Published January 12, 2026
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A new species of the Amynthas corticis group with support from mitogenomic data and a new record of Metaphire agrestis (Goto & Hatai, 1899) (Oligochaeta, Megascolecidae) in northeastern China
Authors/Creators
- 1. Northeast Normal University, Changchun, China|Langfang Normal University, Langfang, China
- 2. Langfang Normal University, Langfang, China
- 3. Mindanao State University at Naawan, Naawan, Philippines
- 4. Northeast Normal University, Changchun, China
- 5. Northeast Normal University, Changchun, China|Northeast Institute of Geography and Agroecology, Chinese Academy of Sciences, Changchun, China
- 6. Langfang Normal University, Langfang, China|Northeast Institute of Geography and Agroecology, Chinese Academy of Sciences, Changchun, China
Description
As part of a series of investigations on earthworm diversity conducted in northeastern China in recent years, a new species, Amynthas dandongensis Han & Zhao, sp. nov., has been discovered. It belongs to the A. corticis species group characterized by having four pairs of spermathecal pores in segments 5/6/7/8/9. The new species mainly differs from the other members of the A. corticis species group by its relatively small body size and the absence of male pores and genital markings in both the spermathecal and male pore regions. The interspecific pairwise COI genetic distances between A. dandongensis Han & Zhao, sp. nov. and the other closely related species ranged from 15% to 22%. Phylogenetic analyses using mitogenomic coding genes further explored the relationships within the A. corticis species group, clarifying the position of A. dandongensis Han & Zhao, sp. nov. in the group. Additionally, a new record of a cosmopolitan earthworm, Metaphire agrestis (= Amynthas agrestis) (Goto & Hatai, 1899) has been investigated in China. New sequences from specimens collected in northeastern China were analyzed together with "M. agrestis" and "A. agrestis" sequences available from GenBank to further explore phylogeographic relationship. The dataset of all M. agrestis sequences showed intraspecific genetic distances of 0–1%, consistent with the observed morphological similarities. Haplotype network analyses also revealed no genetic isolation among different populations of M. agrestis (= A. agrestis). Moreover, re-examination of previous descriptions and illustrations of the male pores of agrestis indicates that this species should be assigned to the genus Metaphire Sims & Easton, 1872, rather than Amynthas Kinberg, 1867.
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References
- Aspe NM (2016) The geographic distribution of the genera in the Pheretima complex (Megascolecidae) in eastern Asia and the Pacific region. Kaiyo Monthly 48: 39–45.
- Aspe NM, James S (2014) New species of Pheretima (Oligochaeta: Megascolecidae) from the Mt. Malindang Range, Mindanao Island, Philippines. Zootaxa 3881(5): 401–439. https://doi.org/10.11646/zootaxa.3881.5.1
- Aspe NM, James SW (2018) Molecular phylogeny and biogeographic distribution of pheretimoid earthworms (Clitellata: Megascolecidae) of the Philippine archipelago. European Journal of Soil Biology 85: 89–97. https://doi.org/10.1016/j.ejsobi.2018.02.001
- Aspe NM, Zhao H, Mapile MRF, Obusan MCM, Zhang Y, Castañares E, Florida EJD, Wu D (2025) Three new species and a new record of pheretimoid earthworms (Crassiclitellata, Megascolecidae) from Misamis Oriental, Philippines, with data from the mitochondrial genome. Zoosystematics and Evolution 101(1): 389–404. https://doi.org/10.3897/zse.101.135702
- Bantaowong U, Chanabun R, Tongkerd P, Sutcharit C, James SW, Panha S (2011) A new species of the terrestrial earthworm of the genus Metaphire Sims & Easton, 1972 from Thailand with redescription of some species. Tropical Natural History 11(1): 55–69. https://doi.org/10.58837/tnh.11.1.103012
- Beddard FE (1900) A revision of the earthworms of the genus Amynthas (Perichaeta). Proceedings of the Zoological Society of London 69(4): 609–652. https://doi.org/10.1111/j.1096-3642.1890.tb01727.x
- Bely AE, Wray GA (2004) Molecular phylogeny of naidid worms (Annelida: Clitellata) based on cytochrome oxidase I. Molecular Phylogenetics and Evolution 30(1): 50–63. https://doi.org/10.1016/S1055-7903(03)00180-5
- Blakemore RJ (2003) Japanese earthworms (Annelida: Oligochaeta): a review and checklist of species. Organisms, Diversity & Evolution 3(3): 241–244. https://doi.org/10.1078/1439-6092-00082
- Blakemore RJ (2004) Checklist of Pheretimoid earthworms after Sims & Easton 1972. In Avances en taxo nomia de lombrices de tierra/Advances in earthworm taxonomy (Annelida: Oligochaeta), edited by A.G. Moreno and S. Borges. Madrid, Spain: Editorial Com plutense, Universidad Complutense: 126–154.
- Blakemore RJ (2007a) Updated checklist of pheretimoids (Oligochaeta: Megascolecidae: Pheretima auct.) taxa. In: Ito MT, Kaneko N, (Eds), A Series of Searchable Texts on Earthworm Biodiversity, Ecology and Systematics from Various Regions of the World, 2nd Edition 2006 and Supplemental. COE soil Ecology Research Group, Yokohama National University, Japan. [CD-ROM Publication]
- Blakemore RJ (2007b) Checklist of megadrile earthworms (Annelida: Oligochaeta) from mainland China plus Hainan Island. Chapter 39 of Blakemore, R.J. A Series of Searchable Texts on Earthworm Biodiversity, Ecology and Systematics from Various Regions of the World (3rd Edition) 47.
- Blakemore RJ (2008) A review of Japanese earthworms after Blakemore 2003. In: Ito MT, Kaneko N (Eds) A Series of Searchable Texts on Earthworm Biodiversity, Ecology and Systematics from Various Regions of the World, 2nd Edition 2006 and Supplemental. COE soil Ecology Research Group, Yokohama National University, Japan. [CD-ROM Publication]
- Blakemore RJ (2009) Cosmopolitan Earthworms—A Global and Historical Perspective. In: Shain DH (Ed.) Annelids in Modern Biology. Wiley and Blackwell, New Jersey, 257–283. https://doi.org/10.1002/9780470455203.ch14
- Blakemore RJ (2010) Saga of Herr Hilgendorf's worms. Zoology in the Middle East 51(sup2): 7–22. https://doi.org/10.1080/09397140.2010.10638453
- Blakemore RJ (2012) Japanese earthworms revisited a decade on. Zoology in the Middle East 58(sup4, suppl 4): 15–22. https://doi.org/10.1080/09397140.2012.10648981
- Blakemore RJ (2013a) Ulleung-do earthworms - Dagelet Island revisited. Journal of Species Research 2(1): 55–68. https://doi.org/10.12651/JSR.2013.2.1.055
- Blakemore RJ (2013b) Jeju-do earthworms (Oligochaeta: Megadrilacea) - Quelpart Island revisited. Journal of Species Research 2(1): 15–54. https://doi.org/10.12651/JSR.2013.2.1.015
- Blakemore RJ (2014) Miscellaneous earthworm types in the Natural History Museum, London (Annelida: Oligochaeta: Megadrilacea: Eudrilidae, Lumbricidae, Megascolecidae, Moniligastridae, Octochaetidae). Opuscula Zoologica (Budapest) 45: 119–155.
- Blakemore RJ, Lee S (2013) Survey of Busan Oligochaeta earthworms supported by DNA barcodes. Journal of Species Research 2(2): 127–144. https://doi.org/10.12651/JSR.2013.2.2.127
- Callaham Jr MA, Hendrix PF, Phillips RJ (2002) Occurrence of an earthworm (Amynthas agrestis) in undisturbed soils of the southern Appalachian Mountains, USA. Pedobiologia 47(5–6): 466–470. https://doi.org/10.1078/0031-4056-00214
- Cameron SL, Miller KB, D'Haese CA, Whiting MF, Barker SC (2004) Mitochondrial genome data alone are not enough to unambiguously resolve the relationships of Entognatha, Insecta and Crustacea sensu lato (Arthropoda). Cladistics: The International Journal of the Willi Hennig Society 20(6): 534–557. https://doi.org/10.1111/j.1096-0031.2004.00040.x
- Chang C-H, Chen J-H (2005) Taxonomic status and intraspecific phylogeography of two sibling species of Metaphire (Oligochaeta: Megascolecidae) in Taiwan. Pedobiologia 49(6): 591–600. https://doi.org/10.1016/j.pedobi.2005.07.002
- Chang C-H, Chen J-H (2008) Earthworm Fauna of Taiwan: Metaphire formosae species group. National Taiwan University Press, Taipei, Taiwan, 60 pp.
- Chang C-H, James S (2011) A critique of earthworm molecular phylogenetics. Pedobiologia 54: S3–S9. https://doi.org/10.1016/j.pedobi.2011.07.015
- Chang C-H, Lin Y-H, Chen IH, Chuang S-C, Chen J-H (2007) Taxonomic re-evaluation of the Taiwanese montane earthworm Amynthas wulinensis Tsai, Shen & Tsai, 2001 (Oligochaeta: Megascolecidae): Polytypic species or species complex? Organisms, Diversity & Evolution 7(3): 231–240. https://doi.org/10.1016/j.ode.2006.06.001
- Chang C-H, Lin S-M, Chen J-H (2008) Molecular systematics and phylogeography of the gigantic earthworms of the Metaphire formosae species group (Clitellata, Megascolecidae). Molecular Phylogenetics and Evolution 49(3): 958–968. https://doi.org/10.1016/j.ympev.2008.08.025
- Chang C-H, Chuang S-C, Wu J-H, Chen J-H (2014) New species of earthworms belonging to the Metaphire formosae species group (Clitellata: Megascolecidae) in Taiwan. Zootaxa 3774(4): 324–332. https://doi.org/10.11646/zootaxa.3774.4.2
- Chang C-H, Snyder B, Szlávecz K (2016) Asian pheretimoid earthworms in North America north of Mexico: An illustrated key to the genera Amynthas, Metaphire, Pithemera, and Polypheretima (Clitellata: Megascolecidae). Zootaxa 4179(3): 495–529. https://doi.org/10.11646/zootaxa.4179.3.7
- Chang C-H, Bartz M, Brown G, Callaham M, Cameron E, Davalos A, Dobson A, Gorres J, Herrick B, Ikeda H, James S, Johnston M, McCay T, McHugh D, Minamiya Y, Nouri Aiin M, Novo M, Ortiz-Pachar J, Pinder R, Szlávecz K (2021) The second wave of earthworm invasions in North America: Biology, environmental impacts, management and control of invasive jumping worms. Biological Invasions 23(11): 1–32. https://doi.org/10.1007/s10530-021-02598-1
- Chen Y, Xu Z (1977) On some new earthworms from China II. Acta Zoologica Sinica 23: 175–181.
- Chen Y, Hsu C-F, Yang T, Fong H-Y (1975) On some new earthworms from China. Acta Zoologica Sinica 21(1): 89–99. [in Chinese with English abstract]
- Davies H (1954) A preliminary list of the earthworms of northern New Jersey with notes. Breviora. Museum of Comparative Zoology 26: 1–13.
- Dong Y, Law MMS, Jiang J, Qiu J (2019) Three new species and one subspecies of the Amynthas corticis-group from Guangxi Zhuang Autonomous Region, China (Oligochaeta, Megascolecidae). ZooKeys 884: 23–42. https://doi.org/10.3897/zookeys.884.30988
- Easton EG (1981) Japanese earthworms: A synopsis of the Megadrile species (Oligochaeta). Bulletin of the British Museum, Natural History. Zoology 40(2): 33–65.
- Edwards CA, Arancon NQ, Bohlen PJ, Hendrix P (2022) Biology and ecology of earthworms. Springer New York, NY, XVI, 567 pp. https://doi.org/10.1007/978-0-387-74943-3
- 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. Molecular Marine Biology and Biotechnology 3: 294–299.
- Gates GE (1953) Further Notes on the Earthworms of the Arnold Arboretum, Boston. Museum of Comparative Zoology 15: 1–9.
- Gates GE (1954) Exotic earthworms of the United States. Bulletin of the Museum of Comparative Zoology at Harvard College 111: 217–258.
- Gates GE (1958) On some species of the Oriental earthworm genus Pheretima Kinberg, 1867: With key to species reported from the Americas. American Museum Novitates 1888: 1–33.
- Gates GE (1963) Miscellanea megadrilogica VII. Greenhouse earthworms. Proceedings of the Biological Society of Washington 76: 9–18.
- Gates GE (1966) Requiem for megadrile utopias. A contribution toward the understanding of the earthworm fauna of North America. Proceedings of the Biological Society of Washington 79: 239–254.
- Gates GE (1982) Farewell to North American Megadriles. Megadrilogica 4: 12–17.
- Görres JH, Melnichuk RDS (2012) Asian invasive earthworms of the genus Amynthas Kinberg in Vermont. Northeastern Naturalist 19(2): 313–322. https://doi.org/10.1656/045.019.0212
- Görres JH, Keller EL, Schall JJ (2017) Genetic structure of two invasive earthworms, Amynthas agrestis and Amynthas tokioensis (Oligochaeta, Megascolecidae), and a molecular method for species identification. Megadrilogica 22: 140–148.
- Goto S, Hatai S (1899) New or imperfectly known species of earthworms. No. 2. Annotationes Zoologicae Japonenses 3: 13–24.
- Han ACN, Zhang Y, Miao P, Wu S, Xiao N, Qin M, Zhao H, Wu D, Aspe NM (2024) Distribution and systematics of the cosmopolitan Amynthas carnosus complex (Crassiclitellata, Megascolecidae) from eastern Asia. Zoosystematics and Evolution 100(3): 1061–1073. https://doi.org/10.3897/zse.100.119292
- Hatai S (1930) Note on Pheretima agrestis (Goto and Hatai), together with the description of four new species of the genus Pheretima. Science Reports of the Tohoku Imperial University, 4th series. Biology (Basel) 5(4): 651–667.
- Heimburger B, Klein A, Roth A, Scheu S, Eisenhauer N, Schaefer I (2023) Population structure and genetic variance among local populations of a non-native earthworm species in Minnesota, USA. Biological Invasions 25(7): 2361–2375. https://doi.org/10.1007/s10530-023-03048-w
- Hong Y, James S (2001) New species of Korean Amynthas Kinberg, 1867 (Oligochaeta, Megascolecidae) with two pairs of spermathecae. Revue Suisse de Zoologie 108: 65–93. https://doi.org/10.5962/bhl.part.79621
- Hong Y, Kim TH (2002) Four new earthworms of the genus Amynthas (Oligochaeta: Megascolecidae) from Korea. Korean Journal of Biological Sciences 6(3): 195–199. https://doi.org/10.1080/12265071.2002.9647651
- Hong Y, Kim TH (2005) The earthworm fauna from Ulleungdo Island, Korea with one new species of the genus Amynthas (Oligochaeta: Megascolecidae). Animal Systematics, Evolution and Diversity 21: 129–136.
- Hong Y, Kim TH (2009) The earthworm composition in agroecosystem of Sunyu Island, Korea. The Korean Journal of Environmental Biology 27(2): 135–139.
- Hong Y, Inkhavilay K, James SW (2024) Three new earthworms of the Amynthas corticis group (Clitellata: Megascolecidae) from the Nam Et-Phouleoi, Laos. Zootaxa 5405(2): 273–280. https://doi.org/10.11646/zootaxa.5405.2.7
- Howell CD (1939) The responses to light in the earthworm, Pheretima agrestis Goto and Hatai, with special reference to the function of the nervous system. The Journal of Experimental Zoology 81(2): 231–259. https://doi.org/10.1002/jez.1400810205
- Huang J, Xu Q, Sun Z, Tang G, Su Z (2007) Identifying earthworms through DNA barcodes. Pedobiologia 51(4): 301–309. https://doi.org/10.1016/j.pedobi.2007.05.003
- Iki S, Oba Y (2022) Records of terrestrial earthworm fauna at Anjo Agricultural and Forestry High School. Mikawa Seibutsu 14: 40–45.
- Ishizuka K (2001) Taxonomic study of the genus Pheretima s. lat. (Oligochaeta, Megascolecidae) from Japan. Bull. Seikei Univ. 33(3): 1–125. [In Japanese]
- Ito TM, Yasuda M, Yamada F (2011) Earthworm fauna (Annelida: Clitellata) of the main campus and Chiyoda Experimental Station of Forestry and Forest Products Research Institute. Bulletin of Forestry and Forest Products Research Institute 10(4): 281–289.
- James SW (2005) Preliminary molecular phylogeny in the Pheretima group of genera (Crassiclitellata: Megascolecidae) using Bayesian analysis. Advances in earthworm taxonomy II (Annelida: Oligochaeta) Cluj University Press, Cluj-Napoca, Romania, 129–142.
- James SW, Shih HT, Chang HW (2005) Seven new species of Amynthas (Clitellata: Megascolecidae) and new earthworm records from Taiwan. Journal of Natural History 39(14): 1007–1028. https://doi.org/10.1080/00222930400001434
- Jeratthitikul E, Bantaowong U, Panha S (2017) DNA barcoding of the Thai species of terrestrial earthworms in the genera Amynthas and Metaphire (Haplotaxida: Megascolecidae). European Journal of Soil Biology 81: 39–47. https://doi.org/10.1016/j.ejsobi.2017.06.004
- Jiang J (2016) Taxonomy and Molecular Phylogeny of the Family Megascolecidae Earthworms from China. Shanghai Jiao Tong University, PhD dissertation. [in Chinese with English abstract]
- Jiang J, Qiu J (2018) Origin and evolution of earthworms belonging to the family Megascolecidae in China. Shengwu Duoyangxing 26(10): 1074–1082. https://doi.org/10.17520/biods.2018105
- Jin Q, Li J, Jiang J, Qiu J (2024) Four new earthworm species of the genera Amynthas and Metaphire (Oligochaeta, Megascolecidae) from Hunan and Anhui provinces, China. ZooKeys 1210: 247–271. https://doi.org/10.3897/zookeys.1210.125963
- Kato J (1972) Earthworms. In: Tochigi Prefecture Animal and Plant Compilation Committee (Еd.) Animals and Plants of Tochigi Prefecture. Shimotsuke Shimbun, Utsunomiya, 545–556.
- Keller EL, Görres JH, Schall JJ (2017) Genetic structure of two invasive earthworms, Amynthas agrestis and Amynthas tokioensis (Oligochaeta, Megascolecidae), and a molecular method for species identification. Megadrilogica 22: 140–148. https://doi.org/10.1007/s10530-021-02598-1
- Kimura M (1980) A simple method for estimating evolutionary rates of base substitutions through comparative studies of nucleotide sequences. Journal of Molecular Evolution 16(2): 111–120. https://doi.org/10.1007/BF01731581
- Kobayashi S (1935) Classification and distribution of Korean terrestrial earthworms (preliminary report). Zoological Journal 47: 128–130.
- Kobayashi S (1938) Earthworms of Korea I. Science Report of the Tohoku Imperial University 13: 89–170.
- Lam D, Nguyen N, Nguyen DA, Nguyen T (2021) A checklist of earthworms (Annelida: Oligochaeta) in southeastern Vietnam. Journal of Threatened Taxa 13(2): 17693–17711. https://doi.org/10.11609/jott.6535.13.2.17693-17711
- Li JL, Jiang J, Jin Q, Qiu J (2024) Three new species of the genus Amynthas (Oligochaeta: Megascolecidae) from Guangxi Zhuang Autonomous Region, China. Zootaxa 5410(1): 49–66. https://doi.org/10.11646/zootaxa.5410.1.2
- Liu M, Li J, Zhang Y, Aspe NM, Wu D, Zhao H, Zheng G (2025) A new species of the Drawida ghilarovi species complex (Oligochaeta, Moniligastridae) in Changbai Mountain, Northeast China. Zoosystematics and Evolution 101(2): 627–641. https://doi.org/10.3897/zse.101.146587
- Meng G, Li Y, Yang C, Liu S (2019) MitoZ: A toolkit for animal mitochondrial genome assembly, annotation and visualization. Nucleic Acids Research 47(11): e63. https://doi.org/10.1093/nar/gkz173
- Michaelsen W (1900) Oligochaeta. Das Tierreich 10: 1–575. https://doi.org/10.5962/bhl.title.11605
- Michaelsen W (1922) Oligochaten aus dem Rijks-Museum van natuurlijke Historie zu Leiden. Capita Zoologica 's Gravenhage 1(3): 1–72.
- Minamiya Y (2014) Encyclopedia of Japanese earthworms. https://japanese-mimizu.jimdofree.com/ [accessed 19 Mar 25]
- Minamiya Y (2017) Fauna of large terrestrial oligochaetes (earthworms) in Nikko. Research Bulletin of Tochigi Prefectural Museum 34: 13–20.
- Nguyen TT, Nguyen NQ, Lam DH, Nguyen AD (2020) Six new species of the genus Metaphire Sims & Easton, 1972 (Annelida: Oligochaeta: Megascolecidae) from southeastern Vietnam. The Raffles Bulletin of Zoology 68: 220–236. https://doi.org/10.26107/RBZ-2020-0019
- Nguyen TT, Ly VV, Lam DH, Nguyen TV, Nguyen AD (2022) A New Earthworm Species of the Genus Metaphire Sims & Easton, 1972 (Oligochaeta: Megascolecidae) from Southern Vietnam. Tropical Natural History 22: 72–84. https://doi.org/10.58837/tnh.22.1.256982
- Ohfuchi S (1937) Description of three new species of the genus Pheretima from northeastern Honshu, Japan. Saito Ho-on Kai Museum of Natural History. Research Bulletin (International Commission for the Northwest Atlantic Fisheries) 12: 13–29.
- Ohfuchi M (1938) On the genus Pheretima in northeastern Honshu. Zoological Journal 50(4): 177–178.
- Ohfuchi S (1957) On a collection of terrestrial Oligochaeta obtained from the Riu-Kiu Islands, together with a note on their geographical distribution (Part 2). Journal of Agricultural Science, Tokyo 3: 243–261.
- Owen PC, Wilson KA (2024) Molecular Survey of Earthworms (Annelida: Clitellata) in Southwestern Ohio, with Identification of a New Non-Native Species (Family Moniligastridae). Ohio Biological Survey Notes 13: 1–5.
- Qin M, Zhao H, Han ACN, Wu D, Zhang Y, Shekhovtsov SV, Aspe NM (2025) Description of three new subspecies of Eisenia nordenskioldi (Annelida: Crassiclitellata) from Northeastern China. Zootaxa 5642(3): 325–344. https://doi.org/10.11646/zootaxa.5642.3.2
- Reynolds JW (1978) The earthworms of Tennessee (Oligochaeta). IV. Megascolecidae, with notes on distribution, biology and a key to the species in the state. Megadrilogica 3: 117–129.
- Reynolds JW (2010) The earthworms (Oligochaeta: Acanthodrilidae, Lumbricidae, Megascolecidae and Sparganophilidae) of northeastern United States, revisited. Megadrilogica 14: 101–157.
- Reynolds JW (2011) The earthworms (Oligochaeta: Acanthodrilidae, Eudrilidae, Glossoscolecidae, Komarekionidae, Lumbricidae, Lutodrilidae, Ocnerodrilidae, Octochaetidae, Megascolecidae and Sparganophilidae) of southeastern United States. Megadrilogica 14: 175–318.
- Reynolds JW (2014) A checklist by counties of earthworms (Oligochaeta: Lumbricidae, Megascolecidae and Sparganophilidae) in Ontario, Canada. Megadrilogica 16: 111–135.
- Reynolds JW (2015a) A checklist by counties of earthworms (Oligochaeta: Lumbricidae, Megascolecidae and Sparganophilidae) in the states of Connecticust, Massachusetts and Rhode Island, USA. Megadrilogica 17(8): 113–123.
- Reynolds JW (2015b) A checklist by counties of earthworms (Oligochaeta: Lumbricidae, Megascolecidae and Sparganophilidae) in New Jersey, USA. Megadrilogica 17(12): 179–190.
- Reynolds JW (2022) The Earthworms (Lumbricidae, Megascolecidae and Sparganophilidae) in Canada. Canada Food Inspection Agency, Ottawa, 1–179.
- Reynolds JW (2023) Current status of Asian pheretimoid earthworms (Oligochaeta: Megascolecidae) in Canada. Megadrilogica 28: 1–20.
- Reynolds JW, Wetzel MJ (2004) Terrestrial Oligochaeta (Annelida: Clitellata) in North America north of Mexico. Megadrilogica 9: 71–98.
- Ronquist F, Teslenko M, van der Mark P, Ayres DL, Darling A, Hohna S, Larget B, Liu L, Suchard MA, Huelsenbeck JP (2012) MrBayes 3.2: Efficient Bayesian phylogenetic inference and model choice across a large model space. Systematic Biology 61(3): 539–542. https://doi.org/10.1093/sysbio/sys029
- Rozas J, Ferrer-Mata A, Sánchez-DelBarrio JC, Guirao-Rico S, Librado P, Ramos-Onsins SE, Sánchez-Gracia A (2017) DnaSP 6: DNA Sequence Polymorphism Analysis of Large Datasets. Molecular Biology and Evolution 34(12): 3299–3302. https://doi.org/10.1093/molbev/msx248
- Sato C, Nendai N, Nagata N, Okuzaki Y, Ikeda H, Minamiya Y, Sota T (2023) Origin and diversification of pheretimoid megascolecid earthworms in the Japanese Archipelago as revealed by mitogenomic phylogenetics. Molecular Phylogenetics and Evolution 182: 107735. https://doi.org/10.1016/j.ympev.2023.107735
- Schall JJ, Nouri-Aiin M, Görres J (2023) Apolocystis bosanqueti n. sp. (Apicomplexa: Eugregarinorida) from the invasive earthworm Amynthas agrestis (Annelida: Megascolecidae), with significance for the monophyly of the family Monocystidae. The Journal of Parasitology 109(2): 56–64. https://doi.org/10.1645/22-66
- Shekhovtsov S, Sundukov YN, Blakemore R, Gongalsky K, Peltek SE (2018) Identifying earthworms (Oligochaeta, Megadrili) of the southern Kuril islands using DNA barcodes. Animal Biodiversity and Conservation 41(1): 9–17. https://doi.org/10.32800/abc.2018.41.0009
- Shekhovtsov S, Rapoport I, Leonov P, Poluboyarova T, Yurlova G, Shipova A, Vasiliev G, Golovanova E, Roohi Aminjan A, Latif R (2024) Phylogeny of Lumbricidae based on mitochondrial genomes with emphasis on the genus Dendrobaena. Organisms, Diversity & Evolution 24(4): 477–487. https://doi.org/10.1007/s13127-024-00656-9
- Shen H-P (2012) Three new earthworms of the genus Amynthas (Megascolecidae: Oligochaeta) from eastern Taiwan with redescription of Amynthas hongyehensis Tsai and Shen, 2010. Journal of Natural History 46(37–38): 2259–2283. https://doi.org/10.1080/00222933.2012.716867
- Shen H-P (2018) Biogeography meets taxonomy: Distribution-based inferences on the accuracy of identification and synonymization of East Asian earthworms. Zootaxa 4496(1): 313–336. https://doi.org/10.11646/zootaxa.4496.1.26
- Shen H-P, Chang C-H, Li C-L, Chih W-J, Chen J-H (2013) Four new earthworm species of the genus Amynthas (Oligochaeta: Megascolecidae) from Kinmen, Taiwan. Zootaxa 3599(5): 471–482. https://doi.org/10.11646/zootaxa.3599.5.4
- Shen H-P, Chang C-H, Chih W-J (2014) Five new earthworm species of the genera Amynthas and Metaphire (Megascolecidae: Oligochaeta) from Matsu, Taiwan. Journal of Natural History 48(9–10): 495–522. https://doi.org/10.1080/00222933.2013.826742
- Shen H-P, Chang C-H, Chih W-J (2019) Two new earthworm species of the genus Amynthas (Oligochaeta: Megascolecidae) from central Taiwan, with comments on some recent species assignments in Amynthas and Metaphire. Zootaxa 4658(1): 101–123. https://doi.org/10.11646/zootaxa.4658.1.4
- Sims RW, Easton EG (1972) A numerical revision of the earthworm genus Pheretima auct. (Megascolecidae: Oligochaeta) with the recognition of new genera and an appendix on the earthworms collected by the Royal Society North Borneo Expedition. Biological Journal of the Linnean Society. Linnean Society of London 4(3): 169–268. https://doi.org/10.1111/j.1095-8312.1972.tb00694.x
- Sjöstrand AE, Sjödin P, Jakobsson M (2014) Private haplotypes can reveal local adaptation. BMC Genetics 15(1): 61. https://doi.org/10.1186/1471-2156-15-61
- Song MJ, Paik KY (1969) Preliminary survey of the earthworms from Dagelet Isl., Korea. Tongmul Hakhoe Chi 12(1): 13–21.
- Song MJ, Paik KY (1970a) Earthworms from Chejoo-do Isl., Korea. Tongmul Hakhoe Chi 1(3): 9–14.
- Song MJ, Paik KY (1970b) On a small collection of earthworms from Geo-je Isl., Korea. Tongmul Hakhoe Chi 4(13): 101–111.
- Song MJ, Paik KY (1971) Earthworms from Mt. Jiri, Korea. Tongmul Hakhoe Chi 14: 192–198.
- Song MJ, Paik KY (1973) Earthworms from Mt. Sopaik, Korea. Tongmul Hakhoe Chi 16(1): 5–12.
- Stamatakis A (2014) RAxML version 8: A tool for phylogenetic analysis and post-analysis of large phylogenies. Bioinformatics (Oxford, England) 30(9): 1312–1313. https://doi.org/10.1093/bioinformatics/btu033
- Sun J, Jiang J, Qiu J (2012) Four new species of the Amynthas corticis-group (Oligochaeta: Megascolecidae) from Hainan Island, China. Zootaxa 3458(1): 149–158. https://doi.org/10.11646/zootaxa.3458.1.8
- Sun J, Zhao Q, Jiang J, Qiu J (2013) New Amynthas species (Oligochaeta: Megascolecidae) from south and central Hainan Island, China and estimates of evolutionary divergence among some corticis-group species. Journal of Natural History 47(17–18): 1143–1160. https://doi.org/10.1080/00222933.2012.743613
- Sun J, Jiang J, Bartlam S, Qiu JP, Hu F (2018) Four new Amynthas and Metaphire earthworm species from nine provinces in southern China. Zootaxa 4496(1): 287–301. https://doi.org/10.11646/zootaxa.4496.1.24
- Tamura K, Peterson D, Peterson N, Stecher G, Nei M, Kumar S (2011) MEGA5: Molecular evolutionary genetics analysis using maximum likelihood, evolutionary distance, and maximum parsimony methods. Molecular Biology and Evolution 28(10): 2731–2739. https://doi.org/10.1093/molbev/msr121
- Tsai C-F (1964) On some earthworms belonging to the genus Pheretima Kinberg collected from Taipei area in North Taiwan. Quarterly Journal of the Taiwan Museum 17: 1–35.
- Tsai C-F, Shen H-P, Tsai S-C (2000) Native and exotic species of terrestrial earthworms (Oligochaeta) in Taiwan with reference to Northeast Asia. Zoological Studies Taipei 39: 285–294.
- Tsai C-F, Shen H-P, Tsai S-C (2001) Some new earthworms of the genus Amynthas (Oligochaeta: Megascolecidae) from Mt. Hohuan of Taiwan. Zoological Studies Taipei 40: 276–288.
- Tsai C-F, Tsai S-C, Shen H-P (2004) A new gigantic earthworm of the genus Metaphire Sims and Easton (Megascolecidae: Oligochaeta) from Taiwan with reference to evolutional trends in body sizes and segment numbers of the Pheretima genus group. Journal of Natural History 38(7): 877–887. https://doi.org/10.1080/0022293021000053867
- Tsai C-F, Shen H-P, Tsai S-C, Lee H-H (2007) Four new species of terrestrial earthworms belonging to the genus Amynthas (Megascolecidae: Oligochaeta) from Taiwan with discussion on speculative synonyms and species delimitation in oligochaete taxonomy. Journal of Natural History 41(5–8): 357–379. https://doi.org/10.1080/00222930701201279
- Tsai C-F, Shen H-P, Tsai S-C, Lin K-J, Hsieh H-L, Yo S-P (2009) A checklist of Oligochaeta (Annelida) from Taiwan and its adjacent islands. Zootaxa 2133(1): 33–48. https://doi.org/10.11646/zootaxa.2133.1.3
- Tsai C-F, Shen H-P, Tsai S-C (2010) Four new species of Amynthas earthworms (Oligochaeta: Megascolecidae) from the Central Mountain Range of southern Taiwan. Journal of Natural History 44(21–22): 1251–1267. https://doi.org/10.1080/00222931003624788
- Uchida T, Ihara M (2003) Soil macrofauna of Mt. Kinka. Bulletin of Environmental Education Research. Miyagi University of Education 6: 31–37.
- Uchida T, Kaneko N (2004) Life history of Megascolecidae earthworms in forest soils at Kanagawa, Japan. Edaphologia 74: 35–45.
- Uehira Y (1978) Ecological studies of terrestrial oligochaetes in Hakodate. Biological Materials 7: 43–51.
- Vabeiryureilai M, Zothansanga C, Lalchhanhima M, Kumar NS, Lalthanzara H (2020) Study on the Amynthas (Kinberg, 1867) earthworm (Megascolecidae: Oligochaeta) diversity through DNA barcoding from Northeast India. Journal of Environmental Biology 41(4): 867–873. https://doi.org/10.22438/jeb/4(SI)/MS_1919
- Wang Y-H, Shih H-T (2010) Earthworm fauna of eastern Taiwan, with descriptions of two new species (Oligochaeta: Megascolecidae). Zootaxa 2341(1): 52–68. https://doi.org/10.11646/zootaxa.2341.1.2
- Xu Q, Xiao N (2011) Terrestrial Earthworms (Oligochaeta, Opisthopora) of China. China Agriculture Press, Beijing, 395 pp.
- Yamaguchi H (1962) On earthworms belonging to the genus Pheretima, collected from the southern part of Hokkaido. Journal of Hokkaido Gakugei University 13: 1–21.
- Zhang L, Jiang J, Dong Y, Qiu J (2015) Complete mitochondrial genome of four pheretimoid earthworms (Clitellata: Oligochaeta) and their phylogenetic reconstruction. Gene 574(2): 308–316. https://doi.org/10.1016/j.gene.2015.08.020
- Zhang L, Sechi P, Yuan M, Jiang J, Dong Y, Qiu J (2016) Fifteen new earthworm mitogenomes shed new light on phylogeny within the Pheretima complex. Scientific Reports 6(1): 20096. https://doi.org/10.1038/srep20096
- Zhao Q, Cluzeau D, Jiang J, Petit EJ, Briard C, Sun J, Pringzing A, Qiu J (2015) Molecular phylogeny of pheretimoid earthworms (Haplotaxina: Megascolecidae) based on mitochondrial DNA in Hainan Island, China. Molecular Biology (Los Angeles, Calif. ) 4(4): 138. https://doi.org/10.4172/2168-9547.1000138
- Zhao H, Fan S, Aspe NM, Feng L, Zhang Y (2022) Characterization of 15 earthworm mitogenomes from Northeast China and its Phylogenetic implication (Oligochaeta: Lumbricidae, Moniligastridae). Diversity 14(9): 714. https://doi.org/10.3390/d14090714
- Zhao H, Han ACN, Liu M, Zhang Y, Aspe NM, Miao P, Wu D (2025) A new pheretimoid earthworm of the genus Metaphire Sims & Easton, 1972 (Oligochaeta, Megascolecidae) from northeastern China with data from the mitochondrial genome. Zoosystematics and Evolution 101(1): 81–89. https://doi.org/10.3897/zse.101.136027