Length-wright relationships of invasive exotic earthworms in the Huron Mountains forests, Michigan

Length-wright relationships of invasive exotic earthworms in the Huron Mountains forests, Michigan | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Research Article Length-wright relationships of invasive exotic earthworms in the Huron Mountains forests, Michigan Xiaoyong CHEN, Timothy Gsell, John Yunger, Mary Carrington, Heng Li This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-5620138/v1 This work is licensed under a CC BY 4.0 License Status: Published Journal Publication published 26 Sep, 2025 Read the published version in Biological Invasions → Version 1 posted 5 You are reading this latest preprint version Abstract Invasive exotic earthworms have been identified as a major ecological threat to forest ecosystems in the Great Lakes region, affecting soil structure, nutrient cycling, and biodiversity. Understanding biological parameters such as body length, weight, growth rate, and regeneration patterns of these earthworms is crucial first step in comprehending their life cycle and their impacts on ecological processes in terrestrial ecosystems. In this study, an earthworm survey was conducted in the Huron Mountains of Michigan's Upper Peninsula. The earthworms were captured from sampling quadrats, identified to species, and their body mass (dry weight) and body length were measured. The purpose of the study was to develop the length-weight relationship (LWR) for the invasive exotic earthworm species that appeared in this area. Results showed that three invasive earthworm species were found in the studied forests, and they were: Dendrobaena octaedra , Aporrectodea longa , and Lumbricus terrestris . Among them, L. terrestris exhibited the largest size (mean length: 60.4 mm; mean weight: 0.254 g), while D. octaedra was the smallest (mean length: 25.5 mm; mean weight: 0.012 g). The LWR regression equations were: W = 0.00001L 2.135 for D. octaedra ; W = 0.000001L 2.816 for A. longa ; and W = 0.000003L 2.618 for L. terrestris . These findings highlighted differences in growth patterns that may be influenced by forest composition and micro-environmental factors. This study provided valuable insights into the biological characteristics of invasive exotic earthworms and their potential impacts on North American forest ecosystems. Earthworms Invasion Body size North American Forests Length-Weight Relation Figures Figure 1 Figure 2 Figure 3 Introduction Invasive species, particularly exotic earthworms, have emerged as a significant concern for North American forest ecosystems, including those in the Great Lakes region (Tiunov et al. 2006 ; Mathieu et al. 2024 ). Historically, native earthworms were largely absent from many areas of the northern United States due to glacial activity, and their reintroduction via human activities has had profound ecological effects (Hale et al. 2005 ; Craven et al. 2017 ; Baumann et al. 2024 ). The introduction of these earthworms, such as Dendrobaena octaedra , Aporrectodea longa , and Lumbricus terrestris , has altered soil structure and nutrient cycling processes (Lejoly et al. 2021 ). These invasive species, which are not native to the region, disrupt soil structure, alter nutrient cycling processes, and reduce biodiversity in plant and animal communities (Singh et al. 2019 ). These impacts are particularly pronounced in previously earthworm-free ecosystems, where the native flora and fauna are poorly adapted to the presence of these species (Phillips et al. 2019 ; Chang et al. 2021 ). To better understand the impacts of these species, it is essential and critical step to study their biological parameters including their body size, growth patterns and regeneration patterns (Chown and Gaston 2010 ; Zinger et al. 2017 ). Among the biological parameters critical to understanding invasive earthworm impacts are their length-weight relationship (LWR) (Hale et al. 2004 ). Examining the correlation between body length and weight can reveal insights into growth rates, resource allocation, and ecological adaptability. Such data are invaluable for predicting the spread and persistence of invasive populations, as well as for informing forest management strategies aimed at mitigating the consequences of earthworm invasions (Blouin et al. 2013 ; Baumann et al. 2024 ). The current study, as a part of research to examine the influence of invasive earthworms on soil structure and function in forests of the Huron Mountains, Michigan, focused on assessing the LWR of invasive exotic earthworm species in the Huron Mountains forests. By developing species-specific LWR equations, this study contributes to the broader understanding of how invasive earthworms influence soil ecosystems and forest health in northern America. Methods The study was conducted in the Huron Mountains Preserve, located in Michigan’s Upper Peninsula, USA (46°45′20″N 88°09′21″W). The average annual precipitation is about 900 mm, which is consistently spread throughout the year. Monthly temperatures vary from − 10.9°C in January to 18.3°C in July (NOAA 2009). The frost-free period lasts between 90 and 120 days. The bedrock in the area consists of Precambrian granitic, gneiss and sandstone formations (Woods 2000 ). The soils are typically shallow, acidic, and derived from rocky glacial till, though some areas feature deeper till and alluvial deposits. This preserve encompasses a range of diverse forest ecosystems, including mixed hardwoods, coniferous stands, and riparian zones. The hardwood forests are primarily composed of tree species such as eastern hemlock ( Tsuga canadensis ), sugar maple ( Acer saccharum ), yellow birch ( Betula alleghaniensis ), red maple ( Acer rubrum ), and basswood ( Tilia americana ). The study period spanned from May 2021 to October 2022, allowing for seasonal variability in earthworm activity to be captured. Three 20m × 20m plots were randomly established across main forest types in the preserve. At each plot, a 0.25 m 2 woody quadrat (0.5m × 0.5m) was used for standardized sampling earthworm. Quadrats were spaced to minimize potential edge effects and ensure independence of sampling units. Earthworms were extracted using a mustard solution, which induces the earthworms to surface for collection (Lawrence and Bowers 2002 ). All earthworms were identified to species level, and both their body length (in millimeters) and dry weight (in grams) were measured. The LWR was analyzed using regression models for each species individually. Statistical analysis included analysis of variance (ANOVA) to assess the significance of the regression models. Results and Discussion Three invasive exotic earthworm species were found in the study sites, each exhibiting significant differences in body size (Table 1 ). Among these, L. terrestris had the largest average body length (60.4 ± 35.5 mm) and weight (0.254 ± 0.344 g), followed by A. longa , which showed intermediate values for both metrics (50.3 ± 24.5 mm and 0.118 ± 0.108 g, respectively). The smallest species was D. octaedra , with a mean length of 25.5 ± 9.3 mm and a weight of 0.012 ± 0.009 g. The observed size differences have important ecological implications. The larger body size of L. terrestris suggests a higher potential for soil disturbance and nutrient cycling, as larger earthworms can ingest and process greater amounts of soil and organic matter per individual (Blouin et al. 2013 ). This aligns with studies that have reported enhanced soil aeration in the presence of large earthworm species (Bertrand et al. 2015 ). Conversely, the smaller body size of D. octaedra may reflect a different ecological role, such as focusing on leaf litter decomposition and fine-scale soil processes (Tiunov et al. 2006 ). The intermediate size of A. longa likely indicates a versatile ecological function, balancing traits of both large and small earthworms in terms of soil processing and nutrient redistribution (Coleman and Wall 2015 ; Wardle and Peltzer 2017 ; Ferlian et al. 2022 ). Table 1 Variation of body size of three invasive earthworm species in the study region* Body size L. terrestris A. longa D. Octaedra Mean length (mm) 60.4 ± 35.5 50.3 ± 24.5 25.5 ± 9.3 Length range (mm) 24.5-119.5 20.0–90.0 8.0-44.5 Mean weight (g) 0.254 ± 0.344 0.118 ± 0.108 0.012 ± 0.009 Weight range (g) 0.017–0.948 0.004–0.291 0.001–0.042 *mean ± SD. The regression models for the LWR of the three species revealed significant associations between body length and weight (Fig. 1 , 2 , and 3 ). Analysis of variance (ANOVA) confirmed the robustness of these models, with p-values less than 0.0001 for all species (Table 2 , 3 , and 4 ). For D. octaedra , the regression model explained 84.9% of the variation in body weight (F = 292.54, p < 0.00001), indicating a consistent relationship between length and weight across the population. Similarly, A. longa exhibited a strong LWR with an R² value of 93.4% (F = 214.27, p < 0.00001). L. terrestris , while having the smallest sample size, also showed a significant relationship, with 90.5% of the variance in weight explained by length (F = 85.53, p < 0.00001). Table 2 Analysis of variance for regression model of mass-length for earthworm D. octaedra Source DF SS MS F-stat p-value Model 1 43.66091 43.66091 292.53858 < 0.00001 Error 52 7.76092 0.14925 Total 53 51.42183 Table 3 Analysis of variance for regression model of mass-length for earthworm A. longa Source DF SS MS F-stat p-value Model 1 39.72614 39.72614 214.26663 < 0.00001 Error 15 2.78108 0.18541 Total 16 42.50722 Table 4 Analysis of variance for regression model of mass-length for earthworm L. terrestris Source DF SS MS F-stat p-value Model 1 26.13383 26.13383 85.52588 < 0.00001 Error 9 2.75010 0.30557 Total 10 28.88393 The differences in LWR patterns among the species likely reflect variations in energy allocation strategies (Rinke et al. 2008 ; Ehrenfeld 2011 ). For instance, L. terrestris may prioritize growth and energy storage, enabling it to dominate environments with high organic matter availability. In contrast, D. octaedra may allocate resources toward reproduction or survival in less favorable conditions, consistent with its smaller size and broader environmental tolerance (Ehrenfeld 2011 ). A. longa appears to occupy an intermediate niche, suggesting adaptability to a range of soil types and conditions (Chown and Gaston 2010 ; Rakel et al. 2020 ; Potapov 2022 ). The presence and functional traits of invasive earthworms can significantly impact soil ecosystems. Large-bodied species like L. terrestris are known to accelerate nutrient cycling, potentially altering plant community composition and forest floor dynamics. This has been documented in other studies, where invasive earthworms have facilitated shifts in nutrient availability and microbial communities (Ferlian et al. 2018 ). On the other hand, smaller species like D. octaedra may contribute more subtly by focusing on fine-scale decomposition processes and enhancing soil microstructure (Zinger et al. 2017 ). Environmental factors such as soil type, moisture, and temperature are likely to influence these growth patterns and functional roles. For example, differences in soil organic matter content and moisture availability across the Huron Mountains could explain the observed variability in body size and LWR. Similar findings have been reported by other previous studies who noted that earthworm activity and distribution are highly dependent on microhabitat conditions (Hale et al. 2005 ; Gutierrez-Lopez et al. 2010; Ahmed and Al-Mutairi 2022 ). Further studies should investigate the interaction between invasive earthworm species and forest ecosystem dynamics, particularly focusing on their combined effects on nutrient cycling, microbial communities, and plant succession. Understanding how environmental gradients influence earthworm traits and their ecological roles could provide insights into managing invasive species and mitigating their impacts on native ecosystems. Conclusion This study highlights the body size variation, LWR and ecological roles of three invasive earthworm species, L. terrestris , A. longa and D. octaedra in the Huron Mountains, revealing distinct ecological functions and impacts on soil ecosystems. L. terrestris , the largest species, demonstrated significant potential for soil disturbance and nutrient cycling, while D. octaedra , the smallest, contributed to fine-scale decomposition processes and soil microstructure. The intermediate-sized A. longa exhibited a versatile role, balancing traits of the other species. Robust LWR models (R² > 84%) reflect species-specific energy allocation strategies influenced by environmental factors such as soil type, moisture and temperature. These findings underscore the ecological implications of invasive earthworms, including their effects on nutrient cycling, microbial activity, and plant community composition. Future research should examine their combined impacts on forest ecosystems and explore how environmental gradients shape their traits and functions to inform management strategies. Declarations Acknowledgements We are grateful to Dr. Kerry Woods and Mr. Brock Francis for their assistance with the field trip and site selection at the Huron Mountains. We would also like to thank Paula Arroyo, Madeleine Naliwko, Quincy Santomieri, Lynda Randa, Dr. Yuanying Peng and Andren Yunger for their assistance with field and laboratory measurements. Authors’ contribution Conceptualization: Xiaoyong Chen, Mary Carrington; Methodology: Xiaoyong Chen, Timothy Gsell, John Yunger, Mary Carrington; Investigation Xiaoyong Chen, Timothy Gsell, John Yunger; Formal analysis: Xiaoyong Chen, Heng Li; Writing - original draft preparation: Xiaoyong Chen, Timothy Gsell, John Yunger; Funding acquisition: Xiaoyong Chen, Mary Carrington. All authors read, edited, and approved the final manuscript. Funding This work was supported by Huron Mountain Wildlife Foundation Grant (2021-2023). Data availability The data will be provided upon request on corresponding author. Conflict of interest The authors have no relevant financial or non-financial interests to disclose. Ethical approval Not applicable. References Ahmed N, Al-Mutairi KA (2022) Earthworms effect on microbial population and soil fertility as well as their interaction with agriculture practices. 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Soil Biol Biochem 171:108730 Gutiérrez-López M, Jesús JB, Trigo D, Fernández R, Novo M, Díaz-Cosín DJ (2010) Relationships among spatial distribution of soil microarthropods, earthworm species and soil properties. Pedobiologia 53:381-389 Hale CM, Reich PB, Frelich LE (2004) Allometric equations for estimation of ash-free dry mass from length measurements for selected European earthworm species ( Lumbricidae ) in the western Great Lakes region. Am Midl Nat 151:179-185 Hale CM, Frelich LE, Reich PB (2005) Exotic European earthworm invasion dynamics in northern hardwood forests of Minnesota, USA. Ecol Apps 15:848-860 Lawrence AP, Bowers MA (2002) A test of the “hot” mustard extraction method of sampling earthworms. Soil Biol Biochem 34:549-552 Lejoly J, Quideau S, Laganière J (2021) Invasive earthworms affect soil morphological features and carbon stocks in boreal forests. Geoderma 404:115262 Mathieu J, Reynolds JW, Fragoso C, Elizabeth H (2024) Multiple invasion routes have led to the pervasive introduction of earthworms in North America. Nat Ecol Evol 8:489-499 NOAA (National Oceanic and Atmospheric Administration) (2009) Climatological data annual summary-Wisconsin. Environmental Data Service, National Climatic Center, Asheville, North Carolina, USA. Phillips HR, Guerra CA, Bartz ML, Briones MJ, Brown G, Crowther TW, Orgiazzi A (2019) Global distribution of earthworm diversity. Science 366:480-485 Potapov AM (2022) Multifunctionality of belowground food webs: resource, size and spatial energy channels. Biol Rev Camb Philos Soc 97:1691-1711 Rakel KJ, Preuss TG, Gergs A (2020) Individual-based dynamic energy budget modelling of earthworm life-histories in the context of competition. 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BioRxiv 2017:154278 Cite Share Download PDF Status: Published Journal Publication published 26 Sep, 2025 Read the published version in Biological Invasions → Version 1 posted Reviewers agreed at journal 31 Jan, 2025 Reviewers invited by journal 30 Dec, 2024 Editor invited by journal 13 Dec, 2024 Editor assigned by journal 11 Dec, 2024 First submitted to journal 10 Dec, 2024 You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. As a division of Research Square Company, we’re committed to making research communication faster, fairer, and more useful. We do this by developing innovative software and high quality services for the global research community. Our growing team is made up of researchers and industry professionals working together to solve the most critical problems facing scientific publishing. Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-5620138","acceptedTermsAndConditions":true,"allowDirectSubmit":false,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":395900273,"identity":"4288cead-c220-4f98-9c77-6646061690f3","order_by":0,"name":"Xiaoyong CHEN","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAAuklEQVRIiWNgGAWjYFACHiCugHHYiNZyBsRgJkULYxspWgyOnz34uHBerZzB+fMHGD6UHSasRbInL9l45rbjxgY3khkYZ5wjQgu/BI+ZNO+2Y4kzZzAzMPO2EaGFTYLH/DfvHKCW/sMMzH+J0QKyhZm3oSaxnyGZgZmRGC2SPTnG0jzHDhjzSyQbHOw5l05Yi8HxM4afeWrq5Nj4Dz588KPMmrAWKIC45wDR6oGgjhTFo2AUjIJRMNIAAKrfNHS3aGiuAAAAAElFTkSuQmCC","orcid":"https://orcid.org/0000-0003-0810-9863","institution":"Governors State University","correspondingAuthor":true,"prefix":"","firstName":"Xiaoyong","middleName":"","lastName":"CHEN","suffix":""},{"id":395900274,"identity":"b4e0c413-0f85-4c2f-9233-3df1c6d6a46c","order_by":1,"name":"Timothy Gsell","email":"","orcid":"","institution":"Governors State University","correspondingAuthor":false,"prefix":"","firstName":"Timothy","middleName":"","lastName":"Gsell","suffix":""},{"id":395900275,"identity":"77cdd2cd-977c-4d2a-8ea9-21123f03af80","order_by":2,"name":"John 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06:25:28","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":27000,"visible":true,"origin":"","legend":"\u003cp\u003eLength-weight relationship for \u003cem\u003eA. longa\u003c/em\u003e in the study site\u003c/p\u003e","description":"","filename":"2.png","url":"https://assets-eu.researchsquare.com/files/rs-5620138/v1/ea614b7341f04a8c9e014828.png"},{"id":72731063,"identity":"698e40ee-cfe2-4b99-8596-3e5abac9a298","added_by":"auto","created_at":"2025-01-01 06:25:28","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":24006,"visible":true,"origin":"","legend":"\u003cp\u003eLength-weight relationship for \u003cem\u003eL. terrestris\u003c/em\u003e in the study site\u003c/p\u003e","description":"","filename":"3.png","url":"https://assets-eu.researchsquare.com/files/rs-5620138/v1/c20f59dec545729f2ee6353b.png"},{"id":92430667,"identity":"b8ae3cd4-68f4-4f92-b113-48309c8bc3e8","added_by":"auto","created_at":"2025-09-29 16:07:30","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":493709,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-5620138/v1/03b05002-0415-4b98-b092-5eaddbf3b957.pdf"}],"financialInterests":"","formattedTitle":"Length-wright relationships of invasive exotic earthworms in the Huron Mountains forests, Michigan","fulltext":[{"header":"Introduction","content":"\u003cp\u003eInvasive species, particularly exotic earthworms, have emerged as a significant concern for North American forest ecosystems, including those in the Great Lakes region (Tiunov et al. \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2006\u003c/span\u003e; Mathieu et al. \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). Historically, native earthworms were largely absent from many areas of the northern United States due to glacial activity, and their reintroduction via human activities has had profound ecological effects (Hale et al. \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2005\u003c/span\u003e; Craven et al. \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Baumann et al. \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). The introduction of these earthworms, such as \u003cem\u003eDendrobaena octaedra\u003c/em\u003e, \u003cem\u003eAporrectodea longa\u003c/em\u003e, and \u003cem\u003eLumbricus terrestris\u003c/em\u003e, has altered soil structure and nutrient cycling processes (Lejoly et al. \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). These invasive species, which are not native to the region, disrupt soil structure, alter nutrient cycling processes, and reduce biodiversity in plant and animal communities (Singh et al. \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). These impacts are particularly pronounced in previously earthworm-free ecosystems, where the native flora and fauna are poorly adapted to the presence of these species (Phillips et al. \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Chang et al. \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2021\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eTo better understand the impacts of these species, it is essential and critical step to study their biological parameters including their body size, growth patterns and regeneration patterns (Chown and Gaston \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Zinger et al. \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). Among the biological parameters critical to understanding invasive earthworm impacts are their length-weight relationship (LWR) (Hale et al. \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2004\u003c/span\u003e). Examining the correlation between body length and weight can reveal insights into growth rates, resource allocation, and ecological adaptability. Such data are invaluable for predicting the spread and persistence of invasive populations, as well as for informing forest management strategies aimed at mitigating the consequences of earthworm invasions (Blouin et al. \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Baumann et al. \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2024\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThe current study, as a part of research to examine the influence of invasive earthworms on soil structure and function in forests of the Huron Mountains, Michigan, focused on assessing the LWR of invasive exotic earthworm species in the Huron Mountains forests. By developing species-specific LWR equations, this study contributes to the broader understanding of how invasive earthworms influence soil ecosystems and forest health in northern America.\u003c/p\u003e"},{"header":"Methods","content":"\u003cp\u003eThe study was conducted in the Huron Mountains Preserve, located in Michigan\u0026rsquo;s Upper Peninsula, USA (46\u0026deg;45\u0026prime;20\u0026Prime;N 88\u0026deg;09\u0026prime;21\u0026Prime;W). The average annual precipitation is about 900 mm, which is consistently spread throughout the year. Monthly temperatures vary from \u0026minus;\u0026thinsp;10.9\u0026deg;C in January to 18.3\u0026deg;C in July (NOAA 2009). The frost-free period lasts between 90 and 120 days. The bedrock in the area consists of Precambrian granitic, gneiss and sandstone formations (Woods \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2000\u003c/span\u003e). The soils are typically shallow, acidic, and derived from rocky glacial till, though some areas feature deeper till and alluvial deposits. This preserve encompasses a range of diverse forest ecosystems, including mixed hardwoods, coniferous stands, and riparian zones. The hardwood forests are primarily composed of tree species such as eastern hemlock (\u003cem\u003eTsuga canadensis\u003c/em\u003e), sugar maple (\u003cem\u003eAcer saccharum\u003c/em\u003e), yellow birch (\u003cem\u003eBetula alleghaniensis\u003c/em\u003e), red maple (\u003cem\u003eAcer rubrum\u003c/em\u003e), and basswood (\u003cem\u003eTilia americana\u003c/em\u003e). The study period spanned from May 2021 to October 2022, allowing for seasonal variability in earthworm activity to be captured.\u003c/p\u003e \u003cp\u003eThree 20m \u0026times; 20m plots were randomly established across main forest types in the preserve. At each plot, a 0.25 m\u003csup\u003e2\u003c/sup\u003e woody quadrat (0.5m \u0026times; 0.5m) was used for standardized sampling earthworm. Quadrats were spaced to minimize potential edge effects and ensure independence of sampling units. Earthworms were extracted using a mustard solution, which induces the earthworms to surface for collection (Lawrence and Bowers \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2002\u003c/span\u003e). All earthworms were identified to species level, and both their body length (in millimeters) and dry weight (in grams) were measured. The LWR was analyzed using regression models for each species individually. Statistical analysis included analysis of variance (ANOVA) to assess the significance of the regression models.\u003c/p\u003e"},{"header":"Results and Discussion","content":"\u003cp\u003eThree invasive exotic earthworm species were found in the study sites, each exhibiting significant differences in body size (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). Among these, \u003cem\u003eL. terrestris\u003c/em\u003e had the largest average body length (60.4\u0026thinsp;\u0026plusmn;\u0026thinsp;35.5 mm) and weight (0.254\u0026thinsp;\u0026plusmn;\u0026thinsp;0.344 g), followed by \u003cem\u003eA. longa\u003c/em\u003e, which showed intermediate values for both metrics (50.3\u0026thinsp;\u0026plusmn;\u0026thinsp;24.5 mm and 0.118\u0026thinsp;\u0026plusmn;\u0026thinsp;0.108 g, respectively). The smallest species was \u003cem\u003eD. octaedra\u003c/em\u003e, with a mean length of 25.5\u0026thinsp;\u0026plusmn;\u0026thinsp;9.3 mm and a weight of 0.012\u0026thinsp;\u0026plusmn;\u0026thinsp;0.009 g. The observed size differences have important ecological implications. The larger body size of \u003cem\u003eL. terrestris\u003c/em\u003e suggests a higher potential for soil disturbance and nutrient cycling, as larger earthworms can ingest and process greater amounts of soil and organic matter per individual (Blouin et al. \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). This aligns with studies that have reported enhanced soil aeration in the presence of large earthworm species (Bertrand et al. \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). Conversely, the smaller body size of \u003cem\u003eD. octaedra\u003c/em\u003e may reflect a different ecological role, such as focusing on leaf litter decomposition and fine-scale soil processes (Tiunov et al. \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2006\u003c/span\u003e). The intermediate size of \u003cem\u003eA. longa\u003c/em\u003e likely indicates a versatile ecological function, balancing traits of both large and small earthworms in terms of soil processing and nutrient redistribution (Coleman and Wall \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Wardle and Peltzer \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Ferlian et al. \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2022\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab1\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 1\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eVariation of body size of three invasive earthworm species in the study region*\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"4\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eBody size\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eL. terrestris\u003c/em\u003e\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eA. longa\u003c/em\u003e\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003e\u003cem\u003eD. Octaedra\u003c/em\u003e\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eMean length (mm)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e60.4\u0026thinsp;\u0026plusmn;\u0026thinsp;35.5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e50.3\u0026thinsp;\u0026plusmn;\u0026thinsp;24.5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e25.5\u0026thinsp;\u0026plusmn;\u0026thinsp;9.3\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eLength range (mm)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e24.5-119.5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e20.0\u0026ndash;90.0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e8.0-44.5\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eMean weight (g)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0.254\u0026thinsp;\u0026plusmn;\u0026thinsp;0.344\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.118\u0026thinsp;\u0026plusmn;\u0026thinsp;0.108\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0.012\u0026thinsp;\u0026plusmn;\u0026thinsp;0.009\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eWeight range (g)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0.017\u0026ndash;0.948\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.004\u0026ndash;0.291\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0.001\u0026ndash;0.042\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cp\u003e*mean\u0026thinsp;\u0026plusmn;\u0026thinsp;SD.\u003c/p\u003e \u003cp\u003eThe regression models for the LWR of the three species revealed significant associations between body length and weight (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e, \u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e, and \u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003e). Analysis of variance (ANOVA) confirmed the robustness of these models, with p-values less than 0.0001 for all species (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e, \u003cspan refid=\"Tab3\" class=\"InternalRef\"\u003e3\u003c/span\u003e, and \u003cspan refid=\"Tab4\" class=\"InternalRef\"\u003e4\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eFor \u003cem\u003eD. octaedra\u003c/em\u003e, the regression model explained 84.9% of the variation in body weight (F\u0026thinsp;=\u0026thinsp;292.54, p\u0026thinsp;\u0026lt;\u0026thinsp;0.00001), indicating a consistent relationship between length and weight across the population. Similarly, \u003cem\u003eA. longa\u003c/em\u003e exhibited a strong LWR with an R\u0026sup2; value of 93.4% (F\u0026thinsp;=\u0026thinsp;214.27, p\u0026thinsp;\u0026lt;\u0026thinsp;0.00001). \u003cem\u003eL. terrestris\u003c/em\u003e, while having the smallest sample size, also showed a significant relationship, with 90.5% of the variance in weight explained by length (F\u0026thinsp;=\u0026thinsp;85.53, p\u0026thinsp;\u0026lt;\u0026thinsp;0.00001).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab2\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 2\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eAnalysis of variance for regression model of mass-length for earthworm \u003cem\u003eD. octaedra\u003c/em\u003e\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"6\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eSource\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eDF\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSS\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eMS\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003eF-stat\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\"\u003e \u003cp\u003ep-value\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eModel\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e43.66091\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e43.66091\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e292.53858\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e\u0026lt;\u0026thinsp;0.00001\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eError\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e52\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e7.76092\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0.14925\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eTotal\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e53\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e51.42183\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab3\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 3\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eAnalysis of variance for regression model of mass-length for earthworm \u003cem\u003eA. longa\u003c/em\u003e\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"6\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eSource\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eDF\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSS\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eMS\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003eF-stat\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\"\u003e \u003cp\u003ep-value\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eModel\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e39.72614\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e39.72614\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e214.26663\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e\u0026lt;\u0026thinsp;0.00001\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eError\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e15\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e2.78108\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0.18541\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eTotal\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e16\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e42.50722\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab4\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 4\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eAnalysis of variance for regression model of mass-length for earthworm \u003cem\u003eL. terrestris\u003c/em\u003e\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"6\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eSource\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eDF\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSS\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eMS\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003eF-stat\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\"\u003e \u003cp\u003ep-value\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eModel\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e26.13383\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e26.13383\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e85.52588\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e\u0026lt;\u0026thinsp;0.00001\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eError\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e9\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e2.75010\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0.30557\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eTotal\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e10\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e28.88393\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cp\u003eThe differences in LWR patterns among the species likely reflect variations in energy allocation strategies (Rinke et al. \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; Ehrenfeld \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2011\u003c/span\u003e). For instance, \u003cem\u003eL. terrestris\u003c/em\u003e may prioritize growth and energy storage, enabling it to dominate environments with high organic matter availability. In contrast, \u003cem\u003eD. octaedra\u003c/em\u003e may allocate resources toward reproduction or survival in less favorable conditions, consistent with its smaller size and broader environmental tolerance (Ehrenfeld \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2011\u003c/span\u003e). \u003cem\u003eA. longa\u003c/em\u003e appears to occupy an intermediate niche, suggesting adaptability to a range of soil types and conditions (Chown and Gaston \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Rakel et al. \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Potapov \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2022\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThe presence and functional traits of invasive earthworms can significantly impact soil ecosystems. Large-bodied species like \u003cem\u003eL. terrestris\u003c/em\u003e are known to accelerate nutrient cycling, potentially altering plant community composition and forest floor dynamics. This has been documented in other studies, where invasive earthworms have facilitated shifts in nutrient availability and microbial communities (Ferlian et al. \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). On the other hand, smaller species like \u003cem\u003eD. octaedra\u003c/em\u003e may contribute more subtly by focusing on fine-scale decomposition processes and enhancing soil microstructure (Zinger et al. \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2017\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eEnvironmental factors such as soil type, moisture, and temperature are likely to influence these growth patterns and functional roles. For example, differences in soil organic matter content and moisture availability across the Huron Mountains could explain the observed variability in body size and LWR. Similar findings have been reported by other previous studies who noted that earthworm activity and distribution are highly dependent on microhabitat conditions (Hale et al. \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2005\u003c/span\u003e; Gutierrez-Lopez et al. 2010; Ahmed and Al-Mutairi \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2022\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eFurther studies should investigate the interaction between invasive earthworm species and forest ecosystem dynamics, particularly focusing on their combined effects on nutrient cycling, microbial communities, and plant succession. Understanding how environmental gradients influence earthworm traits and their ecological roles could provide insights into managing invasive species and mitigating their impacts on native ecosystems.\u003c/p\u003e"},{"header":"Conclusion","content":"\u003cp\u003eThis study highlights the body size variation, LWR and ecological roles of three invasive earthworm species, \u003cem\u003eL. terrestris\u003c/em\u003e, \u003cem\u003eA. longa\u003c/em\u003e and \u003cem\u003eD. octaedra\u003c/em\u003e in the Huron Mountains, revealing distinct ecological functions and impacts on soil ecosystems. \u003cem\u003eL. terrestris\u003c/em\u003e, the largest species, demonstrated significant potential for soil disturbance and nutrient cycling, while \u003cem\u003eD. octaedra\u003c/em\u003e, the smallest, contributed to fine-scale decomposition processes and soil microstructure. The intermediate-sized \u003cem\u003eA. longa\u003c/em\u003e exhibited a versatile role, balancing traits of the other species. Robust LWR models (R\u0026sup2; \u0026gt; 84%) reflect species-specific energy allocation strategies influenced by environmental factors such as soil type, moisture and temperature. These findings underscore the ecological implications of invasive earthworms, including their effects on nutrient cycling, microbial activity, and plant community composition. Future research should examine their combined impacts on forest ecosystems and explore how environmental gradients shape their traits and functions to inform management strategies.\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003e\u003cstrong\u003eAcknowledgements\u003c/strong\u003e\u0026nbsp; \u0026nbsp;We are grateful to Dr. Kerry Woods and Mr. Brock Francis for their assistance with the field trip and site selection at the Huron Mountains. We would also like to thank Paula Arroyo, Madeleine Naliwko, Quincy Santomieri, Lynda Randa, Dr. Yuanying Peng and Andren Yunger for their assistance with field and laboratory measurements.\u003c/p\u003e\n\n\u003cp\u003e\u003cstrong\u003eAuthors\u0026rsquo; contribution\u003c/strong\u003e\u0026nbsp; \u0026nbsp;Conceptualization: Xiaoyong Chen, Mary Carrington; Methodology: Xiaoyong Chen, Timothy Gsell, John Yunger, Mary Carrington; Investigation Xiaoyong Chen, Timothy Gsell, John Yunger; Formal analysis: Xiaoyong Chen, Heng Li; Writing - original draft preparation: Xiaoyong Chen, Timothy Gsell, John Yunger; Funding acquisition: Xiaoyong Chen, Mary Carrington. All authors read, edited, and approved the final manuscript.\u003c/p\u003e\n\n\u003cp\u003e\u003cstrong\u003eFunding\u003c/strong\u003e\u0026nbsp; \u0026nbsp;This work was supported by Huron Mountain Wildlife Foundation Grant (2021-2023).\u003c/p\u003e\n\n\u003cp\u003e\u003cstrong\u003eData availability\u003c/strong\u003e\u0026nbsp; \u0026nbsp;The data will be provided upon request on corresponding author.\u003c/p\u003e\n\n\n\n\u003cp\u003e\u003cstrong\u003eConflict of interest\u003c/strong\u003e\u0026nbsp; The authors have no relevant financial or non-financial interests to disclose.\u003c/p\u003e\n\n\u003cp\u003e\u003cstrong\u003eEthical approval\u003c/strong\u003e\u0026nbsp; \u0026nbsp;Not applicable.\u003c/p\u003e\n"},{"header":"References","content":"\u003col\u003e\n\u003cli\u003eAhmed N, Al-Mutairi KA (2022) Earthworms effect on microbial population and soil fertility as well as their interaction with agriculture practices. Sustainability. 14:7803\u003c/li\u003e\n\u003cli\u003eBaumann TT, Frelich LE, Van Riper LC, Yoo K (2024) Anthropogenic transport mechanisms of invasive European earthworms: a review. Biol Invasions 26:3563-3586\u003c/li\u003e\n\u003cli\u003eBertrand M, Barot S, Blouin M, Whalen J, de Oliveira T, Roger-Estrade J (2015) Earthworm services for cropping systems. A review. Agron Sustain Dev 35:553-567\u003c/li\u003e\n\u003cli\u003eBlouin M, Hodson ME, Delgado EA, Baker G, Brussaard L, Butt KR, Dai J, Dendooven L, Peres G, Tondoh JE, Cluzeau D, Brun JJ (2013) A review of earthworm impact on soil function and ecosystem services: earthworm impact on ecosystem services. Eur J Soil Sci 64:161-182\u003c/li\u003e\n\u003cli\u003eChang CH, Bartz MLC, Brown G et al (2021) The second wave of earthworm invasions in North America: biology, environmental impacts, management and control of invasive jumping worms. 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Soil Org 91:114-138\u003c/li\u003e\n\u003cli\u003eTiunov AV, Hale CM, Holdsworth, AR, Vsevolodova-Perel, TS (2006) Invasion patterns of \u003cem\u003eLumbricidae\u003c/em\u003e into the previously earthworm-free areas of northeastern Europe and the western Great Lakes region of North America. Biol Invasions 8:1223-1234\u003c/li\u003e\n\u003cli\u003eWardle DA, Peltzer DA (2017) Impacts of invasive biota in forest ecosystems in an aboveground-belowground context. Biol Invasions 19:3301-3316\u003c/li\u003e\n\u003cli\u003eWoods KD (2000) Dynamics in late-successional hemlock-hardwood forests over three decades. Ecology 81:110-126\u003c/li\u003e\n\u003cli\u003eZinger L, Taberlet P, Schimann H et al (2017) Soil community assembly varies across body sizes in a tropical forest. BioRxiv 2017:154278\u003c/li\u003e\n\n\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":false,"highlight":"","institution":"","isAcceptedByJournal":true,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"biological-invasions","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"binv","sideBox":"Learn more about [Biological Invasions](https://www.springer.com/journal/10530)","snPcode":"10530","submissionUrl":"https://submission.nature.com/new-submission/10530/3","title":"Biological Invasions","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false},"keywords":"Earthworms, Invasion, Body size, North American Forests, Length-Weight Relation","lastPublishedDoi":"10.21203/rs.3.rs-5620138/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-5620138/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eInvasive exotic earthworms have been identified as a major ecological threat to forest ecosystems in the Great Lakes region, affecting soil structure, nutrient cycling, and biodiversity. Understanding biological parameters such as body length, weight, growth rate, and regeneration patterns of these earthworms is crucial first step in comprehending their life cycle and their impacts on ecological processes in terrestrial ecosystems. In this study, an earthworm survey was conducted in the Huron Mountains of Michigan's Upper Peninsula. The earthworms were captured from sampling quadrats, identified to species, and their body mass (dry weight) and body length were measured. The purpose of the study was to develop the length-weight relationship (LWR) for the invasive exotic earthworm species that appeared in this area. Results showed that three invasive earthworm species were found in the studied forests, and they were: \u003cem\u003eDendrobaena octaedra\u003c/em\u003e, \u003cem\u003eAporrectodea longa\u003c/em\u003e, and \u003cem\u003eLumbricus terrestris\u003c/em\u003e. Among them, \u003cem\u003eL. terrestris\u003c/em\u003e exhibited the largest size (mean length: 60.4 mm; mean weight: 0.254 g), while \u003cem\u003eD. octaedra\u003c/em\u003e was the smallest (mean length: 25.5 mm; mean weight: 0.012 g). The LWR regression equations were: W\u0026thinsp;=\u0026thinsp;0.00001L\u003csup\u003e2.135\u003c/sup\u003e for \u003cem\u003eD. octaedra\u003c/em\u003e; W\u0026thinsp;=\u0026thinsp;0.000001L\u003csup\u003e2.816\u003c/sup\u003e for \u003cem\u003eA. longa\u003c/em\u003e; and W\u0026thinsp;=\u0026thinsp;0.000003L\u003csup\u003e2.618\u003c/sup\u003e for \u003cem\u003eL. terrestris\u003c/em\u003e. These findings highlighted differences in growth patterns that may be influenced by forest composition and micro-environmental factors. This study provided valuable insights into the biological characteristics of invasive exotic earthworms and their potential impacts on North American forest ecosystems.\u003c/p\u003e","manuscriptTitle":"Length-wright relationships of invasive exotic earthworms in the Huron Mountains forests, Michigan","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2025-01-01 06:25:24","doi":"10.21203/rs.3.rs-5620138/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"reviewerAgreed","content":"","date":"2025-01-31T20:56:15+00:00","index":0,"fulltext":""},{"type":"reviewersInvited","content":"","date":"2024-12-30T16:07:34+00:00","index":"","fulltext":""},{"type":"editorInvited","content":"Biological Invasions","date":"2024-12-13T15:24:45+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2024-12-12T03:36:02+00:00","index":"","fulltext":""},{"type":"submitted","content":"Biological Invasions","date":"2024-12-10T21:01:12+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"biological-invasions","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"binv","sideBox":"Learn more about [Biological Invasions](https://www.springer.com/journal/10530)","snPcode":"10530","submissionUrl":"https://submission.nature.com/new-submission/10530/3","title":"Biological Invasions","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false}}],"origin":"","ownerIdentity":"53c9bf58-5aa1-4d7a-8d91-69787b0b5cce","owner":[],"postedDate":"January 1st, 2025","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"published-in-journal","subjectAreas":[],"tags":[],"updatedAt":"2025-09-29T16:04:54+00:00","versionOfRecord":{"articleIdentity":"rs-5620138","link":"https://doi.org/10.1007/s10530-025-03680-8","journal":{"identity":"biological-invasions","isVorOnly":false,"title":"Biological Invasions"},"publishedOn":"2025-09-26 15:57:22","publishedOnDateReadable":"September 26th, 2025"},"versionCreatedAt":"2025-01-01 06:25:24","video":"","vorDoi":"10.1007/s10530-025-03680-8","vorDoiUrl":"https://doi.org/10.1007/s10530-025-03680-8","workflowStages":[]},"version":"v1","identity":"rs-5620138","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-5620138","identity":"rs-5620138","version":["v1"]},"buildId":"8U1c8b4HqxoKbykW_rLl7","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}