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. 2026 Jul 27;10:100420. doi: 10.1016/j.crpvbd.2026.100420

Comparative effectiveness of UV-LED light and CO2 using a 3D-printed suction trap for Culicoides surveillance and bluetongue virus detection in central Thailand

Bunlue Kornmatitsuk a, Yuki Fujisawa a, Thanyaporn Homat b, Sirichai Eardmusic c, Ladawan Sariya d, Sudsaijai Kornmatitsuk a,⁎
PMCID: PMC13469804  PMID: 42597933

Abstract

This study aimed to (i) compare the effectiveness of three attractants (UV-LED light, CO2-only, and UV-LED light + CO2) using a 3D-printed suction trap for Culicoides surveillance, and (ii) determine the presence and circulating serotypes of bluetongue virus (BTV) in Culicoides species in central Thailand. A total of 12474 Culicoides specimens representing 21 species were collected over nine nights. Overall, UV-LED light and UV-LED light + CO2 traps yielded comparable abundance and species diversity, and both outperformed the CO2-only trap (P < 0.001). The five most abundant species were C. oxystoma, C. imicola, C. guttifer, C. peregrinus, and C. huffi. All five species, except C. guttifer, were also more abundant in UV-LED light-based traps than in the CO2-only trap (P < 0.001), and this pattern was consistent for nulliparous and parous females (P < 0.05). Only C. imicola showed a higher parity rate in UV-LED light traps than in the CO2-only trap (P < 0.05). Of 154 pools tested, 53 (34.4%) were BTV-positive; 27 from C. imicola, 20 from C. oxystoma, 5 from C. peregrinus, and 1 from C. huffi. One pool of C. imicola exhibited a relatively high virus titer (Ct value = 25.7). Five serotypes (BTV-2, BTV-4, BTV-7, BTV-12, and BTV-15) were identified, with BTV-15 newly reported. In conclusion, a 3D-printed suction trap baited with UV-LED light alone effectively collected Culicoides spp. without reducing abundance or species diversity, while the co-circulation of multiple BTV serotypes highlights the need to strengthen bluetongue control measures in the country.

Keywords: Culicoides, Vector surveillance, UV-LED light, CO2, 3D-printed suction trap, Bluetongue virus

Graphical abstract

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Highlights

  • •

    First assessment of UV-LED light and CO2 as attractants for BTV vectors in Thailand.

  • •

    No additional effect of CO2 on Culicoides spp. abundance and diversity was observed.

  • •

    First evidence of a direct association between BTV and Culicoides imicola in Southeast Asia.

  • •

    BTV-2, BTV-4, BTV-7, BTV-12, and BTV-15 co-circulated in Thailand, with the first detection of BTV-15.

1. Introduction

Bluetongue virus (BTV), which belongs to the genus Orbivirus (family Sedoreoviridae), is the causative agent of bluetongue (BT), an enzootic, infectious, non-contagious, arthropod-borne viral disease affecting ruminant and camelid species. Cattle are considered the primary reservoir of BTV because infections in this species are typically subclinical and associated with prolonged viremia of several months (MacLachlan et al., 2009; Brown-Joseph et al., 2017). Bluetongue can result in high morbidity and mortality in susceptible animals, such as sheep. In Thailand, no clinical BT cases or outbreaks among ruminant populations have been reported, but high seroprevalence of BTV infection was commonly observed at the herd and individual levels (Homat et al., 2025). However, even subclinical infections can have substantial economic impacts, including a loss of body condition, reduced milk yield, decreased fertility, and animal trade restriction (Saminathan et al., 2020).

Bluetongue virus is transmitted exclusively by certain species of Culicoides (Diptera: Ceratopogonidae) biting midges. Approximately 30 of the 1254 species of Culicoides across the world have been incriminated to varying degrees in the transmission (Meiswinkel et al., 2004). Distinct BTV serotypes are also vectored by specific Culicoides species and occur in specific geographical regions (Federici et al., 2019). In Malaysia and Indonesia, several potential Culicoides vector species have been reported, including C. fulvus, C. actoni, C. wadai, and C. brevitarsis (Daniels et al., 2004). More recently, Fujisawa et al. (2021) reported C. imicola as a potential BTV vector in Thailand, along with C. orientalis, C. oxystoma, and C. fulvus. Nevertheless, information on effective attractants for surveillance of livestock-associated Culicoides species remains limited in Thailand and across the region.

Ultraviolet light-emitting diode (UV-LED) light traps have been widely used to collect Culicoides biting midges because of their low cost, practicality, and efficiency (Hope et al., 2015). Recently, Fujisawa et al. (2023) demonstrated that a 3D-printed suction trap (Mahidol University (MU) UV-LED light trap) showed good performance compared with the standard UV-LED Centers for Disease Control and Prevention (CDC) light trap (BioQuip 2770, BioQuip Inc., Rancho Dominguez, CA, USA). However, many studies have shown that trap types or attractants (with different light sources such as UV tube or UV-LED, or CO2 alone) can greatly influence the abundance and diversity of Culicoides species (Venter et al., 2009; Becker et al., 2021; Brown-Joseph et al., 2022). Previous studies reported that the addition of CO2 increased the abundance of Culicoides biting midges collected (Sloyer et al., 2019; Becker et al., 2021; Zhang and Mathias, 2022). To assess species richness and diversity, Sloyer et al. (2019) suggested that UV-LED light traps with or without CO2 can provide a representative sample of Culicoides species. Bellis and Reid (1996) observed that UV light traps were shown to produce a realistic representation of the age structure of C. wadai but introduced bias in collections of C. brevitarsis. This suggests that estimates of parity, as reflected by parity rates in light-trap collections, may need to be assessed on a species-specific basis.

Furthermore, Becker et al. (2021) reported that Culicoides specimens of different species infected with BTV were not repelled by a UV-light trap; however, the data did not exclude the possibility that BTV may cause unanticipated behavioral effects or alter the light orientation of BTV-infected midges of species other than C. sonorensis (McDermott et al., 2015). Therefore, investigation of whether other potential vector species exhibit altered responses to UV light remains warranted. In addition, the importance of using sensitive surveillance methods for both Culicoides sampling and virus detection in epidemiological studies has also been emphasized (Mayo et al., 2012).

Given these knowledge gaps, the present study aimed to (i) compare the effectiveness of three attractants (UV-LED light, CO2-only, and UV-LED light + CO2) using a 3D-printed suction trap for Culicoides surveillance, and (ii) determine the presence of BTV and circulating serotypes in Culicoides species collected in the study area.

2. Materials and methods

2.1. Experimental design and study area

The study area was selected based on the high bluetongue (BT) prevalence reported by local veterinary services. Therefore, three herd locations (sampling sites 1–3) were chosen (Fig. 1). A field experiment using three replicates of a 3 × 3 Latin square design was conducted to compare three attractants for collecting Culicoides species. The experiment was carried out over nine nights between 28 November and 15 December 2022 (Table 1), with attractants rotated systematically among the three sites to minimize potential confounding effects. Meteorological data, including ambient temperature (°C), relative humidity (%), wind speed (m/s), and rainfall (mm), were obtained at 3-h intervals during the present study from the Hua Hin Weather Observing Station, located approximately 11 km from the sampling sites (Supplementary Fig. S1).

Fig. 1.

Fig. 1

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Map of the Culicoides sampling sites (1–3) in the present study and the BTV-prevalent area: Cha-am and Tha Yang Districts, Phetchaburi Province (light gray) and Hua Hin District, Prachuap Khiri Khan Province (dark gray).

Table 1.

Details of sampling sites (1–3) and collection dates.

Sampling site Location Latitude/Longitude Altitude (m) Collection date Farm animals
1 Cha-am Sub-district, Cha-am District, Phetchaburi Province 12°40′25.5714″N, 99°55′24.4776″E 32 28 Nov.-15 Dec. 2022 28 dairy cattle and 30 goats
2 Sam Phraya Sub-district, Cha-am District, Phetchaburi Province 12°39′33.8322″N, 99°54′44.8200″E 52 28 Nov.-15 Dec. 2022 100 dairy cattle
3 Sam Phraya Sub-district, Cha-am District, Phetchaburi Province 12°40′5.1204″N, 99°53′36.4482″E 76 28 Nov.-15 Dec. 2022 100 dairy cattle and 30 goats
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2.2. Collection methods

Culicoides biting midges were collected using a 3D-printed suction trap (MU UV-LED light trap; Supplementary Fig. S2). Its design and fabrication were previously described by Fujisawa et al. (2023). Three attractants were evaluated: (i) UV-LED light; (ii) CO2-only (without a light source); and (3) UV-LED light + CO2. One trap with an assigned attractant was set up per farm per night from 17:00 to 7:00 h, approximately one hour before sunset to one hour after sunrise. Each trap was suspended from the roof of an open-sided barn at a height of 1.5 m above ground, adjacent to stables, and powered by a 12-V battery. For CO2-based traps, 1 kg of dry ice was placed in a thermos flask above the trap with the spout open for gradual CO2 release. Specimens were preserved in 70% ethanol and stored at −20 °C throughout sorting and morphological identification.

2.3. Sorting and morphological identification

Culicoides specimens were separated from other insects, identified, and counted. Each specimen was identified to species level based on wing patterns and gross morphological features using relevant keys (Wirth and Hubert, 1989; Dyce et al., 2007; Nandi and Mazumdar, 2014). The specimens were then classified according to physiological status based on abdominal characteristics as blood-fed, nulliparous (unpigmented), parous (pigmented), gravid females, and males (Dyce, 1969). Unpigmented and pigmented individuals belonging to the subgenera Hoffmania and Trithecoides are indistinguishable under a stereomicroscope; thus, they were simply classified as non-blood-fed females. To avoid the inclusion of nulliparous individuals from these subgenera in the BTV analysis, they were excluded. Sorting and morphological identification were completed within 3 months after collection. In the absence of transovarial transmission for BTV (Nunamaker et al., 1990; Osborne et al., 2015), only parous and gravid females were selected, transferred to phosphate-buffered saline (PBS), and stored at −80 °C until RNA extraction and subsequent viral analysis.

2.4. Sample pooling and RNA extraction

Parous and gravid midges of morphologically identified species were selected from the relatively abundant species to form insect pools. The pool size varied from 19 to 23 insects depending on the obtained samples (Mayo et al., 2012). Each pool was transferred into a 500 μl PBS solution containing 20 μg/ml of gentamicin and 2.5 μg/ml of amphotericin B, and ground with a mortar and pestle. Total RNA was extracted with TRIzol™ Reagent (Life Technologies Corporation, Carlsbad, CA, USA) following the manufacturer’s instructions.

2.5. Pan-reactive qRT-PCR for BTV detection

The extracted RNA samples were tested for the presence of BTV by pan-reactive qRT-PCR using specific primers and probe, according to Hofmann et al. (2008). The assay was performed following the standard protocol described by Homat et al. (2025). A commercially synthesized double-stranded oligonucleotide (Bionics Co., Ltd., Seoul, South Korea) was used as a positive control. Results were expressed as cycle threshold (Ct) values, defined as the cycle at which the amplification curve crossed the 0.05 threshold. Samples with Ct values ≥ 40 or no Ct value were considered negative (Batten et al., 2015; Kirkland et al., 2022; Homat et al., 2025).

2.6. BTV serotyping

To obtain sufficient sample volume, BTV-positive samples detected by pan-reactive qRT-PCR (Ct value < 40) were pooled according to the attractant used and then subjected to BTV serotyping using two serotype-specific qRT-PCR assays. An in-house singleplex qRT-PCR assay was used to detect 7 serotypes (BTV-2, BTV-3, BTV-4, BTV-7, BTV-12, BTV-16, and BTV-21) according to Homat et al. (2025). In addition, a multiplex serotype-specific qRT-PCR assay was employed to detect 11 serotypes (BTV-1, BTV-2, BTV-3, BTV-5, BTV-7, BTV-9, BTV-12, BTV-15, BTV-20, BTV-21, and BTV-23) across four multiplex panels, with BTV-4 detected by singleplex qRT-PCR (Kirkland et al., 2022). The primers and probes for both serotype-specific qRT-PCR assays are listed in Supplementary Tables S1 and S2. Each positive control for a specific BTV serotype consisted of a commercially synthesized double-stranded oligonucleotide (Bionics Co., Ltd., Seoul, South Korea). Pools with Ct values ≥ 40 or no Ct value were considered negative.

2.7. Statistical analysis

Data analyses were performed using R v.4.5.3 (R Core Team, 2026) with RStudio v.2026.1.2.418 (Posit Team, 2026). For visualization, all Culicoides spp. abundance data were log-transformed [log(count + 1)]; however, raw abundance data were used for statistical analyses.

To investigate the effect of attractant on total Culicoides spp. abundance and species distribution, generalized linear mixed models (GLMMs) with a log link function were fitted using the “glmer” and “glmer.nb” functions in the lme4 package (v.2.0-1) (Bates et al., 2015), with Poisson distribution and negative binomial distribution applied depending on the level of dispersion in the data. The models were fitted by maximum likelihood using the Laplace approximation. Day of collection was included as a random effect, while attractant and location were modeled as fixed effects. Meteorological variables were evaluated but not retained in the final models, as their variation was effectively accounted for by the day of collection. Final models were selected by comparing candidate models with different combinations of explanatory (fixed) variables using the Akaike information criterion (AIC), and the model showing the lowest AIC was chosen for each data group. Differences in the number of Culicoides collected among attractants were subsequently assessed using Tukey-adjusted pairwise comparisons with the “glht” function in the multcomp package (v.1.4-30) (Hothorn et al., 2008). Statistical significance was considered at P < 0.05.

Species diversity indices were calculated using the Shannon’s index (H′) and Simpson’s index of diversity (1 − D), based on species abundance data using the diversity function in the vegan package (v.2.7-3) (Oksanen et al., 2026). Differences in diversity among attractants were assessed using linear mixed models (LMMs) with an identity link, fitted using the “lmer” function in the lme4 package (v.2.0-1) (Bates et al., 2015). Model structure, selection, and pairwise comparisons followed the same approach as for the abundance analyses. Model assumptions were assessed by examining residual diagnostics, including the Shapiro-Wilk test for normality.

Parity rates were modeled using binomial GLMMs with a logit link function, fitted using the “glmer” function in the lme4 package (v.2.0-1) (Bates et al., 2015). The response variable was specified as the counts of parous and non-parous individuals, where parous counts were defined as the sum of parous, gravid, and blood-fed females. Model structure, selection, and pairwise comparisons followed the same approach as for the abundance analyses. Predicted parity rates were obtained using the “emmeans” function in the emmeans package (v.2.0.2) (Lenth and Piaskowski, 2026), with results back-transformed from the link scale to the probability scale.

3. Results

3.1. Culicoides abundance, species richness, and diversity

A summary of Culicoides species collected using a 3D-printed suction trap with three attractants over nine nights is presented in Table 2. A total of 12,474 specimens of Culicoides from 27 collections were captured. The total collection comprised predominantly female specimens, accounting for 84.3% (n = 10,511), while males comprised 15.7% (n = 1963). Of the total females, nulliparous (n = 5221; 49.7%) and parous (n = 2737; 26.0%) were the two most frequently observed physiological groups. The UV-LED light trap collected the largest number of specimens (n = 6221; 49.9%), followed by the UV-LED light + CO2 trap (n = 5549; 44.5%), whereas the CO2-only trap yielded the lowest number (n = 704; 5.6%). No significant difference of the total number was observed between the UV-LED light and UV-LED light + CO2 traps, while the CO2-only trap collected significantly fewer Culicoides specimens than both UV-LED light-based traps (Tukey-adjusted pairwise comparison, CO2-only vs UV-LED light: Z = −9.834, P < 0.0001; CO2-only vs UV-LED light + CO2: Z = −8.990, P < 0.0001) (Fig. 2).

Table 2.

Summary for species of Culicoides collected using a 3D-printed suction trap with three attractants over nine nights in Cha-am District, Phetchaburi Province, Thailand (28 November – 15 December 2022).

Species N
 Nulliparous
 Parous
 Non-blood-fed
 Blood-fed
 Gravid
 Male
 Total
 Grand total (%)
A B C A B C A B C A B C A B C A B C A B C A B C
C. oxystoma 9 8 9 688a 36b 1091a 449a 45b 363a 108a 15b 127a 472a 12b 294a 375 148 262 2092a 256b 2137a 4485 (36.0)
C. imicola 9 7 9 1000a 56b 894a 650a 12b 295d 202a 11b 212a 17 11 25 2 37 1894a 81b 1449a 3424 (27.4)
C. guttifer 9 9 9 82 94 172 45 53 76 1 1 3 1 3 190 150 290 319 298 544 1161 (9.3)
C. peregrinus 9 5 9 207a 2b 112c 162a 1b 71e 42a 3b 47a 142a 1b 68d 76 3 42 629a 10b 340c 979 (7.8)
C. huffi 9 4 9 242a 6b 200a 165a 1b 154a 4 3 9 10 71 1 76 491a 8b 443a 942 (7.6)
C. palpifer 8 3 6 185 4 123 17 13 8 7 12 6 222 4 149 375 (3.0)
C. shortti 7 3 9 92 1 99 61 2 35 23 1 20 2 1 10 5 188 4 160 352 (2.8)
C. arakawae 7 5 9 14 5 27 14 7 12 3 1 1 3 83 15 53 114 28 96 238 (1.9)
C. anophelis 7 1 8 85 1 61 3 9 88 1 70 159 (1.3)
C. fulvus 7 2 6 24 1 31 14 2 16 2 5 24 12 3 4 67 3 68 138 (1.1)
C. innoxius 8 0 7 23 38 6 6 2 6 3 3 34 53 87 (0.7)
C. orientalis 3 0 4 13 4 10 1 4 3 5 32 8 40 (0.3)
C. homotomus 5 2 2 7 2 2 1 1 2 3 1 6 2 1 18 6 4 28 (0.2)
C. brevitarsis 4 1 3 3 5 9 1 1 2 14 1 6 21 (0.2)
C. actoni 4 0 4 1 1 1 1 1 2 2 5 4 9 (0.1)
C. clavipalpis 2 2 1 1 1 1 1 1 2 2 1 5 (<0.1)
C. halonostictus 2 0 2 1 1 1 1 2 2 4 (<0.1)
C. jacobsoni 2 0 1 1 1 2 2 2 4 (<0.1)
C. asiana 1 0 0 2 2 2 (<0.1)
C. flavipunctatus 0 0 1 1 1 1 (<0.1)
C. negripes 0 0 1 1 1 1 (<0.1)
Unidentified 3 2 4 2 3 1 1 3 3 1 1 2 1 1 6 2 11 19 (0.2)
Total 2376a 201b 2644a 1585a 125b 1027a 293a 5b 222a 417a 33b 455a 693a 17b 418a 857 323 783 6221a 704b 5549a 12474
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Abbreviations: N, number of days in which each Culicoides species was found; Attractants used: A, UV-LED light; B, CO2-only; C, UV-LED light + CO2.

Notes: Species diversity indices for three attractants (A-C): Shannon’s index (H′) = 1.83, 1.40, and 1.78; Simpson’s index (1-D) = 0.772, 0.673, and 0.761, respectively. Different superscript letters denote statistically significant differences among attractants of the five most abundant species (P < 0.001), unless otherwise indicated (a-d, P < 0.01; a-e, P < 0.05).

Fig. 2.

Fig. 2

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Box-and-whisker plots illustrating total abundance and the numbers of the five most abundant species of Culicoides collected using a 3D-printed suction trap with three attractants (A, UV-LED light; B, CO2-only; C, UV-LED light + CO2) over nine nights in Cha-am District, Phetchaburi Province, Thailand (28 November to 15 December 2022). Each box represents the interquartile range (25th–75th percentiles) with the median; whiskers indicate the minimum and maximum values, and the cross denotes the mean. Different superscript letters denote statistically significant differences among attractants (P < 0.001).

Across three attractants, 21 species were identified. Species richness was highest in the UV-LED light + CO2 trap (20 species), followed by the UV-LED light trap (19 species) and the CO2-only trap (13 species). The UV-LED light and UV-LED light + CO2 traps did not differ significantly in species diversity, with Shannon indices (H′) of 1.83 and 1.78, and Simpson indices of diversity (1 − D) of 0.772 and 0.761, respectively. In contrast, the species diversity was significantly lower in the CO2-only trap than in both UV-LED light-based traps, with a Shannon index (H′) of 1.40 (Tukey-adjusted pairwise comparison, CO2-only vs UV-LED light: Z = −4.843, P < 0.0001; CO2-only vs UV-LED light + CO2: Z = −5.222, P < 0.0001) and a Simpson index of diversity (1 − D) of 0.673 (Tukey-adjusted pairwise comparison, CO2-only vs UV-LED light: Z = −3.310, P = 0.003; CO2-only vs UV-LED light + CO2: Z = −3.892, P = 0.0003) (Table 2).

3.2. Numbers and physiological status of the most abundant species

The numbers and physiological status of the five most abundant Culicoides species collected using a 3D-printed suction trap with three attractants over nine nights are presented in Table 2 and Fig. 2.

The five most abundant species accounted for 88.1% of the total specimens, comprising C. oxystoma (n = 4485; 36.0%), C. imicola (n = 3424; 27.4%), C. guttifer (n = 1161; 9.3%), C. peregrinus (n = 979; 7.8%), and C. huffi (n = 942; 7.6%), while each of the remaining 16 species contributed less than 3.0% of total abundance. Across three attractants, the UV-LED light and UV-LED light + CO2 traps captured significantly higher numbers of C. oxystoma (Tukey-adjusted pairwise comparison, UV-LED light vs CO2-only: Z = 10.500, P < 0.0001; UV-LED light + CO2 vs CO2-only: Z = 9.059, P < 0.0001), C. imicola (Tukey-adjusted pairwise comparison, UV-LED light vs CO2-only: Z = 7.450, P < 0.0001; UV-LED light + CO2 vs CO2-only: Z = 6.954, P < 0.0001), C. peregrinus (Tukey-adjusted pairwise comparison, UV-LED light vs CO2-only: Z = 12.976, P < 0.0001; UV-LED light + CO2 vs CO2-only: Z = 11.056, P < 0.0001), and C. huffi (Tukey-adjusted pairwise comparison, UV-LED light vs CO2-only: Z = 8.389, P < 0.0001; UV-LED light + CO2 vs CO2-only: Z = 8.288, P < 0.0001) than the CO2-only trap. However, the numbers of C. oxystoma, C. imicola, and C. huffi did not differ significantly between the two UV-LED light-based traps, whereas C. peregrinus was significantly more abundant in the UV-LED light trap than in the UV-LED light + CO2 trap (Tukey-adjusted pairwise comparison, Z = 7.653, P < 0.0001).

Regarding female physiological status, the numbers of nulliparous and parous females of C. oxystoma (Tukey-adjusted pairwise comparison, UV-LED light vs CO2-only: nulliparous, Z = 9.118, P < 0.0001; parous, Z = 6.666, P < 0.0001; UV-LED light + CO2 vs CO2-only: nulliparous, Z = 8.310, P < 0.0001; parous, Z = 6.332, P < 0.0001), C. imicola (Tukey-adjusted pairwise comparison, UV-LED light vs CO2-only: nulliparous, Z = 4.904, P < 0.0001; parous, Z = 10.410, P < 0.0001; UV-LED light + CO2 vs CO2-only: nulliparous, Z = 4.730, P < 0.0001; parous, Z = 10.396, P < 0.0001), C. peregrinus (Tukey-adjusted pairwise comparison, UV-LED light vs CO2-only: nulliparous, Z = 6.247, P < 0.0001; parous, Z = 4.816, P < 0.0001; UV-LED light + CO2 vs CO2-only: nulliparous, Z = 5.488, P < 0.0001; parous, Z = 4.317, P < 0.0001), and C. huffi (Tukey-adjusted pairwise comparison, UV-LED light vs CO2-only: nulliparous, Z = 6.250, P < 0.0001; parous, Z = 4.904, P < 0.0001; UV-LED light + CO2 vs CO2-only: nulliparous, Z = 6.386, P < 0.0001; parous, Z = 4.698, P < 0.0001) were significantly higher in both UV-LED light-based traps compared with the CO2-only trap. Additionally, nulliparous C. peregrinus (Tukey-adjusted pairwise comparison, Z = 3.246, P = 0.002), as well as parous C. imicola (Tukey-adjusted pairwise comparison, Z = 3.036, P = 0.006) and C. peregrinus (Tukey-adjusted pairwise comparison, Z = 2.657, P = 0.017) were significantly more abundant in the UV-LED light trap than in the UV-LED light + CO2 trap. In the parity rate analysis, only the UV-LED light trap showed a higher parity rate for C. imicola than the CO2-only trap (0.443 vs 0.264; Tukey-adjusted pairwise comparison, Z = 2.753, P = 0.015), while no other species showed significant differences across three attractants (Table 3). Additionally, both UV-LED light-based traps collected significantly more blood-fed C. oxystoma (Tukey-adjusted pairwise comparison, UV-LED light vs CO2-only: Z = 7.762, P < 0.0001; UV-LED light + CO2 vs CO2-only: Z = 7.798, P < 0.0001), C. imicola (Tukey-adjusted pairwise comparison, UV-LED light vs CO2-only: Z = 7.549, P < 0.0001; UV-LED light + CO2 vs CO2-only: Z = 8.555, P < 0.0001), and C. peregrinus (Tukey-adjusted pairwise comparison, UV-LED light vs CO2-only: Z = 4.257, P < 0.0001; UV-LED light + CO2 vs CO2-only: Z = 4.458, P < 0.0001), and gravid C. oxystoma (Tukey-adjusted pairwise comparison, UV-LED light vs CO2-only: Z = 7.625, P < 0.0001; UV-LED light + CO2 vs CO2-only: Z = 6.432, P < 0.0001), than the CO2-only trap, whereas gravid C. peregrinus were significantly more abundant in the UV-LED light than in the UV-LED light + CO2 and CO2-only traps (Tukey-adjusted pairwise comparison, UV-LED light vs UV-LED light + CO2: Z = 3.443, P = 0.001; UV-LED light vs CO2-only: Z = 4.812, P < 0.0001).

Table 3.

Observed and predicted parity rate per collection of the five most abundant species of Culicoides collected using a 3D-printed suction trap with three attractants over nine nights in Cha-am District, Phetchaburi Province, Thailand (28 November – 15 December 2022).

Species Total no. of females Observed parity rate
 Predicted parity rate
A B C A B C
C. oxystoma 3700 0.599 0.667 0.418 0.651 0.732 0.634
C. imicola 3360 0.465 0.291 0.367 0.443a 0.264b 0.369ab
C. guttifer 531 0.364 0.365 0.323 0.358 0.364 0.312
C. peregrinus 858 0.626 0.714 0.624 0.701 0.738 0.723
C. huffi 794 0.424 0.143 0.455 0.470 0.178 0.422
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Abbreviations: Attractants used: A, UV-LED light; B, CO2-only; C, UV-LED light + CO2.

Note: Different superscript letters denote statistically significant differences among attractants (P < 0.05).

For C. guttifer, no significant differences among attractants were observed in the total number and all female physiological groups (P ≥ 0.05).

3.3. BTV qRT-PCR detection in Culicoides species

Details of BTV detection in Culicoides species are summarized in Table 4. A total of 154 pools, comprising 3086 specimens, were analyzed for BTV. These included 75 pools of C. oxystoma, 43 of C. imicola, 20 of C. peregrinus, 11 of C. huffi, and 5 of C. guttifer. BTV was detected in 53 pools (34.4%), 27 from C. imicola, 20 from C. oxystoma, 5 from C. peregrinus, and 1 from C. huffi. Of these positive pools, 41 were from the UV-LED light trap, 10 from the UV-LED light + CO2 trap, and 2 from the CO2-only trap. In addition, one positive pool of C. imicola exhibited a Ct value of 25.7, whereas the remaining positive pools had Ct values ranging from 32.2 to 39.5.

Table 4.

Pan-reactive BTV qRT-PCR results from five dominant species of Culicoides collected using a 3D-printed suction trap with three attractants over nine nights in Cha-am District, Phetchaburi Province, Thailand (28 November – 15 December 2022).

Species No. of specimens No. of positive pools (total no. of analyzed pools)
 Ct-value range Pool size
A B C Total
C. oxystoma 1502 15 (44) 2 (3) 3 (28) 20 (75) 36.9–39.9 20–21
C. imicola 865 23 (28) 0 (0) 4 (15) 27 (43) 25.7–39.5 20–23
C. peregrinus 400 3 (14) 0 (0) 2 (6) 5 (20) 35.7–39.3 19–21
C. huffi 220 0 (5) 0 (0) 1 (6) 1 (11) 39.1 20
C. guttifer 99 0 (1) 0 (1) 0 (3) 0 (5) – 19–20
Total 3086 41 (92) 2 (4) 10 (58) 53 (154)
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Abbreviations: Attractants used: A, UV-LED light; B, CO2-only; C, UV-LED light + CO2.

3.4. BTV serotype detection

Details of BTV-serotype distribution are provided in Table 5. A total of 25 pools were analyzed for BTV serotypes, including 14 from C. imicola, 9 from C. oxystoma, and 2 from C. peregrinus. Five serotypes (BTV-2, BTV-4, BTV-7, BTV-12, and BTV-15) were successfully identified, all of which were from C. imicola. Among these, BTV-7 and BTV-12 were the most frequently detected, and notably, BTV-15 was observed in one pool.

Table 5.

BTV serotyping results from three dominant species of Culicoides collected using a 3D-printed suction trap with three attractants over nine nights in Cha-am District, Phetchaburi Province, Thailand (28 November – 15 December 2022).

Species No. of pools tested (attractants used) BTV-serotype distribution (n)
In-house qRT-PCR Multiplex qRT-PCR
C. imicola 12 (A), 2 (C) 2 (1), 7 (4), 12 (1) 4 (1), 7 (2), 12 (3), 15 (1)
C. oxystoma 7 (A), 1 (B), 1 (C) – –
C. peregrinus 1 (A), 1 (C) – –
Total 20 (A), 1 (B), 4 (C) 2 (1), 7 (4), 12 (1) 4 (1), 7 (2), 12 (3), 15 (1)
Open in a new tab

Abbreviations: Attractants used: A, UV-LED light; B, CO2-only; C, UV-LED light + CO2.

4. Discussion

In this study, the effectiveness of a 3D-printed suction trap using UV-LED light and CO2 as attractants was compared. The surveillance capabilities of these attractants were evaluated on the basis of Culicoides abundance, species richness or diversity, and physiological status. In addition, the presence of BTV and the circulating serotypes in Culicoides species in the study area were determined.

Overall, the total abundance of Culicoides did not differ significantly between the two UV-LED light-based traps. Both traps also showed comparable patterns across the five most abundant species. Consistent with these findings, Zhang et al. (2023) reported that UV light traps without CO2 collected greater numbers of Culicoides and higher abundances of most species than CO2-only traps. Walgama and Lysyk (2019) also found no positive effect of CO2 addition at sites with low midge abundance. In contrast, many previous studies have shown that the addition of CO2 to UV light traps – across different models such as the Onderstepoort and miniature CDC light traps – increases the abundance of Culicoides biting midges (Sloyer et al., 2019; Becker et al., 2021; Zhang et al., 2023).

The discrepancy in the effects of CO2 observed among studies may be related to the abundance of host-seeking females, including both nulliparous and parous individuals, at the sampling sites. In addition, UV light trap placement in relation to livestock proximity may also greatly influence trap performance. Similar observations have been reported by Venter et al. (2013), who found no additional effect of CO2 on total Culicoides catches in a UV light trap (Triple trap). Likewise, Venter et al. (2016) reported only minimal effects of CO2, with slight increases in abundance and generally low catches observed when traps were placed in the absence of animal hosts. Furthermore, de Beer et al. (2021) indicated that UV light traps operated near cattle or other livestock have a limited attraction range of less than 4 m, whereas in the absence of livestock, this range may extend to 30–50 m. Additionally, background CO2 levels, distance from oviposition habitats, and environmental conditions such as wind and temperature may also influence CO2 dispersion and the overall trap performance (Venter et al., 2016; Zhang and Mathias, 2022).

No significant differences were observed in the total abundance of female physiological groups among the five most abundant species between the two UV-LED light-based traps. However, significantly higher numbers of parous females of C. imicola and C. peregrinus were recorded in the UV-LED light trap compared with the UV-LED light + CO2 trap. In contrast, only nulliparous C. peregrinus was significantly more abundant in the UV-LED light + CO2 trap. Furthermore, the addition of CO2 was associated with a significant reduction in gravid females, particularly in C. peregrinus. No clear effect was observed on the number of blood-fed females across species. This is supported by Bellis and Reid (1996), who indicated that parity estimates may need to be assessed on a species-specific basis. These findings also suggested that CO2 may vary in its attractiveness to different physiological groups (Anderson and Linhares, 1989; Venter et al., 2016). Therefore, the responses of Culicoides vector species across physiological groups to CO2 warrant further investigation.

In the CO2-only trap, the total abundance of Culicoides was relatively low compared with that in both UV-LED light-based traps. This suggests that CO2 alone, unlike UV-LED light, may be insufficient to effectively attract Culicoides females, including the potential vector species in the study area. Additionally, collections using the CO2-only trap were associated with significantly lower parity rate of C. imicola, compared with using the UV-LED light trap. An additional effect of CO2 on species richness was found, with higher richness recorded in the UV-LED light + CO2 trap compared with the UV-LED light trap. This is consistent with previous studies (Sloyer et al., 2019; Becker et al., 2021), particularly by improving the detection of rare Culicoides species (Walgama and Lysyk, 2019). However, species diversity did not differ significantly between the UV-LED light and UV-LED light + CO2 traps, whereas it was significantly lower in the CO2-only trap. This suggests that the UV-LED light trap without CO2 can be used for Culicoides surveillance without a reduction in species diversity.

Of the 21 Culicoides species identified in the present study, several have been reported to be widely distributed in Malaysia and Indonesia, and four species (C. actoni, C. brevitarsis, C. fulvus, and C. wadai) have been confirmed as BTV vectors in Australia (Daniels et al., 2004). These findings could suggest regional similarity in Culicoides fauna and potential vector assemblages. In Thailand, four species, C. orientalis, C. imicola, C. oxystoma, and C. fulvus, have been suggested as potential vectors of BTV (Fujisawa et al., 2021). In the present study, C. peregrinus and C. huffi were additionally identified as potential BTV vector species. Moreover, BTV-positive pools were predominantly detected in C. imicola, supporting its potential role as a principal vector species in local BTV transmission. Notably, one pool of C. imicola showed a low Ct value (25.7), indicating a relatively high viral load and the presence of a BTV infection in at least one individual (Veronesi et al., 2013; Van der Saag et al., 2015). To our knowledge, this represents the first evidence of a direct association between BTV and C. imicola in Southeast Asia. The UV-LED light trap also yielded more BTV-positive pools, likely due to the higher number of parous C. imicola captured and the higher BTV-positivity rate of this species. Nevertheless, whether UV-LED light exerts a repellent effect on BTV-infected Culicoides vector species requires further investigation. In the BTV-serotype analysis, the co-circulation of BTV-2, BTV-4, BTV-7, BTV-12, and BTV-15 was detected. Most of these serotypes have previously been identified in goat populations in central and western Thailand (Homat et al., 2025), with BTV-15 detected for the first time in Thailand in the present study. The co-circulation of multiple BTV serotypes provides strong evidence of active and ongoing transmission within ruminant populations in the study area.

5. Conclusions

A 3D-printed suction trap baited with UV-LED light alone effectively collected Culicoides biting midges without a significant loss of abundance or species diversity and can be a suitable tool for large-scale BTV surveillance programmes in Thailand. The presence of BTV and the co-circulation of multiple serotypes highlight the need for continued efforts to strengthen bluetongue control measures in the country.

Ethical approval

The animal use and ethics protocol for this study was approved by the Mahidol University-Institute Animal Care and Use Committee (MUVS-ACUC-2020-02-06 and MUVS-ACUC-2024-03-23) in accordance with the Ethical Principles and Guidelines for the Use of Animals of the National Research Council of Thailand.

CRediT authorship contribution statement

Bunlue Kornmatitsuk: Conceptualization, Funding acquisition, Methodology, Formal analysis, Writing - original draft, Writing - review & editing. Yuki Fujisawa: Investigation, Methodology. Thanyaporn Homat: Investigation, Methodology. Sirichai Eardmusic: Investigation, Methodology. Ladawan Sariya: Methodology, Writing - review & editing. Sudsaijai Kornmatitsuk: Conceptualization, Funding acquisition, Methodology, Formal analysis, Writing - review & editing.

Statement on the use of AI-assisted technologies

During the preparation of this work, the authors did not use any artificial intelligence (AI) or AI-assisted technologies to generate, analyze, or interpret scientific content. ChatGPT (OpenAI) was used only to support English-language polishing and formatting prior to submission; the authors reviewed and edited all resulting content and take full responsibility for the content of the published article.

Funding

This research project was supported by Mahidol University (Strategic Research Fund): 2024. Financial support at the initial stage of this study was partially provided by MU-MiniRC 2022, Mahidol University.

Declaration of competing interests

The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper.

Acknowledgments

We would like to thank Mr Aeknarin Saechin, Monitoring and Surveillance Center for Zoonotic Diseases in Wildlife and Exotic Animals, Faculty of Veterinary Science, Mahidol University, and Mr Kandit Kornmatitsuk, for their valuable assistance with statistical analyses.

Footnotes

Appendix A

Supplementary data to this article can be found online at https://doi.org/10.1016/j.crpvbd.2026.100420.

Appendix A. Supplementary data

The following are the Supplementary data to this article:

Multimedia component 1
mmc1.pdf (297.5KB, pdf)
Multimedia component 2
mmc2.pdf (141.2KB, pdf)

Data availability

All data generated or analyzed during this study are included in this published article and its supplementary files.

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Associated Data

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Supplementary Materials

Multimedia component 1
mmc1.pdf (297.5KB, pdf)
Multimedia component 2
mmc2.pdf (141.2KB, pdf)

Data Availability Statement

All data generated or analyzed during this study are included in this published article and its supplementary files.

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