different intensities giving rise to a macro-pattern of various plant communities. Besides this macro-pattern a micro-pattern developed in the grazed area, which was absent under hay-making. In the micro-pattern short, heavily grazed areas alternated with taller, lightly grazed patches, both having the same species composition. The heavily grazed area was characterized by equal amounts of monocots and dicots. The lightly grazed patches were dominated by Agrostis tenuis, and had a large amount of litter which probably causes the absence of mosses. The protein percentage of green material is higher in the heavily grazed areas than in the lightly grazed patches. Sequential charting indicated that the micro-pattern was more or less stable. An interaction between the vegetation micro-pattern and grazing patterns is suggested. Heavy grazing results in forage with a high protein content and hence attracts animals. Light grazing results in forage with a relatively low protein content, animals avoid the area and litter accumulates.
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Free‐ranging large grazers, such as cattle and horses, are increasingly reintroduced to former agricultural areas in Western Europe in order to restore natural and diverse habitats. In this review we outline mechanisms by which large grazers induce and maintain structural diversity in the vegetation (mosaics of grasslands, shrub thickets and trees). This variation in vegetation structure is considered to be important for the conservation of biodiversity of various plant and animal groups. The process of spatial association with unpalatable plants (as‐sociational resistance) enables palatable plants to establish in grasslands maintained by large grazers. In this way, short unattractive (thorny, low quality or toxic) species facilitate taller unattractive shrubs, which facilitate palatable trees, which in turn outshade the species that facilitated their recruitment. Established trees can, therefore, not regenerate under their own canopy, leading to cyclic patch dynamics. Since this cyclic dynamic occurs on a local scale, this contributes to shifting mosaics. The mechanisms involved in creating and maintaining the resulting shifting mosaics are described for temperate flood‐plain and heathland ecosystems, including the effects on nutrient transport within grazed landscapes. How grazing leads to shifting mosaics is described in terms of plant functional types, allowing potential generalisation to other ecosystems. The resulting interaction web of grasses, unpalatable forbs and shrubs, palatable light‐demanding trees and shade‐tolerant trees is discussed, and was found to contain various interesting direct and indirect effects. The key process contributing to spatial diversity in vegetation structure is the alternation of positive (facilitation) interactions between plant species at one life cycle stage, and competitive displacement at another stage. Grazing thus causes directional successional sequences to change to shifting mosaics. The implications of this theory for nature conservation are discussed, including the relevant management problems, possible choices and practical solutions. We conclude that the theoretical framework outlined in this review provides helpful insights when coping with nature conservation issues in temperate woodland habitats.
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Alvars in the Baltic region, particularly on the Swedish islands of Öland and Gotland and in western Estonia, are well‐known for their plant species richness and extensive populations of rare species. Grasslands make up most of alvar vegetation. The extent of these grasslands decreases because of bush enchroachment which takes place in most alvars when traditional land use practice changes, notably when grazing is ceased. The main threat for the alvar grassland is formed by the rapid expansion of Juniperus communis and Potentilla fruticosa in dry and wet sites, respectively. Applied research has been carried out during several years in order to develop plans for the restoration of alvar grassland. Thus it was shown that species richness in areas with a mixture of grassland and shrubland decreases in relation to the increased cover by shrubs. In the case of Juniperus there is a distinct drop in species number as soon as the cover exceeds 75‐80%, while the decrease is more gradual with increased cover of Potentilla. The seed banks under dense stands of these shrubs only contain ca. 20% of the species found in dry and wet alvar meadows. This was confirmed by clearing experiments. Long‐term recordings have shown that establishment of juniper seedlings takes place mainly in half‐open areas between already existing junipers. Intermediate‐sized junipers expanded faster than old and big shrubs. Potentilla shrubs recover fast from cutting or burning. After 2‐4 yr they have almost attained their former size. This recovery can be prevented when cattle grazing is introduced. In the framework of a local alvar restoration project on Öland, covering 7000 ha, grazing regimes are re‐established, fences erected and large‐scale clearings carried out. Recently established juniper shrubs are being cleared, intermediate sized junipers (30‐50 yr old) are selectively removed while creating a mosaic landscape with high biological diversity. Older dense juniper stands are left alone or are only partly cleared. Potentilla stands in moist areas are cut to create moist meadows which are breeding grounds for waders, and to establish corridors between remaining open areas. Three items are discussed (1) the importance of the Stora Alvar area; (2) re‐introduction of grazing and (3) re‐introduction of species. The outstanding importance of the area regarding species richness and endemism should be recognized through a ‘golden list’ to be used along with red and blue lists.
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Species-rich grassland and heathland communities still occur in low-intensity farming systems in many European countries. Gradually, such systems have either been abandoned or more intensively exploited, with a subsequent decrease in species numbers. Until recently, it was thought that restoration of these communities would be straightforward. However, abiotic constraints (with respect to eutrophication and acidification) have hampered restoration more than previously thought. Moreover, very recent research has revealed that biotic constraints can also be important: many plant species are not present in the soil seed bank and their dispersal is limited in the present fragmented landscape.