Introduction

Insects play a critical role in ecosystems and are widely used as indicators of biodiversity (Basset et al., 2012; Samways, 2007; Thomas, 2005). Their distribution and number of individuals are influenced by various factors, including season, forest type, climate, and environmental change, all of which serve as important indicators for evaluating ecosystem health and stability (Kremen et al., 1993). Therefore, accurate assessment and monitoring of insect diversity are essential for establishing policies for conservation and ecosystem management (Andersen, 1999; Didham et al., 2012).

Among various methods for surveying insects, light traps are commonly used to capture nocturnal insects because of their efficiency (Intachat & Woiwod, 1999). Light traps utilize the phototactic behavior of insects and are particularly useful in studies on community structure and seasonal variation of nocturnal species (Frank, 1988). However, survey results may be influenced by trap placement, survey timing and location, and duration of operation (Yela & Holyoak, 1997). Therefore, standardization of trap setup and collection protocols is necessary for clarify the differences in insect diversity by season and forest type (Thomas, 2005).

The bucket light trap is advantageous because of its lightweight design, allowing for easy installation and high sampling efficiency across various environments (Brown et al., 2000). This study was conducted as part of a methodological improvement initiative for the terrestrial insect components of the National Ecosystem Survey. It aimed to compare insect diversity according to season and forest type using bucket light traps and to provide guidelines for optimal trap deployment in terms of timing and location. The results of this study are expected to serve as a valuable foundation for standardizing insect monitoring and evaluating ecosystem changes.

Materials and Methods

Survey period and sites

Surveys were conducted in spring (May), summer (July), and autumn (September). Forest types were categorized as coniferous, deciduous, and mixed. The study was conducted in the areas surrounding Moaksan and Guseongsan in Gimje, Jeollabuk-do. For each forest type, three surveys were conducted per season, totaling nine surveys (27 traps; Table 1).

Survey method

The surveys followed the National Ecosystem Survey guidelines. Bucket light traps (BL, 12 W) were operated using timers set from 20:00 to 24:00, and samples were collected the following morning. The collected specimens were stored on-site and transported to the laboratory for identification.

Sample processing and statistical analysis

The specimens were identified by taxonomic experts and were categorized into the orders Lepidoptera, Coleoptera, Hymenoptera, Diptera, and Hemiptera. One-way analysis of variance (ANOVA) was used to compare the number of species and individuals by season and forest type, followed by Scheffe’s post hoc test. Statistical significance was set at P<0.05, and SPSSWIN v.28.0 (IBM Co., Armonk, NY, USA) was used for all analyses.

Results

Overall summary

In total, 13,531 individuals representing 934 species from 129 families and 14 orders were recorded. The mixed forest yielded 720 species and 5,139 individuals across 13 orders and 99 families; the deciduous forest yielded 645 species and 4,559 individuals from 11 orders and 97 families; and the coniferous forest yielded 580 species and 3,833 individuals from 11 orders and 87 families.

In the coniferous forest, 222 species and 815 individuals were recorded in spring, 341 species and 1,673 individuals in summer, and 280 species and 1,345 individuals in autumn. In the mixed forest, 333 species and 1,112 individuals were recorded in spring, 440 species and 2,452 individuals in summer, and 336 species and 1,575 individuals in autumn. In the deciduous forest, 238 species and 1,091 individuals were found in spring, 398 species and 2,090 individuals in summer, and 306 species and 1,378 individuals in autumn. In all seasons, the mixed forest had the highest numbers of species and individuals, whereas the coniferous forest had the lowest numbers (Table 2).

Seasonal status of the taxonomic groups

In spring, 432 species and 3,018 individuals were recorded, with Lepidoptera being dominant (315 species and 2,413 individuals), followed by Coleoptera (59 species and 315 individuals), Hymenoptera (28 species and 70 individuals), Hemiptera (12 species and 32 individuals), and Trichoptera (6 species and 130 individuals).

In summer, 567 species and 6,215 individuals were recorded, with Lepidoptera being dominant (427 species and 5,037 individuals), followed by Coleoptera (59 species and 317 individuals), Hymenoptera (34 species and 661 individuals), Hemiptera (25 species and 134 individuals), and Neuroptera (7 species and 20 individuals).

In autumn, 474 species and 4,298 individuals were recorded, with Lepidoptera being dominant (310 species and 3,391 individuals), followed by Hymenoptera (63 species and 400 individuals), Diptera (28 species and 108 individuals), Coleoptera (28 species and 76 individuals), and Hemiptera (18 species and 73 individuals; Table 3).

Forest type status of the taxonomic groups

In the coniferous forest, 580 species and 3,833 individuals were recorded. Lepidoptera was dominant (406 species and 2,915 individuals), followed by Coleoptera (68 species and 202 individuals), Hymenoptera (42 species and 425 individuals), Hemiptera (21 species and 104 individuals), and Diptera (16 species and 41 individuals).

In the mixed forest, 720 species and 5,139 individuals were recorded. Lepidoptera was dominant (540 species and 4,207 individuals), followed by Coleoptera (68 species and 276 individuals), Hymenoptera (48 species and 417 individuals), Hemiptera (22 species and 55 individuals), and Trichoptera (15 species and 83 individuals).

In the deciduous forest, 645 species and 4,559 individuals were recorded. Lepidoptera was dominant (471 species and 3,719 individuals), followed by Coleoptera (68 species and 230 individuals), Hymenoptera (39 species and 289 individuals), Hemiptera (26 species and 80 individuals), and Diptera (17 species and 52 individuals; Table 4).

Statistical analysis

Seasonal comparison

The average number of species was highest in summer (265.67), followed by autumn (204.33) and spring (156.22). Similarly, the average number of individuals was highest in summer (690.56), followed by autumn (477.56) and spring (335.33; Table 5).

One-way ANOVA indicated significant differences among seasons in the number of species (F=22.319, P<0.05) and individuals (F=25.476, P<0.05; Table 6).

Scheffe’s post hoc tests revealed significant differences in the number of species between spring and summer (P<0.001), spring and autumn (P<0.025), and summer and autumn (P<0.004). Significant differences were also observed in the number of individuals between spring and summer (P<0.001), spring and autumn (P<0.031), and summer and autumn (P<0.001; Table 7).

Comparison among forest types

The average number of species was highest in the mixed forest (243.89), followed by deciduous forests (203.89) and coniferous forests (178.44). Similarly, the average number of individuals was highest in mixed forest (571.00), followed by deciduous forests (506.56) and coniferous forests (425.89; Table 8).

One-way ANOVA showed significant difference in the number of species by forest type (F=3.691, P<0.040); however, no significant differences were noticed in the number of individuals (F=1.520, P=0.239; Table 9).

Post hoc tests for the number of species indicated significant difference between coniferous and mixed forests (P<0.042), whereas no significant differences were observed between coniferous and deciduous forests or deciduous and mixed forests. No significant differences were observed in the number of individuals among any of the forest-type pairs (Table 10).

Discussion

Seasonal analysis showed that summer had the highest number of species (567 species) and individuals (6,215 individuals), followed by autumn (474 species and 4,298 individuals) and spring (432 species and 3,018 individuals). Compared to spring and autumn, summer showed approximately 31.2% and 19.6% higher number of species and 105.9% and 44.6% higher number of individuals than those in autumn and spring, respectively.

By forest type, the mixed forest had the highest number of species (720 species) and individuals (5,139 individuals), followed by the deciduous forest (645 species and 4,559 individuals) and coniferous forest (580 species and 3,833 individuals). Compared to coniferous and deciduous forests, the mixed forest showed approximately 24.1% and 11.6% more species and 34.1% and 12.7% more individuals respectively.

Statistical analysis by season revealed significant differences in both the number of species and number of individuals. Specifically, significant differences were observed between spring and summer (P<0.001), spring and autumn (P<0.025), and summer and autumn (number of species: P<0.004; number of individuals: P<0.001). These findings suggest that surveys conducted in summer are more effective for identifying insect diversity. Therefore, it is advisable to include summer sampling in biodiversity monitoring programs (Samways, 2007; Yela & Holyoak, 1997). In particular, when conducting multi-season surveys, including summer as a core period can enhance compliance with the National Ecosystem Survey guidelines and improve the monitoring of ecosystem changes (Didham et al., 2012; Kremen et al., 1993).

Regarding forest types, a significant difference in the number of species was observed between coniferous and mixed forests, but no significant differences were found between other forest types. These findings suggests that mixed forests are more favorable for surveying insect communities and securing higher diversity (Andersen, 1999; Brown et al., 2000). Consequently, mixed forests with both coniferous and deciduous components are recommended as priority sites for insect diversity assessments.

Additionally, variations in the number of species and individuals were observed even within the same forest type, depending on the diversity of vegetation and understory development. These results align with previous studies emphasizing the importance of understory vegetation for insect diversity (Thomas, 2005). Therefore, in mixed forests, survey sites with well-developed understory vegetation, including shrubs and herbaceous plants, should be prioritized for more effective biodiversity assessment.

In conclusion, this study clearly demonstrated differences in nocturnal insect diversity by season and forest type. The findings provide a scientific basis for biodiversity monitoring and ecosystem assessment. In particular, the effectiveness of surveys conducted in mixed forests among forest types and during summer among seasons was validated, highlighting the potential for standardizing insect monitoring protocols and improving national survey guidelines.

However, this study has several limitations. First, the results were based on data collected during a single year, which may not capture inter-annual variability in insect diversity. Second, forest types were classified without accounting for fine-scale habitat variables such as understory structure, canopy cover, and vegetation age, which could influence insect assemblages.

Future studies should incorporate multi-year data and include more detailed assessments of microhabitat characteristics. Additionally, identifying indicator species for each forest type would strengthen biodiversity assessment frameworks and contribute to more effective insect-based conservation strategies. These efforts would support ongoing improvements to national biodiversity monitoring guidelines and facilitate evidence-based conservation planning.

Author Contributions

Conceptualization: YGH, EJH. Data curation: YGH, STY. Formal analysis: YGH. Funding acquisition: EJH, STY. Investigation: YGH, STY. Methodology: YGH. Project administration: EJH, STY. Resources: EJH, STY. Supervision: EJH. Validation: YGH, EJH. Visualization: YGH. Writing – original draft: YGH. Writing – review & editing: YGH, EJH.

Conflict of Interest

The authors declare that they have no competing interests.

Funding

This work was supported by a grant from the National Institute of Ecology (NIE) funded by the Ministry of Environment (MOE) of the Republic of Korea (NIE-A-2023-01).

References

1 

Andersen, A.N. (1999) My bioindicator or yours? Making the selection Journal of Insect Conservation, 3, 61-64 .

2 

Basset, Y., Cizek, L., Cuénoud, P., Didham, R.K., Guilhaumon, F., Missa, O., et al. (2012) Arthropod diversity in a tropical forest Science, 338, 1481-1484 .

3 

Brown, N., Jennings, S., Wheeler, P., & Nabe-Nielsen, J. (2000) An improved method for the rapid assessment of forest understorey light environments Journal of Applied Ecology, 37, 1044-1053 .

4 

Didham, R.K., Kapos, V., & Ewers, R.M. (2012) Rethinking the conceptual foundations of habitat fragmentation research Oikos, 121, 161-170 .

5 

Frank, K.D. (1988) Impact of outdoor lighting on moths: an assessment Journal of the Lepidopterists' Society, 42, 63-93 .

6 

Intachat, J., & Woiwod, I.P. (1999) Trap design for monitoring moth biodiversity in tropical rainforests Bulletin of Entomological Research, 89, 153-163 .

7 

Kremen, C., Colwell, R.K., Erwin, T.L., Murphy, D.D., Noss, R.F., & Sanjayan, M.A. (1993) Terrestrial arthropod assemblages: their use in conservation planning Conservation Biology, 7, 796-808 .

8 

Samways, M.J. (2007) Insect conservation: a synthetic management approach Annual Review of Entomology, 52, 465-487 .

9 

Thomas, J.A. (2005) Monitoring change in the abundance and distribution of insects using butterflies and other indicator groups Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences, 360, 339-357 . Article Id (pmcid)

10 

Yela, J.L., & Holyoak, M. (1997) Effects of moonlight and meteorological factors on light and bait trap catches of noctuid moths (Lepidoptera: Noctuidae) Environmental Entomology, 26, 1283-1290 .

Tables
Table 1

Survey period and sites of bucket light traps

Survey round Survey date Season Forest type Coordinates
1st May 29, 2023 Spring Coniferous N 35.72082, E 127.0663
Mixed N 35.72187, E 127.0639
Deciduous N 35.71802, E 127.0655
2nd May 30, 2023 Spring Coniferous N 35.72537, E 127.0648
Mixed N 35.72651, E 127.0658
Deciduous N 35.72775, E 127.0682
3rd May 31, 2023 Spring Coniferous N 35.71260, E 127.0567
Mixed N 35.71271, E 127.0590
Deciduous N 35.71223, E 127.0600
4th July 11, 2023 Summer Coniferous N 35.72082, E 127.0663
Mixed N 35.72187, E 127.0639
Deciduous N 35.71802, E 127.0655
5th July 12, 2023 Summer Coniferous N 35.72537, E 127.0648
Mixed N 35.72651, E 127.0658
Deciduous N 35.72775, E 127.0682
6th July 30, 2023 Summer Coniferous N 35.74209, E 127.0274
Mixed N 35.74589, E 127.0296
Deciduous N 35.74667, E 127.0272
7th September 14, 2023 Autumn Coniferous N 35.72082, E 127.0663
Mixed N 35.72187, E 127.0639
Deciduous N 35.71802, E 127.0655
8th September 15, 2023 Autumn Coniferous N 35.72537, E 127.0648
Mixed N 35.72651, E 127.0658
Deciduous N 35.72775, E 127.0682
9th September 16, 2023 Autumn Coniferous N 35.74209, E 127.0274
Mixed N 35.74589, E 127.0296
Deciduous N 35.74667, E 127.0272
Table 2

Overall summary of the survey results

Season Taxa Coniferous forest Mixed forest Deciduous forest Total
Spring Order 9 9 11 12
Family 53 57 62 78
Species 222 333 238 432
Individual 815 1,112 1,091 3,018
Summer Order 10 10 9 12
Family 53 62 57 84
Species 341 440 398 567
Individual 1,673 2,452 2,090 6,215
Autumn Order 8 8 8 9
Family 46 47 41 65
Species 280 336 306 469
Individual 1,345 1,575 1,378 4,298
Total 11 orders, 87 families, 580 species, and 3,833 individuals 13 orders, 99 families, 720 species, and 5,139 individuals 11 orders, 97 families, 645 species, and 4,559 individuals 14 orders, 129 families, 934 species, and 13,531 individuals
Table 3

Overview of taxonomic groups by season

Taxa Spring Summer Autumn Total




No. of
species
No. of
individuals
No. of
species
No. of
individuals
No. of
species
No. of
individuals
No. of
species
No. of
individuals
Ephemeroptera 1 2 2 9 - - 3 11
Mantodea - - - - 2 2 2 2
Plecoptera 2 21 2 14 - - 4 35
Orthoptera 1 2 3 4 6 79 10 85
Phasmatodea 1 1 1 1 - - 2 2
Embioptera - - 1 1 - - 1 1
Hemiptera 12 32 25 134 18 73 45 239
Megaloptera 1 3 - - - - 1 3
Neuroptera 1 1 7 20 2 11 7 32
Coleoptera 59 315 59 317 28 76 110 708
Hymenoptera 28 70 34 661 63 400 87 1,131
Diptera 5 28 3 7 28 108 32 143
Trichoptera 6 130 3 10 17 158 23 298
Lepidoptera 315 2,413 427 5,037 310 3,391 607 10,841
Total 432 3,018 567 6,215 474 4,298 934 13,531
[i]

-, not applicable.

Table 4

Overview of taxonomic groups by forest type

Taxa Coniferous forest Mixed forest Deciduous forest Total




No. of species No. of
individuals
No. of
species
No. of
individuals
No. of
species
No. of
individuals
No. of
species
No. of
individuals
Ephemeroptera 1 2 - - 2 9 3 11
Mantodea - - 2 2 - - 2 2
Plecoptera 4 20 3 5 2 10 4 35
Orthoptera 7 38 2 32 6 15 10 85
Phasmatodea - - 2 2 - - 2 2
Embioptera - - 1 1 - - 1 1
Hemiptera 21 104 22 55 26 80 45 239
Megaloptera 1 1 1 1 1 1 1 3
Neuroptera 3 11 3 8 5 13 7 32
Coleoptera 68 202 68 276 68 230 110 708
Hymenoptera 42 425 48 417 39 289 87 1,131
Diptera 16 41 13 50 17 52 32 143
Trichoptera 11 74 15 83 8 141 23 298
Lepidoptera 406 2,915 540 4,207 471 3,719 607 10,841
Total 580 3,833 720 5,139 645 4,559 934 13,531
[i]

-, not applicable.

Table 5

Descriptive statistics of the number of species and individuals by season

Factors Mean Standard deviation Standard
error
Minimum Maximum
Number of species Spring 156.22 26.00 8.67 121.00 206.00
Summer 265.67 48.27 16.09 199.00 347.00
Autumn 204.33 25.17 8.39 180.00 253.00
Total 208.74 56.60 10.89 121.00 347.00
Number of individuals Spring 335.33 57.12 19.04 246.00 415.00
Summer 690.56 152.44 50.81 473.00 991.00
Autumn 477.56 85.87 28.62 363.00 598.00
Total 501.15 180.42 34.72 246.00 991.00
Table 6

Analysis of variance for number of species and individuals by season

Factors Sum of squares df Mean square F P-value
Number of species Between groups 54,163.630 2 27,081.815 22.319 <0.05
Within groups 29,121.556 24 1,213.398 - -
Total 83,285.185 26 - - -
Number of individuals Between groups 575,336.963 2 287,668.481 25.476 <0.05
Within groups 271,000.444 24 11,291.685 - -
Total 846,337.407 26 - - -
[i]

-, not applicable.

Table 7

Scheffe’s post hoc multiple comparison for number of species and individuals by season

Dependent variable (I) Season (J) Season Mean difference
(I–J)
Std. error P-value 95% Confidence interval

Lower bound Upper bound
Number of species Spring Summer –109.44444 16.42084 <0.001 –152.2826 –66.6063
Autumn –48.11111 16.42084 <0.025 –90.9492 –5.2730
Summer Spring 109.44444 16.42084 <0.001 66.6063 152.2826
Autumn 61.33333 16.42084 <0.004 18.4952 104.1714
Autumn Spring 48.11111 16.42084 <0.025 5.2730 90.9492
Summer –61.33333 16.42084 <0.004 –104.1714 –18.4952
Number of individuals Spring Summer –355.22222 50.09255 <0.001 –485.9019 –224.5426
Autumn –142.22222 50.09255 <0.031 –272.9019 –11.5426
Summer Spring 355.22222 50.09255 <0.001 224.5426 485.9019
Autumn 213.00000 50.09255 <0.001 82.3203 343.6797
Autumn Spring 142.22222 50.09255 <0.031 11.5426 272.9019
Summer –213.00000 50.09255 <0.001 –343.6797 –82.3203
Table 8

Descriptive statistics of number of species and individuals by forest type

Factors Mean Standard deviation Standard error Minimum Maximum
Number of species Coniferous forest 178.44 36.64 12.21 121.00 228.00
Mixed forest 243.89 62.30 20.77 165.00 347.00
Deciduous forest 203.89 52.33 17.44 132.00 290.00
Total 208.74 56.60 10.89 121.00 347.00
Number of individuals Coniferous forest 425.89 146.84 48.95 246.00 622.00
Mixed forest 571.00 218.93 72.98 327.00 991.00
Deciduous forest 506.56 156.23 52.08 328.00 788.00
Total 501.15 180.42 34.72 246.00 991.00
Table 9

Analysis of variance for number of species and individuals by forest type

Factors Sum of squares df Mean square F P-value
Number of species Between groups 19,591.185 2 9,795.593 3.691 <0.040
Within groups 63,694.000 24 2,653.917 - -
Total 83,285.185 26 - - -
Number of individuals Between groups 95,152.296 2 47,576.148 1.520 0.239
Within groups 751,185.111 24 31,299.380 - -
Total 846,337.407 26 - - -
[i]

-, not applicable.

Table 10

Scheffe’s post hoc multiple comparison for number of species and individuals by forest type

Dependent
variable
(I) Forest type (J) Forest type Mean
difference
(I–J)
Standard
error
P-value 95% Confidence interval

Lower
bound
Upper
bound
Number of species Coniferous forest Mixed forest –65.44444 24.28496 <0.042 –128.7982 –2.0907
Deciduous forest –25.44444 24.28496 0.585 –88.7982 37.9093
Mixed forest Coniferous forest 65.44444 24.28496 <0.042 2.0907 128.7982
Deciduous forest 40.00000 24.28496 0.277 –23.3537 103.3537
Deciduous forest Coniferous forest 25.44444 24.28496 0.585 –37.9093 88.7982
Mixed forest –40.00000 24.28496 0.277 –103.3537 23.3537
Number of individuals Coniferous forest Mixed forest –145.11111 83.39915 0.240 –362.6798 72.4576
Deciduous forest –80.66667 83.39915 0.632 –298.2354 136.9021
Mixed forest Coniferous forest 145.11111 83.39915 0.240 –72.4576 362.6798
Deciduous forest 64.44444 83.39915 0.745 –153.1243 282.0132
Deciduous forest Coniferous forest 80.66667 83.39915 0.632 –136.9021 298.2354
Mixed forest –64.44444 83.39915 0.745 –282.0132 153.1243