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Flora and lepidoptera fauna adversely affected by invasive Lupinus polyphyllus along road verges

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An increasing number of invasive species are changing ecosystems around the world. Road verges have commonly become the first footholds of non-native species in the new environments. Regularly mown road verges also offer habitats for meadow flora and fauna, which in Europe have suffered from the radical decline of semi-natural biotopes due to the agricultural modernization. We studied impacts of an invasive plant Lupinus polyphyllus on the plant and Lepidoptera species composition along road verges. The plant species composition was studied on 15 sites (with 1 m2 quadrats) and butterflies and diurnal moths along 15 transects (with weekly censuses) in SE Finland, each site and transect representing equally lupine invaded verge and an adjacent non-lupine verge. The species richness and diversity of flora and the cover and species richness of low growing (<20 cm) species, in particular, was lower in lupine verges compared to non-lupine verges. Also, the abundance of butterflies was lower in lupine verges. As the lupine cover approached 100%, fewer butterflies were observed in lupine transects compared to the adjacent non-lupine transects and a higher proportion of individuals were flying. Our results suggest that the changes in plant species assemblages and lower plant species richness in lupine invaded areas had “bottom-up” effects on higher trophic levels. Further studies on the control of lupine are urgently needed, but meanwhile we suggest regular mowing before the lupines have shed their seeds, together with the removal of the cuttings, to be the best management option.
Flora and lepidoptera fauna adversely affected by invasive
Lupinus polyphyllus along road verges
Anu Valtonen*, Juha Jantunen, Kimmo Saarinen
South-Karelia Allergy and Environment Institute, La
¨ritie 15, FIN-55330 Tiuruniemi, Finland
Article history:
Received 23 March 2006
Received in revised form
14 June 2006
Accepted 30 June 2006
Available online 17 August 2006
Alternative habitat
Diurnal moth
Non-native species
Plant species
An increasing number of invasive species are changing ecosystems around the world. Road
verges have commonly become the first footholds of non-native species in the new envi-
ronments. Regularly mown road verges also offer habitats for meadow flora and fauna,
which in Europe have suffered from the radical decline of semi-natural biotopes due to
the agricultural modernization. We studied impacts of an invasive plant Lupinus polyphyllus
on the plant and Lepidoptera species composition along road verges. The plant species
composition was studied on 15 sites (with 1 m
quadrats) and butterflies and diurnal moths
along 15 transects (with weekly censuses) in SE Finland, each site and transect representing
equally lupine invaded verge and an adjacent non-lupine verge. The species richness and
diversity of flora and the cover and species richness of low growing (<20 cm) species, in par-
ticular, was lower in lupine verges compared to non-lupine verges. Also, the abundance of
butterflies was lower in lupine verges. As the lupine cover approached 100%, fewer butter-
flies were observed in lupine transects compared to the adjacent non-lupine transects and
a higher proportion of individuals were flying. Our results suggest that the changes in plant
species assemblages and lower plant species richness in lupine invaded areas had ‘‘bottom-
up’’ effects on higher trophic levels. Further studies on the control of lupine are urgently
needed, but meanwhile we suggest regular mowing before the lupines have shed their
seeds, together with the removal of the cuttings, to be the best management option.
2006 Elsevier Ltd. All rights reserved.
1. Introduction
Species introductions have become more common due to the
increased transport, trade, travel and tourism, which provide
vectors and pathways permitting living organisms to cross
biogeographical barriers that would usually block their way
(Vitousek et al., 1996). Some of the non-native species become
invasive and have negative effects on native species, the econ-
omy and public health (Mack et al., 2000). From the ecological
perspective the invasive species can influence all ecosystem
levels from individuals and populations to communities and
ecosystem processes. Within biogeographical regions, the
replacement of local biota with non-indigenous, human intro-
duced species results in the homogenization of ecosystems
(McKinney and Lockwood, 1999). Road verges, in particular,
have often formed footholds for invasive plant species (Trom-
bulak and Frissell, 2000), and in some parts of the world the
majority of road verge flora comprises introduced species
(Wester and Juvik, 1983). Periodic disturbances common on
road verges may eliminate or reduce the cover of competitors
and/or increase resource levels and therefore facilitate inva-
sions (Davis et al., 2000).
Despite the harsh climate, considerable numbers of alien
plant species have spread and become established in the
0006-3207/$ - see front matter 2006 Elsevier Ltd. All rights reserved.
*Corresponding author: Tel.: + 358 5 432 8629; fax: + 358 5 432 8625.
E-mail addresses:, (A. Valtonen),, (J. Jantunen),, (K. Saarinen).
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Nordic countries (Jonsell, 2004). Plant species have arrived
both intentionally and unintentionally along with traditional
agricultural practices and by escaping from gardens into the
wild. Some garden introductions, such as the lupine (Lupinus
polyphyllus), are so competitive that they are considered a
threat to other plant species. The lupine originates in western
North America (Jalas, 1965). It had already been introduced in
central Europe when the species was described in 1827. Over
the last few decades the lupine has been recorded as an inva-
sive species in Britain and in central Europe, where it has both
increased in frequency and advanced its altitudinal limits
(Rich and Woodruff, 1996; Kowarik, 2003; Becker et al., 2005).
In Finland, the species had escaped into the wild in four
southern and western provinces by 1965 (Jalas, 1965). Two
decades later the species had spread almost 400 km north-
wards (Lahti et al., 1995). Nowadays, the lupine is spreading
rapidly along road verges and other disturbed habitats, but
there is a clear indication that the species can spread also
to semi-natural grasslands and natural environments such
as groves of trees (The Finnish Environment Institute, 2005).
Meadow species have found important refuges in regularly
mown verges (Jantunen et al., 2006) but lupine represents a
potential threat for generally low growing plants adapted to
nutrient-poor conditions. Construction work required on road
verges every 20–30 years (Mahosenaho and Pirinen, 1999)
disturbs vegetation and provides suitable areas for invasive
species. Lupines fix nitrogen and their litter fertilizes the
nutrient poor soil (Davis, 1991) which can alter competitive
interactions by enabling colonization by, and the rapid growth
of, tall species that shade and hinder low growing species and
thus reduce plant species richness (Hobbs and Huenneke,
1992; Maron and Connors, 1996; Gosling, 2005). The height
of lupine varies between 50 and 120 cm (Mossberg and Sten-
berg, 2005) and thus a dense lupine canopy can block the light
effectively from other species. Based on other Lupinus species
it is also possible that the lupine alkaloids hinder the germi-
nation of other plant species (Muzquiz et al., 2004). The plant
species on road verges, in turn, provide resources for a wide
variety of insects (Free et al., 1975; Munguira and Thomas,
1992; Eversham and Telfer, 1994; Saarinen et al., 2005). Thus,
changes in the plant communities are likely to have effects
on higher trophic levels (Knops et al., 1999). Globally, invasive
plant species pose a threat to the existence of some rare
butterfly and other insect species (Braby and Douglas, 2004;
Samways and Taylor, 2004), but may also offer resources for
native species in the form of host-plants or nectar (Graves
and Shapiro, 2003).
The objective of this study was to determine the potential
threat of lupine on plant and diurnal Lepidoptera communi-
ties on road verges. We compared the plant and Lepidoptera
species richness, cover, abundance, diversity and assem-
blages between road verges already invaded by lupine and
adjacent non-invaded verges. Plant, butterfly and diurnal
moth species were divided into ecological groups to test
whether meadow species and low growing plant species are
particularly vulnerable to lupine invasion. Furthermore, we
compared the vegetation characteristics important to the
Lepidoptera (i.e. vegetation height providing shelter for diur-
nal moths and nectar availability, mainly used by butterflies)
between the two groups. Finally, we studied the impact of the
lupine on butterfly and diurnal moth individuals by compar-
ing their behavior in lupine and non-lupine verges.
2. Materials and methods
2.1. Vegetation study
We studied effects of lupine on flora on 15 road verge sites in
SE Finland (Fig. 1). In each site, half of the area represented lu-
pine invaded verges, while the rest represented a non-lupine
verge with either no lupines or only sporadic stands. The two
parts were located adjacently on the same side of the road
and it was assumed that the lupine stands were random col-
onizations. On each site we established ten 1 m ·1 m quad-
rats, five on the lupine verge and five on the non-lupine
verge. The quadrats were placed symmetrically in the two
parts. The distance from quadrats to the pavement edge ran-
ged between 1 and 3 m, depending on the verge width, and
the distance between quadrats ranged between 3 and 20 m,
depending on the length of the study area. We surveyed the
plant species composition between 18 and 27 July 2005, before
the verges were mown. We estimated the abundance of plant
species in each quadrat with projection cover (percentage
scale 0–100%) and measured the average vegetation height.
In data analyses, we used average number and coverage of
species of the five quadrats in each site. Additionally, we
determined the total plant species richness in the five quad-
rats. Five species groups were analysed separately: positive
indicators of semi-natural grasslands (‘‘meadow species’’)
and negative indicators of semi-natural grasslands (‘‘weed
species’’) (mainly following Pyka
¨, 2001), low growing species
(average height <20 cm), medium height species (20–50 cm)
and tall species (>50 cm).
2.2. Lepidoptera study
We studied butterflies (Hesperioidea, Papilionoidea) and other
day-active Lepidoptera (Zygaenoidea, Lasiocampoidea, Bomb-
ycoidea, Geometroidea, Noctuoidea) on 15 road verge tran-
sects, ten of which coincided with the vegetation study sites
(Fig. 1). This was due to different criteria: the transects had to
be in sunshine at midday in order to conduct the Lepidoptera
Fig. 1 – Location of vegetation study sites and Lepidoptera
transects in SE Finland.
390 BIOLOGICAL CONSERVATION 133 (2006) 389396
counts, whereas the vegetation study sites could not be located
on road verges mown before 27 July 2005. Transects varied be-
tween 100 and 560 m in length (average 265 m). In each tran-
sect, the first half represented lupine invaded verge (average
cover of L. polyphyllus 86%) and the second half represented
non-invaded verge (average cover 4%). In 2005, we censused
14 transects weekly between late May (week 22) and late Au-
gust (week 35); one transect was censused during weeks 23–
35. At each site we walked through the transects and counted
all individuals within a 5 m ·5 msquare in front of the recorder
(Pollard and Yates, 1993). We also recorded the behavior (flying,
nectaring, basking, hiding) of each individual at first sight. All
behavior related to lupine was recorded separately.
We minimised the differences in censusing time, temper-
ature, wind speed and sunshine between the two parts of
each transect by counting the parts sequentially in varying or-
der. Starting times ranged between 9:50 and 16:15; all cen-
suses were carried out in good weather conditions, the
temperatures being P14 C, the median wind speed on the
Beaufort scale being 3 (gentle breeze), and the median sun-
shine percentage (estimated as 0%, 25%, 50%, 75% and 100%)
being 100%. Due to the adjacent location of the respective lu-
pine and non-lupine verges, the verge and road width, time
since the last soil disturbance, mowing regime, and surround-
ing environment were similar in the transect pairs. Vegeta-
tion height and nectar abundance on the study transects
were estimated once a month in June, July and August. Vege-
tation height was measured from the middle of the transect
and the average of three months was used in the statistical
analyses. Correspondingly, nectar abundance was estimated
once a month on an ordinal scale (1 = low, only few sporadic
nectar plant stands; 2 = moderate, sporadic stands through-
out the transect and 3 = high, nectar flowers abundant
throughout the transect) and the median of the three months
was used in the analyses. In addition, the nectar species rich-
ness was calculated as total number of plant species in flower
(except graminoids) recorded in three monthly evaluations.
In the data analyses, we compared the species richness
(total number of species observed) and abundance (sum of
individuals of all censuses) of butterflies and diurnal moths
separately between the lupine and non-lupine transects.
Summing up the weekly censuses captured the range of sea-
sonal activity in the Lepidoptera. Additionally, we divided the
butterflies into species typical of meadows (‘‘meadow spe-
cies’’), forest edges and clearings (‘‘forest edge species’’), and
fields, farmyards and wasteland (‘‘field margin species’’),
according to the classification by Pitka
¨nen et al. (2001). One
unclassified species Cupido argiades was included in the field
group of butterflies. Correspondingly, we separated the diur-
nal moths into meadow species and other species according
to Kuussaari et al. (2003). Four unclassified species Siona line-
ata,Thetidia smaragdaria,Cerapteryx graminis and Eriopygodes
imbecilla were included in the meadow group based on the pri-
mary habitats and host plants of their larvae.
2.3. Data analyses
We compared the paired samples of plant, Lepidoptera, vege-
tation height and nectar variables of the lupine verge and
non-lupine verge of each site with a non-parametric Sign test.
The Shannon’s diversity index of plant, butterfly and diurnal
moth communities calculated by Pc-Ord 4.0 (McCune and
Mefford, 1999) were accordingly compared. We calculated a
sequential Bonferroni correction for all test sets to lower the
risk of significant differences by chance, and used an error
rate of 0.10, as suggested by Chandler (1995). We also calcu-
lated a ratio of species richness and abundance of Lepidop-
tera in lupine vs. non-lupine verges and used it when
comparing how similar the adjacent lupine and non-lupine
verges were. The association between the ratio and the aver-
age cover of lupine in lupine verges was tested with Spearman
rank correlation. Sign tests and Spearman correlations were
performed by the program SPSS 12.0 for Windows.
Changes in composition of the plant species between the
lupine and non-lupine parts were compared by non-metric
multidimensional scaling (NMS), using Sørensen (Bray Curtis)
distance measure and the slow, through autopilot settings of
Pc-Ord 4.0 (McCune and Mefford, 1999). We further tested the
differences in the species assemblages of plants, butterflies
and diurnal moths in the lupine and non-lupine verges with
a non-parametric multi-response permutation procedure
(MRPP) using a Euclidean distance measure (Zimmerman
et al., 1985). Plant or Lepidoptera species preferring either lu-
pine or non-lupine verges were determined with an indicator
species analysis using the Monte Carlo test of significance
with 1000 runs (Dufrene and Legendre, 1997). Both MRPP
and indicator species analysis were performed using Pc-Ord
4.0 (McCune and Mefford, 1999).
3. Results
3.1. Effect of lupine on plant species
The vegetation was 39 cm taller, on average, and lupine cover
markedly higher, in lupine verges, while the cover of other
plant species was higher in non-lupine verges (Table 1). In lu-
pine verges the cover of L. polyphyllus ranged between 56%
and 78%, while only some lupines appeared in non-lupine
quadrats (cover range between 0 and 7%). A total of 139 plant
species were recorded in 15 sites (Supplementary data A).
Along with lupine, the plant species included 32 meadow spe-
cies, 23 weed species and 17 herbaceous forest species, the
rest being tree species (10) and various species of open envi-
ronments (56). The total species richness, the number of spe-
cies per quadrat (m
) and the species diversity were higher in
non-lupine verges (Tab le 1).
The cover of meadow species, weed species, low growing
and medium height species was significantly lower in lupine
compared to non-lupine verges, but no significant differences
emerged in tall growing species (Table 1). Furthermore, the
species richness of low growing species was significantly
lower in lupine verges, whereas the difference in meadow
species, weed species, medium height and tall growing spe-
cies was not significant.
Non-metric multidimensional scaling identified a two-
dimensional solution for the vegetation data (final
stress = 18.96; instability = 0.00001). The lupine verges were
tightly clustered, whereas the non-lupine verges were scat-
tered over the ordination space (Fig. 2). MRPP indicated a sig-
nificant difference between the lupine and non-lupine verges
BIOLOGICAL CONSERVATION 133 (2006) 389396 391
(T=19.6, p< 0.0005). According to the indicator species
analysis, one species was indicative of lupine verges (L. poly-
phyllus,p= 0.001), while five species were indicative of the
non-lupine verges (Trifolium pratense,p= 0.001; Trifolium re-
pens,p= 0.003; Leontodon autumnalis,p= 0.009; Fragaria vesca,
p= 0.012; Vicia sepium,p= 0.030).
3.2. Effect of lupine on lepidoptera
We recorded a total of 93 Lepidoptera species and 2344 indi-
viduals along the transects: 45 (48%) species and 1512 (65%)
individuals of butterflies and 48 (52%) species and 832 (35%)
individuals of diurnal moths (Supplementary data B). The
majority of diurnal moths were geometrid moths (21 spe-
cies/472 individuals) and noctuid moths (16/324).
We found no differences in the species richness or diver-
sity of Lepidoptera between the lupine and non-lupine verges
(Tab le 2 ). However, the abundance of all individuals and but-
terfly individuals was significantly higher in non-lupine
verges compared to lupine verges. There was a mid-summer
peak of abundance both in butterflies and diurnal moths. In
non-lupine verges, the abundance of butterflies was higher
throughout the summer, whereas diurnal mothswere slightly
less abundant during the peak flight period (weeks 25–27) and
more abundant during the latter half of the study period.
There was a tendency towards higher nectar abundance in
non-lupine verges. No differences were found in diurnal moth
numbers between lupine and non-lupine verges, yet the veg-
etation was significantly higher in lupine verges.
The cover of L. polyphyllus on lupine verges was inversely
associated with the ratio of butterfly individuals in lupine
vs. non-lupine verges (Spearman rank correlation;
=0.520, p= 0.046), i.e. the higher the cover of lupine the
lower the amount of butterfly individuals found on the lupine
verges compared to the adjacent non-lupine verges. No asso-
ciation was found between the lupine cover and ratios in but-
terfly species richness (r
=0.190, p= 0.499) or abundance
=0.038, p= 0.893) and species richness (r
p= 0.337) of diurnal moths.
According to the primary habitat groups of butterflies, 61%
of all recorded individuals represented meadow species, 30%
were forest edge species, and 9% belonged to field margin
species. Correspondingly, 85% of all diurnal moth individuals
belonged to species typical to meadows. Species richness in
meadow butterflies and the abundance of both meadow but-
terflies and diurnal moths were higher in the non-lupine
verges compared to the lupine verges, but the differences
were not significant after the Bonferroni correction (Table 2).
There was little difference in the proportions of the three
primary habitat groups of butterflies between the lupine
(62% meadow/27% forest edge/11% field) and non-lupine
(59%/32%/9%) verges. Correspondingly, the proportions of
meadow diurnal moths were almost the same in the lupine
(83%) and non-lupine verges (86%). The MRPP analysis indi-
cated a non-significant difference in species assemblages be-
tween the two groups for both butterflies (T= 0.358, p= 0.508)
and diurnal moths (T=0.514, p= 0.241). The only species
with a significant indicator value was the burnet moth Zyga-
ena viciae (p= 0.024), which was observed only in the non-
lupine transects.
Butterfly behavior was similar in lupine and non-lupine
verges and included flying (62% of individuals in lupine
verges/65% of individuals in non-lupine verges), nectaring
Fig. 2 – NMS ordination of the vegetation data showing
sample scores of the two axes. The lupine and non-lupine
verges of each study site are connected with a line. The
species scores of Lupinus polyphyllus and 19 other species
positively or negatively affected by lupine are overlayed. The
names of species are abbreviated to the first four letters (see
full names in Supplementary data A).
Table 1 – Differences in vegetation between the lupine
and non-lupine verges; mean, standard deviation (SD)
and significance of Sign test are reported
Lupine Non-lupine Sign test
Mean SD Mean SD p
Vegetation height (cm) 77.9 16.3 38.7 14.2 <0.0005
Total species richness 26.9 6.1 34.7 11.0 0.006
11.7 2.6 15.4 3.9 0.001
Shannon’s diversity index 1.5 0.2 2.6 0.4 <0.0005
Cover of lupine (%) 69.1 7.7 1.4 1.9 <0.0005
Cover of other species (%) 38.1 9.2 97.3 11.2 <0.0005
Meadow species
Total species richness 4.7 2.5 5.8 3.2 0.180
Cover 2.1 1.9 12.7 12.1 0.001
Weed species
Total species richness 5.2 3.2 6.0 3.4 0.549
Cover 7.7 7.3 19.2 19.1 0.002
Low growing species
Total species richness 7.1 2.7 12.1 5.0 <0.0005
Cover 12.6 5.2 35.6 13.5 <0.0005
Medium height species
Total species richness 10.4 3.1 11.7 4.2 0.180
Cover 10.6 3.9 32.6 15.7 <0.0005
Tall species
Total species richness 7.6 3.2 8.7 3.3 0.423
Cover 14.4 7.7 26.9 20.7 0.118
* Difference is significant after Bonferroni correction.
392 BIOLOGICAL CONSERVATION 133 (2006) 389396
(21%/20%), and basking (17%/15%). No egg laying was ob-
served. The higher the cover of lupine in lupine verges the lar-
ger the proportion of individuals flying (Spearman rank
correlation, r
= 0.518, p= 0.048) and the smaller the propor-
tion of nectaring individuals (r
=0.690, p= 0.004) (Fig. 3).
In only one case (which was a lupine verge) the proportion
of individuals nectaring was higher than the proportion of
individuals in flight. In the non-lupine verges the proportion
of individuals in flight ranged between 47% and 84% and the
proportion of individuals nectaring ranged between 0% and
37%. Eight butterfly individuals of the species Gonepteryx
rhamni,Callophrys rubi,Nymphalis io,Polyommatus icarus and
Aporia crataegi were observed visiting the flowers of lupine
once or more. Butterfly visits to lupine flowers represented
6% of all flower visits on lupine verges (n= 129) recorded dur-
ing the study and individuals basking on lupine represented
5% of all basking butterfly individuals on lupine verges
(n= 103) during the study.
Diurnal moth behavior included hiding in the vegetation
(83% of individuals in lupine verges/86% of individuals in
Table 2 – Differences in Lepidoptera numbers and vegetation characteristics important to Lepidoptera; the total, average
(with standard deviation SD), or frequency is given and the significance of Sign test is reported
Lupine Non-lupine Sign test p
Species richness (total) 75 76 0.581
Butterfly species richness (total) 35 42 0.424
Diurnal moth species richness (total) 40 34 0.607
Abundance (total) 1009 1335 0.007
Butterfly abundance (total) 613 899 0.007
Diurnal moth abundance (total) 396 436 0.057
Shannon’s diversity index
Butterflies (average ± SD) 1.9 ± 0.5 2.0 ± 0.5 1.000
Diurnal moths (average ± SD) 1.6 ± 0.5 1.7 ± 0.3 0.607
Meadow species richness
Butterflies (total) 14 17 0.035
Diurnal moths (total) 19 14 0.774
Meadow species abundance
Butterflies (total) 383 532 0.013
Diurnal moths (total) 347 389 0.035
Vegetation height, cm (average ± SD) 55.0 ± 17.5 36.0 ± 15.2 <0.0005
Nectar abundance (1/2/3) (frequency) 6/8/1 2/9/4 0.016
Species richness of nectar plants (total) 73 79 0.118
* Difference is significant after Bonferroni correction.
Fig. 3 – The percentage of individuals flying or nectaring corresponding to the lupine cover of the lupine transects.
BIOLOGICAL CONSERVATION 133 (2006) 389396 393
non-lupine verges), flying (13%/10%), nectaring (2%/2%), and
basking (2%/2%). One individual of Euclidia glyphica was ob-
served visiting the flowers of lupine, which represented 14%
of all flower visits (n= 7) of diurnal moths on lupine verges.
4. Discussions and conclusions
4.1. Lupine decreases the plant species richness
The invasion of lupine decreased the available area and cover
of other plant species, which also lowered the plant species
richness and diversity. In lupine verges, the total species rich-
ness was almost eight species less than in the non-invaded
verges, on average. Although the invasion of nitrogen-fixing
plants may increase the originally low species richness on
barren soils by encouraging further invasive species to settle
(Vitousek and Walker, 1989), we found no such effect. The
cover of other non-indigenous species was very low, including
only Artemisia campestris and Melilotus albus (present on one
non-lupine site).
The higher vegetation on lupine verges suggests that the
lupine has filled the unused space above the original verge
vegetation and the low growing species have been outcom-
peted due to the shading. Since low growing species de-
creased in both species richness and cover and medium
height species only in cover, whereas tall species were not sig-
nificantly affected, shading rather than other characteristics
was the most likely cause of the decline. Yet, the majority
of the meadow species (56%) recorded in the sites were low
growing, a typical adaptation to regular mowing or grazing
(Grime, 2001). Weed species included a smaller proportion of
low growing species (27%), but in both groups the cover was
smaller in lupine verges. Three of the five indicator species
of non-lupine verges were low growing (F. vesca, L. autumnalis
and T. repens), the other two being of medium height. In addi-
tion, according to the indicator species analysis and the NMS
ordination, low and medium height nitrogen fixing plants
(T. pratense,T. repens,T. hybridum and V. sepium) were particu-
larly decreased by the lupine.
By contrast, tall growing species such as Phleum pratense,
Angelica sylvestris and Alopecurus pratensis, located close to lu-
pine verges in the NMS ordination, can compete more effi-
ciently with lupine for light. Half of the weed species (50%)
were tall growing and many of the recorded weed species
are likely to benefit from the nitrogen enhancement by lupine
(Ellenberg et al., 1991). Lupinus species are commonly used in
land reclamation processes, but the problem has been their
vigorous growth and the effects of nitrogen transferred to
other fast growing species, which often become dominant
(Bradshaw, 2000).
4.2. Lupine decreases the lepidoptera abundance
Lupine did not provide food resources for Lepidoptera. By con-
trast, it decreased the cover and species richness of potential
host plants for larvae and nectar plants for adults. The de-
creased quantity and quality of resources both adversely af-
fect the butterflies (Summerville and Crist, 2001). We
recorded, on average, 22 Lepidoptera individuals (19 butterfly
individuals) less in lupine transects compared to the adjacent
non-lupine transects. As this pattern remained throughout
the summer, individuals in flight during different times of
the summer and different phases of the lupine growth, from
vegetative growth in early summer to flowering in mid sum-
mer and withering in late summer, were all adversely
In congruence with Knops et al. (1999), our results reveal
how the loss of biodiversity in basal species (plants) in grass-
land ecosystems can impact the insect communities. For but-
terflies invasions of non-native species have been found to be
alternatively insignificant (Fleishman et al., 2005), positive if
the species have been able to switch their hosts or use the
invasive species as nectar sources (Graves and Shapiro,
2003), or negative if females oviposit on introduced plants
but the larvae are unable to complete development on the
new host plants (Schlaepfer et al., 2005). We found no evi-
dence of Lepidoptera using lupine as a host plant. Instead,
as the cover of lupine approached 100%, fewer individuals ap-
peared to be interested in the vegetation and were merely fly-
ing. On the other hand, no egg laying was observed on any
plant species growing on road verges, probably due to the
short observation period of each individual.
The butterflies actively used the nectar resources on road
verges. Nectar is an important resource for butterflies by
increasing both longevity and fecundity (Murphy et al.,
1983). In lupine invaded verges the butterflies found nectar
from plants growing in the gaps within the lupine stands,
explaining why the nectaring decreased as the cover of lupine
approached 100%. The discrepancy between the high cover of
lupine and the few visitors to lupine flowers on lupine verges
suggests that the lupine is a poor nectar plant or does not
serve nectar, and the few butterfly and moth visits to its flow-
ers were abortive attempts at obtaining nectar.
Nectar has a marked influence on the microdistribution of
butterflies in their habitats (Loertscher et al., 1995), but no
such effect on diurnal moths was found. Many of these spe-
cies do not visit flowers as adults, yet the only Lepidoptera
indicator of non-lupine sites, the burnet moth Z. viciae, visits
flowers actively. The recorded diurnal moths form a contin-
uum of species flying solely during the day to species, which
fly mostly as a result of being disturbed and are more active at
night. Therefore tall lupine stands provided hiding places for
the less active diurnal moths during the day. The lupine can-
opy grows tall in the early summer and provides more hiding
places compared to the adjacent lower vegetation. The rapid
withering of lupine explains the decline of diurnal moths on
lupine verges in the late summer.
4.3. How to control the spread of lupine?
Although the lupine may have positive effects on a limited
range of species, e.g. bumblebees and other insects collecting
lupine pollen, its uncontrolled spread possess a real threat to
meadow plant and Lepidoptera species living on road verge
environments. These species were common on semi-natural
grasslands until the modernization of agriculture during the
20th century.As only fragments of semi-natural grasslands still
remain, many of their species are more often found on alterna-
tive habitats, in particular road verges under regular manage-
ment (Jonsell, 2004; Jantunen et al., 2006). Several studies
394 BIOLOGICAL CONSERVATION 133 (2006) 389396
have addressed the possibility of improving the management
of road verges to increase their suitability for the species of
semi-natural grasslands (Persson, 1995; Schaffers, 2000), but
to our knowledge studies concerning the management of road
verges invaded by non-indigenous plant species are lacking.
To constrain the impact of non-indigenous species, eradi-
cation, or if this is not possible, maintenance control of spe-
cies at acceptable levels, should be organised (Mack et al.,
2000). In New Zealand several plant pathogens which could
serve as biological control agents for the invasive L. polyphyllus
have been identified (Harvey et al., 1996). Mechanical and
manual removal techniques coupled with the removal of litter
and duff have been successful in the control of another inva-
sive species, Lupinus arboreus, but even when the population
of the invasive species has been successfully controlled the
fertilised soil may lead to changes in the plant species assem-
blages (Pickart et al., 1998a,b; Maron and Jefferies, 2001).
In unproductive environments, such as on road verges, an
increase in disturbance and reduction in productivity are
likely to be the best management plan for the control of inva-
sive species (Huston, 2004). We thus suggest that for as long
as studies on the control of L. polyphyllus are lacking, the con-
trol should be based on two mowings in mid and late summer
to prevent the re-flowering, with removal of the cuttings, on a
regular basis. In our study sites the lupine was often missing
in the proximity of the road under more frequent mowing
than the rest of the verge. This indicates that the species is
not well adapted to mowing, losing a large part of its biomass
during mowing and eventually withering away. Regular mow-
ing also prevents other tall species, including many weeds,
possibly invading in the wake of lupine. Unfortunately, the
intensive mowing is harmful to butterflies (Erhardt, 1985),
but these will benefit in the long run via more diverse vegeta-
tion. Eradicating the lupine by mowing may take several
years, but this is not a problem on road verges, which are
mown in any case to ensure traffic safety. In addition, manual
eradication on or close to sites where rare or endangered spe-
cies occur may be needed. The removal of cuttings, on the
other hand, can prevent the fertilising effect of lupine on
the soil. Finally, as the lupine has imposing flowers, it is pos-
sible that some people deliberately spread lupine seeds for
decorational purposes along road verges. Thus, greater public
awareness of the effects of lupine and other invasive species
is needed.
This study was financially supported by the Finnish Road
Administration, Finland’s Ministry of the Environment, Maj
and Tor Nessling Foundation and The Finnish Cultural Foun-
dation’s South Karelia Regional Fund. We also thank Leigh
Plester for the English revision and Heikki Roininen, Sanna
Saarnio, Mikko Kuussaari, Juha Po
¨yry and the anonymous ref-
erees for providing valuable comments and suggestions.
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... According to its wide distribution and local dominance, L. polyphyllus is recognized as a globally invasive plant species (Daehler, 1998;Valtonen et al., 2006). The non-native, synanthropic distribution range comprises roughly seven main areas of different size on all continents except Africa and Antarctica. ...
... From here westwards via Sverdlovsk, Arkhangelsk, and Murmansk to Scandinavia (Figs. 3B, 3C-4). In Europe, L. polyphyllus is specifically considered invasive in Norway (Fremstad, 2010), Sweden (Tyler et al., 2015), Finland (Valtonen et al., 2006), Lithuania (Vyšniauskiené et al., 2011), Germany (Kowarik, 2003), the Czech Republic (Hejda et al., 2009), Switzerland (InfoFlora, 2020, Ukraine, and Central Russia (Vinogradova et al., 2009). Due to historic and recent propagation for soil improvement at roadsides and in coniferous forests (e.g., Volz, 2003), the European area has become the largest range part worldwide. ...
... Sunhalf-shade (5) Drymesic (3) Moderately acidic strongly acidic (2) Very N-poormoderately N-poor (2) Not salt tolerant (1) Sweden Tyler et al. (2021) Sunhalf-shade (7) Drymoist (5) Weakly acidic -acidic (4) (8) Moist (5) Weakly acidic -acidic (4) Intermediate -N-rich (6) Not salt-tolerant (0) Italy Domina et al. (2018) Wissman et al. (2015) road verges and in the road network (J4.2) (Valtonen et al., 2006;Ramula and Pihlaja, 2012;Wissman et al., 2015), but it may also occur in low and medium altitude hay meadows (E2.2) (species-rich Fenno-Scandian boreal and sub-boreal meadows) and abandoned pastures and fields (Rolf Lutz Eckstein, personal observations). ...
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The invasive herb Lupinus polyphyllus has been focus of a number of fact sheets worldwide but a comprehensive summary of the species’ taxonomy and morphology, distribution, habitat requirements, and biology has been lacking. This paper gives a thorough account of the species’ systematic position and taxonomy, highlighting the difficulties to delimit taxa, which is related to interbreeding among members of this genus. However, L. polyphyllus var. polyphyllus is apparently the taxon that has naturalized and is regionally invasive in temperate-humid climates worldwide. We also present an updated distribution map of L. polyphyllus in the native and invaded ranges, which highlights seven regions in the world where the species has been established. We show that the climatic niche of L. polyphyllus in the invaded range shifts towards higher summer precipitation and lower isothermality, probably because the invaded range includes subcontinental regions of eastern Europe and western Siberia. The habitats of L. polyphyllus range from rather dry to wet, have moderately acidic to strongly acidic soils, and the species’ indicator values across Europe suggest that it occurs along a gradient from very nutrient poor sites to intermediate to rich sites from northern to southern Europe. The species shows high resistance to both drought and frost. In Central Europe, the species has a stronghold in alpic mountain hay meadows, abandoned meadows and pastures, low and medium altitude hay meadows, anthropogenic herb stands and temperate thickets and scrubs. In northern Europe, the species occurs in anthropogenic herb stands along roads and railroads as well as in abandoned pastures and fields. We also found some doubtful information about L. polyphyllus in the literature. This refers to its description as “rhizomatous perennial” although it lacks rhizomes; an apparently very high longevity of its seeds, which may only be true under artificial conditions in an ex situ seed repository; and a very deep rooting depth, which may not represent the average rooting depth but rather an extreme value. Knowledge about the interrelationships between the species’ future population dynamics and spread and ongoing climate warming is lacking. Finally, our review points out that there is currently no evidence-based strategy for a cost-efficient management of L. polyphyllus although it is among the most problematic non-native plant species in Europe due to its environmental and socio-economic impacts.
... This species typically flowers relatively early and invades different habitats ranging from road verges to forest understories and different types of mountain meadows (Wissman et al. 2015;Ramula 2017;Klinger et al. 2019). Dominant stands of L. polyphyllus have significantly negative effects on local plant diversity (Valtonen et al. 2006;Hejda et al. 2009;Ramula and Pihlaja 2012), plant community composition (Hansen et al. 2021), and arthropod abundance (Valtonen et al. 2006). In contrast to other invasive neophytes in Europe, L. polyphyllus can, in particular, colonise nutrient-poor habitats of high nature conservation value (Hejda et al. 2009). ...
... This species typically flowers relatively early and invades different habitats ranging from road verges to forest understories and different types of mountain meadows (Wissman et al. 2015;Ramula 2017;Klinger et al. 2019). Dominant stands of L. polyphyllus have significantly negative effects on local plant diversity (Valtonen et al. 2006;Hejda et al. 2009;Ramula and Pihlaja 2012), plant community composition (Hansen et al. 2021), and arthropod abundance (Valtonen et al. 2006). In contrast to other invasive neophytes in Europe, L. polyphyllus can, in particular, colonise nutrient-poor habitats of high nature conservation value (Hejda et al. 2009). ...
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Plant phenology, i. e. the timing of life cycle events, is related to individual fitness and species distribution ranges. Temperature is one of the most important drivers of plant phenology together with day length. The adaptation of their phenology may be important for the success of invasive plant species. The present study aims at understanding how the performance and the phenology of the invasive legume Lupinus polyphyllus vary with latitude. We sampled data across a >2000 km latitudinal gradient from Central to Northern Europe. We quantified variation in phenology of flowering and fruiting of L. polyphyllus using >1600 digital photos of inflorescences from 220 individual plants observed weekly at 22 sites. The day of the year at which different phenological phases were reached, increased 1.3-1.8 days per degree latitude, whereas the growing degree days (gdd) required for these phenological phases decreased 5-16 gdd per degree latitude. However, this difference disappeared, when the day length of each day included in the calculation of gdd was considered. The day of the year of the earliest and the latest climatic zone to reach any of the three studied phenological phases differed by 23-30 days and temperature requirements to reach these stages differed between 62 and 236 gdd. Probably, the invasion of this species will further increase in the northern part of Europe over the next decades due to climate warming. For invasive species control, our results suggest that in countries with a large latitudinal extent, the mowing date should shift by ca. one week per 500 km at sites with similar elevations.
... The cover of lupine varied from 1 to 30% (Table 2), which is a rather low level. It has been recorded that in Central Russia, as well as in higher latitudes in Finland, lupine is capable of forming monodominant stands [8,37]. ...
... The cover of lupine varied from 1 to 30% (Table 2), which is a rather low level. It has been recorded that in Central Russia, as well as in higher latitudes in Finland, lupine is capable of forming monodominant stands [8,37]. All of the studied plant communities (Table 2) were located along dirt roads within settlements and suffered major anthropogenic disturbances. ...
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... Tento problém je široce sledován a diskutován (Chytry et al. 2008, Pysek et al. 2009), přesto existuje jen minimum studií, zaměřených na analýzu dopadů invazních rostlin na motýly a jiný hmyz (např. Valtonen et al. 2006, Hanula & Horn 2011. Porosty trnovníku akátu (Robbinia pseudoacacia) eliminují přirozený vegetační kryt, s nímž pak mizí i cenné druhové spektrum ogranismů (např. ...
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... Consequently, the protection of endemic biodiversity is tightly linked to the preservation of native natural habitats, which are highly threatened by climate change, fragmentation, urbanization, and other landscape modifications, including plantations of non-native flora and the introduction of alien fauna. The latter is facilitated by globalization and the multifaceted transportation of diverse goods and merchandise onto these islands [49,[93][94][95]. ...
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Abstract: This work presents the first outcomes resulting from a DNA barcode reference library of lepidopteran species from Malta. The library presented here was constructed from the specimens collected between 2015 and 2019 and covers the genetic barcodes of 146 species (ca. 25% of lepidopterous Maltese fauna), including four newly recorded Lepidoptera species from the Maltese islands: Apatema baixerasi, Bostra dipectinialis, Oiketicoides lutea, and Phereoeca praecox. The DNA reference barcode library constructed during this study was analyzed in conjunction with publicly available DNA barcodes and used to assess the ability of the local DNA barcodes to discriminate species. Results showed that each species occupies a different BOLD BIN; therefore, DNA barcoding was able to discriminate between the studied species. Our data led to the formation of 12 new BOLD BINs—that is, OTUs that were identified during this work—while nearly 46% of the barcodes generated during this study were never recorded on conspecifics, further indicating the uniqueness of genetic diversity on these central Mediterranean islands. The outcomes of this study highlight the integrative taxonomic approach, where molecular taxonomy plays an important role for biodiversity investigation in its entirety.
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Introducción y objetivos: El cambio climático, el ganado y el creciente uso turístico favorecen la dispersión de plantas exóticas, amenazando la conservación de los ecosistemas altoandinos. Estudiamos los patrones de distribución de plantas nativas y exóticas en senderos recreativos de montaña. M&M: Implementamos el protocolo MIREN en seis senderos (2400-3600 m s.n.m.) en dos áreas protegidas de los Andes centrales de Mendoza en las Cordilleras Frontal y Principal. Resultados: Encontramos 180 especies nativas y 41 exóticas. La riqueza de especies nativas fue máxima a elevaciones intermedias, mientras que la riqueza de exóticas disminuyó con la elevación. La riqueza regional de nativas fue mayor en la Cordillera Frontal que en la Principal (114 versus 71 nativas, respectivamente) mientras que la riqueza regional de exóticas fue menor en la Frontal que en la Principal (20 versus 28 exóticas, respectivamente). La riqueza de exóticas por parcela fue mayor en la Cordillera Frontal que en la Principal. El rango de distribución altitudinal de exóticas fue mayor en la Cordillera Frontal. Dos exóticas abundantes, Taraxacum officinale y Cerastium arvense, estuvieron a lo largo de todo el gradiente en la Cordillera Frontal, alcanzando los 3600 m s.n.m. Encontramos siete exóticas no citadas anteriormente. Conclusiones: Los nuevos registros amplían los rangos de distribución conocidos para algunas especies exóticas. A pesar de que el número de especies exóticas fue similar cerca y lejos de los senderos, cinco especies sólo estuvieron en los bordes de los mismos, lo que sugiere que los senderos favorecen los procesos de invasión.
The article presents experimental data on the species composition of fungal and viral pathogens in the conditions of the secondary range of Lupinus polyphyllus . Tobacco mosaic virus, Bean yellow mosaic virus, Bean common mosaic virus and Pea enation mosaic virus were diagnosed on Lupinus polyphyllus for the first time. The issues related to the peculiarities of the adaptability of viruses to invasive plant species are discussed. The preventive role of vectors ( Aphididae ) in the expansion of pathogens and the widening of the spectrum of host plants (susceptible species) is emphasized. Interaction with vectors, including their non-specific species, is one of the mechanisms of virus adaptability, their expansion into new regions and the formation of new pathosystems with invasive plant species. It is concluded that based on the analysis of trophic connections of vectors, it is possible to prognosticate the search for the most effective variants of harmful organisms for the biocontrol of L. polyphyllus .
This datasheet on Lupinus polyphyllus covers Identity, Overview, Distribution, Dispersal, Hosts/Species Affected, Diagnosis, Biology & Ecology, Environmental Requirements, Natural Enemies, Impacts, Uses, Prevention/Control, Further Information.
Technical Report
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• Det finns minst 190 000 hektar gräsmarker som sköts i anslutning till infrastrukturer, varav 164 000 hektar finns i vägkanter. • Det finns minst 240 000 hektar buskmarker som röjs regelbundet i miljöerna, varav majoriteten finns uteslutande i kraftledningsgator. • Möjligheterna att använda infrastrukturmiljöer i det praktiska naturvårdsarbetet uppmärksammas alltmer, men fler analyser behövs framför allt av vilka biologiska kvaliteter markerna rymmer.
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This paper presents a new and simple method to find indicator species and species assemblages characterizing groups of sites. The novelty of our approach lies in the way we combine a species relative abundance with its relative frequency of occurrence in the various groups of sites. This index is maximum when all individuals of a species are found in a single group of sites and when the species occurs in all sites of that group; it is a symmetric indicator. The statistical significance of the species indicator values is evaluated using a randomization procedure. Contrary to TWINSPAN, our indicator index for a given species is independent of the other species relative abundances, and there is no need to use pseudospecies. The new method identifies indicator species for typologies of species releves obtained by any hierarchical or nonhierarchical classification procedure; its use is independent of the classification method. Because indicator species give ecological meaning to groups of sites, this method provides criteria to compare typologies, to identify where to stop dividing clusters into subsets, and to point out the main levels in a hierarchical classification of sites. Species can be grouped on the basis of their indicator values for each clustering level, the heterogeneous nature of species assemblages observed in any one site being well preserved. Such assemblages are usually a mixture of eurytopic (higher level) and stenotopic species (characteristic of lower level clusters). The species assemblage approach demonstrates the importance of the 'sampled patch size,' i.e., the diversity of sampled ecological combinations, when we compare the frequencies of core and Satellite species. A new way to present species-site tables, accounting for the hierarchical relationships among species, is proposed. A large data set of carabid beetle distributions in open habitats of Belgium is used as a case study to illustrate the new method.
The heart of the PC-ORD system is a group of Fortran programs for multivariate analysis on the MS-DOS family of microcomputers. Analytical features include ordinations (detrended correspondence analysis, Bray-Curtis ordination, principal components analysis, and reciprocal averaging), descriptive statistics, and diversity indices. -from Author
We conducted an experimental study to assess the effects of habitat fragmentation on patch use by butterflies and skippers. Five fragmentation treatments (20%, 40%, 60%, 80%, and 100% habitat remaining) were replicated five times by selectively mowing 15 × 15 m patches of an old field. Community and species responses to fragmentation were tested against several alternative predictions: (1) the proportional area hypothesis, in which species richness or abundance declines linearly with habitat area, (2) the patch heterogeneity hypothesis, in which richness or abundance peaks at intermediate levels of fragmentation, (3) the critical threshold hypothesis, in which a nonlinear decrease in richness or abundance occurs below some critical level of habitat loss, and (4) the patch quality hypothesis, in which declines in richness or abundance with increasing habitat loss are partially offset by the resource value of the remnant habitat. We found that fragmentation treatment significantly affected species richness and total number of patch visits for the overall community, as well as the numbers of patch visits by individual species. Community and species responses were linearly related to the amount of habitat remaining, as predicted by the proportional area model of fragmentation. We also found that plots with a higher cover of flowering forbs received greater patch use. Thus, the effects of habitat quantity (patch area) and habitat quality (flower cover) were independent and additive, suggesting that moderately sized patches of high resource quality may be equivalent to large patches of lower quality. Therefore, to maintain local population densities of Lepidoptera, it may be possible to compensate for losses in habitat area with improvements in habitat quality. Species did not uniformly respond to fragmentation. Rare species were disproportionately affected by fragmentation and did not occur in patches with
Species-rich grasslands that become enriched with nitrogen often suffer decreases in species richness, increases in plant biomass, and invasion by weedy exotic species. Suitable techniques to restore enriched grasslands and reestablish native communities are increasingly needed. Here we report results of a 5-yr experiment in enriched coastal prairie grasslands (Bodega Marine Reserve, Bodega Bay, California, USA), to determine the combined effects of mowing and biomass removal on total soil nitrogen, net rates of mineralization, nitrogen retention, and species richness and biomass. We mowed and removed plant biomass from plots in areas where the N-fixing shrub, bush lupine (Lupinus arboreus), had greatly enriched the soil, and where the community was composed of weedy introduced plants. Our goal was to facilitate the establishment of the native grassland assemblage such as was found at nearby low soil nitrogen sites. Mowing and biomass removal resulted in a dramatic change in the species assemblage, from exotic annual grasses to a mixed exotic/native forb community composed primarily of perennials. Species richness was significantly greater in treated plots than in control plots; weedy exotic grasses diminished in abundance, and both native and exotic forb species increased. In mowed vs. control plots, there was significantly less mean aboveground biomass, but significantly greater belowground biomass. This shift in species composition had significant impacts on nitrogen retention. In late fall and winter when plant-available N was highest, much nitrogen leached from the effectively fallow control plots where germination of annual grasses did not peak until midwinter. In contrast, mowed plots retained substantially greater amounts of nitrogen, due to the presence of perennial plants possessing large amounts of belowground biomass early in the season. Despite the cumulative removal of 22 g N/m2 in biomass over 5 yr, there was no difference between mowed and control plots in total soil N, pool sizes of inorganic N, or net rates of N mineralization. The results indicate that removal of plant biomass by mowing shifted this plant community from an annual grass to a perennial forb assemblage. However, in doing so, N retention by vegetation was increased, making it more difficult to reduce soil N.
Roadside plant communities on the windward slope of Mauna Loa volcano were sampled over an altitudinal range from 500 to 2500 m, and within a zone 3 m wide along the roadway. A total of sixty-nine plant species were found to inhabit the roadside verge. Introduced plant species comprised 74% of the roadside flora. Species with temperate affinities gradually replaced those of tropical origin at elevations above 1000 m. Overall species diversity decreased with elevation. Within the roadside vegetation a distinctive zonation was found to occur with distance from the roadway. Introduced plant species were dominant on the fine-grained road-fill substrate abutting the roadway, while native plant species occupied adjacent, recent lava flow substrates. There was little spatial overlap between these distinctive communities.
(1) In the subalpine region of Central Switzerland (Tavetsch Valley) studies were made of the day-active Lepidoptera faunas in different types of cultivated grassland, in various stages in the development of abandoned grassland and in woodlands (climax vegetation). (2) The results show a close correlation between bufferfly fauna and vegetation type concerning species composition and species richness of Lepidoptera. (3) Species richness of butterflies is highest in early abandoned stages and falls rapidly with the arrival of shrubs and trees. It is also high in traditionally lightly cultivated grassland (unfertilized mown and lightly grazed meadows), but it declines drastically with increasing intensity of cultivation. (4) In general, species richness of butterflies is closely correlated with species richness of vascular plants, but cultivation methods and the short time since abandonment (<5 years) of lightly cultivated grassland have strongly different influences on species richness of Lepidoptera and vascular plants. (5) The results are in contrast to former studies in Central Europe and parallel studies in England. (6) The natural primary habitats of butterflies living in the anthropogenetic types of vegetation investigated and the consequences for nature conservation are discussed.