Biological Conservation 97 (2001) 319±330

www.elsevier.com/locate/biocon

Succession from farmland to heathland: a case for conservation

of nature and historic farming methods
H.J. Degn
Skolevej 44, DK-6950 Ringkùbing, Denmark

Received 13 October 1997; received in revised form 5 July 2000; accepted 7 July 2000

Abstract
The succession of vascular plants was recorded in an old ®eld during 22 years after farming ceased in 1975. The soil is poor, acid
and sandy. Adjoining areas are Calluna±Empetrum heathland. During the ®rst 2 years a few ruderal weedy species dominated
completely. In years 3±5 Jasione montana and Rumex acetosella became dominant. More persistent perennials such as Hieracium
pilosella and Deschampsia ¯exuosa slowly colonised, but after years 13±14 they decreased again. During the last 10 years Calluna
vulgaris and Festuca ovina increased. Calluna vulgaris is now so abundant that the area is recognised as heathland, according to the
de®nition in the Danish Nature Protection Act. The advantages of restoring heathland vegetation by natural succession are com-
pared with more intensive actions. Rarer plant species than those occurring in either farmland or heathland are found, the process
gives a more varied vegetation, and it is historically authentic. # 2001 Elsevier Science Ltd. All rights reserved.
Keywords: Succession; Farmland; Heathland; Restoration; Calluna vulgaris

1. Introduction heathland to woodland or intensively used farmland.

Until about 200 years ago heathland dominated by
heather (Calluna vulgaris) covered more than a quarter
of the area of Jutland, Denmark (Hansen, 1970). In
large parts of the province <10% of the land area was
cultivated.
The fact that the extent of heathlands was due to
human activities is supported historically (e.g. Matthiesen,
1939) as well as biologically (e.g. Odgaard, 1994). After
the primeval forest had been cleared, extensive land use
practice ensured that vast areas remained heathland. It
was grazed by sheep and cattle, cut for fodder and fuel,
burnt to improve grazing, and turves were used to collect
manure and burnt for heating and to ash as a fertilizer.
Old farming traditions sometimes involved cultivation
of remote ®elds in the heathland for only 2±5 years.
When the pool of available nutrients was spent, the ®eld
was abandoned. After a few years Calluna vulgaris
would cover the area again.
After World War II it has become exceptional to observe
®elds returning to heathland, and the general trend has
been in the opposite direction, namely development from
E-mail address: lanhjd@ringamt.dk (H.J. Degn)
0006-3207/01/$ - see front matter # 2001 Elsevier Science Ltd. All rights reserved.
PII: S0006-3207(00)00131-2
The area of heathland in Denmark decreased from about
800,000 ha in 1822 to about 70,000 ha in 1991, half of
which is dune heath (Emsholm, 1992). Most of the
remaining heathland is now protected against active
changes like cultivation, planting, etc., but not against
natural succession, and for only a small part is there an
obligation for management.
The purpose of the present paper is, ®rst, to describe a
case history of natural invasion by vascular plants into an
abandoned sandy ®eld in a heathland area in Jutland over
a period of 22 years after cultivation ceased. Second, to
show that this succession was formerly a regular feature
on abandoned ®elds in heathland areas. And third, to
consider how experience from this study can be used in
nature conservation today. The latter question is perti-
nent, now that agricultural land is taken out of produc-
tion in many Northwestern European countries. Can
some of these areas be converted into natural habitats
as heathland ?
The answer is important in Denmark, as heathlands
with their ¯ora and fauna have a high conservation
priority. For some types of heathlands a high proportion
of the total area is found in Denmark (Diemont et al.,
1996), which gives the country an obvious responsibility.
320 H.J. Degn / Biological Conservation 97 (2001) 319±330
At the same time the cultural values are high: heath-
lands play a signi®cant roÃle in Danish art, literature,
and general identity.
2. Study area
The location of the study is on Air Station Karup
(9 30 E, 56 190 N), which lies on the Karup Melt-water
Plain from the Weichsel Ice Age (Fig. 1). The soil is
coarse outwash sand (49% in the fraction 0.5±0.25 mm).
It is a highly mineral soil, only about 1.5% of the upper
25 cm was organic matter at the start of the study. At
that time the soil was thoroughly mixed to ploughing-
depth (c. 30 cm) due to agricultural use, which ended
the year before.
The study area was a 2350 m plot making up the
western tenth of an unusually narrow ®eld only 23±25 m
wide (Fig. 1). It had been cultivated for more than 60
years (T. Andersen, pers. commun., 1975; topographical
map from 1916). During the ®nal years of cultivation the
crop was spring-sown barley (Hordeum vulgare). After
the last crop in 1975 the soil was ploughed, so that there
was no plant cover present at the start of the study.
Farming methods had been normal except that fertilizing
and control of weeds was inecient during the last years
of cultivation, so that couch-grass (Elytrigia repens)
occurred abundantly.
The adjacent areas along the longer edges of the ®eld
were heathland during the whole study period (Fig. 1).
Calluna vulgaris was dominant, and other common species
Fig. 1. Location of the study area in Denmark (upper left) and in the
landscape (below).
were wavy hair-grass (Deschampsia ¯exuosa), crowberry
(Empetrum nigrum), and cowberry (Vaccinium vitis-idaea).
The vegetation corresponds to the heather±crowberry±
cowberry heath type (5.1.1.3) in the Scandinavian classi-
®cation of vegetation types (PaÊhlsson, 1994). The vege-
tation was cut in 1979 along the northern edge, after
which the vegetation went through the pioneer, building,
and mature phase (Gimingham, 1972). During the last 5±
10 years of the study mountain pine (Pinus mugo)
increased on the northern side, and on the southern side
Calluna vulgaris began to die from old age.
3. Methods
3.1. Species composition of vegetation
From 1976 to 1997 the vegetation of the study area
developed without any direct human interference. Graz-
ing was infrequent and only by hares (Lepus capensis).
The species composition of the vegetation was investi-
gated in 100 circular plots of 0.1 m2 (radius 17.8 cm) at 2 m
intervals along four ®xed transects. From 1994 the num-
ber of sample plots was reduced to 92 due to a military
construction at the western end of the area. The total area
of the plots was about 1% of the study area. Because the
vegetation was approximately uniform over the study
area, the data gave a good description of the vegetation.
All vascular plants rooted or with perennating buds
within the plots were recorded (Raunkiñr, 1934), and
frequency percentages fn was calculated for each species
n as the number of occurrences relative to the total
number of plots.
Diversity was calculated using species presence and
species frequency for each year. The following diversity
measures were used: species number S, frequency sum F ˆ
fn and Simpson's diversity index Sdiv ˆ 1 ÿ  f=F†2 .
Simpson's index ranges from 0 when all individuals
belong to the same species, towards a maximum value of 1
when many species are found, but only a few times each.
Data were collected from 1976 to 1997, always in late
June or in July each year. At this time the latest perennials
(especially Asteraceae) were in bloom, while the dead
winter±annual plants were still present and could be iden-
ti®ed. Species nomenclature follows Hansen (1981).
A TWINSPAN analysis was run, based on the species
with a mean frequency >5% (McCune and Me€ord,
1997). Nine pseudospecies levels were considered on a
linear scale. By this the entire frequency scale is repre-
sented equally, and the pseudospecies levels correspond
roughly to frequency/10.
3.2. Chemical analysis of soil mineral nutrients
Analysis of the soil was made every year 1976±1982 and
then in 1985, 1987, 1989, 1991, and 1996 by an authorized
 H.J. Degn / Biological Conservation 97 (2001) 319±330
commercial laboratory by Danish standard methods used
in agriculture (Anon., 1994). Five subsamples of the soil
down to a depth of 20 cm were collected in March or April
in a ®xed pattern on the study area. These subsamples
were thoroughly mixed before the soil sample was taken.
The concentration of the following elements were mea-
sured: Phosphorus (P) (inorganic phosphates extracted
in 0.1 M H2SO4), sodium (Na) and potassium (K)
(extraction in 0.5 M NH4OOCCH3), calcium (Ca) and
magnesium (Mg) (extraction in 1 M NH4Cl), manga-
nese (Mn) [extraction in 0.5 M Mg(NO3)2], and pH
(measured after suspension in 0.01 M CaCl2).
The concentration of organic matter was determined in
April 1980 and March 1997 down to ploughing depth (c.
30 cm) by ignition loss (heating for 2 h to 550 C). During
the 22 year period the uppermost few centimetres of the
soil pro®le became darker than the lower part, possibly
indicating an accumulation of organic matter near the
surface. Therefore, at the end of the study (1997) soil
pro®les were separated into two samples 0±3 and 3±25
cm depth, which were analyzed separately.
4. Results
4.1. Vegetation
A total of 74 plant species was recorded during the per-
iod 1976±1997. The number increased rapidly from 10
species in 1976 to a maximum of 41 species 8 years later
(Fig. 2). During the last 13 years the number slowly
decreased to about half the number. Annual forbs
Fig. 2. Number of plant species of di€erent growth forms found dur-
ing the period 1976±1997. The total number of plant species (S) is
shown by the upper limit of the shaded area.
321
dominated in the ®rst 3 years, and reached their max-
imum number of 16 species in 1980. They were mostly
weed species from the former agricultural plant commu-
nity. They subsequently decreased to two species in 1997.
When the vegetation had closed after 3 years the sig-
ni®cance of this group became even lower than species
numbers indicate. The plants were as a rule very few
and weakly developed, only germinating in small pat-
ches where the mineral soil was exposed, e.g. by animal
activity.
Annual graminoids and biennial forbs were negligible,
the total species number of these two growth forms
combined did not exceed four in any year.
Perennial forbs and perennial graminoids showed
parallel patterns. A rapid increase was observed from one
in each group in 1976 [sheep's sorrel (Rumex acetosella)
and Elytrigia repens] to maximum species numbers in
1982±1987. From then on numbers decreased slowly to
1997, to about half this maximum.
The ®rst ericaceous dwarf-shrub (Calluna vulgaris)
appeared in 1981, and the second species (Empetrum
nigrum) 4 years later. No other dwarf shrubs invaded in
the last part of the period, not even Vaccinium vitis-
idaea, which is very common in the adjacent heathland
Species numbers give only limited information about
the importance of the di€erent species in the vegetation
since a species counts equally in Fig. 2 whether it is
recorded in one or in 100 circles. In Fig. 3 the frequency
sums are transformed to percentage values which illus-
trate more clearly the dominance of particular growth
forms: annual forbs dominated in the early years, while
dwarf shrubs became more important in the later part
Fig. 3. The percentage distribution of growth forms during the period
1976±1997 according to frequency sums.
322 H.J. Degn / Biological Conservation 97 (2001) 319±330
of the period, and the frequency sum for perennial gra-
minoids was twice as great as for perennial forbs.
Vegetation diversity, as described by Simpson's
diversity index (Fig. 4), rose rapidly to high values in
1980±1983 and then decreased again, as a few species
became dominant in the vegetation.
A high proportion of the total 74 species occurred
sparsely in the vegetation (Table 1). For example 16
species were never found in more than one plot per year
out of the 100 (92) plots investigated. Fig. 5 shows the
frequency percentage during the period for 12 species
that obtained a value of >50% in at least one year. The
species are arranged according to the year, when they
had their maximum values, i.e. early immigrants ®rst
and late dominants last.
The results of the TWINSPAN analysis are shown in
Table 2. Seven stages could be recognised (A±G) and six
species groups were revealed (a±f). Stage A (1976±1977)
is dominated by weeds from group a, Polygonum con-
volvulus, Polygonum aviculare and Elytrigia repens.
Stage B (1978±1981) is characterised by group b pioneer
species (Rumex acetosella, Trifolium arvense and Trifo-
lium campestre) together with group c (only consisting
of Jasione montana).
Stage C (1982±1984) appear as a stage of declining
group b species and establishment of group d and e
species. These two groups mostly comprise common
acid grassland species, namely Agrostis tenuis, Hypo-
choeris radicata, Festuca rubra, Teesdalia nudicaulis,
Fig. 4. Diversity of the vegetation during the period 1976±1997
expressed as Simpson's index of diversity: Sdiv ˆ 1 ÿ  fn =F†2 .
Achillea millefolium, Deschampsia ¯exuosa, Hieracium
pilosella, and Carex arenaria. Group d seems to di€er
from group e mainly in having a shorter persistence.
Stage D (1985±1987) has both transitory and persistent
grassland species from group d and e, whereas in stage
E (1988±1991) group d declines and terrain is slowly
gained by late immigrants of group f (Luzula campes-
tris, Calluna vulgaris and Festuca ovina). Stage F (1992±
1993) and stage G (1994±1997) are very similar and
show a consolidation of dominance of group f species,
forming the heath-like physiognomy of the vegetation.
4.2. Soil chemistry
Some of the soil chemical parameters changed during
the period of the study (Fig. 6). There was a steady
decrease in the phosphorus content of the soil during
the period from about 8 mg to about 2±3 mg per 100 g
soil. Sodium showed a steady increase for the ®rst 6
years, but after a peak in 1989 decreased to original
levels. Calcium content showed an initial decrease dur-
ing the ®rst 6 years, followed by a relatively stable value
for the rest of the period. Soil pH showed a linear
decrease of 0.5 unit over the period. The organic matter
content was 1.5% at the start. In 1997 the concentration
in the uppermost 3 cm had increased to 3.7%, while in
the lower 3±25 cm it was only 1.2%.
5. Discussion
5.1. The succession compared to other reports
Continuous records of the succession described above
have not been published before, but several of the stages
are known from static descriptions. Details vary, but the
results of Warming (1897), Bùrgesen and Jensen (1904),
Ferdinandsen (1918), BoÈcher (1941, 1970), Holst (1987),
Mogensen (1994), and Rehfeldt (1999) all ®t into the
succession described in the present paper. Characteristic
species as well as combinations of species in these
descriptions are the same as found in the present study
(Table 2 and Fig. 7), so the general trend of the succes-
sion described in this paper is normal.
5.2. Factors a€ecting the process
The succession from farmland to heathland is slower
to-day than in earlier times. Old descriptions of farming
practices in Middle and Western Jutland reported that
Calluna vulgaris began to dominate 3±5 years after ces-
sation of cultivation (Begtrup, 1808±1812; Blicher, 1839;
Dalgas, 1830; Aagaard, 1811). A botanical analysis in
the early years of the 20th century reported that in ®elds
uncultivated for 1, 4, 5, 6, and 10 years the frequencies
of Calluna vulgaris were already as high as 88, 96, 28,
Table 1
Frequency percentages for all vascular plant species found during the period 1976±1997

Year

Plant species 1976 1977 1978 1979 1980 1981 1982 1983 1984 1985 1986 1987 1988 1989 1990 1991 1992 1993 1994 1995 1996 1997

Achillea millefolium 5 4 7 18 8 11 19 16 19 14 14 17 15 11 9 7 5 9 10
Agrostis stricta 2 6 2 6 5 3 2 2 1
Agrostis tenuis 3 2 7 12 9 10 21 12 13 21 8 8 11 5 4 4 3 3
Aira praecox 1 1 2 2 3 1 1
Anthoxanthum odoratum 1 1 2 3 2 2
Anthyllis vulneraria 1 1 1 1 4 1 2
Apera spica-venti 8 18 12 3 1
Arabidopsis thaliana 10 2 4 1 1 1
Arenaria serpyllifolia 1 1

H.J. Degn / Biological Conservation 97 (2001) 319±330

Armeria maritima 1 1 4 2 2
Brassica cf. campestris 1
Calluna vulgaris 1 2 5 6 10 15 17 27 45 55 59 73 66 69 85 85 86
Campanula rotundifola 1
Capsella bursa-pastoris 1 5
Carex arenaria 5 9 7 14 15 12 10 9 4 9 7 10 6 2 1 2 4 4 1
Cerastium fontanum ssp. triviale 5 2 3 3 7 7 9 2 9 8 2 2 2 1 1 1 1 1
Chamaenerion angustifolium 1 2 2 1 2 3 2 2 3 1 2 6 2 2 2 2 5 4 3 4 5
Chenopodium album 22 26 2
Cirsium arvense 1
Conyza canadensis 1
Corynephorus canescens 2 6 5 5 4 7 3 3 2
Crepis capillaris 1
Dactylis glomerata 1 1 1 1
Deschampsia ¯exuosa 2 18 30 53 67 72 91 89 90 87 94 84 76 72 61 62 66 58 63 73
Elytrigia repens 62 92 96 99 99 94 45 15 16 11 3 3 1 1
Empetrum nigrum 1 1 2 1 3 2 3 5 1 1 5 3
Erigeron acer 1 10 14 12 6 11 10 5 4 1 1
Erodium cicutarium 4 8 5 8 1 1
Erophila verna 13 1
Erysimum cheiranthoides 3 2
Festuca ovina 2 3 1 7 6 8 19 29 34 54 60 42 71 76 65 81 85 71 91
Festuca rubra 1 4 6 15 30 25 24 29 25 24 21 20 18 11 3 5 2 8 5
Filago minima 18 1 9 2 1
Galium saxatile 1
Helichrysum arenarium 1 3 1 1 3 2 1 4 2 1 2 1 1 1 1 1 1 1
Hieracium pilosella 11 18 39 65 86 96 85 100 99 99 99 99 99 94 87 74 68 59 46 38 40
Hieracium umbellatum 1 1 1 3 8 8 7 4 11 7 5 7 1 2 2 1 2
Holcus mollis 1 3 5 7 9 7 6 4 2 4 3 1
Hypochoeris radicata 2 5 25 50 66 68 65 31 8 7 4 6 3 1 1
Jasione montana 13 62 81 31 40 33 39 26 16 14 13 11 28 27 17 13 19 27 34 12 18

323
(continued on next page)
 324
Table 1 (continued)
Year

Plant species 1976 1977 1978 1979 1980 1981 1982 1983 1984 1985 1986 1987 1988 1989 1990 1991 1992 1993 1994 1995 1996 1997

Knautia arvensis 2 1 1 1 2 1 1 1 1 2
Lapsana communis 1
Linaria vulgaris 3 2 10 20 25 15 2 6 2 1 2 1
Leontodon autumnalis 3 1 1
Lotus corniculatus 2 3 3 4 5 5 4 2 1 2 2 1 2 1 1 3
Luzula campestris 1 4 7 9 17 14 11 27 15 7 13 9 11 12

H.J. Degn / Biological Conservation 97 (2001) 319±330

Luzula multi¯ora ssp. multi¯ora 1 2 3 2 1
Myosotis arvensis 2
Plantago lanceolata 2 1 1 1 1 1
Poa pratensis 1 1 1 3 1 1 1 1
Polygonum aviculare 49 51 16 10 4 2 1
Polygonum convolvulus 50 54 15 24 8 1 1 3 1 1 1 2
Polygonum lapathifolium 1
Rhinanthus minor 1 1 1
Rumex acetosa 1
Rumex acetosella 2 2 30 65 78 80 35 9 14 10 9 15 2 2 5 3 2 3 7 6 2 3
Scleranthus annuus ssp. annuus 3 7 5 2 4 1 1 2
Scleranthus perennis 3 6 13 2 4 5 6 7 3 1 2 1 3 1 1
Senecio jacobaea 1 1
Senecio vernalis 4 39 26 7
Silene vulgaris 1
Solidago virgaurea 2 1 3 3 3 2 1 2 3 2 5 1 4
Spergula arvensis 30 14 10 1
Taraxacum sp. 3 1 2
Teesdalia nudicaulis 38 7 3 8 13 10 5 7 2 3 1 1 7 5 5 5 3
Thymus serpyllum 1 1 2 1 3 1 2 3 2 1 2
Trifolium arvense 1 58 62 12 79 15 19 72 7 2 1 7 1 1
Trifolium campestre 16 84 23 16 23 19 30 14 10 3 1 1 10 6 1 3
Veronica opaca 1 2 1 1
Vicia hirsuta 2 1
Vicia lathyroides 1
Vicia sativa ssp. angustifolia 1 1 3 5 3 1 1 5 1 1 1 1 5 1 2
Viola canina 1 1
Viola tricolor ssp. tricolor 16 23 7 1 2 1 2 1 1 2 4 1 1
 H.J. Degn / Biological Conservation 97 (2001) 319±330 325

Fig. 5. The frequency percentages of all 12 species with a frequency >50% at least in 1 year during the period 1976±1997. The species are placed
according to the year, when they had their maximum values. The time sequence is from the upper left corner downwards in the columns, ®nally to
end in the lower right corner.
 326
Table 2
TWINSPAN table for 18 most important species (mean frequency >5%). Numbers in the table are frequency percentages. The stages (A±G) and species groups (a±f) are those recognised by the
programme and described in the text. The ``trees'' beneath and to the right of the table indicate the relationships within the stages and within the species groupsa

H.J. Degn / Biological Conservation 97 (2001) 319±330

a
Key to the species: polcon, Polygonum convolvulus; polavi, Polygonum aviculare; elyrep, Elytrigia repens; rumacl, Rumex acetosella; triarv, Trifolium arvense; tricam, Trifolium campestre; jas-
mon, Jasione montana; agrten, Agrostis tenuis; hyprad, Hypochoeris radicata; fesrub, Festuca rubra; teenud, Teesdalia nudicaulis; achmil, Achillea millefolium; des¯e, Deschampsia ¯exuosa; hiepil,
Hieracium pilosella; carare, Carex arenaria; luzcam, Luzula campestre; calvul, Calluna vulgaris; fesovi, Festuca ovina.
 H.J. Degn / Biological Conservation 97 (2001) 319±330 327

mineral nutrients, these observations may point to an

explanation why the process formerly was faster.
The nutrient status of modern ®elds could also be the
reason why the process to-day does not always end with
dominance by Calluna vulgaris. Many recently aban-
doned ®elds in the Jutland landscape have developed
into grassland, frequently with tall herbs. This has also
been described by Holst (1987) and Mogensen (1994).
Grassland communities can develop even in close
proximity to stands of seed-producing Calluna vulgaris,
which rules out the lack of seeds as an explanation.

5.3. Consequences for restoration of heathland

Several attempts have been made to restore heathland

on abandoned agricultural land in more intensive ways.
Techniques include spreading harvested seed capsules,
litter, topsoil, turves, or even segments (1±2 m2) of
heathland (Gimingham, 1992; Pywell et al., 1995). To
improve soil conditions for heathland plants (Pywell et
al., 1994), turf-cutting (Smith et al., 1991) and even
acidi®cation of the soil (Owen et al., 1999) have been
used. The success has in several cases been questionable.
If a seed bank or adjacent seed source is present,
heathland can be restored on appropriate soils by uti-
lizing natural succession. This would have several
advantages:

. A more varied heathland vegetation can be the

Fig. 6. Values for some of the most important nutrients and pH in the
soil in the period 1976±1996.
64, and 100% respectively (Ferdinandsen, 1918). Bùr-
gesen and Jensen (1904) found Calluna vulgaris present
in the second year after the last crop of rye (Secale
cereale).
In addition, some descriptions indicated that the pro-
cess was faster on ®elds that had not been manured or
marled (Blicher, 1795; Ferdinandsen, 1918; BoÈcher,
1970). In one case, Ferdinandsen (1918) made analyses
in two neighbouring ®elds 10 years after abandonment.
In the marled ®eld Calluna vulgaris was found in 1% of
the samples compared to 88% in the unmarled ®eld. As
modern ®elds have a higher pH and content of major
result, as a greater variety of seeds may spread to
the area. Active application of seeds from Calluna
vulgaris will favour this species, as most other
heathland plants will have shed their seeds at the
time of ripening and harvest of Calluna vulgaris
seeds in October±November (Putwain and Rae,
1988; Pywell et al., 1996). If active application is
successful, it means that the stages with a high
species number (Fig. 2) and a high index of
diversity (Fig. 4) are left out, as the process moves
faster to a more uniform Calluna-dominated vege-
tation.
Also, at the landscape level, areas in di€erent
stages from farmland to heathland will contribute
to a higher variation.
. Current management rotates many heathlands in
the cycle: young Calluna vulgaris ! old Calluna
vulgaris ! (by management) ! young Calluna
vulgaris ! etc. In Jutland there is a historical tra-
dition of temporary ®elds scattered in the heath-
land and used only for a few years. They followed
the cycle: heathland ! farmland ! fallow land !
(by invasion of Calluna vulgaris) ! heathland !
etc.
. During the successional stages from farmland to
heathland a number of plants and animals, which
328 H.J. Degn / Biological Conservation 97 (2001) 319±330

Fig. 7. The area photographed towards the east, showing characteristic species. 1976: Polygonum convolvulus, Chenopodium album; 1979: Rumex
acetosella, Jasione montana; 1981: Trifolium arvense; 1987: Deschampsia ¯exuosa, Hieracium pilosella; 1995: Calluna vulgaris, Deschampsia ¯exuosa.

are not found either in the farmland or in the
heathland, ®nd a place to live for some years.
Some of these species are more rare than those of
both agricultural ®elds and heathland proper (in
this study e.g. Agrostis stricta, Filago minima and
Veronica opaca). A mosaic of communities or
successional stages of di€erent ages would ensure
biotopes for far more species than if the whole
landscape was covered by Calluna vulgaris.
. More subjectively it could be argued that nature
restoration should be done in a natural way, not
necessarily in the most ecient way. Focusing too
 H.J. Degn / Biological Conservation 97 (2001) 319±330
much on e€ectiveness could lead to a ``farming''
attitude, giving priority to Calluna vulgaris, or even
regarding Calluna vulgaris as a crop. Then success
would be to create cover of Calluna vulgaris as fast
and complete as possible.
6. Conclusion
The aim of heathland restoration in the region of
Calluna-heathland in northwestern Europe will most
often be dominance by Calluna vulgaris. The maximum
cover in the present study was not achieved after 22
years, but will occur in years to come.
This is not to say that succession will stop. In 5±10
years the oldest Calluna vulgaris plants will begin to die.
If it should be decided that the area should be domi-
nated by Calluna vulgaris, management will be neces-
sary (Gimingham, 1972, 1992).
Today the vegetation of the study area corresponds to
the heather-heath type (5.1.1.5) in the Scandinavian
classi®cation of vegetation types (PaÊhlsson, 1994). The
vegetation on the adjacent heathland, which bears no signs
of ever having been ploughed, corresponds to the heather±
crowberry±cowberry±heath type (5.1.1.3), because species
like Vaccinium vitis-idaea and bear-berry (Arctostaphylos
uva-ursi) are common. Their dispersal is very slow, and
propagation from seeds is rarely observed (Mentz, 1909;
Ritchie, 1955).
The results presented above show that heathland of
great value can be created on former arable land simply
by letting natural succession proceed for 22 years. But
the principle is recommended only in two situations.
The ®rst is where there are abandoned ®elds adjacent
to existing heathland. Such locations will get a high
priority, as it has been shown that large heathlands have
a higher quality than smaller ones (Webb and Hopkins,
1984; Webb and Vermaat, 1990). Large heathlands have
a lower diversity of plants and invertebrates, which is to
be preferred in this case as an increase in diversity indi-
cates a loss of heathland characteristics due to invasion
by non-heathland species. Marrs (1993) has also
recommended the creation of new heathland nature
reserves on land adjacent to existing ones, because cul-
tivated land in such areas is likely to be at the low end
of the agricultural fertility spectrum, and because it may
receive propagules from the nature reserve helping
colonization.
Secondly, natural succession could be used on small
areas in the heathland that are cultivated for a few years
and then abandoned again as in former times. There is a
risk in Denmark that ploughing of protected heathland
would be understood as destruction, because all heath-
lands >2500 m2 have been protected by law since 1992
precisely to prevent them being ploughed for farmland
or planted with trees. But the arguments in point 1±3
329
above are all in favour of the principle. It gives a greater
diversity of plants and animals on the area during the
succession, it gives a greater variation of the landscape,
and it is historically authentic. However, the principle
should only be used on a small part of an existing
heathland as formerly. Today further restrictions would
be added: only on soils which have been ploughed
before, and only when no special values were found on
the area cultivated. These two demands can be ful®lled
on many areas.
Acknowledgements
The Danish Air Force is thanked for permisson to
work on Air Station Karup, which is normally closed to
the public. I am very grateful to Assistant Professor
Henning Adsersen for highly valuable contributions
about statictics as well as many useful suggestions and
comments on the manuscript. I am also very grateful to
Professor Nigel Webb for many useful comments on
two earlier drafts of the manuscripts.
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