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Article

Determinants of Butterfly Community Structure and Composition at the Local Habitat Level: Importance of Neighboring Vegetation and Management Status: A Case Study

by
Masahiko Kitahara
* and
Taisuke Yasuda
Division of Natural Environment Research, Mount Fuji Research Institute, Yamanashi Prefectural Government, Kenmarubi, Fujiyoshida 403-0005, Japan
*
Author to whom correspondence should be addressed.
Diversity 2024, 16(6), 310; https://doi.org/10.3390/d16060310
Submission received: 15 April 2024 / Revised: 14 May 2024 / Accepted: 16 May 2024 / Published: 21 May 2024
(This article belongs to the Special Issue Biodiversity, Ecology and Conservation of Lepidoptera)
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Abstract

:
Exploring the determinants of biological community structure is important not only for understanding the formation of the community, but also for promoting its biodiversity conservation. We monitored butterfly communities in a grassland and woodland area of Mount Fuji, Japan, and explored the environmental factors that influenced them. Multivariate regression tree (MRT) analysis generated a 3-leaved MRT (G1, G2, and G3) and showed the total explained variation of 64.4% in the species composition. The explanatory variables discriminating between the two branches in the first node and in the second node of the MRT were neighboring plant community and management status, respectively. The neighboring plant community was related to the distribution of butterfly dietary resources and contributed a lot in determining the species composition of the community while the management status was largely related to the amount of their dietary resources, affecting the species richness and diversity. The PCA results revealed that the three key groups (G1, G2, and G3) were formed along the gradients of these two factors. Consequently, maintaining mowing management and the diversity of neighboring vegetation is very important for the continuance of butterfly community diversity and composition and the conservation of Red Listed species in the area.

1. Introduction

Exploring the factors that determine the structure and composition of biological communities has been one of the important themes of community ecology [1,2,3]. This kind of approach is also vital for the conservation of biodiversity [4]. In particular, the conservation of living organisms is often practiced at the level of local ecosystems, and in this respect, exploring the determinants of community structure at the local habitat level is very important not only for understanding the formation of local communities, but also for promoting biodiversity conservation in the area [5,6].
In butterfly communities, many studies have been conducted and discussed to date on the determinants of the community structure and composition at the relatively broad geographic level [7,8,9,10,11,12,13,14,15,16,17,18,19,20,21]. From the results of these studies, it is generally accepted that climate change and habitat alteration are two key determinants of butterfly community structure and composition at the relatively broad geographic level [7,9,11,12,13,14,18,20]. Meanwhile, at the local habitat level, various environmental factors have been reported to determine the structure and composition of butterfly communities. For example, physical environmental conditions (including climate) have been reported as determinants of the structure and composition of local butterfly communities [22,23,24,25]. Similarly, vegetation and landscape structure [25,26,27], natural or anthropogenic disturbance and management [22,28,29,30,31], and food resources [23,32] have also been reported as determinants of the structure and composition of local butterfly communities. Thus, unlike the case of relatively broad geographical levels, various cases have been reported for the determinants at the local habitat level, and it is assumed that the determination and formation process of the butterfly community structure will differ depending on the situation in each region. On the other hand, few studies have clarified the relationship between local butterfly community structure and the determinants, that is, how the determinants influence and function to determine the community structure and composition, while clarifying the mechanism and function of the determinants also leads to the conservation of local butterfly communities [33].
In the present study, we chose a grassland and woodland area at the foot of Mount Fuji in central Japan as the study site. Previous studies [34,35] have shown that the area has a highly diverse butterfly fauna including several Red Listed species. Moreover, it was an area where we could categorize various environmental factors (habitat types, neighboring plant community, management status, trampling pressure, and distance from the central part of the grassland) that are thought to affect the community structure of butterflies. Therefore, it was a very suitable area for exploring the environmental factors that determine the butterfly community structure. Under these conditions, we monitored butterfly communities in the area in 2009, and attempted to analyze the butterfly community structure and composition. The goals of this study were (1) to clarify what environmental factors are most relevant to the determination of local butterfly community structure and composition in the area, (2) to explore how these environmental factors function in determining the local butterfly community structure and composition (i.e., the mechanism of community structure and composition determination), and (3) to clarify what kind of species groups the local butterfly community is composed of along the gradients of these environmental determinants. As above-mentioned, the analysis in this study was based on data from a one-year sampling period. Therefore, this study is reported as a case study from a locality in central Japan.

2. Materials and Methods

2.1. Study Area

The study area is located in a grassland and woodland area (980 m a.s.l.) at the northwestern foot of Mt. Fuji, central Japan (35°26′54″ N, 138°36′46″ E). The terrain in this area is almost horizontal, but has irregular undulations with an elevation difference of about 8 m. The surface layer of this area is composed of scoria-like lava and volcanic ash resulting from past volcanic eruptions in the Mt. Fuji area. The study area consists mainly of landscapes such as grasslands, forests, and firebreak belts at the edges (Figure 1).
The grassland was used as a source of grass for fuel and forage until 60 years ago [34]. After that, it was abandoned, but part of it became plantations for afforestation. However, the afforestation was not successful, although mowing management was carried out. This is probably due to severe weather conditions such as strong winds and extremely low temperatures in winter. Alternatively, this may be related to severe soil conditions due to winter freezing [36]. The management of the plantation (mowing) continued until 2005, but the plantation has been abandoned since then. The grassland at the time of this study (i.e., in 2009) was mostly dominated by poaceous grasses such as Miscanthus sinensis, Arundinella hirta, and Spodiopogon sibiricus. Various other herbaceous plant species were also present including Red Listed plants such as Tephroseris flammea, Platanthera hologlottis, Swertia pseudochinensis, and Vincetoxicum pycnostelma. Furthermore, in the grassland, several shrub trees such as Rhamnus davurica, Malus toringo, and Euonymus sachalinensis were scattered due to the progress of secondary succession [34].
The forest adjacent to the grassland consisted of a mixture of deciduous and coniferous trees and larch (Larix kaempferi) plantations. The forest was separated from the grassland by a firebreak belt that was about 10 m wide (2 km long) and was established in 1959 [37]. Since 1961, all grasses and herbs within the firebreak belt have been mowed and removed by the land managers each year in late autumn to prevent the fire from spreading to adjacent areas if a fire were to break out [34].
In the study area, it was difficult to set many replicates in the same habitat type because the area of each habitat type discrimination was small (see Figure 1). In the traditional ‘Pollard walk’ monitoring protocol [38], transect lengths typically range from 1.5 to 2 km. However, in recent years, an increasing number of studies have used a 50-m long transect or divided a long transect into sections of a 50-m transect for survey and analysis, depending on the conditions of the study site, and have obtained effective results (i.e., Modified Pollard transects) [39,40,41]. In particular, Barkmann et al. [42] demonstrated the effectiveness of a 50-m long transect for both population monitoring at the regional scale and for local site comparison regarding diversity and overall habitat quality. However, it is necessary for the survey of butterflies to set up the 50 m transects apart from each other. As the result, we set up two or three 50 m transects in the same habitat type. If possible, it would be desirable to set up three or more replicates in each habitat type; however, in this study, two 50 m transects were set up in most habitat types due to the small area of each habitat type, as described above. As discussed below, this did not cause any problems in the analysis of the results. The multiple transects of the same habitat type were located 20 m to 80 m away from other habitat types of the transects. However, all transects were located within the range of about 550 m × 250 m (Figure 1), which is within the range of movement, even for sedentary species [43,44].
In total, we set up 11 transects in five habitat types (A to E) (Figure 1). Type A (two transects of Nos. 5 and 6) was a firebreak belt with mowing in the autumn once a year, surrounded on both sides by a mixed forest of deciduous trees and conifers with a height of 10 m and more (treated as “forest–forest”). Type B (two transects of Nos. 7 and 8) was a firebreak belt with mowing in the autumn once a year, surrounded on one side by the mixed forest stated above and on the other side by shrubs 3–4 m high (treated as “forest–forest”). Type C (two transects of Nos. 3 and 4) was a firebreak belt with mowing in the fall once a year, and surrounded on one side by a similar mixed forest to those stated above and on the other side by an abandoned grassland (treated as “grassland–forest”). Type D (two transects of Nos. 1 and 2) was an abandoned grassland that was mowed every year (1998 until 2005) up to 4 years before the present survey started (treated as “grassland–grassland”). Type E (three transects of Nos. 9, 10, and 11) was an abandoned grassland that has not been managed (mown) for several decades (treated as “grassland–grassland”). Type D and E transects were at least over 20 m away from the edge of the nearest forest (Figure 1).

2.2. Butterfly Survey

In the present study, we used the transect counts [33,45,46], and recorded all adult butterflies observed within about 5 m on both sides and in front of each transect. The transects were visited under sunny and light wind weather conditions between 10:00 and 14:00. Each transect was visited twice a month from May to September, the main emergence period for the adults of each butterfly species in 2009. Individuals that could not be identified immediately were captured by net, identified, and released. In the field, it is impossible to distinguish between Pieris melete and P. japonica. Therefore, these two congeneric species complexes were treated as Pieris spp. for the analysis.

2.3. Explanatory Variables

We selected five categorical environmental factors in each transect as explanatory variables that were thought to affect the butterfly community structure such as habitat types (habitat), neighboring plant community (nei.com), management status (manag), trampling pressure (tramp), and distance from the central part of the grassland (dist). The reasons for choosing each factor as an explanatory variable and their categorization are as follows.
(1) Habitat type: Since butterflies utilize species-specific dietary resources and breeding sites, different habitat types such as grasslands and forests may affect the butterfly community structure. The habitat type was divided into five categories: Rank 1—grassland where mowing continued until recently, Rank 2—grassland where mowing has not been performed for a long time, Rank 3—one side of the transect was grassland, and the other side was woodland, Rank 4—one side of the transect was shrubs, and the other side was woodland, and Rank 5—both sides of the transect were woodland.
(2) Neighboring plant community: Butterflies usually depend on species-specific plants for food resources, so differences in plant communities near their habitat may affect the butterfly community structure. The neighboring plant community was divided into three categories: Rank 1—both sides of the transect were grassland, Rank 2—one side of the transect was grassland, and the other side was woodland, and Rank 3—both sides of the transect were woodland.
(3) Management status: Since the presence or absence of management directly influences the conditions of vegetation in habitats, which are dietary resources for butterflies, differences in management status may lead to differences in butterfly community structure. The management status in each transect was divided into three categories: Rank 1—mowing once a year, Rank 2—mowing stopped several years ago, and Rank 3—mowing stopped several decades ago.
(4) Trampling pressure: Since the trampling pressure directly influences the conditions of vegetations in habitats, which are dietary resources for butterflies, differences in trampling pressure may lead to differences in butterfly community structure. The trampling pressure was divided into two ranks based on whether there was a path used by people along each transect: Rank 0—low trampling pressure, and Rank 1—high trampling pressure).
(5) Distance from the central part of the grassland: As the distance from the center of the grassland increases, the surrounding forest becomes closer, resulting in differences in vegetation structure and landscape structure. As a result, it is expected that the community structure of butterflies, which normally use different dietary plants and breeding sites, will be changed. The distance from the central part of the grassland was classified into four ranks (0, 1, 2, and 3) from the near side to the far side. Table 1 shows the rank values of the five explanatory variables in each transect.

2.4. Data Analysis

To explore the environmental factors driving the butterfly community structure and composition, we used multivariate regression tree analysis (MRT [47]) based on butterfly species abundance data in 11 transects (species-by-sites data set, see Appendix A) as a response variable and the five categorical environmental variables (habitat types, neighboring plant community, management status, trampling pressure, and distance from the central part of the grassland, see Table 1) as explanatory variables. In the MRT settings, the one standard error (SE) rule was used for pruning, and 10 group cross-validation was performed 100 times. MRT is characterized by the ability to handle multiple categorical variables as explanatory variables and are reported to be more accurate than canonical correspondence analysis (CCA) and redundancy analysis (RDA), which are direct gradient analysis methods [47], while MRT is a distribution-free and fully non-parametric data analysis tool, immune from spatial autocorrelation [48,49]. MRT analysis was performed on 19 July 2019 in R software (version 2.12.0 [50]) using the MVPART package [51].
To examine the validity of the species groups discriminated by MRT analysis and the ordination and arrangement of all component species of the butterfly community, we performed principal component analysis (PCA) based on the results of the MRT analysis using R software (version 2.12.0 [50]) using the MVPART package [51]. Eleven survey transects and all butterfly species observed were plotted jointly in a PCA biplot to evaluate the habitat preferences of the species and identify representative species of the groups identified in MRT. In addition, we examined the contribution to the variance in the community of each component of the PCA.
In order to compare the utilization patterns of dietary resources of the representative species of the groups identified in the MRT and PCA analyses, we examined both the type of larval host plants and the species number of adult nectar source plants based on information in the literature [52,53,54,55,56,57,58,59]. Larval host plants were divided into two types (herbaceous and woody plants) based on Unno and Aoyama [52]. The species number of adult nectar source plants was determined based on those described in Fukuda et al. [53,54,55,56]. The representative species of the butterfly groups were examined for Red Listed species based on the Red List 2019 of Japan [60]. The Tukey test, which is a method of multiple comparisons of means to compare three or more groups simultaneously, was used using R to compare the utilization patterns of adult nectar plants among the characteristic species of the three groups (G1, G2, and G3) identified by MRT analysis.

3. Results

In the present study, we recorded a total of 1226 individuals of 48 butterfly species during the whole study period. The annual total number of individuals in each transect in all butterfly species recorded (corresponding to response variables) is shown in Appendix A. The two community parameters (the total numbers of species and individuals) in each transect are shown in Figure 2.
MRT analysis generated a 3-leaved MRT with the residual error (Error) = 0.356, the cross-validation error (CV Error) = 0.795, and the standard error (SE) = 0.166, and showed the total explained variation of 64.4% in the species composition (Figure 3). The explanatory variable discriminating between the two branches in the first node of the 3-leaved MRT was neighboring plant community (nei.com). The group of the first leaf (G1) was characterized by the transect surrounded on both sides by forest (Rank 3), while the other group was characterized by the transect surrounded on one side by grassland and on the other side by forest (Rank 2), or the transect surrounded on both sides by grassland (Rank 1). The explanatory variable discriminating between the two branches in the second node of the MRT was management status (manag). The group of the second leaf (G2) was characterized by the transect with the continuation of mowing once a year (Rank 1) or where mowing was stopped two or three years ago (Rank 2), while the group of the third leaf (G3) was characterized by the transect with mowing stopping at least 10 or more years ago (Rank 3). Thus, transects 5, 6, 7, and 8 belonged to group G1, transects 1, 2, 3, and 4 belonged to the group G2, and transects 9, 10, and 11 belonged to group G3, and the positional relationship between the transects is shown in Figure 4.
The groups (G1, G2, and G3) identified by MRT in Figure 3 and the 48 butterfly component species were plotted jointly in a PCA biplot (Figure 5). The first axis of the PCA explained 42.6% of the total community variation (Table 2), mainly separating the communities surrounded by forests from those surrounded by grasslands. The second axis of the PCA explained 36.1% of the variation (Table 2), mainly separating the communities associated with management from the communities with no management. The first two axes of the PCA accounted largely for 78.8% of the variance in the community. The group G1 discriminated by MRT was distributed on the right side of the first axis (forest-oriented), and the other groups (G2 and G3) were distributed on the left side of the first axis (grassland-oriented). On the other hand, group G3 was distributed on the upper side of the second axis (no management-oriented), and the other groups (G1 and G2) were distributed on the underside of the second axis (with management-oriented). Thus, the PCA also distinctly separated the three groups discriminated by MRT. Pieris spp., Eurema mandarina, Argynnis paphia, Argynnis ruslana, and Argynnis anadyomene were characteristic species of group G1 and are not Red Listed species (Table 3). Minois dryas, Leptalina unicolor, Fabriciana adippe, Plebejus argus, Parnara guttata, Ochlodes venatus, and Gonepteryx maxima were characteristic species of group G2, three of which (L. unicolor, P. argus, and G. maxima) are Red Listed species (Table 3). Fixsenia mera, Brenthis daphne, Ypthima argus, and Inachis io were characteristic species of group G3, one of which (Brenthis daphne) is a Red Listed species (Table 3). According to Figure 2 and Figure 5, the number of butterfly component species was higher in groups G1 and G2 than in group G3.
The type of larval host plants and the number of herbaceous or woody species of adult nectar source plants in each of the characteristic species of the three butterfly groups (G1, G2, and G3) discriminated in the MRT analysis are shown in Table 3. In terms of the larval host plants, the characteristic species of all groups were almost herb (grass)-feeders while in terms of the adult nectar source plants, marked differences were detected among the characteristic species of the three groups. That is, the characteristic species of group G1 used a higher percentage of woody nectar source plants than those of the other two groups, G2 and G3 (Figure 6 and the results of the Tukey test (Table 4)).

4. Discussion

4.1. Determinants of Butterfly Community Structure and Composition

In the present study, we showed through the MRT analysis that the first important determinant of the butterfly community structure is the neighboring plant community. That is, it was found that the differences in the surrounding plant communities of the transects most influenced the determination of the structure and composition of the butterfly community. This result seems to be valid, because many butterflies depend on specific plants for their dietary resources during both the larval and adult stages. In fact, many studies have already shown that plant related factors are the determinants of butterfly community structure and composition at the local level [15,23,25,26,32,61,62,63]. However, although it has been well-investigated that the structure and composition of butterfly communities change with different vegetation (the above studies), few attempts have been made to clarify its function and mechanism. In our study, due to the first determinant (neighboring plant community), the butterfly community was divided into two groups: one of the transects surrounded by forests (G1), and the other of the transects surrounded mainly by grasslands (G2 and G3). Moreover, when the adult dietary habits of the characteristic species of those groups were analyzed, it was found that the species of G2 and G3 mostly utilized herbaceous nectar plants, while those of G1 utilized more woody nectar plants (Figure 6, Table 3 and Table 4). Pocewicz et al. [64] argued for the importance of butterfly resource plant spatial distributions in determining the butterfly population densities. Thus, a mechanism was suggested whereby the difference in the neighboring plant communities causes the difference in the distribution of the nectar plant species of butterflies, and in turn, the difference in the species composition of the butterfly community (i.e., a species group that uses more woody nectar plants (G1) vs. a species group that uses herbaceous nectar plants (G2 or G3)). That is, from our results, it can be considered that the neighboring plant communities functioned in determining the species composition of the butterfly community. In previous studies [34,35,65], it was known that nectar plants for butterflies in and around our study area were used with a bias toward herbs rather than woody plants, and this was linked to the species richness of the species group (G2), which will be discussed below.
Next, we showed that the second important determinant of the butterfly community structure is the management status. There have also been many studies on how human management and the degree of natural or human-caused disturbance greatly influence the determination of the butterfly community structure and composition at the local level [9,22,24,27,29,30,31,66,67,68,69,70,71]. In our study, due to the second determinant (management status), the butterfly community was divided into two groups, one of the transects with the continuation of mowing once a year, or those where mowing was stopped a few years ago (G2) and the other of the transects with mowing stopping quite a long time ago (G3). It is known that the optimal timing and frequency of management promotes plant diversity and richness (e.g., [71,72]). In previous studies at the present study site [34,35], it was found that the differences in the management status of each transect caused those in the number of nectar plants for adult butterflies. That is, we know that the transects with management (mowing) had a larger number of flowering plants (adult nectar resources) than those without management (mowing). Thus, it was suggested that the differences in the management status of the transects made the differences in the number of adult nectar plants among the transects surrounded mainly by grasslands, and therefore, they caused the differences in the structure of the butterfly community. It is noteworthy that the number of butterfly species and their total population abundance were high in group G2 and extremely low in group G3 (Figure 2 and Figure 5). Generally, it is accepted that one of the local factors strongly affecting butterfly diversity and richness is the number of flowering nectar producing plants (e.g., [32,73,74,75]). From these, a mechanism was suggested whereby the difference in management status causes the difference in the abundance of the flowering nectar source plants of butterflies, and in turn, the difference in the species richness and abundance of the butterfly community (i.e., a species group with relatively high species richness and abundance (G2) vs. a species group with relatively low species richness and abundance (G3)). That is, from our results, it can be considered that the management status functioned in determining the species richness and abundance (diversity) of the butterfly community. However, although this study was able to show that annual management (mowing) in late autumn increases butterfly diversity and abundance, it was unable to determine the optimal mowing style such as timing and frequency of mowing, which affects the increase in butterfly diversity. Future research is needed on this point.
The results of the PCA (Figure 5) also strongly support the results and discussion of the MRT analysis above: the first axis of the PCA (PC1: neighboring plant community) and the second axis (PC2: management status) accounted for a cumulative 78.8% of the total community variation, providing evidence that these two factors are key determinants of the butterfly community. The first axis reflected an environmental gradient from grassland to forest, clearly suggesting that it functions to change the species composition of the butterfly community (from grassland to forest species). On the other hand, the second axis reflected an environmental gradient from managed to unmanaged, with more species in each of the G1 and G2 groups and fewer species in the G3 group, clearly suggesting that it serves to change the species richness (diversity) of the butterfly community.
Overall, it can be greatly emphasized that our study explored the determinants of butterfly community structure at the local habitat level and clarified their mechanism of action, and each determinant had a different role (function) in determining the local butterfly community structure. In addition, our results demonstrate that at the local habitat level, the environmental factors that strongly control the distribution and abundance of butterfly food resources such as the present neighboring plant community and management status are the most contributing factors that determine the local butterfly community structure.

4.2. Characteristics of the Butterfly Groups

In the present study, it was found that the butterfly community was composed of three groups (G1, G2, and G3) with different characteristics. The characteristic species of group G1 were featured by larval dietary resources being mostly herbaceous plants, and adult ones being relatively more woody plants (Table 3). The characteristic species of group G2 were featured by both the larval and adult dietary resources being mostly herbaceous plants (Table 3). The characteristic species of group G3 were featured by larval dietary resources being mostly herbaceous plants, and adult ones being slightly woody plants, as the values of P in the Tukey test showed (Table 4).
Based on the above, it was confirmed that the butterfly community in the study area was composed of group G2, which was associated with the grassland in the early successional stage maintained by management, to which representative species such as L. unicolor, F. adippe, and P. argus belonged; group G3, which was associated with the grassland in the late successional stage with no management, to which representative species such as F. mera, B. daphne, and Y. argus belonged, and group G1, which was associated with mixed areas of grassland and forests, to which representative species such as Pieris spp., E. mandarina, and A. paphia belonged. In addition, there may be a group associated only with forest habitats. However, in the present study, it was not possible to set up a survey transect associated only with forests, so the existence of this group is unclear.
Up until now, the species groupings in butterflies have been mainly made and discussed based on their life history characteristics and ecology [12,66,67,70,76,77,78]. However, in comparison to this, there have not been many studies on species groupings based on their habitat structure and successional stage. In this regard, our study revealed the existence of butterfly groups that establish on the gradient of secondary succession from grassland to forest habitats. In Japan, Inoue [79] and Kobayashi et al. [80] reported in detail that the species composition and grouping of butterflies changed clearly along the successive stages of deciduous forest development, supporting the results of this study. In the future, more research is needed to verify whether the species groupings of butterflies along these secondary successional stages have a generality. Furthermore, the analysis of which groups contain many endangered species and are linked to endangered characteristics (e.g., oligo-voltinism, narrow dietary breadth, narrow geographic distribution [5,6]) is an essential issue for the conservation of local biodiversity. In the present study, the largest number of Red Listed butterfly species was present among the characteristic species of group G2, established in the early stage of secondary succession (Table 3). This was in good agreement with the tendency of Japanese Red Listed butterfly species, that is, many of the Red Listed butterfly species are grassland species in rural areas of Japan [60]. As above-mentioned, the G2 species group is strongly linked to management and had a large number of species, so the maintenance of the secondary grassland environment through optimal management is of utmost importance for the conservation of the major endangered butterfly species and butterfly diversity in the region.

4.3. Conservation Implications

As mentioned at the end of the previous section, it was considered essential to maintain the grassland habitats corresponding to the initial stage of secondary succession in order to conserve the Red Listed species and the species richness of the butterfly community, and for that purpose, it is very important to continue with mowing management. Otherwise, as group G3 of this study shows, abandonment of this management promotes secondary succession, reducing Red Listed species and reducing the butterfly species richness (cf. [61]). In general, it is well-known that human activities and management are important for maintaining the high diversity of butterflies in semi-natural grassland (e.g., [69]). On the other hand, the maintenance of vegetation landscape diversity (existence of both grasslands and forests) is considered important for the conservation of butterfly community diversity (species composition). That is, the homogenization of vegetation landscapes will lead to a simplification of butterfly species composition (cf. [81,82]). Overall, it can be concluded that continued mowing management and the maintenance of vegetation landscape diversity are paramount to the conservation of the diversity (species composition and species richness) and endangered species in the local butterfly communities in the region.

5. Conclusions

In conclusion, our study demonstrates that the neighboring plant community and management status are very important environmental factors for determining the structure and composition of butterfly communities at the local habitat level. In particular, the neighboring vegetation was related to the distribution of butterfly dietary resources and contributed a lot in determining the species composition of the community, while the management status was largely related to the amount of dietary resources, affecting the species richness and diversity. Furthermore, it became clear that characteristic species groups were formed along the gradients of these two factors.
Our research was a case study from a locality in Central Japan. Thus, it is hoped that further similar studies will be conducted to explore the generality and consistency of the results and patterns obtained in this study.

Author Contributions

Conceptualization, M.K.; Methodology, M.K. and T.Y.; Software, T.Y.; Validation, M.K. and T.Y.; Data analysis, T.Y.; Investigation, M.K. and T.Y.; Writing—original draft preparation, M.K.; Writing—review and editing, M.K. and T.Y.; Supervision, M.K.; Project administration, M.K.; Funding acquisition, M.K. and T.Y. All authors have read and agreed to the published version of the manuscript.

Funding

The research was supported in part by both Grants-in-Aid for Scientific Research (B) (no. 17310138) and for Scientific Research (C) (no. 20510221) from the Japan Society for the Promotion of Science (JSPS) to M. Kitahara (represents the applicant).

Data Availability Statement

The data presented in this study are available in the manuscript and Appendix A.

Acknowledgments

We thank the members of the Mount Fuji Research Institute of Yamanashi Pref. for their suggestions, help, and cooperation for this study.

Conflicts of Interest

The authors declare no conflicts of interest.

Appendix A. List of Butterfly Species Observed in the Present Study, and the Annual Total Number of Individuals of Each Species in Each Transect, Corresponding to Response Variables

Table A1. List of butterfly species observed in the present study, and the annual total number of individuals of each species in each transect, corresponding to the response variables.
Table A1. List of butterfly species observed in the present study, and the annual total number of individuals of each species in each transect, corresponding to the response variables.
Symbol in the PCATransect (Habitat Type)
Species1 (D)2 (D)3 (C)4 (C)5 (A)6 (A)7 (B)8 (B)9 (E)10 (E)11 (E)Total
Hesperiidae
Daimio tethysb26010 111010005
Choaspes benjaminiib30000 000100001
Leptalina unicolorb2110120 1789121020594
Ochlodes venatusb222513 3245500342
Ochlodes ochraceusb25001 012010016
Potanthus flavusb28100 100000002
Aeromachus inachusb24505 1001010013
Thoressa variab31000 010000001
Pelopidas mathiasb29011 000000002
Pelopidas jansonisb27201 200000005
Parnara guttatab23862 10432511042
Papilionidae
Parnassius citrinariusb1102 2233000013
Papilio xuthusb3000 000020002
Papilio protenorb4000 000010001
Papilio dehaaniib2020 0230310011
Papilio maackiib5000 001000001
Pieridae
Eurema mandarinab18012 518167700157
Gonepteryx maximab195210 2224300030
Colias erateb20004 3001300112
Pieris (melete or japonica) spp. b17014 03230121520096
Lycaenidae
Artopoetes pryerib9000 003000003
Rapala aratab11000 100000001
Fixsenia merab7073 30004962355
Lampides boeticusb12010 000000001
Zizeeria mahab8000 012010004
Celastrina argiolusb13000 000000011
Everes argiadesb10101 001000003
Plebejus argusb62227 176711334486
Curetis acutab14000 010000001
Nymphalidae
Libythea celtisb47000 010010002
Brenthis daphneb35253 0111623731
Argyronome laodiceb36102 8303530126
Argynnis ruslanab34422 01173320034
Argynnis paphiab33201 7131491310060
Argynnis anadyomeneb37210 2672400024
Damora saganab44000 001000001
Fabriciana adippeb321914 814761231176
Speyeria aglajab41000 010100002
Argyreus hyperbiusb42001 001000002
Limenitis camillab39000 011120005
Limenitis glorificab38111 1250000011
Neptis sapphob43000 020000002
Neptis pryerib45001 000000001
Polygonia c-aureumb40000 120110005
Inachis iob46000 000001001
Ypthima argusb16763 34241073756
Minois dryasb15503144 442220162492114295
Melanitis phedimab48000 010000001
Total 10785168 1421651531061454739691226

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Diversity 16 00310 g001
Figure 1. Aerial view of the study area. The eleven transects set in the area are indicated by red lines. The number and habitat type (in parentheses) of each transect are indicated. See the text for details. The aerial photograph was taken in 2011 by Yamanashi Land Improvement Business Association. The photograph was taken two years after this study, but no significant changes in the landscape or vegetation have been observed during this period.
Figure 1. Aerial view of the study area. The eleven transects set in the area are indicated by red lines. The number and habitat type (in parentheses) of each transect are indicated. See the text for details. The aerial photograph was taken in 2011 by Yamanashi Land Improvement Business Association. The photograph was taken two years after this study, but no significant changes in the landscape or vegetation have been observed during this period.
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Figure 2. The total number of individuals and the number of species of all butterflies recorded in each transect during the study period.
Figure 2. The total number of individuals and the number of species of all butterflies recorded in each transect during the study period.
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Figure 3. The 3-leaved multivariate regression tree (MRT) resulting from the best predictive model of the butterfly species/transect data (see Appendix A), constrained by the explanatory variable data set (see Table 1), selected by cross-validation. Terminal nodes show the number of transects in the node, and the barplots show the frequency distribution of species predicted by the node.
Figure 3. The 3-leaved multivariate regression tree (MRT) resulting from the best predictive model of the butterfly species/transect data (see Appendix A), constrained by the explanatory variable data set (see Table 1), selected by cross-validation. Terminal nodes show the number of transects in the node, and the barplots show the frequency distribution of species predicted by the node.
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Figure 4. The positional relationship between the three groups (G1, G2, and G3) identified by the MRT analysis and the transects belonging to the respective groups. The numbers show each transect. Light blue lines indicate paths.
Figure 4. The positional relationship between the three groups (G1, G2, and G3) identified by the MRT analysis and the transects belonging to the respective groups. The numbers show each transect. Light blue lines indicate paths.
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Figure 5. Principal components biplot of the three group means from the best predictive tree in Figure 3. Each color represents the group of the same color in Figure 3. Large circles are group means and small circles are individual transects. The symbol (b + number) in the scatter diagram is an abbreviation given to each butterfly species (see Appendix A for full names), and indicates its position on the scatter diagram. Only the characteristic species of each group are shown with their full names near the symbol.
Figure 5. Principal components biplot of the three group means from the best predictive tree in Figure 3. Each color represents the group of the same color in Figure 3. Large circles are group means and small circles are individual transects. The symbol (b + number) in the scatter diagram is an abbreviation given to each butterfly species (see Appendix A for full names), and indicates its position on the scatter diagram. Only the characteristic species of each group are shown with their full names near the symbol.
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Figure 6. The proportion of woody species of adult nectar plants in each of the characteristic species of the three groups (G1, G2, and G3) identified by MRT analysis. The original data of this figure are shown in Table 3. The statistical significance between the groups by the Tukey test (Table 4) are shown in the figure.
Figure 6. The proportion of woody species of adult nectar plants in each of the characteristic species of the three groups (G1, G2, and G3) identified by MRT analysis. The original data of this figure are shown in Table 3. The statistical significance between the groups by the Tukey test (Table 4) are shown in the figure.
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Table 1. The rank values of the five explanatory variables in each of the eleven transects. See the text for the criteria that determined the respective rank values.
Table 1. The rank values of the five explanatory variables in each of the eleven transects. See the text for the criteria that determined the respective rank values.
Environmental Factor (Explanatory Variable)
Transect (Type)Habitat Type (Habitat)Neighboring Plant Community (nei.com)Management Status (Manag)Trampling Pressure (Tramp)Distance from the Central Part of the Grassland (Dist)
1 (D)11200
2 (D)11200
3 (C)32111
4 (C)32111
5 (A)53113
6 (A)53113
7 (B)43112
8 (B)43112
9 (E)21300
10 (E)21300
11 (E)21300
Table 2. The values of standard deviation, proportion of variance, and cumulative proportion in each component of the PCA.
Table 2. The values of standard deviation, proportion of variance, and cumulative proportion in each component of the PCA.
Component
PC1PC2PC3PC4PC5PC6
Standard deviation16.830 15.501 8.158 4.805 4.388 3.822
Proportion of variance0.426 0.361 0.100 0.035 0.029 0.022
Cumulative proportion0.426 0.788 0.888 0.923 0.952 0.974
Table 3. Type of larval host plants and the number of species of adult nectar plants in each characteristic species of the three butterfly groups (G1, G2, and G3) discriminated in the MRT analysis. ○ indicates applicable items.
Table 3. Type of larval host plants and the number of species of adult nectar plants in each characteristic species of the three butterfly groups (G1, G2, and G3) discriminated in the MRT analysis. ○ indicates applicable items.
SpeciesType of Larval Host Plants (1)No. of Species of Adult Nectar Plants (2)Red Listed Species (3)
Herbaceous PlantsWoody PlantsHerbaceous PlantsWoody Plants
G1
Pieris spp.○ 19 (79.2%)5 (20.8%)
Eurema mandarina ○21 (77.8%)6 (22.2%)
Argynnis paphia○ 26 (72.2%)10 (27.8%)
Argynnis ruslana○ 18 (78.3%)5 (21.7%)
Argynnis anadyomene○ 13 (54.2%)11 (45.8%)
G2
Minois dryas○ 25 (89.3%)3 (10.7%)
Leptalina unicolor○ 22 (95.7%)1 (4.3%)○
Fabriciana adippe○ 24 (77.4%)7 (22.6%)
Plebejus argus○ 16 (100.0%)0 (0%)○
Gonepteryx maxima ○11 (91.7%)1 (8.3%)○
Ochlodes venatus○ 11 (100.0%)0 (0%)
Parnara guttata○ 79 (91.9%)7 (8.1%)
G3
Fixsenia mera ○11 (84.6%)2 (15.4%)
Brenthis daphne○ 15 (83.3%)3 (16.7%)○
Ypthima argus○ 39 (79.6%)10 (20.4%)
Inachis io○ 36 (94.7%)2 (5.3%)
(1) Based on Unno and Aoyama [52]; (2) Based on Fukuda et al. [53,54,55,56]; (3) Based on the Ministry of the Environment of Japan [60].
Table 4. The results of the Tukey test for comparisons among the mean proportions of woody species of adult nectar plants in the characteristic species of the three groups (G1, G2, and G3) identified by MRT analysis.
Table 4. The results of the Tukey test for comparisons among the mean proportions of woody species of adult nectar plants in the characteristic species of the three groups (G1, G2, and G3) identified by MRT analysis.
Species Groups
Compared		EstimateStd. Errorz Value Pr (>|z|)
G2 − G1 == 0−1.3431 0.3472 −3.868 <0.001***
G3 − G1 == 0−0.8343 0.3687 −2.263 0.0609 .
G3 − G2 == 00.5088 0.3952 1.287 0.4014
Signif. codes: ‘***’ 0.001, ‘**’ 0.01, ‘*’ 0.05, ‘.’ 0.1.
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Kitahara, M.; Yasuda, T. Determinants of Butterfly Community Structure and Composition at the Local Habitat Level: Importance of Neighboring Vegetation and Management Status: A Case Study. Diversity 2024, 16, 310. https://doi.org/10.3390/d16060310

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Kitahara M, Yasuda T. Determinants of Butterfly Community Structure and Composition at the Local Habitat Level: Importance of Neighboring Vegetation and Management Status: A Case Study. Diversity. 2024; 16(6):310. https://doi.org/10.3390/d16060310

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Kitahara, Masahiko, and Taisuke Yasuda. 2024. "Determinants of Butterfly Community Structure and Composition at the Local Habitat Level: Importance of Neighboring Vegetation and Management Status: A Case Study" Diversity 16, no. 6: 310. https://doi.org/10.3390/d16060310

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Kitahara, M., & Yasuda, T. (2024). Determinants of Butterfly Community Structure and Composition at the Local Habitat Level: Importance of Neighboring Vegetation and Management Status: A Case Study. Diversity, 16(6), 310. https://doi.org/10.3390/d16060310

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