There is growing concern over the loss of grassland and forest species worldwide due to land use changes. In Japan, young forest plantations provide important habitats for grassland species. However, the decline in forest logging frequency has led to a decrease in the area of young plantations, which may in turn cause a decline in the number of grassland species. Power line corridors in forest plantations can act as habitats for early and late successional species, as they contain vegetation in diverse stages. This study evaluated the importance of these corridor zones as habitats for early and late successional butterflies in Japan. The species richness and abundance of butterflies were recorded in power line corridors, young plantations, forest roads, and mature plantations. Vegetation height and food resource availability for larvae and adult butterflies were also measured. The species richness and abundance of those of late successional butterflies were highest in power line corridors and young plantations, and lowest in mature plantations; and early successional butterflies and food resource availability were highest in power line corridors, and lowest in mature plantations. The species richness and abundance of butterflies within power line corridors were largely explained by vegetation height and food resource availability. Our results indicate that power line corridors within conifer plantations provide important habitats for early and late successional butterflies.
Implications for insect conservation
Increasing the habitat value of power line corridors through appropriate vegetation management can have an important role in preserving insect species.
In recent decades, land use change has been a significant driver of global biodiversity loss (Sala et al. 2000; Gerstner et al. 2014). For instance, deforestation has led to a decline in forest-dependent species (Hanski et al. 2007; Vergara et al. 2013), while farmland abandonment and agricultural intensification have led to a decline in grassland-dependent species that are supported by regular vegetation management practices, such as logging, mowing, and burning (Uchida and Ushimaru 2015; Seibold et al. 2019).
A wide range of anthropogenic land use practices, such as vegetation clearing, often maintain early successional vegetation, which can therefore function as alternative habitat for early successional animal species, such as grassland-dependent species (Kalarus and Bąkowski 2015; Ohwaki et al. 2018a; Yaida et al. 2019; Steinert et al. 2020). For example, forest roads where road verge vegetation is kept short by mowing can provide important habitats for butterfly and diurnal moth species (Saarinen et al. 2005). Young plantations can also harbor many early successional (and often threatened) species for up to ten years after felling because the low vegetation provides abundant food resources for them (Viljur and Teder 2016; Ram et al. 2020).
Compared to the above mentioned land use types, power line corridors (i.e., open habitat underneath electric transmission lines) are likely to serve as significant habitats for early successional species (Berg et al. 2013, 2016) mainly for the following three reasons. First, vegetation in power line corridors is usually managed once every 5 to 10 years, which allows the maintenance of the habitat for early successional species. Thus, the maintenance of power line corridors is less frequent than that of young plantations, where vegetation is controlled every year for the first 3–5 years; and is also less frequent than that of forest roads or railway tracks where vegetation is controlled several times a year (Tikka et al. 2000; Eldegard et al. 2015; Ohwaki et al. 2018b). Second, the timing of vegetation management interventions often varies among locations in power line corridors, allowing the presence of multiple vegetation types, ranging from grasslands to shrublands, including sections with saplings and smaller trees, and generating habitats for various early successional species (Clarke et al. 2006; Wagner et al. 2014, 2019). Third, as power line corridors are linear and extensive, they also connect natural habitats for meta-populations of early successional species (Berg et al. 2016). Although numerous studies have examined the function of power line corridors as habitats worldwide, the majority have focused on evaluating them as habitats for grassland or early successional species (Berg et al. 2011; Askins et al. 2012; Lampinen et al. 2015). However, power line corridors might also function as habitats for forest or late successional species, due to the variety of vegetation types coexisting in them, including shrubland and early successional forests (Komonen et al. 2013; Plewa et al. 2020).
In Japan, coniferous forest plantations account for approximately 25% of the national territory (Forestry Agency 2018). In particular, mature conifer plantations usually consist of tall trees of a single species, which do not support early successional or late successional species (Makino et al. 2007). Over the last few decades, semi-natural grasslands have declined substantially across the country (Fukamachi et al. 2002; Suka et al. 2011), and coppice logging—which once maintained a suitable habitat for early successional species—has also ceased in many parts of the country (Okubo et al. 2005; Oono et al. 2020). Therefore, the presence of open land in young conifer plantations may act as an important alternative habitat for early successional species (Yamaura et al. 2012, 2016). However, the decline in Japan’s forest economy, after its peak in the 1980s, has reduced the frequency of logging and planting in coniferous forest plantations (Forestry Agency 2018), leading to the loss of this important habitat type (Yamaura et al. 2012). Power line corridors in conifer plantations may thus function as an alternative to young plantations, providing an important habitat for early successional and late successional species. Nevertheless, this hypothesis has rarely been examined in Japan.
This study aims to assess the importance of power line corridors as habitats for early successional and late successional butterfly species in Japan’s conifer plantation landscapes. Butterflies were chosen because, (1) their abundance and diversity is in rapid decline in many parts of the world (Thomas et al. 2004; Wenzel et al. 2006; Nakamura 2011; Wepprich et al. 2019); (2) these insect species are known to be good indicators of overall biodiversity (Dennis 2004b; Brereton et al. 2011); and (3) knowledge of their food resources is well-established (Fukuda et al. 1982, 1983, 1984a, b; Saito et al. 2016). The species richness and abundance of butterflies was compared in four major habitat types in plantation landscapes: power line corridors, young plantations, mature plantations, and forest roads (Fig. 1). Then, vegetation height and food resources for both larvae and adult butterflies—which are essential determinants of butterfly abundance and richness—were measured (Dennis 2004a; Vanreusel et al. 2007; Berg et al. 2013; Curtis et al. 2015). Finally, the factors that explain the species richness and abundance of butterflies in the four above mentioned habitat types were investigated.
Materials and methods
The study was conducted south of Mt. Fuji (138° 38′–45′ E, 35° 14′–17′ N), in central Japan (Fig. 1). The study area’s average annual temperature is 15.8℃, and the average annual rainfall is 2109.1 mm (JMA 2020). The region is dominated by Japanese cypress (Chamaecyparis obtusa) and Japanese cedar (Cryptomeria japonica), distributed as a mosaic of mature and young plantations. In mature plantations, the understory vegetation is limited by the closed canopy, as reported in a previous study (Ramovs and Roberts 2003). In young plantations, light availability allows the growth of herbs and shrubs among the trees. In the investigated study area, forest roads, which are all paved, form a network of diverse herbs and shrubs growing along their edges. Two power lines corridors run through the study area, with understory vegetation ranging from grassland—which occurs immediately after vegetation clearance by mowing—to grass-shrub mosaic vegetation, shrubland, and early successional woodlands and forests. Grassland communities in the region are dominated by Japanese pampas grass (Miscanthus sinensis) and other grasses, which also grow in semi-natural grasslands (Kubo et al. 2009; Stewart et al. 2009). The shrubland is dominated by deciduous broad-leaved trees such as Chinese sumac (Rhus javanica), Japanese bladdernut (Staphylea bumalda), and Weigela decora. The early successional forest community is dominated by different deciduous, broad-leaved trees, such as Lindera praecox, Quercus serrata, and Japanese oak (Quercus crispula). No other potential habitats for butterflies (e.g., semi-natural grasslands) were identified in the studied landscape.
Within the study area, 77 study sites were established: 47 in power line corridors, 14 in young plantations, 10 on forest roads, and 6 in mature plantations (Supp. Material 1). The sites were chosen based on the vegetation height in the corridors to include bare ground, grassland, shrub, and early successional forest communities. In the studied region, the variation in vegetation height was much greater in power line corridors than in the planted forests and roads. Therefore, to encompass such an environmental heterogeneity, a greater number of survey sites were established in power line corridors. The corridor widths at the sites ranged from 15 to 65 m (mean, 33 ± 14 m). The length between adjacent sites located within the corridors ranged from 50 to 800 m (mean, 157 ± 129 m). As forest roads within 500 m of power line corridors may have more butterflies than those further away (Berg et al. 2016), the study sites were set on forest roads that were all paved and in young plantations located more than 500 m from any power line corridors. The width of the grassy edges along forest roads at the sites ranged from 3 to 9 m (mean, 6 ± 2 m), and the area of young plantations ranged from 0.2 ha to 9.7 ha (mean, 2.7 ± 2.8 ha). The elevation of the sites ranged from 585 to 945 m (mean, 735 ± 101 m).
Butterflies were surveyed at each site by using the transect method (Pollard 1977), which involved walking a 50 m transect at each site for 5 min and recording the species and abundance (i.e., occurrence) of all butterflies within 5 m on both sides. All butterflies that appeared in front of, and above, the observer were recorded. The survey was based on visual confirmation in principle, and any individuals that were difficult to identify were captured with a net, identified, and immediately released.
The survey was conducted during the daytime (from 08:30 to 16:00) on sunny or slightly cloudy days with little wind. Considering the time-related activity patterns and seasonal appearance of different species, all transects were visited three times: once in the morning (08:30–11:00), once at noon (11:00–13:00), and once in the afternoon (13:30–16:00). The visits were conducted during each of the following periods: May 11–25 (spring), July 13–27 (summer), and September 6–19 (fall), 2018.
The observed butterflies were classified into grassland, ruderal, and forest butterflies based on previous studies by Tanaka (1988), Shirouzu (2006), and Ohwaki (2018c). In this study, grassland butterflies were defined as early successional species that inhabit natural and semi-natural grasslands, but never inhabit modified open areas such as urban parks or residential areas. Ruderal butterflies were defined as early successional species that inhabit open areas, including urban parks and residential areas. Forest butterflies were defined as late successional species associated with forests. In addition, early succession habitats were defined as vegetation such as bare land and grasslands, and late succession habitats as forest composed of tall sun trees or shade trees (cf. Ohwaki 2018c).
Five plots per transect were randomly set at each site to examine vegetation structure. The plots were established at the grassy edge of forest roads, at least 1 m apart from paved roads. Plot size was 1 m × 1 m where the average vegetation height was < 1 m, 3 m × 3 m where the height was 1–5 m, and 5 m × 5 m where the height was > 5 m, given that tall plants cover a large area.
The average vegetation height in each plot was recorded as vegetation height. All vegetation heights above 10 m were recorded as 10 m. Based on Fukuda et al. (1982, 1983, 1984a, b) and Shirouzu (2006), the species richness of host plant of observed butterflies was recorded as potential larval food resources, and the flowering plant coverage was recorded as potential adult food resources. The preference of butterflies for flowering plants generally differs substantially among butterfly species (Tudor et al. 2004). Detailed information on the flowering plants preferred by the butterfly species observed in the present study was limited, therefore all flowering plants were treated as potential adult food resources for butterflies. No sap or fruits, which are also adult food resources, were detected. Flowering plant coverage was recorded at six levels: 0%, 1–10%, 11–25%, 26–50%, 51–75%, and 76–100%. For analysis purposes, the levels were represented as 0%, 5%, 17.5%, 37.5%, 62.5%, and 87.5%, respectively. The mean values of the five plots at each site were used in the analyses.
Considering that seasonal changes affect phenology, vegetation height and flowering plant coverage were recorded in May 17–24 (spring), July 19–27 (summer), and September 16–29 (fall), 2018, while host plant species richness in September 16–29, 2018.
The species richness and abundance of grassland, ruderal, and forest butterflies were analyzed per transect per survey period. In a preliminary analysis, the Moran’s I was calculated to test spatial autocorrelation in the species richness and abundance of butterflies among study sites and it was found to be statistically non-significant (grassland butterflies: richness, P = 0.18; abundance, P = 0.46; ruderal butterflies: richness, P = 0.68; abundance, P = 0.14; forest butterflies: richness, P = 0.27; abundance, P = 0.20), indicating the independence of each site. The Pearson product-moment correlation coefficient between plot size and host plant species richness or flowering plant coverage was also calculated, and no significant correlation was found, which indicates that variable plot size is not associated with the quantity of food resources.
Two analyses were conducted. First, the species richness and abundance of grassland, ruderal, and forest butterflies, vegetation height, host plant species richness, and flowering plant coverage were compared in each habitat type. In this analysis, generalized linear mixed models (GLMMs) were performed using species richness and abundance of butterflies, vegetation height, host plant species richness, and flowering plant coverage as response variables, and habitat types as explanatory variables. Young plantations were used as reference category. In order to address the issue of pseudoreplication, study sites within 500 m were grouped together (77 study sites were divided into 15 groups), and group ID was used as a random effect. This analysis was based on records that combined all study periods, and records of each individual study period.
Second, the relationship between the species richness and abundance of grassland, ruderal, and forest butterflies and environmental variables (corridor width, vegetation height, host plant species richness, and flowering plant coverage) was analyzed using the records of power line corridors. In this analysis, GLMMs were performed using species richness and abundance of butterflies as response variables; corridor width, vegetation height, host plant species richness, and flowering plant coverage as explanatory variables; and study periods (i.e., months) and group ID for each study site as a random effect. Furthermore, to allow for direct comparisons of the effects of the explanatory variables, all continuous variables were scaled to have a mean of zero and a standard deviation of one.
In these analyses, a negative binomial error distribution was assumed, as overdispersion was confirmed by a Poisson regression conducted in a preliminary analysis. All analyses were performed in R v. 3.3.2 (R Core Team 2018).
Comparison of species richness and abundance of butterflies
Across all sites, a total of 2,123 individual butterflies were observed, representing 62 species (410 belonged to ten species of grassland butterflies, 847 to 16 species of ruderal butterflies, and 866 to 36 species of forest butterflies; Supp. Material 2, 3). The species richness and abundance of grassland and ruderal butterflies were highest in the order of power line corridors, young plantations, forest roads, and mature plantations (Supp. Material 4). The parameters were significantly higher in power line corridors than in young plantations (Fig. 2a, b). A similar pattern was found across seasons (Supp. Material 4, 5a, 5b). The species richness and abundance of forest butterflies per transect per survey period was higher in power line corridors and young plantations, and lowest in mature plantations (Fig. 2c, Supp. Material 4). The same pattern was found in May and July; however, both species richness and abundance of forest butterflies were significantly higher in power line corridors than in young plantations in September (Supp. Material 5c).
Comparison of vegetation heights and food resources
Vegetation heights were higher in mature plantations and lower on forest roads, as shown in Fig. 3a, and presented similar trends across all seasons (Supp. Material 6a). Food resource availability tended to be highest in power line corridors and lowest in mature plantations (Fig. 4). Although there were more host plant species for grassland butterflies in young plantations than in the other three habitat types, power line corridors still possessed the largest number of host plant species for ruderal and forest butterflies (Fig. 4). Power line corridors also supported higher flowering plant coverage (Fig. 5). This pattern remained the same across seasons (Supp. Material 6b). The mean and standard deviation (SD) of flowering plant coverage were highest in power line corridors in most seasons (Supp. Material 7).
Factors associated with butterfly species richness and abundance
The species richness and abundance of grassland, ruderal, and forest butterflies within power line corridors were positively correlated with flowering plant coverage, and negatively associated with vegetation height (Table 1). However, there was no significant correlation between the species richness and abundance of the above mentioned butterflies, and corridor width or host plant species richness.
The results of the present study show that power line corridors within conifer plantations in Japan provide important habitats for grassland, ruderal, and forest butterflies. In particular, the corridors support higher species richness and abundance of grassland and ruderal butterflies than young plantations do, which have been recognized as an important habitat type for many butterflies dependent on early successional seral stages in Japan (Yamaura et al. 2012, 2016). More importantly, the results showed a higher diversity and abundance of forest butterflies in power line corridors than in mature plantations, where nectar resources can be limited to absent. These results suggest that power line corridors—human-made open habitats—play an important role in the conservation of both early successional and late successional butterflies.
The high diversity and abundance of butterflies in the three habitat groups (i.e., grassland, ruderal, and forest butterflies) observed in power line corridors can be explained by the greater availability of food resources—especially flowering plants (adult food resource)—found in them than in the other habitat types. In turn, the abundance of food resources in power line corridors can be explained by the infrequent vegetation management occurring in these areas, as reported in a previous study (Russell et al. 2018). In fact, this increases plant species richness over several years, as vegetation grows and stratification develops, thereby diversifying the vegetation structure (Wagner et al. 2019). Thus, many host plants for grassland, ruderal, and forest butterflies are expected to grow in the corridors and to be absent from closed-canopy communities. As a result, more grassland and ruderal butterflies can be supported by power line corridors than by other habitat types. Furthermore, as more diverse vegetation is present in the corridors than along forest roads, the former may represent a more important habitat for forest butterflies. However, there were no significant differences in the species richness or abundance of forest butterflies between power line corridors and young plantations, so, potentially; both habitats had sufficient resources for the larvae and adults of this type of butterflies.
The species richness and abundance of grassland, ruderal, and forest butterflies in mature plantations were extremely low, as reported in a previous study (van Halder et al. 2011). Significantly fewer host plants and flowering plants were found in mature plantations, indicating their unsuitability as butterfly habitats. In recent years, the abandonment of forest management practices has posed a severe threat to biodiversity in Japan (Yamaura et al. 2012). The results of this study suggest that this lack of management would contribute to further expanding the unsuitable habitat area for several species.
In addition, this study showed that the suitability of power line corridors as habitats for butterflies also largely depends on vegetation height and, unsurprisingly, food resource availability. In particular, flowering plant coverage seemed to be an important determinant of the richness or abundance of all three species groups (i.e., grassland, ruderal, and forest butterflies). Host plant species richness (i.e., the availability of food resources for larvae) was not positively associated with the richness or abundance of any of the species groups. This result suggests that power line corridors function as adult habitats, possibly because of their extent and openness which ensure the presence crucial nectar resources (cf. Wagner et al. 2014, 2019), and not as breeding habitats for butterflies. The result showing that all three species groups were negatively associated with vegetation height indicates the importance of vegetation management to create open habitats within the corridors by maintaining low vegetation. None of the species groups were affected by the width of the corridors, which indicates that even small power line corridors can support high species richness and abundance of early and late successional butterflies.
Nonetheless, power line corridors can have some negative impacts on biodiversity (Janss 2000; Vistnes and Nellemann 2001; Richardson et al. 2017). For example, in some cases they can promote forest fragmentation (Goosem and Marsh 1997; Strevens et al. 2008). In fact, it is known that power line corridors can limit the migration of species inhabiting the surrounding forest (Goosem and Marsh 1997). Also, importantly, power line corridors might contribute to the establishment and spread of a number of invasive plants (Bradley and Mustard 2006). In order to understand the conservation function of power line corridors, it is necessary to comprehensively examine both the positive aspects found in the present study and the negative ones.
This study has shown that power line corridors represent important habitats for both early successional and late successional butterfly species in Japan. This finding has significant implications for biodiversity conservation in “underused” countries such as Japan. Indeed, in Japan there has been a substantial and widespread decline in the abundance and species richness of many butterfly species that depend on early or late successional habitats, due to the loss of semi-natural grasslands or deforestation, and to the conversion of natural forests (Nakamura 2011; Uchida and Ushimaru 2015). Although this study focused only on butterflies, power line corridors may act as important habitats for insect species other than butterflies (Villemey et al. 2018). In fact, abundant flowering plants growing within power line corridors may provide food resources for diverse flower-visiting insects (i.e., pollinators), such as bees and moths (Russell et al. 2005; Wojcik and Buchmann 2012; Berg et al. 2016), as their species richness and abundance are known to be positively associated with the amount of flowering plants available (Ebeling et al. 2008; Blaauw and Isaacs 2014; Torné-Noguera et al. 2014). It is also likely that power line corridors have an important function in the reproduction of early successional species (King and Byers 2002). Therefore, increasing the habitat value of power line corridors through appropriate vegetation management (see the above section) can play an important role in the conservation of insect species, which ultimately contributes to preventing the ongoing decline of entomofauna worldwide (Sánchez-Bayo and Wyckhuys 2019; Wagner 2020).
The data and material that support the findings of this study are available from the corresponding author upon reasonable request.
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We thank the Shizuoka District Forest Office for permission to enter the National Forest. This study was partly funded by the Institute of Global Innovation Research, Tokyo University of Agriculture and Technology.
This study was partly funded by the Institute of Global Innovation Research, Tokyo University of Agriculture and Technology.
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Oki, K., Soga, M., Amano, T. et al. Power line corridors in conifer plantations as important habitats for butterflies. J Insect Conserv (2021). https://doi.org/10.1007/s10841-021-00343-6
- Alternative habitat
- Conifer plantation landscape
- Early successional species
- Grassland-dependent species
- Vegetation management