Journal Search Engine

to

Search Advanced Search Adode Reader(link)
Download PDF Export Citation Metrics Korean Bibliography
ISSN : 1225-0171(Print)
ISSN : 2287-545X(Online)
Korean Journal of Applied Entomology Vol.64 No.3 pp.159-168
DOI : https://doi.org/10.5656/KSAE.2025.06.0.019

New Record of the Soil Mite Macrocheles penicilliger (Berlese, 1904) (Acari: Mesostigmata) from Dokdo, Korea

Sehyeon Lim1, Hwalsu Hwang2, Seoyoung Keum3, Kyeong-Yeoll Lee1,2,4*
1Department of Plant Medicine, College of Agriculture and Life Sciences, Kyungpook National University, Daegu 41566, Korea
2Research Institute for Dok-do and Ulleung-do Island, Kyungpook National University, Daegu 41566, Korea
3Silkworm and Insect Management Center, Agricultural Resource Management Institute, Sangju 37110, Korea
4Institute of Plant Medicine, Kyungpook National University, Daegu 41566, Korea
*Corresponding author:leeky@knu.ac.kr
May 6, 2025 June 5, 2025 June 10, 2025

Abstract


The Dokdo islands are volcanic islands with a unique climate and ecosystem and represent the easternmost land area of the Republic of Korea. Although biodiversity studies have been conducted in Dokdo, soil arthropods have rarely been investigated. In this study, we collected soil from both of the main islands of Dokdo in 2023 and 2024 and isolated soil mites using the Berlese–Tullgren method. For species identification, morphological features and mitochondrial cytochrome c oxidase subunit I (COI) gene sequences were analyzed, revealing the presence of Macrocheles penicilliger (Berlese, 1904), a previously unrecorded species in Korea. Morphological comparisons showed that three pairs of the sternal setae of M. penicilliger from Dokdo differed from those in the described literature in that the first pair was plumose distally, while the second and third pairs were simple. In addition, the COI sequences of individuals collected from Dongdo (East island) and Seodo (West island) of Dokdo differed by 2.43%, and the sequence of the Seodo individual was 100% identical to those from Belgian samples. These findings reveal the genetic diversity of M. penicilliger in Dokdo and suggest that its distribution on the islands may be related to its phoretic behavior, which is associated with insects inhabiting the nests of migratory birds.


Biodiversity , Soil mites , Species identification , Geographic distribution , Phoresis

독도의 토양 응애 Macrocheles penicilliger (Berlese, 1904) (응애아강: 중기문목) 미기록종 보고

임세현1, 황활수2, 금서영3, 이경열1,2,4*
1경북대학교 식물의학과
2경북대학교 울릉도독도연구소
3경상북도농업자원관리원 잠사곤충사업장
4경북대학교 식물의학연구소

초록


독도는 대한민국 최동단에 위치한 화산섬으로서 독특한 기후와 생태계를 이룬다. 독도에 대한 생물다양성 연구가 진행되어 왔지만, 토양의 절지동물은 거의 조사된 바가 없다. 본 연구에서는 2023-2024년도에 독도에서 토양을 채취하여 Berlese-Tullgren 방법으로 토양응애를 분리하였다. 종 동정을 위하여 형태학적 특징과 미토콘드리아 cytochrome c oxidase subunit I (COI) 유전자 염기서열 분석한 결과 국내 미기록종인 Macrocheles penicilliger (Berlese, 1904)으로 나타났다. 독도의 M. penicilliger의 흉판모 3쌍은 기존 문헌의 묘사된 것과 달리 첫째쌍은 말단이 깃털 형태이지만, 둘째 및 셋째쌍은 단순한 형태이었다. 또한, 동도와 서도에서 채집한 개체들의 COI 서열은 2.43% 차이가 있었으며 특히 서도 개체의 COI 염기서열은 벨기에 샘플과 100% 일치했다. 본 연구결과는 M. penicilliger의 유전적 다양성을 밝혔으며 독도에 분포한 원인으로서 철새의 둥지에 서식하는 곤충들에 편승하여 이동하는 행동과 관계 있을 것이라고 추측한다.


종다양성 , 토양 응애 , 종 동정 , 지리적 분포 , 편승

    Dokdo is a volcanic island located in the East Sea approximately 87.4 km east of Ulleungdo, and 216.8 km from the Korean mainland. The island consists of two main islets, Dongdo (East Island) and Seodo (West Island), as well as 89 smaller islets and rock formations, and is designated as Natural Monument No. 336 of Korea (Cultural Heritage Administration, 2003). With its volcanic origin dating back 4.6 to 2.5 million years (Jeon, 2005), Dokdo presents a distinctive geological and ecological environment among the Korean islands (Kim et al., 2007).

    Although biotic surveys in Dokdo have documented vascular plants, birds, marine invertebrates, insects, and microorganisms (Hwang et al., 2023), and several studies have investigated soil properties and soil-dwelling organisms, including microbes and nematodes (Kim et al., 2019;Namirimu et al., 2019;Eo et al., 2022;Lee et al., 2024), soil microarthropods, especially mites, have not been cataloged. Notably, acarological studies have been conducted on nearby Ulleungdo, highlighting the lack of comparable data for Dokdo (Keum and Jung, 2012;Kim et al., 2013;Bayartogtokh and Bae, 2024).

    Soil mesofauna, such as mites (Acari) and springtails (Collembola), are abundant in soil ecosystems and are frequently used as bioindicators due to their sensitivity to environmental changes and ease of sampling (Gerlach et al., 2013;George et al., 2017). Among the Acari, the order Mesostigmata contains many predatory mites that contribute to the regulation of other soil invertebrate populations (Gulvik, 2007;George et al., 2017).

    The genus Macrocheles Latreille (Mesostigmata: Macrochelidae) is a globally distributed group often found in dung, decaying organic matter, and animal nests, often exhibiting phoresy on insects such as beetles and flies (Evans and Browning, 1956;Emberson, 1973;Krantz and Whitaker, 1988). In Korea, 10 species of Macrocheles have been recorded: M. aviaton Seo, 1966; M. decoloratus (Koch, 1893); M. glaber (Müller, 1860); M. insignitus Berlese, 1918; M. japonicus Evans and Hyatt, 1963; M. muscaedomesticae (Scopoli, 1772); M. nataliaeBregetova and Koroleva, 1960;M. punctatus Ishikawa, 1967; M. scutatiformis Petrova, 1967; and M. similis Krantz and Filipponi, 1964 (Kontschán et al., 2014;Keum et al., 2016;Ji et al., 2023a, 2023b;National Institute of Biological Resources, 2024). Among them, M. aviaton is the only species originally described from the Korean peninsula, based on material from North Korea. However, due to the lack of available specimens and follow-up studies, its current taxonomic status and distribution remain uncertain.

    In this study, we report the first record of Macrocheles penicilliger in Korea, found in Dokdo and identified based on both morphological and molecular data. We also discuss its genetic variation and provide comments on its identification, which have implications for future taxonomic and ecological research on phoretic mites inhabiting isolated island ecosystems.

    Materials and Methods

    Sample collection

    Soil samples were collected from both main islands of Dokdo, Dongdo and Seodo, in 2023 and 2024. The samples were taken to a depth of 5 cm below the soil surface. Mites were extracted using a Berlese–Tullgren funnel trap with a 30 W halogen lamp over 72 h (Bano and Roy, 2016), and extracted mites were preserved in a 70% ethanol solution at -20°C. For slide preparation, the mites were cleared in lactophenol and mounted on a glass slide with a drop of Hoyer’s medium (Krantz and Walter, 2009).

    Morphological identification

    Species identification of the slide-mounted mite specimens was performed using an Olympus BX50 light microscope (Olympus, Japan) and a Keyence VHX digital microscope (Keyence, Japan). Morphological analysis was carried out using the identification keys of Evans and Browning (1956) and idiosomal chaetotaxy system of Lindquist and Evans (1965), as applied to macrochelids by Hyatt and Emberson (1988). All measurements are in micrometers (μm) and based on Keyence VHX digital microscope software. The length and width of the idiosoma were measured along the midline, from the anterior margin to the posterior end, and at the widest point of the body, respectively. The examined specimens were deposited in the National Institute of Biological Resources (NIBR), Korea.

    DNA extraction and PCR amplification

    The non-mashed method was used for mite DNA extraction (Khaing et al., 2013). A single mite was placed in a 1.5 mL microcentrifuge tube containing 180 μL of genomic digestion buffer and 20 μL of proteinase K solution, both from the PureLink Genomic DNA Mini Kit (Invitrogen, CA, USA). The sample was then heated to 55°C for 12 hours. The exoskeleton was then collected to serve as the voucher specimen, while the remaining solution was processed for DNA extraction according to the PureLink Genomic DNA Mini Kit protocol.

    For amplification, PCRs were performed using 30 μL reaction mixtures consisting of 15 μL of Pre-Mix (Solgent, Daejeon, Korea), 2 μL of each primer (10 pmol/μL), 4 μL of DNA extract, and 7 μL of distilled water. The 658 bp mitochondrial cytochrome c oxidase I (COI) nucleotide sequence was targeted using the primer pair LCO1490 (5'-GGT CAA CAA ATC ATA AAG ATA TTG G-3') and HCO2198 (5'-TAA ACT TCA GGG TGA CCA AAA AAT CA-3') (Folmer et al., 1994). The thermocycler protocol included an initial denaturation at 92°C for 5 min, followed by 35 cycles of denaturation at 92°C for 60 s, annealing at 55°C for 60 s, and extension at 72°C for 60 s, with a final extension step at 72°C for 5 min. The PCR products were confirmed by 1% agarose gel electrophoresis using UltraPure™ low melting point (LMP) Agarose (Invitrogen, CA, USA) and subsequently purified.

    DNA sequence assembly and phylogenetic analyses

    Sequencing was performed at Solgent (Daejeon, Korea) using the BigDye® Terminator v3.1 Cycle Sequencing Kit. The resulting COI sequences were aligned with 16 sequences from 13 species retrieved from the GenBank database of the National Center for Biotechnology Information (NCBI) (Table S1) using the L-INS-i method (Katoh et al., 2005) implemented in MAFFT via the online service (Katoh et al., 2019). Genetic distances within the sequence dataset were calculated based on the p-distance using MEGA X (Kumar et al., 2018). The aligned sequences were analyzed using MrModeltest v2.4 (Nylander, 2004) to determine the optimal substitution model. Based on the Akaike information criterion (AIC), the GTR+I+G model (general time reversible model with a proportion of invariant sites and gamma-distributed rate variation between sites) was selected as the best-fitting model for both maximum likelihood (ML) and Bayesian inference (BI) analyses.

    The phylogenetic tree based on the ML method was constructed using the Windows version of IQ-TREE2 v2.3.6 (Minh et al., 2020) with 1,000 bootstrap resamples. The BI analysis was conducted using the Windows version of MrBayes v3.2.7 (Ronquist et al., 2012). The analysis included four Markov chain Monte Carlo (MCMC) chains and two independent runs with 107 generations each, with sampling every 1,000 generations. The first 50% of the generations were discarded as burn-in. Two Glyptholaspis sp. (Macrochelidae) sequences (MN361179 and MN363457) were used as an outgroup.

    Results

    Systematic accounts

    Family Macrochelidae Vitzthum, 1930

    Genus Macrocheles Latreille, 1829

    Species Macrocheles penicilliger (Berlese, 1904)

    Description of the Korean specimens

    Female: Dorsal shield 830–850 long and 480–500 wide, with a smooth anterior margin and a serrated posterior margin extending from the lateral edges to the terminal region (Figs. 1A, 2A). Dorsal setae plumose, except for z1, z5, j6, z6, J2, and J5, which are simple (Fig. 2A), with z5 occasionally plumose (Fig. 3). Sternal shield bearing three pairs of sternal setae, with the first pair (st1) distally plumose and the second and third pairs (st2 and st3) simple (Figs. 1B, 2B). Metasternal shields paired, each with one simple seta pair st4 located laterally to the genital shield. Ventri-anal shield 290–300 long and 300– 310 wide, bearing three pairs of pre-anal setae, one pair of para-anal setae, and a single post-anal seta. Pre-anal and para-anal setae simple; post-anal seta short and plumose. No platelet present between the genital and ventrianal shields. One pair of elongate, strongly sclerotized metapodal plates present lateral to the ventrianal shield. Epistome with simple lateral projections and a deeply bifurcated median process (Fig. 1C). Fixed and movable cheliceral digits, each with two denticles; ventral seta elongated and densely pilose. In leg I, tarsus 120– 130 long, longer than tibia, which is about 100.

    Material examined: 1♀, soil and litter, Seodo (West Isl.), Dokdo (37°24'06.2"N, 131°86'51.5"E) and 13♀, soil and litter, Dongdo (East Isl.), Dokdo (37°23'88.3"N, 131°86'90.6"E), Ulleung-eup, Ulleung-gun, Gyeongsangbuk-do, Korea, 15. vi. 2023., H. Hwang coll.; 3♀, soil and litter, Dongdo (East Isl.), Dokdo (37°23'88.3"N, 131°86'90.6"E), Ulleung-eup, Ulleunggun, Gyeongsangbuk-do, Korea, 13. viii. 2024., S.-H. Lim coll.

    Global distribution: USA, Europe, Australia, New Zealand, India, Africa, Japan (Takaku, 2000), Russia (Bregetova and Koroleva, 1960), China (Lin and Zhang, 2010), Iran (Hajizadeh et al., 2020), Korea.

    Molecular analyses

    The genetic distanced between the 16 COI sequences from the Dokdo specimens (accession nos. PV188055, PV276880, PV276881, and PV452829–PV452841) were determined (Table 1). The sequence PV188055 was obtained from the mite collected on Seodo, while the other sequences were obtained from mites on Dongdo. There was a sequence dissimilarity of 2.43% (16/658 bp) between PV188055 and the other Dokdo sequences. When the Dokdo mite sequences were compared to 16 publicly available COI sequences from the NCBI GenBank database representing 13 Macrocheles species using the pdistance method (Table 1), the Seodo mite sequence (PV 188055) was 100% identical with four sequences of M. penicilliger from Belgium (accession no. MH983630, MH983673, MH983678, MH983713), registered by Young et al. (2019). In contrast, the other Dokdo sequences (PV276880, PV276881, and PV452829–PV452841) showed 2.43–2.58% dissimilarity with the Belgian sequences. In contrast, interspecific variation between the Dokdo mite sequences and other Macrocheles species ranged from 23% to 27% dissimilarity (Table 1).

    Both ML and BI phylogenetic analyses grouped the Seodo mite sequence (PV188055) with the four Belgian sequences in a sing le n ode, w hereas t he D ong do m ite sequences w ere separated from the Belgian sequences (Fig. 4). The Bayesian posterior probability and bootstrap value for the node within the M. penicilliger branch separating these sequences was 1.00 and 100, respectively, indicating a high degree of node stability in both phylogenetic analyses.

    Discussion

    In this study, we found M. penicilliger in Dokdo, and it is the first report of this species in Korea. Our morphological analysis of the Dokdo specimens revealed minor differences when compared to the redescription of M. penicilliger in Evans and Browning (1956). In our specimens, the first pair of sternal seta (st1) were weakly plumose distally and the second and third setae (st2 and st3) were simple, whereas Evans and Browning (1956) described the first pair of sternal seta as strongly plumose and the second and third pairs of sternal seta as weakly plumose distally. These morphological differences represent genetic variation between populations at the intraspecific level that may be associated with the wide geographic distribution of M. penicilliger (Bregetova and Koroleva, 1960;Takaku, 2000;Lin and Zhang, 2010;Hajizadeh et al., 2020).

    In addition, we analyzed the COI sequences of 16 M. penicilliger individuals from Dokdo and found that the COI sequence of the one individual from Seodo, differed in 2.43% of base pairs from the rest of the individuals, which were from Dongdo, suggesting that the individuals from the two main islands of Dokdo are genetically distinct. Interestingly, the COI sequence of the Seodo individual was 100% identical to those reported from Belgium. This suggests that, although this species is distributed worldwide, some groups are genetically quite conservative.

    Many species in the genus Macrocheles exhibit phoretic behavior, migrating by attaching themselves to the bodies of various flying insects (Evans and Browning, 1956;Cicolani, 1983;Krantz and Whitaker, 1988). Therefore, it is speculated that M. penicilliger on the Dokdo may have arrived via flying insect vectors originating from other regions. Knee (2017) suggested that the genetic divergence observed in M. willowae was more strongly associated with host specificity to various burying beetles (Coleoptera: Silphidae, Nicrophorus) than with geographic isolation. Although M. penicilliger is typically found in soil or decaying leaf litter, it has also been recorded in bird and small mammal nests and in guano, and has been associated with Trox scaber and T. suguyai (Coleoptera: Trogidae), both of which are burying beetles that often congregate around bird nests (Evans and Browning, 1956;Takaku, 2000). Furthermore, M. penicilliger has been found in the nests of Bombus terrestris (Hymenoptera: Apidae) and Dichotomius carolinus (Coleoptera: Scarabaeidae) (Emberson, 1973;Krantz and Whitaker, 1988).

    In addition to insect-mediated dispersal, bird-mediated introduction has also been proposed as a potential pathway for mite colonization of avian nests. Bajerlein et al. (2006) suggested that Mesostigmata mites including M. merdarius found in white stork (Ciconia ciconia) nests may have been introduced either through phoresy on coprophagous beetles or via the incorporation of cow dung by adult storks during nest construction. Similarly, Gwiazdowicz et al. (2006) identified M. glaber, M. merdarius, and other Mesostigmata mites in the nests of the white-tailed sea eagle (Haliaeetus albicilla), and proposed that the presence of mites commonly found in soil and decaying wood could be explained by their passive introduction along with nest-building materials transported by the birds. In the case of M. penicilliger from Dokdo, it is possible that this species was introduced by the black-tailed gull (Larus crassirostris), the island's primary breeding bird. However, these gulls typically construct their nests on bare ground or rocky cliffs using grass-like plants collected from the surrounding area (Lee and Yoo, 2005;Kim et al., 2017). Considering the type of nesting material used, the ecological preference of M. penicilliger for soil and leaf litter habitats, and the geographic isolation of Dokdo, it seems unlikely that M. penicilliger was introduced by black-tailed gulls.

    Taken together, the observed presence of genetically distinct populations on Dongdo (East Island) and Seodo (West Island) suggests that the phoretic hosts may have differed, or that multiple independent introductions from different sources occurred. To clarify the origin and dispersal patterns of M. penicilliger on Dokdo, further genetic studies of populations from other geographic regions, as well as targeted investigations of potential phoretic hosts inhabiting the Dokdo, are needed.

    Acknowledgments

    We thank Jaeseok Oh from Seoul National University for his valuable comments. This research was supported by a research grant (311058-05-3-CG000) from the Ministry for Food, Agriculture, Forestry, and Fisheries and a research grant (NRF-2016R1A6A1A05011910) from the Basic Science Research Program provided by the National Research Foundation of Korea (NRF) and funded by the Ministry of Education.

    Supplementary Information

    Supplementary data are available at Korean Journal of Applied Entomology online (http://www.entomology2.or.kr).

    Statements for Authorship Position & Contribution

    • Lim, S.: Kyungpook National University, Student in MS; Conducted the experiments and wrote the manuscript

    • Hwang, H.: Kyungpook National University, Ph.D; Collected samples

    • Keum, S.: Agricultural Resource Management Institute, Silkworm and Insect Management Center, Researcher; Provided revision and comments on the manuscript

    • Lee, K.-Y.: Kyungpook National University, Professor; Designed the research and wrote the manuscript

    All authors read and approved the manuscript.

    Figure

    KJAE-64-3-159_F1.jpg

    Microphotographs of Macrocheles penicilliger (Berlese, 1904): A, dorsal idiosoma; B, ventral idiosoma; C, epistome.

    KJAE-64-3-159_F2.jpg

    Line-drawing plates of Macrocheles penicilliger (Berlese, 1904): A, dorsal shield; B, ventral shields.

    KJAE-64-3-159_F3.jpg

    Variation in the morphology of z5 seta in Macrocheles penicilliger specimens from Dokdo: A, z5 seta weakly plumose; B, z5 seta simple.

    KJAE-64-3-159_F4.jpg

    Phylogenetic trees of Macrocheles species, including a maximum likelihood tree generated using IQ-TREE (A) and a Bayesian inference tree created using MrBayes (B). Node labels denote bootstrap support values and Bayesian posterior probabilities, respectively. The specimens collected in the current study are denoted by asterisks. The blue and red bars represent Dondo and Seodo, respectively.

    Table

    Genetic distances (p-distances) within Macrocheles penicilliger and between M. penicilliger and other Macrocheles species. *Sequences collected in this study.

    Reference

    1. Bajerlein, D., Błoszyk, J., Gwiazdowicz, D., Ptaszyk, J., Halliday, R., 2006. Community structure and dispersal of mites (Acari, Mesostigmata) in nests of the white stork (Ciconia ciconia). Biologia (Bratisl). 61, 525-530.
    2. Bano, R., Roy, S., 2016. Extraction of soil microarthropods: a low cost Berlese-Tullgren funnels extractor. Int. J. Fauna Biol. Stud. 2, 14-17.
    3. Bayartogtokh, B., Bae, Y.S., 2024. New findings of poronotic oribatid mites (Acari: Oribatida) from Korea. Zootaxa 5405, 151- 184.
    4. Berlese, A., 1904. Acari nuovi. Volume II. Redia 2, 10-32.
    5. Bregetova, N.G., Koroleva, E.V, 1960. The macrochelid mites (Gamasoidea, Macrochelidae) in the USSR. Parazitol. Sb. 19, 32- 154.
    6. Cicolani, B., 1983. Una nuova specie di Macrocheles (Acarina: Macrochelidae) raccolta nella grotta della zinzulusa (Castro Marina-Paglie). Boll. del Mus. Civ. di Stor. Nat. di Verona 9, 151- 159.
    7. Cultural Heritage Administration, 2003. Natural monument white paper. Cultural Heritage Administration, Daejeon.
    8. Emberson, R.M., 1973. Macrochelid mites in NZ (Acarina: Mesostigmata: Macrochelidae). N. Z. Entomol. 5, 118-127.
    9. Eo, J.K., Park, E., Choi, J.W., Shin, H.C., Choi, S.S., Park, S.Y., 2022. Unrecorded soil fungi isolated from the Dokdo, Korea. Proc. Natl. Inst. Ecol. Repub. Korea 3, 172-177.
    10. Evans, G.O., Browning, E., 1956. British mites of the subfamily Macrochelinae Tragardh (Gamasina-Macrochelidae). Bull. br. Mus. nat. Hist. Zool. 4, 1-55.
    11. Folmer, O., Black, M., Hoeh, W., Lutz, R., Vrijenhoek, R., 1994. DNA primers for amplification of mitochondrial cytochrome c oxidase subunit I from diverse metazoan invertebrates. Mol. Mar. Biol. Biotechnol. 3, 294-299.
    12. George, P.B.L., Keith, A.M., Creer, S., Barrett, G.L., Lebron, I., Emmett, B.A., Robinson, D.A., Jones, D.L., 2017. Evaluation of mesofauna communities as soil quality indicators in a national- level monitoring programme. Soil Biol. Biochem. 115, 537-546.
    13. Gerlach, J., Samways, M., Pryke, J., 2013. Terrestrial invertebrates as bioindicators: an overview of available taxonomic groups. J. Insect Conserv. 17, 831-850.
    14. Gulvik, M.E., 2007. Mites (Acari) as indicators of soil biodiversity and land use monitoring: a review. Polish J. Ecol. 55, 415-440.
    15. Gwiazdowicz, D., Bloszyk, J., Bajerlein, D., Halliday, R., Mizera, T., 2006. Mites (Acari: Mesostigmata) inhabiting nests of the white-tailed sea eagle Haliaeetus albicilla (L.) in Poland. Entomol. Fenn. 17, 366-372.
    16. Hajizadeh, J., Ramzi, S., Daghighi, E., 2020. Mesostigmatid mites (Acari: Mesostigmata) associated with tea orchards in Iran. J. Biol. Stud. 3, 30-47.
    17. Hwang, U.W., Rho, H.S., Park, B., Choi, E.H., Shin, C.R., Kim, S.H., Lee, J.R., Kim, H.C., Shin, M.K., Park, T.S., Jun, J.M., Lee, H.G., Lee, J.E., Oh, Y.S., Myoung, J.G., Choi, C.G., Park, J.H., Park, S.J., Lee, J.M., Lee, J.H., Kwon, H.Y., Park, K.T., Lim, C.W., Jung, S.W., Lee, M.J., Lee, Y.C., Shin, Y.H., Choi, H.J., Lee, Y.W., Kil, H.J., Kim, J.H., Kang, M.S., Lee, E.Y., Lee, S.H., Kim, Y.H., Jung, J.W., Jang, K.H., Lim, Y.J., Ryu, S.H., Min, W.G., Park, J.M., Lee, H.J., Woo, M.S., Kim, Y.B., Myoung, S.H., 2023. Comprehensive and synthetic inventory of Dokdo Island, Republic of Korea. J. Species Res. 12, 1-69.
    18. Hyatt, K.H., Embersom, R.M., 1988. A review of the Macrochelidae (Acari: Mesostigmata) of the British Isles. Bull. Br. Museum, Nat. Hist. Zool. 54, 63-125.
    19. Jeon, Y.G., 2005. The regional geomorphology of Dokdo (volcanic island). J. Korean Assoc. Reg. Geogr. 11, 19-28.
    20. Ji, S.J., Jung, C., Bang, H.W., Song, M.O., Jung, J., Yoon, S.M., Lee, S.H., Keum, S.Y., Yang, H.M., Lee, D.M., Lee, G.H., Oh, J.S., Kim, K.C., Park, H.S., Moon, H.J., Joharchi, O., Kang, Y.S., Eom, K.S., Lee, K.J., Eun, Y., Kim, T.H., Karanovic, I., Lee, J.H., Choe, S.J., Min, G.S., 2023a. Unrecorded species of Korean invertebrates discovered through the project of ‘Discovery of Korean Indigenous Species’ II. J. Species Res. 12, 68-89.
    21. Ji, S.J., Jung, J.W., Kim, S.H., Ahn, D.H., Kim, M.S., Lee, J.H., Yang, H.M., Lee, G.H., Nam, E.J., Park, T.S., Jost, A.B., Pham, H.T.M., Park, J.N., Park, J.H., Keum, S.Y., Karanovic, I., Karanovic, T., Park, J.K., Jung, C., Min, G.S., 2023b. Unrecorded species of Korean invertebrates discovered through the project of ‘Discovery of Korean Indigenous Species’ III. J. Species Res. 12, 341-354.
    22. Katoh, K., Kuma, K., Toh, H., Miyata, T., 2005. MAFFT version 5: improvement in accuracy of multiple sequence alignment. Nucleic Acids Res. 33, 511-518.
    23. Katoh, K., Rozewicki, J., Yamada, K.D., 2019. MAFFT online service: multiple sequence alignment, interactive sequence choice and visualization. Brief. Bioinform. 20, 1160-1166.
    24. Keum, E.S., Jung, C., 2012. Distribution of gamasid mites (Acari, Mesostigmata) on Seonginbong, Ulleung Island. Korean J. Soil Zool. 16, 2.
    25. Keum, E.S., Takaku, G., Lee, K.W., Jung, C., 2016. New records of phoretic mites (Acari: Mesostigmata) associated with dung beetles (Coleoptera: Scarabaeidae) in Korea and their ecological implication. J. Asia-Pac. Entomol. 19, 353-357.
    26. Khaing, T.M., Lee, J.H., Lee, W.G., Lee, K.Y., 2013. A new record of Amphitetranychus quercivorus (Acari: Tetranychidae) in Korea and molecular comparison with A. viennensis. J. Asia-Pac. Entomol. 16, 155-160.
    27. Kim, D., Hwang, H.S., Shim, J.K., Yang, J., Pak, J.H., Lee, K.Y., 2019. Molecular biodiversity assessment of plant-parasitic nematodes in Dokdo Island, Korea. Nematology 21, 275-292.
    28. Kim, J., Bayartogtokh, B., Jung, C., 2013. A new record of oribatid mite species, Punctoribates hexagonus Berlese, 1908 (Acari: Oribatida: Mycobatidae) in Korea. Korean J. Appl. Entomol. 52, 79-83.
    29. Kim, M., Kwon, Y., Nam, K., Lee, H., Myeong, H., Noh, H.S., 2017. Breeding population and habitat of black-tailed gulls (Larus crassirostris) on Nando Island, Natural Monument. Korean J. Environ. Biol. 35, 134-142.
    30. Kim, M.H., Oh, Y.J., Kim, C.S., Han, M.S., Lee, J.T., Na, Y.E., 2007. The Flora and Vegetation Distribution in Dokdo. Korean J. Environ. Agric. 26, 85-93.
    31. Knee, W., 2017. New Macrocheles species (Acari, Mesostigmata, Macrochelidae) associated with burying beetles (Silphidae, Nicrophorus) in North America. Zookeys 721, 1-32.
    32. Kontschán, J., Park, S.J., Lim, J.W., Hwang, J.M., Seo, H.Y., 2014. Contribution to the mite (Acari) fauna of the Korean Peninsula. J. Species Res. 3, 63-78.
    33. Krantz, G.W., Walter, D.E., 2009. A manual of acarology, 3rd ed. Texas Tech University Press, Lubbock.
    34. Krantz, G.W., Whitaker Jr, J.O., 1988. Mites of the genus Macrocheles (Acari: Macrochelidae) associated with small mammals in North America. Acarologia 29, 225-259.
    35. Kumar, S., Stecher, G., Li, M., Knyaz, C., Tamura, K., 2018. MEGA X: Molecular evolutionary genetics analysis across computing platforms. Mol. Biol. Evol. 35, 1547-1549.
    36. Lee, D.Y., Jeong, I., Kim, S., Choi, J.W., Won, M.H., Kim, D., Kim, D., Kim, Y.K., Jeon, J., Ryu, J., Bang, W., Chang, J.H., Choi, K.S., 2024. Checklist for the insect fauna of two East Sea Islands (Ulleungdo Is. and Dokdo Is.) in the Republic of Korea. Biodivers. Data J. 12, e129360.
    37. Lee, W., Yoo, J., 2005. Status of the breeding population of Black-tailed gulls on Hongdo is-land. Pacific Seabirds 32, 2.
    38. Lin, J.Z., Zhang, Z.Q., 2010. Macrochelidae of China: a review of progress, with a checklist. Zoosymposia 4, 272-279.
    39. Lindquist, E.E., Evans, G.O., 1965. Taxonomic concepts in the Ascidae, with a modified setal nomenclature for the idiosoma of the Gamasina (Acarina: Mesostigmata). Mem. Entomol. Soc. Canada 97, 5-66.
    40. Minh, B.Q., Schmidt, H.A., Chernomor, O., Schrempf, D., Woodhams, M.D., von Haeseler, A., Lanfear, R., 2020. IQ-TREE 2: new models and efficient methods for phylogenetic inference in the genomic Era. Mol. Biol. Evol. 37, 1530-1534.
    41. Namirimu, T., Kim, J., Zo, Y.G., 2019. Isolation and identification of alkali-tolerant bacteria from near-shore soils in Dokdo island. Microbiol. Biotechnol. Lett. 47, 105-115.
    42. National Institute of Biological Resources (NIBR), 2024, National List of Korea. https://kbr.go.kr/ (Acessed 27 March, 2025).
    43. Nylander, J.A.A., 2004. MrModeltest v2. Program distributed by the author. Evolutionary Biology Centre, Uppsala University, Uppsala.
    44. Ronquist, F., Teslenko, M., Van Der Mark, P., Ayres, D.L., Darling, A., Höhna, S., Larget, B., Liu, L., Suchard, M.A., Huelsenbeck, J.P., 2012. MrBayes 3.2: Efficient Bayesian phylogenetic inference and model choice across a large model space. Syst. Biol. 61, 539-542.
    45. Takaku, G., 2000. Macrochelid mites (Acari: Macrochelidae) associated with Trox sugayai Masumoto and Kiuchi (Coleoptera: Trogidae) on Amami-Oshima Island, Japan. J. Acarol. Soc. Japan 9, 119-127.

    Vol. 40 No. 4 (2022.12)

    Journal Abbreviation Korean J. Appl. Entomol.
    Frequency Quarterly
    Doi Prefix 10.5656/KSAE
    Year of Launching 1962
    Publisher Korean Society of Applied Entomology
    Indexed/Tracked/Covered By