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.
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).
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 | |||
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 | |||
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 | - | - | - | |
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 | - | - | - | |
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 | ||