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Journal of Insect Conservation

, Volume 19, Issue 2, pp 255–264 | Cite as

Abundances and movement of the Scarce Copper butterfly (Lycaena virgaureae) on future building sites at a settlement fringe in southern Sweden

  • Christine Haaland
ORIGINAL PAPER

Abstract

The Scarce Copper (Lycaena virgaureae) is a species that has suffered serious decline in several European countries. In Scandinavia it is still comparatively abundant, but with ongoing losses of flower-rich grasslands near forests further decline is expected. A mark–release–recapture study was carried out in July 2013 at 14 sites on the outskirts of a village located near Malmö, Sweden. The study area comprised in total an 11.4 ha network of abandoned agricultural sites, road verges and forest edges. A private garden was also included. Butterflies were marked individually and the capture position was recorded by GPS. Sex, behaviour and flower visits were also recorded. During the study 852 butterflies were marked and 170 of these were recaptured at least once (recapture rate 20 %), resulting in 1,078 captures (including multiple recaptures). Movement between patches accounted for 41 % of all recaptures and mean distance between recaptures was 112 ± 146 m (n = 226). The number of captures was strongly positively correlated with patch size (ρ = 0.95, p < 0.05), while the emigrant and immigrant fractions were significantly negatively correlated with patch size. Overall, the Scarce Copper was surprisingly abundant in the area, but planned construction of residential areas will result in the loss of most habitat patches.

Keywords

Dispersal Garden Housing development Mark–release–recapture Rumex thyrsiflorus Urbanisation 

Introduction

Many butterfly species have shown a rapid decline in recent decades. This trend is affecting both common and rarer species and is taking place globally (e.g. Thomas and Abery 1995; Maes and Van Dyck 2001; Inoue 2005; Van Swaay et al. 2006; Wenzel et al. 2006; Van Dyck et al. 2009; Swengel et al. 2011). The causes are manifold and include habitat loss (e.g. Nilsson et al. 2013), habitat fragmentation (e.g. Polus et al. 2007), land use change and land management changes (e.g. Dover et al. 2011; Streitberger et al. 2012). Other factors cited include atmospheric nitrogen deposition (Wallisdevries and Van Swaay 2006; Feest et al. 2014), decreased nectar supply (Wallisdevries et al. 2012) and decline of larval food plants (Pleasants and Oberhauser 2013). Climate change (Wallisdevries and Van Swaay 2006; Breed et al. 2013) is another threat to many butterfly species. Loss of butterfly species has also been reported in conservation sites (Nilsson et al. 2013; Filz et al. 2013), indicating that either management is not optimal or that adverse conditions in the surrounding environment cannot be compensated for (Filz et al. 2013).

Urbanisation has been found to have a negative impact on butterfly diversity (e.g. Kitahara and Fujii 1994; Dennis and Hardy 2001; Di Mauro et al. 2007; Meehan et al. 2013; Restrepo and Halffter 2013) and butterfly abundance (Blair and Launer 1997). The number of specialist species is reported to decline from the more semi-natural areas outside urban areas to city centres (Kitahara and Fujii 1994; Clark et al. 2007). Bergerot et al. (2011) were able to distinguish different butterfly species assemblages along such an urban gradient. The reasons suggested for the negative impact of urban development on butterfly diversity are lack of suitable habitat (Wood and Pullin 2002), habitat loss (New and Sands 2002) and isolation of habitat patches (Preston et al. 2012). On the other hand, high quality habitats and the connectivity of habitat patches can lead to butterfly diversity comparable with that in the surroundings, even in urban areas (Öckinger et al. 2009).

The effect of urban development on butterfly populations seems to be less well studied than the effect on overall diversity. Preston et al. (2012) related the extinction of the Quino Checkerspot (Euphydryas editha quino) partly to human population growth in a study area in southern California (US). Marschalek and Deutschman (2008) studied the Hermes Copper (Lycaena hermes) in urbanised areas of San Diego and identified urbanisation and wild fires as severe threats to this indigenous species. Moreover, butterfly movement studies in urban and urbanised areas seem to be rather limited. Movement between gardens has been tracked through mark–release–recapture studies for the Small White (Pieris rapae) in New York (Matteson and Langellotto 2012). Bergerot et al. (2012) compared movement patterns for the Speckled Wood (Pararge aegeria) in different landscapes and found that individuals were more sedentary in a highly fragmented urban landscape (Paris) than in other landscape types. The work of Bergerot et al. (2013) on the Cabbage White (Pieris brassicae) and that of Leidner and Haddad (2010) on Atrytonopsis spec 1 demonstrate the high mobility of certain butterfly species in highly urbanised landscapes, but also highlight certain natural and human barriers to butterfly movement.

Despite all the negative effects of urbanisation on butterfly diversity, abundances and populations, there is also evidence that large numbers of species still occur in urbanised areas (e.g. Konvicka and Kadlec 2011). The results of butterfly monitoring schemes by residents of urban areas provide further evidence of this (Matteson et al. 2012). Studies by e.g. Strausz et al. (2012) also show the importance of urban abandoned sites and waste sites for an endangered butterfly species (Large Copper, Lycaena dispar) in and around Vienna.

Scarce Copper (Lycaena virgaureae L.) populations have been studied, including movement studies, in Norwegian mountain meadows (Fjellstad 1998), agricultural sites in Sweden (Douwes 1975a, b; Schneider et al. 2003), a Mongolian meadow system (Barua et al. 2011) and clearcut areas/meadows in a forested area in Bavaria, Germany (Schlumprecht and Bräu 2013). Several of these studies concluded that the species is relatively mobile, dispersing up to several kilometres.

The aim of the present study was to explore the use by the Scarce Copper of abandoned agricultural sites at a settlement fringe awaiting exploitation for housing development. Specific objectives were to estimate Scarce Copper abundances, identify movement patterns and determine habitat use, such as preferences for certain adult food resources.

Methods

The species

The Scarce Copper has a Eurasian distribution (Kudrna et al. 2011). In Europe, it is found in most countries, but not in the UK, Ireland, Belgium and the Netherlands (Tolman and Lewington 1997). The Scarce Copper occurs in most of Fennoscandia up to the Polar Circle and even further north (Kudrna et al. 2011). It flies up to elevations of 1,000–2,000 m (Tolman and Lewington 1997), especially in its southern distribution (e.g. Spain, Switzerland). However, in Fennoscandia, for example, it also occurs at low elevations, around sea level. In Asia, the Scarce Copper is distributed from Asia Minor to Mongolia (Kudrna et al. 2011).

The habitat of the Scarce Copper is described as flower-rich meadows situated in the vicinity of forest edges (Henriksen and Kreutzer 1982). Males commonly use forest edges as perch sites (Douwes 1975a).

The larval food plant is Rumex spp., with Rumex acetosa or Rumex acetosella being mentioned most often (Ebert 1993; Schlumprecht and Bräu 2013). However, Rumex crispus is cited by Settele et al. (2000) and Rumex thyrsiflorus by Schlumprecht and Bräu (2013, citing Bink 1992). The overwintering stage is the egg and the larvae hatch in spring (Henriksen and Kreutzer 1982). Flying season is one generation from June–September (depending on geographical area).

Based on historical records for Bavaria, Schlumprecht and Bräu (2013) concluded that the Scarce Copper was probably a very common species in parts of its distribution area at the beginning of the last century. However, the species has since experienced a considerable decline in several parts of its distribution area, and in several regions in Central Europe (e.g. in Switzerland and Germany). It is red-listed regionally or nationally (Settele et al. 2000). In fact, in some regions it has become extinct (Ebert 1993; Schlumprecht and Bräu 2013).

In Sweden, the Scarce Copper is still comparatively common, but is under decline (Öckinger et al. 2006; Nilsson et al. 2013). A worrying trend is that the species is disappearing from localities where it has been established for a long time, some of which are nature reserves (Nilsson et al. 2013). The reason for the decline is probably habitat loss and/or non-optimal management of the habitat, e.g. grass swards being cut too short. The Scarce Copper is known to occur in abundance in localities with favourable conditions (Douwes 1975b; Barua et al. 2011).

The study area

The study was carried out on the outskirts of the village of Veberöd, which is located 30 km from Malmö, Sweden’s third largest city. Thus the study area is situated in the most southern part of Sweden, in the county of Scania. The south-western part of this county has experienced a population increase and increased urbanisation of the countryside during the past decade. This development has been accelerating since the construction of the Öresund bridge, which connects the region with Copenhagen in Denmark. Veberöd has 4,000 inhabitants (2010), but there are plans to expand and to double the number of residents within the next 20–40 years. A total of 3,000 new housing units are planned, of which 250–300 will be in the study area (Lund Municipality 2009).

The mark–release–recapture experiment was carried out on 14 patches at the south-eastern edge of the village and in one private garden. Ten of these patches were abandoned grasslands or abandoned arable land and four were road/track verges. All patches were located between the village and the surrounding forest and the size varied between 0.04 and 4 ha. The soil at the site is very sandy and post-farming succession has been very slow in some patches, whereas in others pine trees have managed to establish. Time since abandonment of the former agricultural patches varies from 10 to more than 30 years. The vegetation is characterised by species which like dry, sandy soils, such as sheep’s bit scabious (Jasione montana), thrift (Armeria maritima) and thyrse sorrel (R. thyrsiflorus), but also others such as yarrow (Achillea millefolium) and field scabious (Knautia arvensis), indicating low to moderate soil nitrogen status. R. thyrsiflorus is by far the most abundant Rumex species in the study area, with R. acetosa and R. acetosella being comparatively rare. The road verges are cut once a year. The former agricultural patches are now mostly unmanaged, but in one area management measures have been initiated to prevent further succession and the grass was cut for the first time after the present study had been carried out. The patches were chosen since they form a network of potential habitat patches and were largely unmanaged. Patch boundaries are formed by paths, fences, tracks or the boundary to a different land use or vegetation type. The private garden patch was added to get an indication of whether gardens are used for resource complementation (adult food resources) by the Scarce Copper. To analyse the full extent of garden use, more gardens would need to be studied, of course. For each patch, the flower abundance was estimated in the middle of the experiment (1 = low abundance, 2 = moderate abundance and 3 = high abundance of flowering plants). The abundance of R. thyrsiflorus was estimated in the same way.

Mark–release–recapture experiment

The mark–release–recapture experiment was carried out from 6 to 26 July 2013. Butterflies were caught with a net, individually marked and immediately released at the capture site. Marking involved writing numbers with a permanent pen (Staedler Lumcolor, permanent) on the underside of the left hind wing. The location (coordinates) of capture/release was determined with a global positioning system (GPS) tracker (Garmin etrex 10). The coordinates, date and time of capture, patch number and sex were all recorded for each capture/recapture. Wing status was estimated on a scale from 1 to 3 (1 = fresh with bright colours, 2 = slightly worn, 3 = damaged, worn out). Behaviour just before capture was also recorded (flying, flower visit, resting or creeping in vegetation). In cases of flower visits, the visited flower was identified, mostly to species level but in a few cases to genus level.

At the beginning of the experiment all patches were visited on the same day, but as butterfly numbers increased half the patches were monitored on 1 day and the other half on the following day (in one case 2 days later due to weather conditions). Each patch was visited 12 times during the mark–release–recapture experiment but the garden was visited more often, and thus oversampled compared with the other patches.

Analysis

For statistical analysis, STATISTICA 12.0 (StatSoft 2013) was used, with Arc GIS 10.1 (ESRI 2012) as the geographic information system (GIS). Distance moved was calculated as a straight line between the location of the first capture and the location of the following capture (determined from the coordinates given by the GPS unit).

Distance decay curves were calculated according to Hill et al. (1996) using a ln-plot of inverse cumulative proportion of individuals moving certain distances for the negative exponential function and a double-ln plot for the inverse power function. Distance decay curves are calculated to allow for comparison of dispersal ability between species and to predict long-distance movements, which often are not detected with mark-release-recapture.

The emigration and immigration fractions were also calculated according to Hill et al. (1996) in the following way: Resident fraction (RF) is the number of residents (R) of a patch divided by the sum of residents, emigrants (E) and immigrants (I) (RF = R/R + E+I). The emigrant fraction is E/E + R and the immigrant fraction I/I + R.

Results

Captures and recaptures

During the mark–release–recapture study 852 butterflies were marked, 661 (78 %) males and 191 (22 %) females (Table 1). A further 226 recapture events were recorded, resulting in 1,078 captures in total. Of the males, 159 individuals were recaptured at least once, which represents a recapture rate of 24 %. Of the females, only 11 individuals were recaptured at least once, which represents a recapture rate of just 6 %. Maximum number of recaptures per individual was four. Number of captures increased for both sexes during the study period (Fig. 1), with a linear increase for captures of males (y = 18.65 + 80.53, R2 = 0.77) and an exponential increase for captures of females (y = 1.1585e0.7506x, R2 = 0.92). Wing status, which can indicate the age of butterfly individuals, changed for males, with a decreasing proportion of individuals with fresh wings (Fig. 2a). Caught females, on the other hand, nearly all had fresh wings (Fig. 2b). The time span between recaptures was a maximum of 16 days and mean time between recaptures was 4.6 ± 3.6 days (n = 226).
Table 1

Results of mark–release–recapture data for the Scarce Copper in the study area (Veberöd, Sweden)

 

Total

Males

Females

Number individuals marked

852

661

191

Number of captures

1,078

875

203

Number of recaptures

226

214

12

 Number recaptured once

170

159

11

 Number recaptured twice

44

43

1

 Number recaptured three times

11

11

0

 Number recaptured four times

1

1

0

Recapture rate of individuals

20 %

24 %

6 %

Fig. 1

Number of captures of male (n = 875) and female (n = 203) Scarce Coppers during the survey period in the study area (Veberöd, Sweden)

Fig. 2

Wing status of Scarce Copper individuals captured in different survey periods during July 2013: a males, n = 875, b females, n = 203; whereby 1 = fresh, 2 = slightly worn, 3 = damaged, worn out

In the garden, the first capture was recorded on 18 July, 12 days after the start of the field study. Apart from one farm track verge, all other patches were occupied by the Scarce Copper from the beginning of the field study. While the number of captures generally increased during the field study, this was not the case for road verges, which were cut during this mark–release–recapture experiment. There, number of captures dropped after cutting.

Movement

The mean distance moved between each successive capture was 112 ± 146 m (n = 226; Table 2). The minimum distance recorded between captures was 1 m and the maximum 997 m. This was also the longest distance detected for movement of an individual. The mean distance moved by individuals was 149 ± 180 m (n = 170). Differences between sexes were not analysed due to the low recapture rate of females. Two-thirds of the movements detected were up to 100 m, one-third between 101 and 300 m (Fig. 3). Only 12 movements were beyond 300 m. The distance decay curve fitted a negative exponential function (R2 = 0.90, F1–8 = 75.8, p < 0.001) better than an inverse power function (R2 = 0.78, F1-8 = 27.8, p < 0.001).
Table 2

Results of mark-release recapture experiment on moved distances by the Scarce Copper in the study area (Veberöd, Sweden)

Movement parameters

 

Mean distance moved between successive recaptures (m)

112 ± 146

Mean distance moved by individuals (between first and last recapture) (m)

149 ± 180

Maximum recorded distance moved between recaptures (m)

997

Maximum recorded distance moved by individual (m)

997

Mean residence time between successive recaptures (days)

4.6

Mean residence time between first and last recapture (days)

6.2

Maximum recorded resident timea (days)

17

aBetween successive recaptures, no higher residence time was recorded between first and last recapture

Fig. 3

Movement frequencies of the Scarce Copper (n = 226) in the study area (Veberöd, Sweden)

It was found that 59 % of all captures were recaptures in the same patch as the previous capture, while 41 % of all recaptures were movements between patches (Fig. 4a). Movements to adjacent patches counted for more than half (58 %) of all movements between patches. The number of captures and residents in a patch and the resident fraction were strongly positively correlated with patch area and not with other factors such as flower abundance or abundance of R. thyrsiflorus (Table 3). The emigrant fraction and immigrant fraction were negatively correlated with patch area.
Fig. 4

a Movement of the Scarce Copper in the study area (Veberöd, Sweden). White lines are straight lines between capture and recapture points (=black dots). b Planned housing development in the study area (Veberöd, Sweden). Light areas investigated patches, red areas areas for planned development, P planned area for car parking, ? area under investigation (references to colours apply to the online version only). Numbers are the number of captures per patch (for patch number 15 in the east, numbers are given for both the part to be built on and the undeveloped part. (Color figure online)

Table 3

Spearman rank correlations between number of captures, residents, emigrant, immigrants and their fractions, with patch area, flower abundance and Rumex thyrsiflorus abundance

 

Area

Flower abundance

Abundance of Rumex thyrsiflorus

Captures

0.96***

ns

ns

R

0.93***

ns

ns

E

ns

ns

ns

I

ns

ns

ns

R-fraction

0.82***

ns

ns

E-fraction

−0.78***

ns

ns

I-fraction

−0.75**

ns

ns

ns not significant p > 0.05

*** p < 0.001; ** p < 0.01

Behavioural observations

Of the 1,078 individual observations of butterfly behaviour prior to capture, 44 % were observed while sitting on a flower, 33 % were flying and 23 % were resting or creeping on vegetation. There were significant differences in behaviour between the sexes. While 38 % of males were observed flying prior to capture, only 8 % of females did so (χ2 = 39.59, p < 0.0001; Fig. 5). Moreover, 70 % of females were seen on flowers before capture, but only 38 % of males (χ2 = 23.98, p < 0.0001). The proportions of individuals recorded resting/creeping on vegetation were similar between the sexes (24 % of males, 21 % of females).
Fig. 5

Behavioural observations prior to capture of Scarce Copper individuals (males, n = 875, females n = 203) in the study area (Veberöd, Sweden)

Flower visits

Of the 474 flower visits recorded, almost one-third (n = 154) were observed on yarrow (A. millefolium). Other flowers frequently visited were thrift (Armeria maritime), sheep’s bit scabious ( J. montana), oregano (Origanum vulgare), alfalfa (Medicago sativa) and field scabious (K. arvensis) (Fig. 6). In total, 23 different species/genera were visited. The differences in flower visits between the sexes was significant (Wilcoxon matched pair test, T = 4.5; Z = 3.86, p < 0.001). In particular, the difference in visits to yarrow was significant (males 27 % of all flower visits, females 45 %; χ2 = 6.82, p < 0.01).
Fig. 6

Frequencies of flower visits of the Scarce Copper (males, n = 331 and females, n = 143) in the study area (Veberöd, Sweden)

Larval food plant

Although egg laying was not observed, it can be assumed that the larval food plant species used in the study area is R. thyrsiflorus. This assumption is based on the fact that that the other Rumex species are very rare in the study area and that a butterfly species with such high abundances must use a larval food plant that is found at many places. R. thyrsiflorus is very common in the study area. Females were observed on a number of occasions creeping on the ground vegetation between leaves and stalks of R. thyrsiflorus, which could be interpreted as a search for egg-laying sites. Male Scarce Coppers were frequently observed sitting on leaves of R. thyrsiflorus that had turned red, with the colour of their wings nicely matching the colour of the plant.

Planned housing development

The planned sites for housing development include two-thirds of the habitat patches of the Scarce Copper studied here (Fig. 4b). In the planned development area, 613 Scarce Copper individuals (57 %) were captured. Other areas which were not investigated are also scheduled for development. Some of these areas could not be investigated due to time constraints, but are certainly used by the Scarce Copper (north-eastern corner of the largest planned development site in the study area).

Discussion

Captures, recaptures, behaviour

The number of butterflies captured was much higher than expected prior to the experiment. The densities observed were higher than e.g. in a heterogeneous, small-scale agricultural landscape with very extensive use in another Swedish study (Schneider et al. 2003). The high abundance of Scarce Coppers can be explained by several factors. Firstly, there was a high abundance of larval food plant in the study area. Secondly, the Scarce Copper is known to be particularly attracted to semi-natural meadows and wood glades (Henriksen and Kreutzer 1982; Ebert 1993; Schlumprecht and Bräu 2013). These habitats are characterised by high vegetation of grasses and forbs, high abundance of flowering plants and low management intensity. These conditions were largely met in the habitat patches studied, although flower abundance was variable and not always high. Finally, the study area has sandy soils and the vegetation indicates low nutrient levels in the soil, despite most of the area being previously used as arable land. On more fertile soils, succession and thus vegetation structure and plant species composition would have been very different, and would probably not have resulted in such favourable conditions for the Scarce Copper.

While numbers of captures were relatively high, recapture rates were rather low compared with those reported in other mark–release–recapture studies on the Scarce Copper (Schneider et al. 2003; Barua et al. 2011). This was especially true for the recapture rate of females, which was very low, only 6 %. This low recapture rate of females was mainly caused by two factors: (1) comparatively few females were marked during the study and (2) females emerge later than males and due to various constraints the study was too short to capture late-emerging females. This did not affect the general results and conclusions of the study, but recapture numbers of females were too low for female movements to be analysed and discussed.

In addition, the detectability of females might be lower than that of males. Behaviour differed quite substantially between the sexes regarding the proportion of individuals flying, sitting on flowers or resting in/on vegetation (other than flowers). Males were seen much more often flying and they were observed particularly frequently flying back and forth along the forest edges, which has been interpreted as patrolling behaviour by males in their territory (Douwes 1975a). When recorded as resting on vegetation other than flowers, males were often sitting in a very visible position on other plant parts, while females could be observed searching for egg-laying places by creeping near the ground in dense vegetation, where they are difficult to detect. According to previous literature, female recapture rate of Scarce Copper can be equivalent to male recapture rate (Douwes 1975b), or only half the male recapture rate (Barua et al. 2011).

The spectrum of plant species visited for flower visits was broad, but with a clear preference for yarrow (A. millefolium). The preference of the Scarce Copper for using certain members of the Asteraceae as an adult food resource is well known (Ebert 1993; Schlumprecht and Bräu 2013), in particular members of the genus Achillea (e.g. Barua et al. 2011). Females were observed creeping on Rumex thrysiflorus leaves near the ground, probably in search of egg-laying sites. However, no egg laying could be observed, and in general there are very few recorded observations of egg laying in the literature. Only one such reference is made, to Rumex thrysiflorus as a plant for egg laying (Bink 1992 cit. in Schlumprecht and Bräu 2013). More research on the egg-laying and larval stages of the Scarce Copper is needed to confirm this.

Movement

The mean distance moved between two consecutive captures was 112 ± 146 m, which was lower than in other mark–release–recapture studies with the Scarce Copper (Schneider et al. 2003; Barua et al. 2011). Comparison between study sites is difficult due to the many factors influencing mean distances moved. These include spatial factors (size and shape of the study area, spatial arrangement of habitat patches, isolation; Bonelli et al. 2013), habitat quality (Baguette et al. 2011) and quality of the matrix (Ricketts 2001). Moreover, different methodologies are used, for example focusing on inter- or intra-patch movement. Schneider et al. (2003) worked in a much larger study area and without GPS equipment (no detection of short movements), which resulted in longer mean distances being recorded than in the present study. Barua et al. (2011), on the other hand, carried out their study in a smaller, but contiguous, study area, also using GPS, and found longer movement distances. Nevertheless, the fact that the maximum distance moved recorded was 997 m and that 40 % of the recorded movements were between patches underlines that the Scarce Copper has substantial mobility even in the limited habitat network studied here. As reported by Schneider et al. (2003), the distance decay curve better fitted a negative exponential than an inverse power function. Inverse-power functions generally predict a greater probability for long distance movements than negative exponential functions (Hill et al. 1996). Numbers or fractions of residents, emigrants and immigrants have previously been related to area, isolation (e.g. Bonelli et al. 2013) and habitat quality (Baguette et al. 2011). However, in the present study there was only a correlation with patch area, and not with recorded variables for patch quality. A probable explanation for this is that the patches did not vary sufficiently in quality for this factor to be significant, and thus patch area was the only significant factor.

Overall, it can be seen from the movement data that the Scarce Copper moves frequently between habitat patches, here on the edges of a village. Gardens with attractive flowers are used for resource complementation for adult butterflies, which fly from habitat patches at the urban fringe to the garden and back.

The planned housing development will result in the loss of most habitat patches in the centre of the study area (Fig. 4b), leaving a complex of habitat patches in the west of the study area and one in the east. The distance between these two remaining parts of the study area is within the flight range of the Scarce Copper. However, the question is whether the species is able to cross 1 km of built-up area. If this poses difficulties, movements between the western and eastern part of the area can be expected to decrease.

Implication for conservation

This study shows the importance of undeveloped and unmanaged land at settlement fringes as butterfly habitats. In the absence of semi-natural meadows in particular, the types of unmanaged land found in the study area can be an important habitat for butterfly species such as the Scarce Copper. In the whole of Sweden, no more than 6,000 ha of semi-natural meadows remain due to conversion to ley, abandonment or afforestation. The decline in Scarce Copper in the study region, even in nature reserves (Öckinger et al. 2006; Nilsson et al. 2013), indicates that habitat quality on semi-natural pastures, in particular with strong grazing pressure, is not optimal for this species. However, even if succession progresses slowly in the study area, at some stage management measures will be necessary, but should be of a very extensive nature (late cutting and not every year, preferably rotationally).

It is important to point out that in the study area, road verges alone are not sufficient to ensure long-term survival of the Scarce Copper. It has been described as a species occurring along forest roads, but in low densities, followed by extinction (Ebert 1993). The road verges studied here were cut in mid-July, thus offering no resources in the middle of the main flying season.

The importance of undeveloped land in urban areas has recently been pointed out for another Copper species (Strausz et al. 2012), but the number of butterfly studies focusing on this type of habitat is still limited. Despite the fact that certain butterfly species can be much more abundant in urban fringe habitats than in managed nature reserves, undeveloped urban areas rarely receive protection from development.

Development pressure in the study region is high. Furthermore, arable land still in use must be spared from development according to national policies to ensure a certain level of domestic food production. These two factors are creating a situation where housing development is increasingly taking place on non-arable land, including urban green space, forest, semi-natural pastures and undeveloped land, which most often has higher species diversity than intensively used agricultural land.

In the area studied here, the impact of future housing development could be decreased by decreasing the number of housing units planned, more careful location of the planned housing units and leaving the existing vegetation in certain patches instead of transforming it into typical urban green space areas with low biodiversity. Forest edges in particular could be left undeveloped.

Notes

Acknowledgments

This study was funded by the Swedish University of Agricultural Sciences, Department of Landscape Architecture, Planning and Management, Alnarp. I am thankful to Mary McAfee, who corrected the English.

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Copyright information

© Springer International Publishing Switzerland 2014

Authors and Affiliations

  1. 1.Department of Landscape Architecture, Planning and ManagementSwedish University of Agricultural SciencesAlnarpSweden

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