{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,4,29]],"date-time":"2026-04-29T18:57:31Z","timestamp":1777489051568,"version":"3.51.4"},"reference-count":73,"publisher":"PeerJ","license":[{"start":{"date-parts":[[2022,7,13]],"date-time":"2022-07-13T00:00:00Z","timestamp":1657670400000},"content-version":"unspecified","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/100005825","name":"USDA National Institute of Food and Agriculture","doi-asserted-by":"crossref","award":["1018366"],"award-info":[{"award-number":["1018366"]}],"id":[{"id":"10.13039\/100005825","id-type":"DOI","asserted-by":"crossref"}]},{"DOI":"10.13039\/100010941","name":"University of Vermont","doi-asserted-by":"crossref","id":[{"id":"10.13039\/100010941","id-type":"DOI","asserted-by":"crossref"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"abstract":"<jats:p>The invasive Asian earthworms,<jats:italic>Amynthas tokioensis<\/jats:italic>and<jats:italic>A. agrestis<\/jats:italic>, have been successful in entering North American forests in recent decades, with significant damage to both soils and above-ground environments. This success could be driven in part by a polyploid genetic system and parthenogenetic reproduction, often suggested as benefits for invasive species. Therefore, we assessed the genetic population structure, genetic diversity, and reproductive system of both species using morphological traits and panels of microsatellite markers. A total of 216<jats:italic>A. tokioensis<\/jats:italic>and 196<jats:italic>A. agrestis<\/jats:italic>from six sites in Vermont USA were analyzed. Although all worms were morphologically hermaphroditic, all the<jats:italic>A. agrestis<\/jats:italic>lacked the male pore (the structure allowing pass of sperm between individuals), and only 19% of the<jats:italic>A. tokioensis<\/jats:italic>possessed the male pore. All<jats:italic>A. tokioensis<\/jats:italic>earthworms were triploid (scored for three alleles for at least 1 locus, and usually several), and<jats:italic>A. agrestis<\/jats:italic>was a mix of triploid and diploid individuals. Notable was the high proportion (80%) of<jats:italic>A. agrestis<\/jats:italic>earthworms that were diploid at one site. There was clearly clonal reproduction, with identical seven- locus genotypes observed for earthworms from each site, with as many as 45 individuals with the identical genotype at one site. However, the earthworms were also genetically diverse, with 14 genotypes observed for<jats:italic>A. tokioensis<\/jats:italic>and 54 for<jats:italic>A. agrestis<\/jats:italic>, and with many singleton genotypes (a single individual). Most genotypes (71% for<jats:italic>A. tokioensis<\/jats:italic>and 92% for<jats:italic>A. agrestis<\/jats:italic>) were found at a single site. The greatest number of genotypes was found at a commercial nursery where fully 23\/26<jats:italic>A. agrestis<\/jats:italic>earthworms were singleton genotypes. As expected for the pattern of private clone alleles at sites, several measures of geographic genetic differentiation were positive, and as expected for triploid systems, an AMOVA analysis showed high within-individual genetic diversity. The paradox of clear clonal reproduction, but with a great number of genotypes for each species, and the mix of triploid and diploid individuals could be explained if the worms have been sexually reproductive, with the switch to the uniparental system only recently (or even if sexual reproduction is episodic). Last, a large number of microsatellite loci were recovered for each species and there sequence and suggested PCR primers are provided for free use by other researchers.<\/jats:p>","DOI":"10.7717\/peerj.13622","type":"journal-article","created":{"date-parts":[[2022,7,13]],"date-time":"2022-07-13T07:59:52Z","timestamp":1657699192000},"page":"e13622","source":"Crossref","is-referenced-by-count":6,"title":["Genetic population structure and reproductive system of two invasive Asian earthworms,<i>Amynthas tokioensis<\/i>and<i>Amynthas agrestis<\/i>"],"prefix":"10.7717","volume":"10","author":[{"given":"Maryam","family":"Nouri-Aiin","sequence":"first","affiliation":[{"name":"Plant and Soil Science Department, University of Vermont, Burlington, VT, United States of America"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Samantha","family":"Connolly","sequence":"additional","affiliation":[{"name":"Plant Biology Department, University of Vermont, Burlington, VT, United States of America"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Cheryl","family":"Keough","sequence":"additional","affiliation":[{"name":"Plant and Soil Science Department, University of Vermont, Burlington, VT, United States of America"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Annie Jean","family":"Smigelsky","sequence":"additional","affiliation":[{"name":"Plant and Soil Science Department, University of Vermont, Burlington, VT, United States of America"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Yiyi","family":"Wen","sequence":"additional","affiliation":[{"name":"Environmental Science, Rubinstein School of the Environment, University of Vermont, Burlington, VT, United States of America"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Jeremy","family":"Howland","sequence":"additional","affiliation":[{"name":"Plant and Soil Science Department, University of Vermont, Burlington, VT, United States of America"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Jos. J.","family":"Schall","sequence":"additional","affiliation":[{"name":"Department of Biology, University of Vermont, Burlington, VT, United States of America"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Josef H.","family":"G\u00f6rres","sequence":"additional","affiliation":[{"name":"Plant and Soil Science Department, University of Vermont, Burlington, VT, United States of America"}],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"4443","published-online":{"date-parts":[[2022,7,13]]},"reference":[{"key":"10.7717\/peerj.13622\/ref-1","doi-asserted-by":"crossref","first-page":"24","DOI":"10.1046\/j.1523-1739.2003.02365.x","article-title":"Introduction: population biology, evolution, and control of invasive species","volume":"1","author":"Allendorf","year":"2003","journal-title":"Conservation Biology"},{"key":"10.7717\/peerj.13622\/ref-2","first-page":"147","article-title":"Characteristics and modes of origin of weeds","author":"Baker","year":"1965","journal-title":"The Genetics of Colonizing Species"},{"key":"10.7717\/peerj.13622\/ref-3","doi-asserted-by":"publisher","first-page":"505","DOI":"10.1038\/nature13855","article-title":"Belowground biodiversity and ecosystem functioning","volume":"515","author":"Bardgett","year":"2014","journal-title":"Nature"},{"key":"10.7717\/peerj.13622\/ref-4","doi-asserted-by":"crossref","DOI":"10.4324\/9780429322884","volume-title":"The masterpiece of nature: the evolution and genetics of sexuality","author":"Bell","year":"2019"},{"key":"10.7717\/peerj.13622\/ref-5","doi-asserted-by":"publisher","first-page":"241","DOI":"10.1078\/1439-6092-00082","article-title":"Japanese earthworms (Annelida: Oligochaeta): a review and checklist of species","volume":"3","author":"Blakemore","year":"2003","journal-title":"Organisms Diversity & Evolution"},{"key":"10.7717\/peerj.13622\/ref-6","doi-asserted-by":"publisher","first-page":"2101","DOI":"10.1111\/j.1365-294X.2004.02209.x","article-title":"A simple method for the calculation of microsatellite genotype distances irrespective of ploidy level","volume":"13","author":"Bruvo","year":"2004","journal-title":"Molecular Ecology"},{"key":"10.7717\/peerj.13622\/ref-7","doi-asserted-by":"publisher","first-page":"117","DOI":"10.1007\/978-1-4020-5429-7_11","article-title":"Policy and management responses to earthworm invasions in North America","volume-title":"Biological invasions belowground: earthworms as invasive species","author":"Callaham","year":"2006"},{"key":"10.7717\/peerj.13622\/ref-8","doi-asserted-by":"publisher","first-page":"3291","DOI":"10.1007\/s10530-021-02598-1","article-title":"The second wave of earthworm invasions in North America: biology, environmental impacts, management and control of invasive jumping worms","volume":"23","author":"Chang","year":"2021","journal-title":"Biological Invasions"},{"key":"10.7717\/peerj.13622\/ref-9","doi-asserted-by":"publisher","first-page":"495","DOI":"10.11646\/zootaxa.4179.3.7","article-title":"Asian pheretimoid earthworms in North America north of Mexico: an illustrated key to the genera Amynthas, Metaphire, Pithemera, and Polypheretima (Clitellata: Megascolecidae)","volume":"4179","author":"Chang","year":"2016","journal-title":"Zootaxa"},{"key":"10.7717\/peerj.13622\/ref-10","doi-asserted-by":"publisher","first-page":"1090","DOI":"10.1111\/1755-0998.12639","article-title":"Resolving microsatellite genotype ambiguity in populations of allopolyploid and diploidized autopolyploid organisms using negative correlations between allelic variables","volume":"17","author":"Clark","year":"2017","journal-title":"Molecular Ecology Resources"},{"key":"10.7717\/peerj.13622\/ref-11","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1206\/3947.1","article-title":"Hybridization between whiptail lizards in Texas: Aspidoselis laredoensis and A. gularis, with notes on reproduction of a hybrids","volume":"3947","author":"Cole","year":"2020","journal-title":"American Museum Novitates"},{"key":"10.7717\/peerj.13622\/ref-12","doi-asserted-by":"publisher","first-page":"836","DOI":"10.1038\/nrg1711","article-title":"The advantages and disadvantages of being polyploid","volume":"6","author":"Comai","year":"2005","journal-title":"Nature Reviews Genetics"},{"key":"10.7717\/peerj.13622\/ref-13","doi-asserted-by":"publisher","first-page":"213","DOI":"10.1038\/sj.mp.4000500","article-title":"Molecular heterosis as the explanation for the controversy about the effect of the DRD2 gene on dopamine D2 receptor density","volume":"4","author":"Comings","year":"1999","journal-title":"Molecular Psychiatry"},{"key":"10.7717\/peerj.13622\/ref-14","first-page":"A196","article-title":"Molecular heterosis: implications for psychiatric genetics","volume":"61","author":"Comings","year":"1997","journal-title":"American Journal of Human Genetics"},{"key":"10.7717\/peerj.13622\/ref-15","doi-asserted-by":"publisher","first-page":"19","DOI":"10.1006\/mgme.2000.3015","article-title":"Molecular heterosis: a review","volume":"71","author":"Comings","year":"2000","journal-title":"Molecular Genetics and Metabolism"},{"key":"10.7717\/peerj.13622\/ref-16","doi-asserted-by":"publisher","first-page":"152","DOI":"10.1016\/j.apsoil.2017.05.029","article-title":"A large set of microsatellites for the highly invasive earthworm Amynthas corticis predicted from low coverage genomes","volume":"119","author":"Cunha","year":"2017","journal-title":"Applied Soil Ecology"},{"key":"10.7717\/peerj.13622\/ref-17","doi-asserted-by":"publisher","first-page":"31","DOI":"10.1016\/j.ejsobi.2017.06.003","article-title":"Insights into the origin of parthenogenesis in oligochaetes: strong genetic structure in a cosmopolitan earthworm is not related to reproductive mode","volume":"81","author":"De Sosa","year":"2017","journal-title":"European Journal of Soil Biology"},{"key":"10.7717\/peerj.13622\/ref-18","doi-asserted-by":"publisher","first-page":"69","DOI":"10.1007\/978-3-642-14636-7_5","article-title":"Reproduction of earthworms: sexual selection and parthenogenesis","volume":"24","author":"D\u00edaz Cos\u00edn","year":"2011","journal-title":"Biology of Earthworms"},{"key":"10.7717\/peerj.13622\/ref-19","volume-title":"Biology and ecology of earthworms","author":"Edwards","year":"1996"},{"key":"10.7717\/peerj.13622\/ref-20","doi-asserted-by":"publisher","first-page":"979","DOI":"10.3389\/fgene.2021.686246","article-title":"Genetic diversity and population structure of Metaphire vulgaris based on the mitochondrial COI gene and microsatellites","volume":"12","author":"Fang","year":"2021","journal-title":"Frontiers in Genetics"},{"key":"10.7717\/peerj.13622\/ref-21","doi-asserted-by":"publisher","first-page":"162","DOI":"10.1111\/1365-2656.12746","article-title":"Invasive earthworms erode soil biodiversity: a meta-analysis","volume":"87","author":"Ferlian","year":"2018","journal-title":"Journal of Animal Ecology"},{"key":"10.7717\/peerj.13622\/ref-22","doi-asserted-by":"publisher","first-page":"e02936","DOI":"10.1002\/ecy.2936","article-title":"Soil chemistry turned upside down: a meta-analysis of invasive earthworm effects on soil chemical properties","volume":"101","author":"Ferlian","year":"2020","journal-title":"Ecology"},{"key":"10.7717\/peerj.13622\/ref-23","doi-asserted-by":"publisher","first-page":"2085","DOI":"10.1016\/j.soilbio.2011.06.007","article-title":"A vagrant clone in a peregrine species: phylogeography, high clonal diversity and geographical distribution in the earthworm Aporrectodea trapezoides (Dug\u00e8s, 1828)","volume":"43","author":"Fern\u00e1ndez","year":"2011","journal-title":"Soil Biology and Biochemistry"},{"key":"10.7717\/peerj.13622\/ref-24","doi-asserted-by":"publisher","first-page":"127","DOI":"10.1016\/j.soilbio.2012.10.021","article-title":"Earthworms, good indicators for palaeogeographical studies? Testing the genetic structure and demographic history in the peregrine earthworm Aporrectodea trapezoides (Dug\u00e8s, 1828) in southern Europe","volume":"58","author":"Fern\u00e1ndez","year":"2013","journal-title":"Soil Biology and Biochemistry"},{"key":"10.7717\/peerj.13622\/ref-25","doi-asserted-by":"crossref","DOI":"10.1002\/9780470979365","volume-title":"Molecular ecology","author":"Freeland","year":"2011","edition":"2"},{"key":"10.7717\/peerj.13622\/ref-26","doi-asserted-by":"publisher","first-page":"502","DOI":"10.1002\/fee.2099","article-title":"Side-swiped: ecological cascades emanating from earthworm invasions","volume":"17","author":"Frelich","year":"2019","journal-title":"Frontiers in Ecology and the Environment"},{"key":"10.7717\/peerj.13622\/ref-27","doi-asserted-by":"publisher","first-page":"35","DOI":"10.1007\/978-1-4020-5429-7_5","article-title":"Earthworm invasion into previously earthworm-free temperate and boreal forests","volume-title":"Biological invasions belowground: earthworms as invasive species","author":"Frelich","year":"2006"},{"key":"10.7717\/peerj.13622\/ref-28","first-page":"55","article-title":"The earthworms of Burma-VI","volume":"52","author":"Gates","year":"1954","journal-title":"Records of the Indian Museum"},{"key":"10.7717\/peerj.13622\/ref-29","doi-asserted-by":"publisher","first-page":"571","DOI":"10.1093\/jhered\/esp057","article-title":"Ploidy and the causes of genomic evolution","volume":"100","author":"Gerstein","year":"2009","journal-title":"Journal of Heredity"},{"key":"10.7717\/peerj.13622\/ref-30","first-page":"7","article-title":"Failure of an Amynthas agrestis (Goto & Hatai, 1899) (Oligochaeta: Megascolecidae) population to expand its range within a sugar maple (Acer saccharum) stand","volume":"17","author":"G\u00f6rres","year":"2014","journal-title":"Megadrilogica"},{"key":"10.7717\/peerj.13622\/ref-31","doi-asserted-by":"publisher","first-page":"111","DOI":"10.1016\/j.apsoil.2015.10.001","article-title":"Temperature and moisture variables affecting the earthworms of genus Amynthas Kinberg, 1867 (Oligachaeta: Megascolecidae) in a hardwood forest in the Champlain Valley, Vermont, USA","volume":"104","author":"G\u00f6rres","year":"2016","journal-title":"Applied Soil Ecology"},{"key":"10.7717\/peerj.13622\/ref-32","doi-asserted-by":"publisher","first-page":"1651","DOI":"10.1007\/s10530-018-1663-x","article-title":"Winter hatching in New England populations of invasive pheretimoid earthworms Amynthas agrestis and Amynthas tokioensis: a limit on population growth, or aid in peripheral expansion?","volume":"20","author":"G\u00f6rres","year":"2018","journal-title":"Biological Invasions"},{"key":"10.7717\/peerj.13622\/ref-33","doi-asserted-by":"publisher","first-page":"13","DOI":"10.2307\/2987199","article-title":"Multivariate analysis and multidimensional geometry","volume":"17","author":"Gower","year":"1967","journal-title":"Journal of the Royal Statistical Society. Series D (The Statistician)"},{"key":"10.7717\/peerj.13622\/ref-34","doi-asserted-by":"publisher","first-page":"2017","DOI":"10.1007\/s10530-012-0208-y","article-title":"Impacts of invasive Asian (Amynthas hilgendorfi) and European (Lumbricus rubellus) earthworms in a North American temperate deciduous forest","volume":"14","author":"Greiner","year":"2012","journal-title":"Biological Invasions"},{"key":"10.7717\/peerj.13622\/ref-35","volume-title":"The causes of evolution","author":"Haldane","year":"1990"},{"key":"10.7717\/peerj.13622\/ref-36","doi-asserted-by":"publisher","first-page":"848","DOI":"10.1890\/03-5345","article-title":"Exotic European earthworm invasion dynamics in northern hardwood forests of Minnesota, USA","volume":"15","author":"Hale","year":"2005","journal-title":"Ecological Applications"},{"key":"10.7717\/peerj.13622\/ref-37","doi-asserted-by":"publisher","first-page":"500","DOI":"10.2144\/99273st03","article-title":"Universal linker and ligation procedures for construction of genomic DNA libraries enriched for microsatellites","volume":"27","author":"Hamilton","year":"1999","journal-title":"BioTechniques"},{"issue":"suppl_1","key":"10.7717\/peerj.13622\/ref-38","doi-asserted-by":"publisher","first-page":"S96","DOI":"10.1098\/rsbl.2003.0025","article-title":"Barcoding animal life: cytochrome c oxidase subunit 1 divergences among closely related species","volume":"270","author":"Hebert","year":"2003","journal-title":"Proceedings of the Royal Society of London. Series B: Biological Sciences"},{"key":"10.7717\/peerj.13622\/ref-39","doi-asserted-by":"publisher","first-page":"87","DOI":"10.1007\/978-1-4020-5429-7_9","article-title":"Invasion of exotic earthworms into ecosystems inhabited by native earthworms","volume-title":"Biological invasions belowground: earthworms as invasive species","author":"Hendrix","year":"2006"},{"key":"10.7717\/peerj.13622\/ref-40","doi-asserted-by":"publisher","first-page":"593","DOI":"10.1146\/annurev.ecolsys.39.110707.173426","article-title":"Pandora\u2019s box contained bait: the global problem of introduced earthworms","volume":"39","author":"Hendrix","year":"2008","journal-title":"Annual Review of Ecology, Evolution, and Systematics"},{"key":"10.7717\/peerj.13622\/ref-41","doi-asserted-by":"crossref","first-page":"304","DOI":"10.1016\/S0031-4056(23)03643-0","article-title":"On the evolution of clonal diversity in parthenogenetic earthworms","volume":"23","author":"Jaenike","year":"1982","journal-title":"Pedobiologia"},{"key":"10.7717\/peerj.13622\/ref-42","doi-asserted-by":"publisher","first-page":"196","DOI":"10.1086\/283622","article-title":"Clonal niche structure in the parthenogenetic earthworm Octolasion tyrtaeum","volume":"116","author":"Jaenike","year":"1980","journal-title":"The American Naturalist"},{"key":"10.7717\/peerj.13622\/ref-43","doi-asserted-by":"publisher","first-page":"729","DOI":"10.1093\/icb\/19.3.729","article-title":"Evolution and ecology of parthenogenesis in earthworms","volume":"19","author":"Jaenike","year":"1979","journal-title":"American Zoologist"},{"key":"10.7717\/peerj.13622\/ref-44","doi-asserted-by":"publisher","first-page":"1723","DOI":"10.1007\/s10530-020-02215-7","article-title":"Genetic diversity of an invasive earthworm, Lumbricus terrestris, at a long-term trading crossroad, the Champlain Valley of Vermont, USA","volume":"22","author":"Keller","year":"2020","journal-title":"Biological Invasions"},{"key":"10.7717\/peerj.13622\/ref-45","first-page":"7","article-title":"Genetic structure of two invasive earthworms, Amynthas agrestis and Amynthas tokioensis (Oligochaeta, Megascolecidae), and a molecular method for species identification","volume":"17","author":"Keller","year":"2017","journal-title":"Megadrilogica"},{"key":"10.7717\/peerj.13622\/ref-46","doi-asserted-by":"publisher","first-page":"385","DOI":"10.1023\/B:DOBS.0000046663.14974.4d","article-title":"The mechanism of cold hardiness of egg cocoons of the earthworm Dendrobaena octae dra (Sav.) (Lumbricidae: Oligochaeta)","volume":"398","author":"Leirikh","year":"2004","journal-title":"Doklady Biological Sciences"},{"key":"10.7717\/peerj.13622\/ref-47","doi-asserted-by":"publisher","first-page":"1671","DOI":"10.1007\/s10530-017-1653-4","article-title":"Tracking an invasion: community changes in hardwood forests following the arrival of Amynthas agrestis and Amynthas tokioensis in Wisconsin","volume":"20","author":"Laushman","year":"2018","journal-title":"Biological Invasions"},{"issue":"5","key":"10.7717\/peerj.13622\/ref-48","doi-asserted-by":"publisher","first-page":"478","DOI":"10.1093\/oxfordjournals.molbev.a040620","article-title":"The similarity index and dna fingerprinting","volume":"7","author":"Lynch","year":"1990","journal-title":"Molecular Biology and Evolution"},{"key":"10.7717\/peerj.13622\/ref-49","doi-asserted-by":"publisher","first-page":"150669","DOI":"10.1016\/j.pedobi.2020.150669","article-title":"Tools for monitoring and study of peregrine pheretimoid earthworms (Megascolecidae)","volume":"83","author":"McCay","year":"2020","journal-title":"Pedobiologia"},{"key":"10.7717\/peerj.13622\/ref-50","doi-asserted-by":"publisher","first-page":"239","DOI":"10.1656\/045.024.0302","article-title":"Distribution and habitat of the endemic earthworm Eisenoides lonnbergi (Michaelsen) in the northeastern United States","volume":"24","author":"McCay","year":"2017","journal-title":"Northeastern Naturalist"},{"key":"10.7717\/peerj.13622\/ref-51","doi-asserted-by":"publisher","first-page":"1126","DOI":"10.1111\/1755-0998.13145","article-title":"Genodive version 3.0: easy-to-use software for the analysis of genetic data of diploids and polyploids","volume":"20","author":"Meirmans","year":"2020","journal-title":"Molecular Ecology Resources"},{"key":"10.7717\/peerj.13622\/ref-52","doi-asserted-by":"publisher","first-page":"792","DOI":"10.1111\/j.1471-8286.2004.00770.x","article-title":"Genotype and Genodive: two programs for the analysis of genetic diversity of asexual organisms","volume":"4","author":"Meirmans","year":"2004","journal-title":"Molecular Ecology Notes"},{"key":"10.7717\/peerj.13622\/ref-53","doi-asserted-by":"publisher","first-page":"645","DOI":"10.1111\/j.1755-0998.2011.03003.x","article-title":"Coelomic fluid: a noninvasive source of DNA in earthworms","volume":"11","author":"Minamiya","year":"2011","journal-title":"Molecular Ecology Resources"},{"key":"10.7717\/peerj.13622\/ref-54","doi-asserted-by":"publisher","first-page":"113","DOI":"10.1139\/er-2017-0066","article-title":"Exotic Asian pheretimoid earthworms (Amynthas spp., Metaphire spp.): potential for colonisation of south-eastern Canada and effects on forest ecosystems","volume":"26","author":"Moore","year":"2018","journal-title":"Environmental Reviews"},{"key":"10.7717\/peerj.13622\/ref-55","doi-asserted-by":"publisher","first-page":"298","DOI":"10.1139\/er-2021-0074","article-title":"A review of exotic earthworm observations in the Canadian boreal forest and taiga zones","volume":"30","author":"Moore","year":"2022","journal-title":"Environmental Reviews"},{"key":"10.7717\/peerj.13622\/ref-56","doi-asserted-by":"publisher","first-page":"583","DOI":"10.1093\/genetics\/89.3.583","article-title":"Estimation of average heterozygosity and genetic distance from a small number of individuals","volume":"89","author":"Nei","year":"1978","journal-title":"Genetics"},{"key":"10.7717\/peerj.13622\/ref-57","doi-asserted-by":"crossref","DOI":"10.7312\/nei-92038","volume-title":"Molecular evolutionary genetics","author":"Nei","year":"1987"},{"key":"10.7717\/peerj.13622\/ref-58","doi-asserted-by":"publisher","first-page":"54","DOI":"10.1016\/j.apsoil.2019.05.004","article-title":"Earthworm cocoons: the cryptic side of invasive earthworm populations","volume":"141","author":"Nouri-Aiin","year":"2019","journal-title":"Applied Soil Ecology"},{"key":"10.7717\/peerj.13622\/ref-59","doi-asserted-by":"publisher","first-page":"e11101","DOI":"10.7717\/peerj.11101","article-title":"Biocontrol of invasive pheretimoid earthworms using Beauveria bassiana","volume":"9","author":"Nouri-Aiin","year":"2021","journal-title":"PeerJ"},{"key":"10.7717\/peerj.13622\/ref-60","doi-asserted-by":"publisher","first-page":"103822","DOI":"10.1016\/j.apsoil.2020.103822","article-title":"Identifying the unidentifiable: a PCR multiplex protocol for the diagnosis of invasive pheretimoid earthworm species, verified by morphological and barcode identification","volume":"161","author":"Nouri-Aiin","year":"2021","journal-title":"Applied Soil Ecology"},{"key":"10.7717\/peerj.13622\/ref-61","doi-asserted-by":"publisher","first-page":"557","DOI":"10.1016\/j.ympev.2010.04.010","article-title":"Cryptic speciation of hormogastrid earthworms revealed by mitochondrial and nuclear data","volume":"56","author":"Novo","year":"2010","journal-title":"Molecular Phylogenetics and Evolution"},{"key":"10.7717\/peerj.13622\/ref-62","doi-asserted-by":"publisher","first-page":"89","DOI":"10.1016\/j.soilbio.2015.02.031","article-title":"Multiple introductions and environmental factors affecting the establishment of invasive species on a volcanic island","volume":"85","author":"Novo","year":"2015","journal-title":"Soil Biology and Biochemistry"},{"key":"10.7717\/peerj.13622\/ref-63","doi-asserted-by":"publisher","first-page":"452","DOI":"10.1016\/j.cell.2007.10.022","article-title":"The evolutionary consequences of polyploidy","volume":"131","author":"Otto","year":"2007","journal-title":"Cell"},{"key":"10.7717\/peerj.13622\/ref-64","first-page":"1","article-title":"First earthworm (Annelida: Oligochaeta) species\u2019 collections in Canada and the continental United States","volume":"23","author":"Reynolds","year":"2018","journal-title":"Megadrilogica"},{"key":"10.7717\/peerj.13622\/ref-65","first-page":"83","article-title":"New Asian pheretimoid jumping earthworm records (Oligochaeta: Megascolecidae) in Canada","volume":"26","author":"Reynolds","year":"2021","journal-title":"Megadrilogica"},{"key":"10.7717\/peerj.13622\/ref-66","doi-asserted-by":"publisher","first-page":"305","DOI":"10.1146\/annurev.ecolsys.32.081501.114037","article-title":"The population biology of invasive species","volume":"32","author":"Sakai","year":"2001","journal-title":"Annual Review of Ecology and Systematics"},{"key":"10.7717\/peerj.13622\/ref-67","doi-asserted-by":"publisher","first-page":"133","DOI":"10.1016\/j.pedobi.2010.12.002","article-title":"Parthenogenesis, polyploidy and reproductive seasonality in the Taiwanese mountain earthworm Amynthas catenus (Oligochaeta, Megascolecidae)","volume":"54","author":"Shen","year":"2011","journal-title":"Pedobiologia"},{"key":"10.7717\/peerj.13622\/ref-68","doi-asserted-by":"publisher","first-page":"349","DOI":"10.1007\/s10530-010-9826-4","article-title":"Spatial variability of an invasive earthworm (Amynthas agrestis) population and potential impacts on soil characteristics and millipedes in the Great Smoky Mountains National Park, USA","volume":"13","author":"Snyder","year":"2011","journal-title":"Biological Invasions"},{"key":"10.7717\/peerj.13622\/ref-69","doi-asserted-by":"publisher","first-page":"541","DOI":"10.1038\/nature05178","article-title":"Genome-wide genetic analysis of polyploidy in yeast","volume":"443","author":"Storchov\u00e1","year":"2006","journal-title":"Nature"},{"key":"10.7717\/peerj.13622\/ref-70","doi-asserted-by":"publisher","first-page":"5","DOI":"10.1007\/978-1-4020-5429-7_2","article-title":"Dispersal and clonal diversity of North-European parthenogenetic earthworms","volume-title":"Biological invasions belowground: earthworms as invasive species","author":"Terhivuo","year":"2006"},{"key":"10.7717\/peerj.13622\/ref-71","volume-title":"The geographical distribution of animals: with a study of the relations of living and extinct faunas as elucidating the past changes of the earth\u2019s surface","author":"Wallace","year":"1876"},{"key":"10.7717\/peerj.13622\/ref-72","doi-asserted-by":"publisher","first-page":"1003","DOI":"10.1046\/j.1420-9101.1999.00119.x","article-title":"A pluralist approach to sex and recombination","volume":"12","author":"West","year":"1999","journal-title":"Journal of Evolutionary Biology"},{"key":"10.7717\/peerj.13622\/ref-73","doi-asserted-by":"publisher","first-page":"2070","DOI":"10.1890\/09-0979.1","article-title":"Dietary flexibility aids Asian earthworm invasion in North American forests","volume":"91","author":"Zhang","year":"2010","journal-title":"Ecology"}],"container-title":["PeerJ"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/peerj.com\/articles\/13622.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/peerj.com\/articles\/13622.xml","content-type":"application\/xml","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/peerj.com\/articles\/13622.html","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/peerj.com\/articles\/13622.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2023,11,24]],"date-time":"2023-11-24T14:14:27Z","timestamp":1700835267000},"score":1,"resource":{"primary":{"URL":"https:\/\/peerj.com\/articles\/13622"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2022,7,13]]},"references-count":73,"alternative-id":["10.7717\/peerj.13622"],"URL":"https:\/\/doi.org\/10.7717\/peerj.13622","archive":["CLOCKSS","LOCKSS","Portico"],"relation":{},"ISSN":["2167-8359"],"issn-type":[{"value":"2167-8359","type":"electronic"}],"subject":[],"published":{"date-parts":[[2022,7,13]]},"article-number":"e13622"}}