Range expansion of the invasive hybrid cattail Typha x glauca exceeds that of its maternal plant T. angustifolia in the western Prairie Pothole Region of North America

Range expansion of the invasive hybrid cattail Typha x glauca exceeds that of its maternal plant T. angustifolia in the western Prairie Pothole Region of North America | 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 Range expansion of the invasive hybrid cattail Typha x glauca exceeds that of its maternal plant T. angustifolia in the western Prairie Pothole Region of North America Sanjuti Deb Joyee, Marcel Dorken, Joanna Freeland This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-4607793/v1 This work is licensed under a CC BY 4.0 License Status: Published Journal Publication published 12 Sep, 2024 Read the published version in Biological Invasions → Version 1 posted 5 You are reading this latest preprint version Abstract Wetlands around the world face unprecedented threats, including from invasive species. In North America, the invasive cattail hybrid Typha x glauca dominates wetlands around the Laurentian Great Lakes, and more recently was found in high abundance across the central and eastern Prairie Pothole Region, an area that includes one of the world’s largest wetland complexes. Surveys of the Typha hybrid zone have so far been largely conducted in areas where hybrids are well established, and it therefore remains unclear whether the range expansion of this invasive hybrid occurs after the establishment of its maternal species, T. angustifolia. We surveyed 50 wetlands in the western PPR, and found that while 75% of plants were native T. latifolia , the second most common group was F 1 hybrids: these had greater abundance and occupancy than T. angustifolia despite the fact that T. angustifolia produces relatively few hybrid seeds, and therefore highlights the importance of long-distance dispersal for this hybrid range expansion. The distribution of hybrids combined with the paucity of non-F1 hybrids suggests that the western PPR represents a leading edge of the range expansion by invasive T. x glauca. Our results show that T. x glauca has the capacity for continued range expansion that does not rely on the presence of T. angustifolia , and the impacts of this range expansion should be monitored because of its potential to impede ecosystem services and reduce local biodiversity. Typha Prairie Pothole Region hybrid range expansion wetlands Figures Figure 1 Introduction Wetlands provide essential ecosystem services and accommodate high levels of biodiversity across an estimated 5-10% of the Earth’s surface; however, despite their importance, >70% of natural wetlands have been lost globally since 1900 (Davidson 2014; Kingsford et al. 2016). Threats to wetlands include development, climate change, pollution, and invasive species (Zedler and Kercher 2004; Xu et al. 2019). In wetlands within the region surrounding the Laurentian Great Lakes and St. Lawrence Seaway in North America, one problematic invader is the cattail hybrid complex that comprises Typha x glauca, which is the interspecific F 1 hybrid of native T. latifolia and naturalized T. angustifolia (Grace and Harrison 1986), plus backcrossed and advanced-generation hybrids. In these regions, the hybrid displaces parental species and dominates wetlands (Travis et al. 2010; Freeland et al. 2013; Pieper et al. 2020), and is considered invasive because it reduces biodiversity and alters wetland functions (reviewed in (Bansal et al. 2019). The invasion of wetlands around the Laurentian Great Lakes has led to multiple management programs aimed at controlling or eliminating T. x glauca (e.g. Lawrence et al. 2016; Graham et al. 2022), and has also raised concerns about range expansion by this invasive hybrid, including into the Prairie Pothole Region (PPR) of North America. The PPR occurs within the Great Plains of central North America, and includes one of the world’s largest wetland complexes. Here there are an estimated 5–8 million wetlands commonly referred to as potholes, most of which are depressions < 1 ha in area, < 1 m in depth, and commonly within farmland (Doherty et al. 2016). Wetlands in the PPR provide essential ecosystem services, and harbour substantial levels of biodiversity, for example they enable millions of waterfowl to migrate through, and breed within, the PPR each year (Doherty et al. 2018; Pattison-Williams et al. 2018). Monitoring invasive wetland plants in the PPR is an essential to our understanding of whether these wetlands are at risk from substantial modification following biological invasions. A recent study that surveyed 131 wetlands spread over approximately 350,000 km 2 in the central and eastern PPR found hybrid cattails in over 80% of surveyed wetlands; in contrast, T. latifolia was found in 26% of wetlands, and T. angustifolia in 18% of wetlands (Tangen et al. 2022). Typha x glauca is fertile, and can interbreed to form advanced-generation hybrids, or backcross with either of its parent species to form backcrossed hybrids (Pieper 2017); because advanced-generation and backcrossed hybrids cannot be differentiated, we refer to these collectively as non-F1 hybrids. Non-F1 hybrids comprised the most common taxonomic category in the central/eastern PPR (57% of genets compared to 30% that were F1 hybrids), which suggests that the hybrid zone in the central/eastern PPR is not very recent. While the Tangen et al. study (2022) provided important information about the abundance and occupancy of hybrid Typha throughout wetlands in the central and southeast PPR, some knowledge gaps remain. In this study we expanded the investigation of the Typha hybrid zone into previously unsampled regions of the west/northwest PPR, beyond the range limit of T. angustifolia . F1 T. x glauca hybrids arise following asymmetric hybridization, in which T. angustifolia is normally the maternal parent and T. latifolia is the paternal parent (Pieper 2017). There are almost no records of T. angustifolia from the western PPR, which could preclude the local formation of hybrids, and we therefore addressed the following questions: 1) is the paucity of T. angustifolia records from the western PPR an accurate reflection of T. angustifolia occurrence in this region? 2) If T. angustifolia plants occur in the western PPR, are they producing seeds that are T. x glauca ? 3) Does the range expansion of T. x glauca follow the range expansion of its maternal species, T. angustifolia ? Invasive plant species pose a major threat to biological diversity in some regions of the PPR (Jones et al. 2023), and this study provides additional information on a potentially problematic range expansion of an invasive wetland hybrid across the PPR. Methods Field work Between August 19 and August 24, 2022, we sampled 244 Typha plants from 50 wetlands (34 wetlands in the province of Saskatchewan, 16 wetlands in the province of Alberta) in the western PPR (Figure 1, Table S1). Sampled sites were mostly restricted to small wetlands accessible from roadsides because a substantial portion of the Prairie Pothole Region is privately owned agricultural land. This sampling strategy should not have influenced our findings because an earlier study found that Typha species and hybrids had comparable frequencies in wetlands versus ditches in eastern North America (Pieper et al., 2020). Typha latifolia is native to Saskatchewan and Alberta (Flora of North America 2008) and was thus expected to occur at the majority of sites; we therefore prioritized the sampling of T. angustifolia followed by T. x glauca . We did this by first visually inspecting the stands at each site and identifying plants that appeared to have the narrowest leaves and – for flowering plants - the largest gaps between the staminate and pistillate flowers: T. angustifolia has the narrowest leaves and largest flower gaps, T. latifolia has the widest leaves and typically no discernible flower gap, and F1 hybrids are intermediate in both traits (Smith 1967). Up to eight plants with what appeared to be the narrowest leaves were sampled at each site. From each sampled plant we collected two leaf fragments approximately 10 cm long. Leaf samples were placed into labelled coin envelopes, and the envelopes were placed in a plastic zippered bag containing Sor bead orange silica beads for desiccation. In total, we collected leaf samples from 245 plants across 50 sites. In addition, we collected mature fruits from nine plants (not all plants flower in any given year) that were provisionally identified as T. angustifolia and stored these in paper bags that were enclosed within resealable plastic bags. Genotyping Because the leaf widths and flower gaps of advanced-generation (AGH) and backcrossed hybrids (BCH) can overlap with parental taxa (Snow et al., 2010; Kirk et al., 2011; Geddes et al., 2021; Tangen et al., 2022), we used genetic information to identify the taxonomy of each sampled plant. In brief, we followed the methods described in (Chambers et al. 2024) to extract DNA and genotype each plant at up to five loci with species-specific alleles (one microsatellite locus and four PCR-RFLP loci). Plants with exclusively Typha latifolia alleles (identified as T. latifolia), T. angustifolia alleles (identified as T. angustifolia), or heterozygous loci (one T. latifolia and one T. angustifolia allele at each locus; identified as F 1 hybrids) were genotyped at all five loci. Plants that were heterozygous for Typha latifolia and Typha angustifolia alleles at some but not at all loci (genotyped at between two and five loci) were identified as advanced generation or backcrossed hybrids and pooled into a single category of non-F1 hybrids. Seed germination and identification The nine putative T. angustifolia plants from which we collected seeds were taxonomically confirmed from genetic data. To investigate whether these plants were generating exclusively F1 hybrids, we germinated their seeds and genotyped a subset of their offspring. The seeds were separated from the inflorescence stalk, and from each maternal plant approximately 0.5g of fruit was prepared for germination following the protocol outlined in (Ahee et al. 2015). The seeds were left to germinate in petri dishes filled with DI water in a climate-controlled (15-20 °C) greenhouse at Trent University. After six days the seedlings were transferred into 10 cm pots filled with well-draining soil from Sunshine Professional growing mixture (germination grade; Sun Gro Horticulture, Brantford, Canada) and left in the greenhouse on plastic trays filled with water. Seedlings were fertilized after 45 days using 100 mL of 0.5 % water-soluble 20:20:20 N:P: K general-purpose fertilizer (Plant-Prod, Leamington, Ontario). Ten seedlings from each maternal plant were harvested after approximately 80-90 days, by which time they had reached heights of approximately 8 cm. Harvested seedlings were put into coin envelopes in resealable plastic bags containing orange silica beads (Sor bead, USA) for desiccation. DNA extraction and genotyping followed the same methods described for the plant samples collected in the field. Results Distribution of taxa Overall, the majority of the samples that we genotyped were T. latifolia , followed by F1 T. x glauca, T. angustifolia , and then non-F1 hybrids (Table 1 ). Hybrids were more abundant and identified at more sites than the maternal taxon T. angustifolia (Fig. 1) despite a sampling strategy that prioritized T. angustifolia . T. angustifolia offspring The 90 genotyped offspring of nine T. angustifolia plants comprised 78 (87%) T. angustifolia , 10 (11%) F1 hybrids, and 2 (2%) non-F1 hybrids (advanced-generation or backcrossed hybrids). Seed samples from five of the nine T. angustifolia plants included no hybrid offspring. The remaining four plants had 1/10, 1/10, 2/10, and 8/10 hybrids (either F1 or back-crossed hybrids) among their offspring. Discussion The data collected in this study allow us to address the three questions presented in our introduction. First, the paucity of T. angustifolia records from the western PPR does indeed seem to be a fairly accurate reflection of its occurrence in this region: despite a sampling strategy that prioritized T. angustifolia , we found this species at only four of the 50 sampled sites. At least one record suggests that T. angustifolia has been present in the PPR since the late 1970s (Harms and Ledingham 1986 ). This record was not verified with genetic data but, if accurate, suggests that despite being in the western PPR for several decades T. angustifolia remains rare and thus may not be adapted to conditions in this region. Our second question asked whether T. angustifolia in the western PPR are producing hybrid seeds. Although our sample size was small (reflecting the small number of T. angustifolia in this region – and the even smaller number of flowering T. angustifolia ), we found that T. angustifolia mostly produce T. angustifolia seedlings, with only 13% of our seedling sample comprising hybrids. This was an unexpected finding given the predominance of T. latifolia in the area (75% of plants despite a sampling strategy that favoured other taxa), which in turn should increase the likelihood of T. angustifolia plants being pollinated by heterospecifics and producing mostly hybrids. Previous studies have shown that seedling production by T. angustifolia is comparable when pollinated by T. angustifolia, T. latifolia , or T. x glauca (Pieper et al. 2017 ), and therefore the predominance of T. angustifolia seedlings may be at least partly explained by other forms of reproductive barriers such as different flowering times. Although we did not find T. x glauca in any of the Alberta sites, it was found in 10/34 Saskatchewan wetlands, compared to T. angustifolia in only 3/34 Saskatchewan wetlands, suggesting a westwards expansion. Additionally, unlike in the central/eastern PPR, we found very few non-F1 hybrids, suggesting that hybrids have only recently colonized this area. The western PPR therefore appears to be the leading edge of the hybrid range expansion. Our third question asked whether the range expansion of T. x glauca into the western PPR is following that of its maternal species T. angustifolia , and our data suggest this is not the case: because the hybrid occurred at more sites, and across a larger area, than T. angustifolia , it does not appear reliant on its maternal species for successful establishment at new sites. Although we found that T. angustifolia produce only a small proportion hybrid seeds, these have the potential for dispersal across short and long distances via wind, water, or animal vectors including waterfowl (Soons 2006 ; Soons et al. 2008 ), and thus can colonize sites from which the maternal species is absent. Additionally, backcrossed hybrids, advanced-generation hybrids, and vegetative propagation can perpetuate hybrid Typha in the absence of its maternal species. In eastern Canada and the central/eastern PPR T. x glauca has outcompeted both parental species (Freeland et al. 2013 ; Pieper et al. 2020 ; Tangen et al. 2022 ), and may be able to dominate wetlands even with relatively low propagule pressure. Our results collectively suggest that T. x glauca has the capacity for continued range expansion that is not reliant on the presence of T. angustifolia , and the impacts of this range expansion should be monitored because of the potential for this wetland plant to impede ecosystem services and reduce local biodiversity. Table 1 Typha taxa (number and percentage) identified from 50 wetlands in the western Prairie Pothole Region. SK = Saskatchewan, AB = Alberta. Taxon Number (%) of sampled plants Number (%) of sampled SK wetlands (out of a total of 34) Number (%) of sampled AB wetlands (out of a total of 16) Total number (%) of wetlands Typha latifolia 183 (75%) 24 (71%) 16 (100%) 40 (80%) F1 Typha x glauca 40 (16%) 10 (29%) 0 (0%) 10 (20%) Typha angustifolia 17 (7%) 3 (9%) 1 (6%) 4 (8%) Backcrossed/advanced-generation hybrids 5 (2%) 3 (9%) 0 (0%) 3 (6%) Declarations The following statements must be included in your submitted manuscript under the heading 'Statements and Declarations'. This should be placed after the References section. Please note that submissions that do not include required statements will be returned as incomplete. Funding Funding for this project was provided by Discovery Grants from the Natural Sciences and Engineering Research Council of Canada awarded to J. Freeland (RGPIN-2017-04371) and M. Dorken (RGPIN-2018-04866). Competing Interests The authors have no relevant financial or non-financial interests to disclose. Author Contributions This research was part of Sanjuti Deb Joyee’s MSc thesis, co-supervised by Joanna Freeland and Marcel Dorken. Study conception and design by Joanna Freeland and Marcel Dorken. Field work (sample collection) was performed by Joanna Freeland, laboratory and greenhouse methods by Sanjuti Deb Joyee. The first draft of the manuscript was written by Joanna Freeland and all authors commented on previous versions of the manuscript. All authors read and approved the final manuscript. References Ahee JE, Van Drunen WE, Dorken ME (2015) Analysis of pollination neighbourhood size using spatial analysis of pollen and seed production in broadleaf cattail ( Typha latifolia ). Botany 93:91–100. https://doi.org/10.1139/cjb-2014-0169 Bansal S, Lishawa SC, Newman S, et al (2019) Typha (Cattail) Invasion in North American Wetlands: Biology, Regional Problems, Impacts, Ecosystem Services, and Management. Wetlands 39:645–684 Chambers A, Chambers B, Bhargava D, et al (2024) A simple method to genetically differentiate invasive F1 Typha hybrids (T. × glauca) and advanced-generation/backcrossed hybrids from parent species (T. Latifolia and T. angustifolia) in eastern Canada and northeastern USA. J Great Lakes Res 50:. https://doi.org/10.1016/j.jglr.2023.102257 Davidson NC (2014) How much wetland has the world lost? Long-term and recent trends in global wetland area. 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Aquat Conserv Mar Freshw Ecosyst 26:892–916. https://doi.org/10.1002/aqc.2709 Lawrence BA, Lishawa SC, Rodriguez Y, Tuchman NC (2016) Herbicide management of invasive cattail (Typha × glauca) increases porewater nutrient concentrations. Wetl Ecol Manag 24:. https://doi.org/10.1007/s11273-015-9471-x Pattison-Williams JK, Pomeroy JW, Badiou P, Gabor S (2018) Wetlands, Flood Control and Ecosystem Services in the Smith Creek Drainage Basin: A Case Study in Saskatchewan, Canada. Ecol Econ 147:. https://doi.org/10.1016/j.ecolecon.2017.12.026 Pieper S, Dorken M, Freeland J (2020) Genetic structure in hybrids and progenitors provides insight into processes underlying an invasive cattail (Typha × glauca) hybrid zone. Heredity (Edinb) 124:714–727. https://doi.org/10.1038/s41437-020-0307-y Pieper SJ, Nicholls AA, Freeland JR, Dorken ME (2017) Asymmetric Hybridization in Cattails (Typha spp.) and Its Implications for the Evolutionary Maintenance of Native Typha latifolia. 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CRC Crit Rev Plant Sci 23:431–452. https://doi.org/10.1080/07352680490514673 Supplementary Files TableS1.docx Cite Share Download PDF Status: Published Journal Publication published 12 Sep, 2024 Read the published version in Biological Invasions → Version 1 posted Reviewers agreed at journal 26 Jun, 2024 Reviewers invited by journal 25 Jun, 2024 Editor invited by journal 21 Jun, 2024 Editor assigned by journal 20 Jun, 2024 First submitted to journal 19 Jun, 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. 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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-4607793","acceptedTermsAndConditions":true,"allowDirectSubmit":false,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":318962331,"identity":"5997730a-b7ea-4b50-9ffa-07b105fabc79","order_by":0,"name":"Sanjuti Deb Joyee","email":"","orcid":"","institution":"Trent University - Peterborough Campus: Trent University","correspondingAuthor":false,"prefix":"","firstName":"Sanjuti","middleName":"Deb","lastName":"Joyee","suffix":""},{"id":318962332,"identity":"7068bc8f-a508-4f79-8c40-7e6a8f4affa9","order_by":1,"name":"Marcel Dorken","email":"","orcid":"","institution":"Trent University - Peterborough Campus: Trent University","correspondingAuthor":false,"prefix":"","firstName":"Marcel","middleName":"","lastName":"Dorken","suffix":""},{"id":318962333,"identity":"1cbc71ce-da43-4580-9eeb-77091526036b","order_by":2,"name":"Joanna Freeland","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAABAUlEQVRIiWNgGAWjYBACxgYwJQfEzEAmmw3RWoyhTLY0CYQgfgDXcpiwFub25mePC2oM8vnZGxs/F5SdrzNv7z3+mIfBTh6nw3qOmRvPOGZgObPnYLP0jHO3JWTOnEts5mFINsRlFeOMBDNpHrY/BgY3EhukedtuS0hI5BgCtRzA6TrGGenfpHn+GRjY33/Y/Ju37ZyEhPwbsBZ73FpyzICGGxgYSDC2ARkHgLbwgLUk4tTSc6ZMmrcPqONMYps1z7lkyRk8OYYz5xgkJ+PSYtjevk2a55uBAX/74cO3ecrs+CXYzxh8eFNhZ4tTCw4JAxzqgQBn6I+CUTAKRsEogAMAj4xQWfBYjCEAAAAASUVORK5CYII=","orcid":"https://orcid.org/0000-0002-5251-7680","institution":"Trent University - Peterborough Campus: Trent University","correspondingAuthor":true,"prefix":"","firstName":"Joanna","middleName":"","lastName":"Freeland","suffix":""}],"badges":[],"createdAt":"2024-06-19 19:52:20","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-4607793/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-4607793/v1","draftVersion":[],"editorialEvents":[{"content":"https://doi.org/10.1007/s10530-024-03439-7","type":"published","date":"2024-09-12T15:58:07+00:00"}],"editorialNote":"","failedWorkflow":false,"files":[{"id":60489979,"identity":"49c25771-1c53-4f76-9cce-0daa096dcd21","added_by":"auto","created_at":"2024-07-17 10:26:41","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":413390,"visible":true,"origin":"","legend":"\u003cp\u003eDistribution of \u003cem\u003eTypha\u003c/em\u003e taxa across the 50 sampled wetlands. At each site a pie chart illustrates the percentage of each taxon. Black dots within the pie chart represent the site location, and leader lines were used to connect pie charts to site locations when nearby wetlands were also sampled.\u003c/p\u003e","description":"","filename":"Figure1Map.png","url":"https://assets-eu.researchsquare.com/files/rs-4607793/v1/014fe8b4788e76265cb26934.png"},{"id":64619263,"identity":"c62ab6d6-3708-4f91-9bbe-21432300b134","added_by":"auto","created_at":"2024-09-16 16:13:35","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":724215,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-4607793/v1/f604eb40-c14d-488b-a75e-6616b2a7e16e.pdf"},{"id":60490447,"identity":"9a941980-c72e-4af7-9abd-9ffab61d1072","added_by":"auto","created_at":"2024-07-17 10:34:41","extension":"docx","order_by":1,"title":"","display":"","copyAsset":false,"role":"supplement","size":20259,"visible":true,"origin":"","legend":"","description":"","filename":"TableS1.docx","url":"https://assets-eu.researchsquare.com/files/rs-4607793/v1/9f13de12163382fa75504cdc.docx"}],"financialInterests":"","formattedTitle":"Range expansion of the invasive hybrid cattail Typha x glauca exceeds that of its maternal plant T. angustifolia in the western Prairie Pothole Region of North America","fulltext":[{"header":"Introduction","content":"\u003cp\u003eWetlands provide essential ecosystem services and accommodate high levels of biodiversity across an estimated 5-10% of the Earth\u0026rsquo;s surface; however, despite their importance, \u0026gt;70% of natural wetlands have been lost globally since 1900\u0026nbsp;(Davidson 2014; Kingsford et al. 2016). \u0026nbsp;Threats to wetlands include development, climate change, pollution, and invasive species\u0026nbsp;(Zedler and Kercher 2004; Xu et al. 2019). \u0026nbsp;In wetlands within the region surrounding the Laurentian Great Lakes and St. Lawrence Seaway in North America, one problematic invader is the cattail hybrid complex that comprises \u003cem\u003eTypha\u0026nbsp;\u003c/em\u003ex \u003cem\u003eglauca,\u0026nbsp;\u003c/em\u003ewhich is the interspecific F\u003csub\u003e1\u003c/sub\u003e hybrid of native \u003cem\u003eT. latifolia\u0026nbsp;\u003c/em\u003eand naturalized \u003cem\u003eT. angustifolia\u0026nbsp;\u003c/em\u003e(Grace and Harrison 1986), plus backcrossed and advanced-generation hybrids. \u0026nbsp;In these regions,\u0026nbsp;the hybrid displaces parental species and dominates wetlands\u0026nbsp;(Travis et al. 2010; Freeland et al. 2013; Pieper et al. 2020), and is considered invasive because it reduces biodiversity and alters wetland functions (reviewed in\u0026nbsp;(Bansal et al. 2019). \u0026nbsp;\u003c/p\u003e\n\u003cp\u003eThe invasion of wetlands around the Laurentian Great Lakes has led to multiple management programs aimed at controlling or eliminating \u003cem\u003eT.\u0026nbsp;\u003c/em\u003ex \u003cem\u003eglauca\u0026nbsp;\u003c/em\u003e(e.g. Lawrence et al. 2016; Graham et al. 2022), and has also raised concerns about range expansion by this invasive hybrid, including into the Prairie Pothole Region (PPR) of North America. \u0026nbsp;The PPR occurs within the Great Plains of central North America, and includes one of the world\u0026rsquo;s largest wetland complexes. \u0026nbsp;Here there are an estimated 5\u0026ndash;8 million wetlands commonly referred to as potholes, most of which are depressions \u0026lt; 1 ha in area, \u0026lt; 1 m in depth, and commonly within farmland\u0026nbsp;(Doherty et al. 2016). \u0026nbsp;Wetlands in the PPR provide essential ecosystem services, and harbour substantial levels of biodiversity, for example they enable millions of waterfowl to migrate through, and breed within, the PPR each year\u0026nbsp;(Doherty et al. 2018; Pattison-Williams et al. 2018). \u0026nbsp;Monitoring invasive wetland plants in the PPR is an essential to our understanding of whether these wetlands are at risk from substantial modification following biological invasions. \u0026nbsp;\u003c/p\u003e\n\u003cp\u003eA recent study that surveyed 131 wetlands spread over approximately 350,000 km\u003csup\u003e2\u003c/sup\u003e in the central and eastern PPR found hybrid cattails in over 80% of surveyed wetlands; in contrast, \u003cem\u003eT. latifolia\u0026nbsp;\u003c/em\u003ewas found in 26% of wetlands, and \u003cem\u003eT. angustifolia\u0026nbsp;\u003c/em\u003ein 18% of wetlands\u0026nbsp;(Tangen et al. 2022). \u0026nbsp;\u003cem\u003eTypha\u0026nbsp;\u003c/em\u003ex \u003cem\u003eglauca\u0026nbsp;\u003c/em\u003eis fertile, and can interbreed to form advanced-generation hybrids, or backcross with either of its parent species to form backcrossed hybrids (Pieper 2017); because advanced-generation and backcrossed hybrids cannot be differentiated, we refer to these collectively as non-F1 hybrids. \u0026nbsp; Non-F1 hybrids comprised the most common taxonomic category in the central/eastern PPR (57% of genets compared to 30% that were F1 hybrids), which suggests that the hybrid zone in the central/eastern PPR is not very recent. \u0026nbsp;\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eWhile the Tangen et al. study (2022) provided important information about the abundance and occupancy of hybrid \u003cem\u003eTypha\u003c/em\u003e throughout wetlands in the central and southeast PPR, some knowledge gaps remain. \u0026nbsp; In this study we expanded the investigation of the \u003cem\u003eTypha\u0026nbsp;\u003c/em\u003ehybrid zone into previously unsampled regions of the west/northwest PPR, beyond the range limit of \u003cem\u003eT. angustifolia\u003c/em\u003e. \u0026nbsp;F1 \u003cem\u003eT.\u0026nbsp;\u003c/em\u003ex \u003cem\u003eglauca\u0026nbsp;\u003c/em\u003ehybrids arise following asymmetric hybridization, in which \u003cem\u003eT. angustifolia\u0026nbsp;\u003c/em\u003eis normally the maternal parent and \u003cem\u003eT. latifolia\u0026nbsp;\u003c/em\u003eis the paternal parent (Pieper 2017). \u0026nbsp;There are almost no records of \u003cem\u003eT. angustifolia\u0026nbsp;\u003c/em\u003efrom the western PPR, which could preclude the local formation of hybrids, and we therefore addressed the following questions: 1) is the paucity of \u003cem\u003eT. angustifolia\u0026nbsp;\u003c/em\u003erecords from the western PPR an accurate reflection of \u003cem\u003eT. angustifolia\u0026nbsp;\u003c/em\u003eoccurrence in this region? 2) If \u003cem\u003eT. angustifolia\u0026nbsp;\u003c/em\u003eplants occur in the western PPR, are they producing seeds that are \u003cem\u003eT.\u0026nbsp;\u003c/em\u003ex \u003cem\u003eglauca\u003c/em\u003e? 3) Does the range expansion of \u003cem\u003eT.\u0026nbsp;\u003c/em\u003ex \u003cem\u003eglauca\u0026nbsp;\u003c/em\u003efollow the range expansion of its maternal species, \u003cem\u003eT. angustifolia\u003c/em\u003e? \u0026nbsp;Invasive plant species pose a major threat to biological diversity in some regions of the PPR (Jones et al. 2023), and this study provides additional information on a potentially problematic range expansion of an invasive wetland hybrid across the PPR. \u0026nbsp;\u003c/p\u003e"},{"header":"Methods","content":"\u003ch3\u003eField work\u003c/h3\u003e\n\u003cp\u003eBetween August 19 and August 24, 2022, we sampled 244 \u003cem\u003eTypha\u0026nbsp;\u003c/em\u003eplants\u003cem\u003e\u0026nbsp;\u003c/em\u003efrom 50 wetlands (34 wetlands in the province of Saskatchewan, 16 wetlands in the province of Alberta) in the western PPR (Figure 1, Table S1). \u0026nbsp;\u003cem\u003e\u0026nbsp;\u0026nbsp;\u003c/em\u003eSampled sites were mostly restricted to small wetlands accessible from roadsides because a substantial portion of the Prairie Pothole Region is privately owned agricultural land. \u0026nbsp;This sampling strategy should not have influenced our findings because an earlier study found that\u0026nbsp;\u003cem\u003eTypha\u003c/em\u003e species and hybrids had comparable frequencies in wetlands versus ditches in eastern North America (Pieper et al., 2020). \u0026nbsp;\u003cem\u003eTypha latifolia\u0026nbsp;\u003c/em\u003eis native to Saskatchewan and Alberta\u0026nbsp;(Flora of North America 2008)\u0026nbsp;and was thus expected to occur at the majority of sites; we therefore prioritized the sampling of \u003cem\u003eT. angustifolia\u0026nbsp;\u003c/em\u003efollowed by \u003cem\u003eT.\u0026nbsp;\u003c/em\u003ex \u003cem\u003eglauca\u003c/em\u003e. \u0026nbsp;We did this\u003cem\u003e\u0026nbsp;\u003c/em\u003eby first visually inspecting the stands at each site and identifying plants that appeared to have the narrowest leaves and \u0026ndash; for flowering plants - the largest gaps between the staminate and pistillate flowers: \u0026nbsp;\u003cem\u003eT. angustifolia\u0026nbsp;\u003c/em\u003ehas the narrowest leaves and largest flower gaps, \u003cem\u003eT. latifolia\u0026nbsp;\u003c/em\u003ehas the widest leaves and typically no discernible flower gap, and F1 hybrids are intermediate in both traits\u0026nbsp;(Smith 1967). \u0026nbsp;Up to eight plants with what appeared to be the narrowest leaves were sampled at each site. From each sampled plant we collected two leaf fragments approximately 10 cm long. \u0026nbsp;Leaf samples were placed into labelled coin envelopes, and the envelopes were placed in a plastic zippered bag containing Sor bead orange silica beads for desiccation. \u0026nbsp;\u0026nbsp;\u003cem\u003e\u0026nbsp;\u003c/em\u003eIn total, we collected leaf samples from 245 plants across 50 sites. \u0026nbsp;In addition, we collected mature fruits from nine plants (not all plants flower in any given year) that were provisionally identified as \u003cem\u003eT. angustifolia\u0026nbsp;\u003c/em\u003eand stored these in paper bags that were enclosed within resealable plastic bags. \u0026nbsp; \u0026nbsp;\u003c/p\u003e\n\u003ch3\u003eGenotyping\u003c/h3\u003e\n\u003cp\u003eBecause the leaf widths and flower gaps of advanced-generation (AGH) and backcrossed hybrids (BCH) can overlap with parental taxa (Snow et al., 2010; Kirk et al., 2011; Geddes et al., 2021; Tangen et al., 2022), we used genetic information to identify the taxonomy of each sampled plant. \u0026nbsp;In brief, we followed the methods described in\u0026nbsp;(Chambers et al. 2024)\u0026nbsp;to extract DNA and genotype each plant at up to five loci with species-specific alleles (one microsatellite locus and four PCR-RFLP loci). \u0026nbsp;Plants with exclusively\u0026nbsp;\u003cem\u003eTypha latifolia alleles (identified as T. latifolia), T. angustifolia\u0026nbsp;\u003c/em\u003ealleles\u0026nbsp;\u003cem\u003e(identified as T. angustifolia), or heterozygous loci (one T. latifolia and one T. angustifolia allele at each locus; identified as F\u003csub\u003e1\u0026nbsp;\u003c/sub\u003e\u003c/em\u003ehybrids) were genotyped at all five loci. \u0026nbsp;Plants that were heterozygous for\u0026nbsp;\u003cem\u003eTypha latifolia\u003c/em\u003e and\u0026nbsp;\u003cem\u003eTypha angustifolia\u0026nbsp;\u003c/em\u003ealleles at some but not at all loci (genotyped at between two and five loci) were identified as advanced generation or backcrossed hybrids and pooled into a single category of non-F1 hybrids. \u0026nbsp;\u0026nbsp;\u003c/p\u003e\n\u003ch3\u003eSeed germination and identification\u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp;\u003c/h3\u003e\n\u003cp\u003eThe nine putative \u003cem\u003eT. angustifolia\u0026nbsp;\u003c/em\u003eplants from which we collected seeds were taxonomically confirmed from genetic data. \u0026nbsp;To investigate whether these plants were generating exclusively F1 hybrids, we germinated their seeds and genotyped a subset of their offspring. \u0026nbsp;The seeds were separated from the inflorescence stalk, and from each maternal plant approximately 0.5g of fruit was prepared for germination following the protocol outlined in (Ahee et al. 2015). \u0026nbsp;The seeds were left to germinate in petri dishes filled with DI water in a climate-controlled (15-20 \u0026deg;C) greenhouse at Trent University. \u0026nbsp; After six days the seedlings were transferred into 10 cm pots filled with well-draining soil from Sunshine Professional growing mixture (germination grade; Sun Gro Horticulture, Brantford, Canada) and left in the greenhouse on plastic trays filled with water. Seedlings were fertilized after 45 days using 100 mL of 0.5 % water-soluble 20:20:20 N:P: K general-purpose fertilizer (Plant-Prod, Leamington, Ontario). Ten seedlings from each maternal plant were harvested after approximately 80-90 days, by which time they had reached heights of approximately 8 cm. \u0026nbsp;Harvested seedlings were put into coin envelopes in resealable plastic bags containing orange silica beads (Sor bead, USA) for desiccation. \u0026nbsp;DNA extraction and genotyping followed the same methods described for the plant samples collected in the field.\u003c/p\u003e"},{"header":"Results","content":"\u003cp\u003eDistribution of taxa\u003c/p\u003e \u003cp\u003eOverall, the majority of the samples that we genotyped were \u003cem\u003eT. latifolia\u003c/em\u003e, followed by F1 \u003cem\u003eT.\u003c/em\u003e x \u003cem\u003eglauca, T. angustifolia\u003c/em\u003e, and then non-F1 hybrids (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). Hybrids were more abundant and identified at more sites than the maternal taxon \u003cem\u003eT. angustifolia\u003c/em\u003e (Fig.\u0026nbsp;1) despite a sampling strategy that prioritized \u003cem\u003eT. angustifolia\u003c/em\u003e.\u003c/p\u003e \u003cp\u003e \u003cem\u003eT. angustifolia\u003c/em\u003e offspring\u003c/p\u003e \u003cp\u003eThe 90 genotyped offspring of nine \u003cem\u003eT. angustifolia\u003c/em\u003e plants comprised 78 (87%) \u003cem\u003eT. angustifolia\u003c/em\u003e, 10 (11%) F1 hybrids, and 2 (2%) non-F1 hybrids (advanced-generation or backcrossed hybrids). Seed samples from five of the nine \u003cem\u003eT. angustifolia\u003c/em\u003e plants included no hybrid offspring. The remaining four plants had 1/10, 1/10, 2/10, and 8/10 hybrids (either F1 or back-crossed hybrids) among their offspring.\u003c/p\u003e"},{"header":"Discussion","content":"\u003cp\u003eThe data collected in this study allow us to address the three questions presented in our introduction. First, the paucity of \u003cem\u003eT. angustifolia\u003c/em\u003e records from the western PPR does indeed seem to be a fairly accurate reflection of its occurrence in this region: despite a sampling strategy that prioritized \u003cem\u003eT. angustifolia\u003c/em\u003e, we found this species at only four of the 50 sampled sites. At least one record suggests that \u003cem\u003eT. angustifolia\u003c/em\u003e has been present in the PPR since the late 1970s (Harms and Ledingham \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e1986\u003c/span\u003e). This record was not verified with genetic data but, if accurate, suggests that despite being in the western PPR for several decades \u003cem\u003eT. angustifolia\u003c/em\u003e remains rare and thus may not be adapted to conditions in this region. Our second question asked whether \u003cem\u003eT. angustifolia\u003c/em\u003e in the western PPR are producing hybrid seeds. Although our sample size was small (reflecting the small number of \u003cem\u003eT. angustifolia\u003c/em\u003e in this region \u0026ndash; and the even smaller number of flowering \u003cem\u003eT. angustifolia\u003c/em\u003e), we found that \u003cem\u003eT. angustifolia\u003c/em\u003e mostly produce \u003cem\u003eT. angustifolia\u003c/em\u003e seedlings, with only 13% of our seedling sample comprising hybrids. This was an unexpected finding given the predominance of \u003cem\u003eT. latifolia\u003c/em\u003e in the area (75% of plants despite a sampling strategy that favoured other taxa), which in turn should increase the likelihood of \u003cem\u003eT. angustifolia\u003c/em\u003e plants being pollinated by heterospecifics and producing mostly hybrids. Previous studies have shown that seedling production by \u003cem\u003eT. angustifolia\u003c/em\u003e is comparable when pollinated by \u003cem\u003eT. angustifolia, T. latifolia\u003c/em\u003e, or \u003cem\u003eT.\u003c/em\u003e x \u003cem\u003eglauca\u003c/em\u003e (Pieper et al. \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2017\u003c/span\u003e), and therefore the predominance of \u003cem\u003eT. angustifolia\u003c/em\u003e seedlings may be at least partly explained by other forms of reproductive barriers such as different flowering times.\u003c/p\u003e \u003cp\u003eAlthough we did not find \u003cem\u003eT.\u003c/em\u003e x \u003cem\u003eglauca\u003c/em\u003e in any of the Alberta sites, it was found in 10/34 Saskatchewan wetlands, compared to \u003cem\u003eT. angustifolia\u003c/em\u003e in only 3/34 Saskatchewan wetlands, suggesting a westwards expansion. Additionally, unlike in the central/eastern PPR, we found very few non-F1 hybrids, suggesting that hybrids have only recently colonized this area. The western PPR therefore appears to be the leading edge of the hybrid range expansion. Our third question asked whether the range expansion of \u003cem\u003eT.\u003c/em\u003e x \u003cem\u003eglauca\u003c/em\u003e into the western PPR is following that of its maternal species \u003cem\u003eT. angustifolia\u003c/em\u003e, and our data suggest this is not the case: because the hybrid occurred at more sites, and across a larger area, than \u003cem\u003eT. angustifolia\u003c/em\u003e, it does not appear reliant on its maternal species for successful establishment at new sites. Although we found that \u003cem\u003eT. angustifolia\u003c/em\u003e produce only a small proportion hybrid seeds, these have the potential for dispersal across short and long distances via wind, water, or animal vectors including waterfowl (Soons \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2006\u003c/span\u003e; Soons et al. \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2008\u003c/span\u003e), and thus can colonize sites from which the maternal species is absent. Additionally, backcrossed hybrids, advanced-generation hybrids, and vegetative propagation can perpetuate hybrid \u003cem\u003eTypha\u003c/em\u003e in the absence of its maternal species. In eastern Canada and the central/eastern PPR \u003cem\u003eT.\u003c/em\u003e x \u003cem\u003eglauca\u003c/em\u003e has outcompeted both parental species (Freeland et al. \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Pieper et al. \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Tangen et al. \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2022\u003c/span\u003e), and may be able to dominate wetlands even with relatively low propagule pressure. Our results collectively suggest that \u003cem\u003eT.\u003c/em\u003e x \u003cem\u003eglauca\u003c/em\u003e has the capacity for continued range expansion that is not reliant on the presence of \u003cem\u003eT. angustifolia\u003c/em\u003e, and the impacts of this range expansion should be monitored because of the potential for this wetland plant to impede ecosystem services and reduce local biodiversity.\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\u003e\u003cem\u003eTypha\u003c/em\u003e taxa (number and percentage) identified from 50 wetlands in the western Prairie Pothole Region. SK\u0026thinsp;=\u0026thinsp;Saskatchewan, AB\u0026thinsp;=\u0026thinsp;Alberta.\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"5\"\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 \u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eTaxon\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eNumber (%) of sampled plants\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eNumber (%) of sampled SK wetlands (out of a total of 34)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eNumber (%) of sampled AB wetlands (out of a total of 16)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003eTotal number (%) of wetlands\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eTypha latifolia\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e183 (75%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e24 (71%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e16 (100%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e40 (80%)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eF1 \u003cem\u003eTypha\u003c/em\u003e x \u003cem\u003eglauca\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e40 (16%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e10 (29%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0 (0%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e10 (20%)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eTypha angustifolia\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e17 (7%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e3 (9%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e1 (6%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e4 (8%)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eBackcrossed/advanced-generation hybrids\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e5 (2%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e3 (9%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0 (0%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e3 (6%)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e "},{"header":"Declarations","content":"\u003cp\u003eThe following statements must be included in your submitted manuscript under the heading 'Statements and Declarations'. This should be placed after the References section. Please note that submissions that do not include required statements will be returned as incomplete.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eFunding\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eFunding for this project was provided by Discovery Grants from the Natural Sciences and Engineering Research Council of Canada awarded to J. Freeland (RGPIN-2017-04371) and M. Dorken (RGPIN-2018-04866). \u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eCompeting Interests\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eThe authors have no relevant financial or non-financial interests to disclose.\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAuthor Contributions\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eThis research was part of Sanjuti Deb Joyee’s MSc thesis, co-supervised by Joanna Freeland and Marcel Dorken. \u0026nbsp; Study conception and design by Joanna Freeland and Marcel Dorken. \u0026nbsp;Field work (sample collection) was performed by Joanna Freeland, laboratory and greenhouse methods by Sanjuti Deb Joyee. \u0026nbsp; \u0026nbsp;The first draft of the manuscript was written by Joanna Freeland and all authors commented on previous versions of the manuscript. All authors read and approved the final manuscript.\u003c/em\u003e\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\n\u003cli\u003eAhee JE, Van Drunen WE, Dorken ME (2015) Analysis of pollination neighbourhood size using spatial analysis of pollen and seed production in broadleaf cattail ( \u003cem\u003eTypha latifolia\u003c/em\u003e ). Botany 93:91\u0026ndash;100. https://doi.org/10.1139/cjb-2014-0169\u003c/li\u003e\n\u003cli\u003eBansal S, Lishawa SC, Newman S, et al (2019) Typha (Cattail) Invasion in North American Wetlands: Biology, Regional Problems, Impacts, Ecosystem Services, and Management. Wetlands 39:645\u0026ndash;684\u003c/li\u003e\n\u003cli\u003eChambers A, Chambers B, Bhargava D, et al (2024) A simple method to genetically differentiate invasive F1 Typha hybrids (T. \u0026times; glauca) and advanced-generation/backcrossed hybrids from parent species (T. Latifolia and T. angustifolia) in eastern Canada and northeastern USA. J Great Lakes Res 50:. https://doi.org/10.1016/j.jglr.2023.102257\u003c/li\u003e\n\u003cli\u003eDavidson NC (2014) How much wetland has the world lost? Long-term and recent trends in global wetland area. Mar Freshw Res 65:. https://doi.org/10.1071/MF14173\u003c/li\u003e\n\u003cli\u003eDoherty K, Howerter D, Devries J, Walker J (2016) Prairie Pothole Region of North America. In: Finlayson C, Milton G, Prentice R, Davidson N (eds) The Wetland Book. Springer\u003c/li\u003e\n\u003cli\u003eDoherty K, Howerter D, Devries J, Walker J (2018) Prairie pothole region of North America. In: Finlayson C, Milton R, Prentice R, Davidson N (eds) The Wetland Book II: Distribution, Description and Conservation. Springer, New York, pp 1\u0026ndash;10\u003c/li\u003e\n\u003cli\u003eFlora of North America (2008) Typha latifolia. In: efloras. Missouri Botanical Garden, St. Louis, MO \u0026amp; Harvard University Herbaria, Cambridge, MA\u003c/li\u003e\n\u003cli\u003eFreeland J, Ciotir C, Kirk H (2013) Regional differences in the abundance of native, introduced, and hybrid Typha spp. in northeastern North America influence wetland invasions. Biol Invasions 15:2651\u0026ndash;2665. https://doi.org/10.1007/s10530-013-0481-4\u003c/li\u003e\n\u003cli\u003eGalatowitsch SM, Anderson NO, Ascher PD, Hall A (1999) Invasiveness in wetland plants in temperate north america. Wetlands 19:733\u0026ndash;755\u003c/li\u003e\n\u003cli\u003eGrace J, Harrison J (1986) The biology of Canadian weeds: 73. Typha latifolia L., Typha angustifolia L. and Typha x glauca Godr. Can J Plant Sci 66:361\u0026ndash;379. https://doi.org/10.4141/cjps86-051\u003c/li\u003e\n\u003cli\u003eGraham A, Mudrzynski B, Polzer E, Wilcox DA (2022) Restoration of a Lake Ontario-connected fen through invasive Typha removal. Restor Ecol 30:. https://doi.org/10.1111/rec.13562\u003c/li\u003e\n\u003cli\u003eHarms VL, Ledingham GF (1986) Narrow-leaved cattail, Typha angustifolia, and the hybrid cattail, Typha \u0026times; glauca, newly reported form Saskatchewan. Can F Nat 100:107\u0026ndash;110\u003c/li\u003e\n\u003cli\u003eJones SA, DeKeyser ES, Dixon C, Kobiela B (2023) Invasive Species Change Plant Community Composition of Preserved Prairie Pothole Wetlands. Plants 12:. https://doi.org/10.3390/plants12061281\u003c/li\u003e\n\u003cli\u003eKingsford RT, Basset A, Jackson L (2016) Wetlands: conservation\u0026rsquo;s poor cousins. Aquat Conserv Mar Freshw Ecosyst 26:892\u0026ndash;916. https://doi.org/10.1002/aqc.2709\u003c/li\u003e\n\u003cli\u003eLawrence BA, Lishawa SC, Rodriguez Y, Tuchman NC (2016) Herbicide management of invasive cattail (Typha \u0026times; glauca) increases porewater nutrient concentrations. Wetl Ecol Manag 24:. https://doi.org/10.1007/s11273-015-9471-x\u003c/li\u003e\n\u003cli\u003ePattison-Williams JK, Pomeroy JW, Badiou P, Gabor S (2018) Wetlands, Flood Control and Ecosystem Services in the Smith Creek Drainage Basin: A Case Study in Saskatchewan, Canada. Ecol Econ 147:. https://doi.org/10.1016/j.ecolecon.2017.12.026\u003c/li\u003e\n\u003cli\u003ePieper S, Dorken M, Freeland J (2020) Genetic structure in hybrids and progenitors provides insight into processes underlying an invasive cattail (Typha \u0026times; glauca) hybrid zone. Heredity (Edinb) 124:714\u0026ndash;727. https://doi.org/10.1038/s41437-020-0307-y\u003c/li\u003e\n\u003cli\u003ePieper SJ, Nicholls AA, Freeland JR, Dorken ME (2017) Asymmetric Hybridization in Cattails (Typha spp.) and Its Implications for the Evolutionary Maintenance of Native Typha latifolia. J Hered 108:479\u0026ndash;487. https://doi.org/10.1093/jhered/esx036\u003c/li\u003e\n\u003cli\u003eSmith S (1967) Experimental and natural hybrids in North American Typha (Typhaceae). Am Midl Nat 78:257\u0026ndash;287. https://doi.org/10.2307/2485231\u003c/li\u003e\n\u003cli\u003eSoons MB (2006) Wind dispersal in freshwater wetlands: Knowledge for conservation and restoration. Appl Veg Sci 9:. https://doi.org/10.1111/j.1654-109x.2006.tb00676.x\u003c/li\u003e\n\u003cli\u003eSoons MB, Van Der Vlugt C, Van Lith B, et al (2008) Small seed size increases the potential for dispersal of wetland plants by ducks. J Ecol 96:619\u0026ndash;627. https://doi.org/10.1111/j.1365-2745.2008.01372.x\u003c/li\u003e\n\u003cli\u003eTangen BA, Bansal S, Freeland JR, et al (2022) Distributions of native and invasive Typha (cattail) throughout the Prairie Pothole Region of North America. Wetl Ecol Manag 30:. https://doi.org/10.1007/s11273-021-09823-7\u003c/li\u003e\n\u003cli\u003eTravis SE, Marburger JE, Windels S, Kub\u0026aacute;tov\u0026aacute; B (2010) Hybridization dynamics of invasive cattail (Typhaceae) stands in the Western Great Lakes Region of North America: A molecular analysis. J Ecol 98:7\u0026ndash;16. https://doi.org/10.1111/j.1365-2745.2009.01596.x\u003c/li\u003e\n\u003cli\u003eXu T, Weng B, Yan D, et al (2019) Wetlands of international importance: Status, threats, and future protection. Int J Environ Res Public Health 16:. https://doi.org/10.3390/ijerph16101818\u003c/li\u003e\n\u003cli\u003eZedler JB, Kercher S (2004) Causes and consequences of invasive plants in wetlands: Opportunities, opportunists, and outcomes. CRC Crit Rev Plant Sci 23:431\u0026ndash;452. https://doi.org/10.1080/07352680490514673\u003c/li\u003e\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":"Typha, Prairie Pothole Region, hybrid, range expansion, wetlands","lastPublishedDoi":"10.21203/rs.3.rs-4607793/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-4607793/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eWetlands around the world face unprecedented threats, including from invasive species. In North America, the invasive cattail hybrid \u003cem\u003eTypha\u003c/em\u003e x \u003cem\u003eglauca\u003c/em\u003e dominates wetlands around the Laurentian Great Lakes, and more recently was found in high abundance across the central and eastern Prairie Pothole Region, an area that includes one of the world\u0026rsquo;s largest wetland complexes. Surveys of the \u003cem\u003eTypha\u003c/em\u003e hybrid zone have so far been largely conducted in areas where hybrids are well established, and it therefore remains unclear whether the range expansion of this invasive hybrid occurs after the establishment of its maternal species, \u003cem\u003eT. angustifolia.\u003c/em\u003e We surveyed 50 wetlands in the western PPR, and found that while 75% of plants were native \u003cem\u003eT. latifolia\u003c/em\u003e, the second most common group was F\u003csub\u003e1\u003c/sub\u003e hybrids: these had greater abundance and occupancy than \u003cem\u003eT. angustifolia\u003c/em\u003e despite the fact that \u003cem\u003eT. angustifolia\u003c/em\u003e produces relatively few hybrid seeds, and therefore highlights the importance of long-distance dispersal for this hybrid range expansion. The distribution of hybrids combined with the paucity of non-F1 hybrids suggests that the western PPR represents a leading edge of the range expansion by invasive \u003cem\u003eT.\u003c/em\u003e x \u003cem\u003eglauca.\u003c/em\u003e Our results show that \u003cem\u003eT.\u003c/em\u003e x \u003cem\u003eglauca\u003c/em\u003e has the capacity for continued range expansion that does not rely on the presence of \u003cem\u003eT. angustifolia\u003c/em\u003e, and the impacts of this range expansion should be monitored because of its potential to impede ecosystem services and reduce local biodiversity.\u003c/p\u003e","manuscriptTitle":"Range expansion of the invasive hybrid cattail Typha x glauca exceeds that of its maternal plant T. angustifolia in the western Prairie Pothole Region of North America","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2024-07-17 10:26:36","doi":"10.21203/rs.3.rs-4607793/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"reviewerAgreed","content":"","date":"2024-06-26T16:03:21+00:00","index":0,"fulltext":""},{"type":"reviewersInvited","content":"","date":"2024-06-25T17:08:59+00:00","index":"","fulltext":""},{"type":"editorInvited","content":"Biological Invasions","date":"2024-06-21T18:23:49+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2024-06-20T13:14:32+00:00","index":"","fulltext":""},{"type":"submitted","content":"Biological Invasions","date":"2024-06-19T15:52:00+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":"83fe40ab-5f6f-4edf-830c-612b93bab29c","owner":[],"postedDate":"July 17th, 2024","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"published-in-journal","subjectAreas":[],"tags":[],"updatedAt":"2024-09-16T16:05:13+00:00","versionOfRecord":{"articleIdentity":"rs-4607793","link":"https://doi.org/10.1007/s10530-024-03439-7","journal":{"identity":"biological-invasions","isVorOnly":false,"title":"Biological Invasions"},"publishedOn":"2024-09-12 15:58:07","publishedOnDateReadable":"September 12th, 2024"},"versionCreatedAt":"2024-07-17 10:26:36","video":"","vorDoi":"10.1007/s10530-024-03439-7","vorDoiUrl":"https://doi.org/10.1007/s10530-024-03439-7","workflowStages":[]},"version":"v1","identity":"rs-4607793","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-4607793","identity":"rs-4607793","version":["v1"]},"buildId":"qtupq5eGEP_6zYnWcrvyt","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}