loosestrife (Lythrum salicaria)], perpet- Management and Control Issues for Native, uation of misconceptions regarding the origin of an invasive species [e.g., reed Invasive Species (Reed Canarygrass): canarygrass (Phalaris arundinacea)], as Evaluating Philosophical, Management, and well as theories of invasive spread. This set the background for seminal papers Legislative Issues on invasive species spread by Galato- witsch et al. (1999) and a special issue 1 1 1 of Euphytica for plant breeders during Neil O. Anderson , Alan G. Smith , Andrzej K. Noyszewski , crop domestication (Anderson et al., Emi Ito2, Diana Dalbotten3, and Holly Pellerin3 2006a, 2006b, 2006c). In this review, we examine imp- lications from an inherent bias on the ADDITIONAL INDEX WORDS. cryptogenic species, evolution, herbaceous ornamentals, “ ” Phalaris arundinacea postulated exotic origin of reed can- invasion terminology, arygrass to exemplify how new scientific SUMMARY. The issue of native invasive species management rarely occurs and is findings realigned the origination of fraught with biological, social, and economic challenges as well as posing difficulties midwest U.S. (Minnesota) populations in decision-making for land managers. The terminology for categorization of invasive as “native” and thus whether it should species is examined in the context of their bias(es), which complicates control. An be controlled or preserved by land man- example of a newly determined native species, which is also invasive, is used as an agers, based on potential revised regula- example to navigate control and regulatory issues. Native, invasive reed canarygrass tory (legislative) mandates. This may (Phalaris arundinacea L.) occurs throughout Minnesota and most likely the entire midwest region of central United States and Canadian provinces. The species was have wide-ranging applicability to other previously assumed to be an exotic, nonnative Eurasian import but recent molecular native, invasive plant species. evidence supports its status as a native but invasive species. We address how this change to being a native but highly invasive species modifies approaches to mitigate Invasive species terminology its potential control for state, Tribal, and local authorities. The implications of these and bias(es) new findings will require differential shifts in land managers’ perspectives and Since invasion biology emerged as approaches for control. Particular differences may exist for Tribal Land Managers vs. ’ departments of natural resources and private agencies. Additionally, regulatory a science after Elton s monograph challenges have yet to be decided on how to legislate control for a native invasive (1958), terminology to describe the species that had been previously assumed as exotic or foreign in origin. These involved or affected species has gone opportunities to change attitudes and implement judicial control measures will serve through an interesting evolution, albeit as a template for other invasive species that are native in origin. with occasional inherent bias(es). As mentioned previously, weed science has been fraught with similar bias in judgmental terminology or definitions. lant invasions were recognized as Bureau of Land Management (2006) Rawls (1996) describes “burdens an important biological phenome- posted “The Weed Hall of Shame.” In of judgment” to explain disagreements non more than a century ago invasion biology, bias toward a species P that permeate society. A burden en- may arise from determining (sometimes (Galatowitsch et al., 1999; Zedler and compasses careful consideration of con- erroneously due to the lack of molecu- Kercher, 2005). The modern discipline flicting, complex data because people lar, genomic tools) whether a species is of invasion biology commenced with the with divergent backgrounds (such as publication of the monograph The Ecol- native or exotic (nonnative) to a conti- fi among invasion biology scientists, land ogy of Invasions by Animals and Plants nent or speci c area of actual or potential managers, regulators and legislators fi (Elton, 1958). Although signi cant sci- invasion. Additional bias(es) may result from varying backgrounds, cultures, fi fi enti c research has identi ed theoretical from implicating spread explicitly based and disciplinary foci) have different val- and causal mechanisms of invasive spe- on human-mediated factors and remov- ues, logic, cause and effects, and views cies spread, biotic and abiotic factors, ing humans from species consideration of evidence (Frank, 2019). These may and land management approaches for in natural ecosystems. Vague or judg- cause persistent disagreements in com- their control, invasion biology terminol- mental terminology helps to perpetuate plex fields such as invasion biology. ogy—often with inherent bias(es)—con- these bias(es). Since beginning research For example, reducing the science of tinues to create confusion in the field. on invasion biology in 1989, we have invasion biology to assertions such as Part of the bias arose from defini- found this bias in numerous invasive “natives are beneficial and exotics are tions and views of weeds in cultivated species with long lag phases from intro- not” overlooks the complexities of spe- settings, predominantly defining a weed duction to eventual spread [e.g., purple cies-species interactions and purported in a judgmental, nonscientificmanneras “a plant out of place” (Illinois Industrial University, 1872; Wilson, 1863) that, Units To convert U.S. to SI, To convert SI to U.S., when it escapes cultivation and estab- multiply by U.S. unit SI unit multiply by lishes self-propagating populations in 0.4047 acre(s) ha 2.4711 the wild (Davis et al., 2000; Elton, 2.54 inch(es) cm 0.3937 1958; Mack et al., 2000), becomes an 25.4 inch(es) mm 0.0394 À invasive species. Until recently, the U.S. 2.2417 ton(s)/acre t·ha 1 0.4461
354 August 2021 31(4) axiomatic threats of exotic species to humans” (Iannone et al., 2020). (Williamson, 1996), “exotic” (Green, (Goodenough, 2010). Long (2011) This overlooks other vectors implicated 1997), “imported” (Williamson and argued these burdens caused disagree- in the global spread of invasives, such as Fitter, 1996), “indigenous” (Sauer, ments in other scientific disciplines (e.g., wind, water, natural disasters, birds, 1988), “introduced” (Lonsdale, 1994), climate science), which were “founded and mammals. In addition to divorcing “invasive” (Daehler, 1998), “invasive in difference rather than error.” humans from being a natural species in nonindigenous” (Pimentel et al., Invasive biology terms such as ecosystems, such terminology may 2000), “native” (Meyer and Florence, “native” and “nonnative,”“exotic,” ignore previous evolution of plant 1996), “naturalized” (Hussey et al., “invaders,” or “foreign” denote a communities in ecosystems and that 1992), “nonnative, immigrant” (Baz- human value-added component to such systems are dynamic and ever- zaz, 1986), “nonnative, introduced by their meaning and creates a belief sys- changing, such as responding to the humans” (Iannone et al., 2020), and/ tem that invasives “are a unique eco- effects of climate change. Indeed, in or “weedy” (Fox, 1990). Definitions of logical phenomenon” (Davis and the United Kingdom, about equal these terms to create a more formalized Thompson, 2001). Such restricted numbers of native and nonnative spe- lexicon of terminology are often debat- views regard native plants as being cies are expanding their range such that able (Daehler, 2001; Cuda et al., static or nonspreading outside of their they lack distinguishing characteristics 2014; Richardson et al., 2000, 2008; “native” geographic area, whereas (Davis and Thompson, 2001; Thomp- Warren, 2007) and recent efforts to “exotic” species spread beyond their son et al., 1995). A positive correlation clarify them have been questioned intended or cultivated range. Some among exotic and native “species (Colautti and Richardson, 2009). invasion biologists assert that humans richness” in wetlands was found, For example, the University of Flo- are the sole vector of invasive species wherein exotic invasive species were rida Institute of Food and Agricul- that do or can “cause harm,” creating equally likely to dominate a wetland as tural Sciences, Invasive Species definitions of an invasive species as: “A native ones, sharing similar negative Council defined seven terms as their species that a) is nonnative to a speci- effects on species biodiversity (Houla- lexicon for Floridian invasion biology fied geographic area, b) was introduced han and Findlay, 2004). (“native, nonnative, introduced, by humans (intentionally or uninten- Despite these findings, a native established, invasive, nuisance, range tionally), and c) does or can cause envi- species may be regarded by some to change”), while instructing stake- ronmental or economic harm or harm be desirable in cultivated and wild holderstoavoidtermssuchas“native ecosystems, providing essential eco- invasive” (replace with “nuisance”), Received for publication 11 Jan. 2021. Accepted for system services, whereas an exotic or “invasive exotic,”“invasive weed,” publication 15 June 2021. foreign species may be viewed as “alien,”“foreign,” and “indigenous” Published online 28 July 2021. undesirable outside of its native range. (Iannone et al., 2020). 1Department of Horticultural Science, University of Minnesota, St. Paul, MN 55108 Of course these static and restrictive Additional connotations with definitions are reinforced by research invasive species’ terminology outside of 2Department of Earth Sciences, University of Min- nesota, Minneapolis, MN 55455 area and/or commodity-based artifi- the science relate to their impact (Davis “ 3St. Anthony Falls Laboratory, University of Minne- cial demarcation of silos of knowl- and Thompson, 2001). For example, sota, Minneapolis, MN 55414 edge.” Iannone et al. (2020) would in 1999, U.S. President Bill Clinton This article was presented as part of the Invasive classify spreading nonnatives as issued the landmark Executive Order Plants Research Professional Interest Group work- “nuisance” and/or “established” spe- on Invasive Species (Order 13112, 3 shop entitled “Progress in Identification and Con- trol of Weedy to Invasive Plants both Domestic and cies exhibiting “range change,” but Feb. 1999) wherein an “‘invasive spe- Abroad” held on 12 Aug. 2020 during the virtual not “invasive” since they are native. cies’ means an alien species whose American Society for Horticultural Science’s annual conference. The fact remains that plant species introduction does or is likely to cause Funding in support of this publication was from the spread regardless of political, geo- economic or environmental harm or Minnesota Invasive Terrestrial Plants and Pests Cen- graphical, or social boundaries via var- harm to human health.” Numerous ter; the Minnesota Agricultural Experiment Station; ious vectors across time and space, U.S. federal and state agencies, such as Czech–US American Science Information Center (AMVIS); Ministry of Education, Czech Republic, although human-mediated spread by the U.S. Fish and Wildlife Service University of South Bohemia in Cesk e Bud ejovice; a naturally occurring species, humans (2012) and the U.S. Department of European Union Education for Competitiveness (Homo sapiens L.), is a common, but Agriculture, Animal and Plant Health Operational Programme; and the J. William Ful- bright Scholar Program. not the only, mode of dispersal. Inspection Service (2020), as well as N.O.A. and A.G.S. are Professors. Resultant vagueness of terminol- international agencies, such as the “ ” “ ” A.K.N. is a Postdoctoral Research Associate; currently ogy between native and nonnative European Union (2010) and the Inter- Quality Control Analyst III, Aldevron, Fargo, ND. species has been stipulative (Frank, national Union for the Conservation of E.I. is a Professor and Director of the Limnological 2019), requiring assumptions that are Nature (2021) define invasive species Research Center. arbitrary (Chew and Hamilton, 2011; in a similar manner. D.D. is an Associate Director of Diversity and Simberloff, 2012) or subjective to retain Similar to the classic definition of a Broader Impacts. flexibility (Colautii and Richardson, weed, species dichotomization into dis- H.P. is a Research Scientist, Elder, and Instructor. 2009). Terminology for the same phe- tinct classes of “native,”“nonnative” N.O.A. is the corresponding author. E-mail: nomena was not restricted to these invoke normative claims of inalienable [email protected]. “native” and “nonnative” terms (Davis rights of native species within their This is an open access article distributed under the CC BY-NC-ND license (https://creativecommons. et al., 2000), since an invasive species range with an amoral superiority com- org/licenses/by-nc-nd/4.0/). couldbereferredtoas:“alien” (Crawley plex (Chew and Hamilton, 2011). https://doi.org/10.21273/HORTTECH04796-21 et al., 1996), “colonizer” or “endemic” Introduction of “nonnative” species
August 2021 31(4) 355 into a region has been proposed to be that arise for affected invasion biology native North American types and reed “tantamount to trespassing” (Chew interest groups and clientele for spe- canarygrass of European origin. and Hamilton, 2011). cies categorized as native, noninvasive; HISTORIC MISCONCEPTIONS LEAD Oddly enough, pre-20th-century native, invasive; exotic, noninvasive; TO BIAS. Reed canarygrass is a cool-sea- Europeans viewed introduced species and exotic, invasive (Table 1). Reed son riparian grass with large populations more positively (Frank, 2019), particu- canarygrass is used as the example to in Europe, Asia, and North America larly since New World food species such illustrate its current status in each along fresh watercourses (Ambros as corn (Zea mays L.), potato (Solanum category. and Stykar, 1999) and floodplains tuberosum L.), tomato (Solanum lyco- (Hroudov a et al., 2009) growing to persicum L.), and pepper (Capsicum Reed canarygrass water depths of 30 cm. It is also an annuum L.) fed the post-Columbian IDENTIFICATION OF NORTH important commercial crop for forage world and were integrated into various AMERICAN NATIVE VERSUS EXOTIC and has been cultivated as an ornamen- cultures worldwide (Weatherfield, EUROPEAN. The question naturally tal perennial since the 18th century 1989). In the United States, there were (Alway, 1931; Carlson et al., 1996; Cas- “ ” arises whether there are qualitative, acclimatization societies that intro- diagnostic trait (visual trait) differences ler, 2010; Schoth, 1938), as well as for ducedOldWorldspeciesintotheconti- at the macroscopic and/or micro- revegetation (Figiel et al., 1995), shore- nent and the official U.S. government scopic levels that distinguish between line restoration (Figiel et al., 1995), policy supported new PIs until recently North American native and exotic phytoremediation (Chekol et al., (Coates, 2006). It was not until the late European individuals of reed canary- 2002), nitrogen removal in wastewater 20th century that the realization of det- grass. The standard dichotomous key treatment facilities, and for biofuels rimental environmental and ecosystem (which denotes two or more differ- (Vymazal, 2001, 2013). Reed canary- impacts, such as extinction, created reg- ences for a particular trait to distin- grassisapredominantforbincentral ulatory acts, endangered species lists, guish types) for reed canarygrass European ecosystems but is viewed as noxious weed lists, invasive species lists, an archaeophyte (predating the Roman separate the species from the others in and gave rise to environmental groups Empire) instead of being classified as the genus (Anderson, 1961). The spe- and nongovernmental agencies [NGOs invasive (Anderson et al., 2016). cies is characterized as follows: “14. (Frank, 2019)]. In fact, Elton (1958) Reed canarygrass is a widespread Plants perennial from scaly rhizomes; provided a militaristic view of this as invasive species across the globe inflorescence usually more than 5 cm. ecological “invasions.” In 1964, a (Holm et al., 1991; Kercher and long, lobed or branched at the base; monograph was produced by Baker and Zedler, 2004). The species predomi- caryopsis 2.0 mm. long or less, mostly Stebbins (1965) on the detrimental, ” nates in large wetland areas of the evolutionary results of “nonnative” spe- northern hemisphere (Anderson, United States (Zedler and Kercher, cies introductions. Invasion biology 1961). However, within reed canary- 2004) because it tolerates nutrient came together as a whole in the 1980s, grass itself, the only differential traits fluctuations in riparian environs as based on articulations by the Scientific separate two types, based on leaf color- well as transportation corridors (Jaku- Committee on Problems of the Envi- ation, into ribbon grass [P. arundina- bowski et al., 2010) and produces “ À ronment (SCOPE) regarding intro- cea f. picta (L.) Arcang; 15. Leaves – 1 ı ” high biomass [4 14 t·ha (C zkova duced species/environmental properties longitudinally striped with white 4a. ] et al., 2015; Gyulai et al., 2003)]. that create invasive or noninvasive and reed canarygrass (P. arundinacea Reed canarygrass is the most effective “ species (Davis, 2006), as well as the dedi- f. arundinacea; 15. Leaves uniformly grass species to extract nitrogen from … ” cation by the Organization of Con- green 4b. ) (Anderson, 1961). soils and waterways (Marten and servation Biology to biodiversity Although minor quantitative variation Heath, 1985), thus providing ecosys- conservation (Soul e, 1985; Takacs, exists for plant height and biomass, all tem services in areas with high nutri- 1996). Most recently, invasion science plants still possess the diagnostic traits, ent levels. Its impact on biodiversity has been used to encompass biological including ligules (Anderson, 1961) in decreasing native plant species invasions as a multidisciplinary field and Floret Type 4 (Voshell et al., exceeds that of both flowering-rush including sociology and management 2015). Thus, there is no differentia- (Butomus umbellatus L.) and purple (Ricciardi and Cohen, 2007) wherein tion between North American native loosestrife, other riparian invasives data from scientists may be directly and exotic European individuals (Lavoie et al., 2003, 2005). translated into land management con- (Akiyama et al., 2015). Importation of reed canarygrass trol methods. Biogeographic origin is integral by agrarian and horticultural Euro- Because the focus of this review is for risk assessment of PIs (Frank, pean settlers into North America to illustrate the challenges and oppor- 2019), although it comes with caveats occurred repeatedly for cultivation as tunities in decision-making regarding such as native species may be invasive a forage and ornamental crop (Harris, native yet invasive species and their or cause “harm” and many nonnative 1835; Lavergne and Molofsky, 2004; controversial terminology (Anderson species may cause no harm. Genetic New England Farmer, 1834; Schoth, et al., 2020a, 2020b), we are not pro- markers have recently been found to 1938). The extent of spread by the posing new, less-biased terms for inva- distinguish among these types. A imported reed canarygrass germplasm sive species. Rather, we categorize the review of the history leading to this in extant populations is largely dilemmas using standard terminology discovery follows. However, land unknown and is in the process of being and their connotations to illustrate the managers cannot use any visual, diag- determined (Noyszewski et al., 2019, issues for control and management nostic trait to distinguish between 2020, 2021). However, multiple
356 August 2021 31(4) Table 1. Potential interest group response options for decision-making, risk assessment, control, and management of four native/invasive and noninvasive/invasive species combinations using current reed canarygrass risk assessments as an example. Interest group Native, noninvasive Native, invasive Exotic, noninvasive Exotic, invasive Monitoring and Not listed, unless an Not categorized; no If it affects commercial If it affects commercial regulatory U.S. endangered or threatened listing for reed agriculture and agriculture and interstate federal agenciesz species [Convention on canarygrass interstate commerce: commerce: added to International Trade in added to noxious noxious weed lists, import/ Endangered Species weed lists, import/ export lists of allowable (CITES)]. export lists of species. Reed canarygrass Reed canarygrass is not allowable species. not listed. endangered or Reed canarygrass For importation: enforcement threatened. not listed. by USDA-APHIS-PPQ only For importation: For importation: if listed. Reed canarygrass is enforcement by [U.S. enforcement by not listed Department of Agriculture USDA-APHIS-PPQ (USDA), Animal and only if listed. Reed Plant Health Inspection canarygrass is not Service (APHIS), Plant listed. Protection Quarantine (PPQ)] only if listed. Reed canarygrass is not listed. Tribal Land Native species models; reed Native noninvasive Risk assessment of Risk assessment of invasive Managers [Great canarygrass is not listed species models are invasive species to species to Tribal resources: Lakes Indian not proposed; reed Tribal resources: Invasive species and invasive Fish and Wildlife canarygrass is not invasive  native co-  native co-occurrence Commission, listed occurrence models: models: “Quantify the risk (GLIFWC); i.e., “Quantify the risk of of nonnative invasive plants states nonnative invasive to culturally important surrounding the plants to culturally native plants”; reed Great Lakes] important native canarygrass listing as an plants.” exotic, invasive species but no invasive species model created Public land Minnesota Department of MNDNR: nonexistent MNDNR: nonexistent MNDNR: invasive, nonnative managers (state Natural Resources category; “invasive” is category; MNDOT: plant list; invasive terrestrial of Minnesota)x (MNDNR): Native Plant exclusive to “native”: not categorized; reed plant list; reed canarygrass Encyclopedia; MNTaxa: reed canarygrass not canarygrass not listed listing as a “nuisance The State of Minnesota listed; Minnesota species” because it Vascular Plant Checklist; Department of “outcompetes most native “native” is incorrectly Transportation species in natural wetlands”; listed as being exclusive (MNDOT) Minnesota MNTaxa lists reed to “invasive” Noxious Weeds list: canarygrass as “invasive” specially regulated MNDOT Minnesota Noxious category; reed Weeds list has four canarygrass not listed; classifications: prohibited, native and invasive eradicate; prohibited, poison ivy is “specially control; restricted noxious regulated” weeds; specially regulated; reed canarygrass not listed Minnesota Department of Agriculture (MNDA): Invasive species programs for plant protection, pest detection and management unit, weed team, noxious weed law, early detection Private Land The Nature Conservancy in The Nature Conservancy in Managers (State Minnesota: protects Minnesota: no reed of Minnesota) native plant communities canarygrass listing
(Continued on next page)
August 2021 31(4) 357 Table 1. (Continued). Interest group Native, noninvasive Native, invasive Exotic, noninvasive Exotic, invasive Nongovernmental Minnesota Native Plant EEDMapS (Early University of Minnesota EEDMapS invasive species: organizations Society; Restore Your Detection and Andersen Reed canarygrass listing; (state of Shore: Native Plant Distribution Mapping Horticulture Library Minnesota Invasive Species Minnesota) Encyclopedia; University System) invasive Plant Information Advisory Council (MISAC): of Minnesota Andersen species: reed Online database: under guidelines of the Horticulture Library canarygrass listing as: includes exotic, Minnesota State Plant Information Online “Nativity of this plant cultivated species Management Plan for database: includes native, is debated; it is native Invasive Species cultivated species to Europe and possibly parts of Asia, but it may also be native to NW U.S.” zMonitoring U.S. federal agency listings arise across governmental agencies (not included here are the 50 U.S. states and territories, local or municipal listings; Canadian listings would include the 10 Canadian provinces and three territories and local or municipal listings): the USDA, USDA-APHIS, USDA-APHIS-PPQ, U.S. Army Corps of Engineers, U.S. Bureau of Land Management, U.S. Department of Defense, U.S. Federal Highway Administration, and the U.S. Fisheries and Wildlife Service (see text for specifics). yGLIFWC provides examples of potential responses. Not included here are the separate risk assessment procedures of the 11 Minnesota Tribal Nations: Bois Forte Band of Chippewa, Fond Du Lac Reservation, Grand Portage Band of Chippewa Indians, Leech Lake Band of Ojibwe, Lower Sioux Indian Community, Mille Lacs Band of Ojibwe, Prairie Island Indian Community, Red Lake Band of Chippewa Indians, Shakopee Mdewakanton Sioux (Dakota) Community, Upper Sioux Community, and White Earth Reservation. xAlthough the number and type of state agencies with Public Land Managers for invasive species enforcement varies by each U.S. state and Canadian province, in the example State of Minnesota, the involved agencies are MNDNR, MNDA, and MNDOT. reports have surfaced in recent research Cartographic evidence also sup- lack of diagnostic trait(s) as well as documenting native North American ports the historic existence of one or molecular marker(s) at that time, reed canarygrass populations. more native reeds. For example, some which could unequivocally differenti- NATIVE AMERICAN USES: of the earliest maps of Verendrye ate native vs. European ecotypes POTENTIAL SPREAD OR SPECIES (circa 1737), Thompson (circa 1814), (Dore and McNeill, 1980), resulted ’ fi PERPETUATION. Most important to Long (circa 1823), and Pope (circa in reed canarygrass classi cation as a understanding reed canarygrass’ native 1849) named “Ga-shashagunushko- cryptogenic species (Carlton, 1996). history is the widespread, yet tragically kawi-sibi” or “the-place-of-rushes-riv- Anderson (2019) reported on ignored or overlooked, use of this er” in Ojibwe for the Roseau River the discovery of “an unplowed, pris- grass by Native American Tribal (French for “reed”)inMinnesota tine field (part of the ancient Lake Nations. Across the North American (Northwest Regional Development Agassiz lakebed) was used to harvest continent, reed canarygrass was used Commission, 2014; Prud’homme, hay during the Dust Bowl era (1930s) extensively for weaving, clothing, fish- 1916). It is unknown which “reed- and sold throughout the midwestern ing weirs, basket making, or thatching like” grass this referred to; i.e., reed U.S.” Transport of hay harvested for wigwam (wickiup, wetu) roofs canarygrass or common reed [Phrag- from this native, unplowed field of (Densmore, 1974; Kindscher and mites australia (Cav.) Trin. ex pure reed canarygrass in the 1930s Noguera, 2002; Turner et al., 1980). Steud.], both of which are native. Dust Bowl era may have enhanced its Seeds of related species, maygrass COROLLARY EVIDENCE OF NATIVE spread via highway corridors on land, (Phalaris caroliniana Walters) and lit- RANGE. Analysis in the 1970s of shore- instead of waterways, across the mid- tle seed canarygrass (Phalaris minor line reed canarygrass in the Ottawa western United States and Canada Retz.), were also important food sour- and French Rivers in Ontario, Can- (Anderson, 2019). Building interstate ces (Rea, 1991). Collection of leaves ada, as well as the northern Great and interprovincial highways after for weavings occurred along river Lakes, concluded it was native (Dore 1950 increased the area of disturban- edges and trails (Steltzer, 1976). and McNeill, 1980). Marten and ces along highway corridors and Transport of live plant material and Heath (1985) classified it as “indi- drainage ditches favorable to reed can- seeds in and among ceded tribal terri- genous” to all five continents’ tem- arygrass to invade (Anderson, 2019). tories throughout the 1800s along perate climates and later, Merigliano In summation, clearly there are lakes and rivers, which were major and Lesica (1998) determined popu- numerous reports of native popula- transportation corridors until roads lations of the interior western North tions throughout North America pre- and railroads were constructed, may America were also native. Remote po- dating settlements by European have aided in the spread of reed canar- pulations in Lake Mistassini and Anti- colonizers as well as thereafter. These ygrass across the continent. Both costi Island of Quebec (Lavoie et al., have been confirmed with more Native Americans as well as the French 2005) were also postulated to be recent genetic evidence that follows. and English colonizers connected with native. Historic herbaria plant speci- GENETIC EVIDENCE OF ITS the Hudson Bay Company forts in mens from as early as 1825 matched NATIVE STATUS. Reviews of early Canada and the United States used corsican reed canarygrass [P. arundi- genetic and molecular studies of reed these for transportation well into the nacea ssp. rotgesii (Husn.) Kergu elen canarygrass (Anderson, 2019; Anderson early 20th century (Newman, 1985). (Merigliano and Lesica, 1998)]. A et al., 2016; Casler and Undersander,
358 August 2021 31(4) 2006; Noyszewski et al., 2019) provide et al., 2009; Jakubowski et al., 2011). 2001), and Roseau, MN (Seed Today, additional evidentiary support for the Cross-pollination among native North 2012), as well as ornamental clones species as a North American native. American and European types would worldwide. Its displacement of other Not surprisingly for individuals within be possible since grasses are wind pol- native species is undisputed (Miller thesamespecies,sharedgenetic linated and reed canarygrass is self- and Zedler, 2003; Zedler and Kercher, markers among native North American incompatible (Casler and Hovin, 2004). Significant costs have been and European individuals were com- 1984; Jakubowski et al., 2013). This realized in its control and correspond- monly reported for biochemical and would alter the North American gene ing restoration efforts (Reinhardt molecular nuclear genetic markers. pool (Vila et al., 2000). An increase in Adams and Galatowitsch, 2005). Biochemical markers, such as allozymes reed canarygrass’ North American Ornamental ‘Picta’ ribbon grass or alloenzymes (enzyme forms range is not a result of hybridization also naturally occurs as wild popula- encoded by different alleles that differ among European, forage, and North tions in North America (Barkworth structurally but not functionally from American individuals (Jakubowski et al., 2007). Lavoie et al. (2005) pro- other allozyme alleles at the same et al., 2011), particularly from SNP posed an invasion window during genetic locus) showed a high level of data (Noyszewski et al., 2018, 2019, 1923–43 for reed canarygrass in Can- similarity among North American and 2020, 2021), despite unsubstantiated ada, which was the result of native European types (Gifford et al., 2002; theories to the contrary (Lavergne ‘Picta’ being sold in Montr eal nurseries Lavergne and Molofsky, 2007); similar and Molofsky, 2007). in 1834 onwards (Guilbaut, 1834). findings were reported with single Despite these evidential data on ‘Picta’ is, thus, one example of the sequence repeats (SSRs) of DNA (Ket- the native status of at least some popu- native North American yet invasive spe- tenring et al., 2019). Molecular genetic lations throughout North America, cies’ spread. This first invasion window markers showed shared sequences but many invasion biology researchers cannot be attributed to early Canadian also genetic differences among conti- continued to posit that reed canary- forage cultivars, since they were not nents; e.g., for intersimple sequence grass is exotic and native only to Eur- licensed until years later (in 1959) and repeats of DNA [ISSRs (Anderson asia as well as being introduced to distribution occurred even later from et al., 2016)], amplified fragment North America. These encompass a 1965 onwards (Bittman et al., 1980; length polymorphisms [AFLPs (Ander- range of hypotheses from the importa- Goplen et al., 1964). A second Cana- son et al., 2018; Jakubowski et al., tion of invasive forage types from Eur- dian invasion window was during 2013)], single nucleotide polymor- asia (Dore and McNeill, 1980; 1963–78, this time with pure, native phisms [SNPs (Noyszewski et al., Lavergne and Molofsky, 2004), cross reed canarygrass, most likely due to 2018, 2019, 2020, 2021)], as well as hybridization between Eurasian and high nutrient runoff from agricultural internal transcribed spacer regions of North American populations (Lav- fields, increased production of low alka- the nuclear ribosomal cistron, 18S- ergne and Molofsky, 2007), as well as loid cultivars in commercial fields, 5.8S-26S [ITS (Graper et al., 2021; the release of individuals with hybrid human disturbance such as the build- Noyszewski et al., 2021; Quintanar vigor (Merigliano and Lesica, 1998). ing of highway corridors, and water et al., 2007; Voshell and Hilu, 2014; However, if ecologists were familiar- level changes (Lavoie et al., 2005). Voshell et al., 2011)]. Nelson et al. ized with all scientific literature, it An additional native, invasive (2014) found that populations of wild, would have become clear that early spread hypothesis includes the mid- forage, and ornamental North Ameri- 1940s reed canarygrass breeding in western United States dispersal of reed can and European reed canarygrass had Canada and the United States had canarygrass (Anderson, 2019). Our high levels of genetic diversity within, used native North American land races recent discovery of the distribution of rather than among, populations; this (Carlson et al., 1996). ‘Auburn’, hay across the midwestern United was confirmed in additional studies ‘Superior’,and‘Ioreed’ forage types States from the unplowed field in (Anderson et al., 2018; K avov aetal., were selected directly from land races Roseau, MN, during the Dust Bowl era 2017; Nelson and Anderson, 2015). (Casler, 2010) and large cultivated warrants further investigations. In her- At least one European forage cultivar populations were derived from native baria across North America, reed canar- [e.g., Chrastava (Caga s, 2008)] was stands in the Rocky Mountain states ygrass specimens predating the Dust genetically similar to wild European and southern Canadian provinces Bowl (1930–40) are numerous, as they populations (Anderson et al., 2016). (Schoth, 1938). These facts, along were frequently collected by explorer- All extant and historic herbaria speci- with the substantive use of reed canar- botanists (Jakubowski et al., 2011, mens of reed canarygrass in Minnesota ygrass by Native Americans through 2013). All Minnesota and Wisconsin were found to be native and genetically the centuries (Densmore, 1974; herbaria specimens predating the 1930s distinct from European samples, based Kindscher and Noguera, 2002; Turner dustbowl are native (Jakubowski et al., on SNP and ITS data (Graper et al., et al., 1980), provide additional assur- 2011, 2013; Noyszewski et al., 2019). 2021; Noyszewski et al., 2018, 2019, ances that many populations of reed Thus, we propose that an additional 2020, 2021). canarygrass in North America were invasion window occurred in the INVASIVE SPREAD. The theorized and are native. United States from the 1930s to the role of forage and ornamental culti- Additional propagule pressure present day, which overlaps with the vars in increasing the spread of inva- may have arisen from the primary reed first invasion window in Canada. How- sive reed canarygrass across North canarygrass producers of commercial ever, the question remains what the America (Merigliano and Lesica, forage seed production in Saskatche- impact of the spread of native, Euro- 1998) is unsubstantiated (Casler wan, Canada (Statistique Canada, pean, or their hybrids means for
August 2021 31(4) 359 agencies, regulators, tribal land manag- Such species would be added to exist- (GLIFWC; Falk et al., 2015) provides ers, public land managers, private land ing noxious weed lists and import/ for invasive species reporting, species managers, and NGOs (Table 1). Deci- export lists of allowable species by the account listings, risks assessment mod- sion making and new risk assessment following agencies. USDA-APHIS els, and species distribution models for models or invasive species predictive (2010) maintains the Federal Noxious tribal land managers in states and prov- schemes (Whitney and Gabler, 2008) Weed List, which controls import/ inces surrounding the Great Lakes, wouldberequiredtoreevaluatethesta- export through USDA-APHIS-PPQ. which includes portions of the State of tus of native, invasive reed canarygrass The U.S. Department of Defense Minnesota (Table 1). The 11 Minne- for all organizations (Table 1). (2019) maintains DENIX. The U.S. sota Tribal Nations, namely the Bois Army Corps of Engineers previously Forte Band of Chippewa (2021), Fond Monitoring and regulatory U.S. published an Army Corps Noxious Du Lac Band of Lake Superior Chip- federal agencies and Nuisance Plant Management pewa (2021), Grand Portage Band of U.S. federal agencies and pro- Information System (website no lon- Lake Superior Chippewa (2021), grams that work on reed canarygrass ger available) but now has a nation- Leech Lake Band of Ojibwe (2021), monitoring include the U.S. Depart- wide Invasive Species Policy, which Lower Sioux Indian Community ment of Agriculture (USDA); the applies to all Civil Works projects (2021), Mille Lacs Band of Ojibwe Animal and Plant Health Inspection (U.S. Department of the Navy and (2021), Prairie Island Indian Commu- Service (APHIS); the Federal Noxious U.S. Army Corps of Engineers, 2009) nity (2021), Red Lake Nation (2019), Weed List; Plant Protection and (Table 1). The National Invasive Spe- Shakopee Mdewakanton Sioux Quarantine (PPQ); the interagency cies Act (U.S. Congress, 1996) cojoins (Dakota) Community (2021), Upper Aquatic Nuisance Species Taskforce; with the U.S. Army Corps of Engi- Sioux Community (2021), and the “ the U.S. Army Corps of Engineers neers to either prevent or reduce White Earth Nation (2021) may also Noxious and Nuisance Plant Manage- establishment of invasive and nonna- have specific recommendations and ment Information System; the U.S. tive species.” The U.S. Bureau of reporting for invasive species control. Bureau of Land Management; the Land Management (2006) published Reed canarygrass is listed and U.S. Department of Defense Environ- “The Weed Hall of Shame” and also searchable as an “exotic, invasive” spe- mental Network and Information provides information on weeds and cies in GLIFWC (2019), although no Exchange (DENIX); the U.S. Federal invasives (U.S. Bureau of Land Man- invasive species model has been cre- Highway Administration; and the agement, n.d.) (Table 1). The USFWS ated (Table 1). Initial discussions with U.S. Fish and Wildlife Service (2015) provides an Invasive Species some of the Minnesota Tribal Nations (USFWS) Invasive Species Program Program (2015). The U.S. Federal revealed no current knowledge of (Table 1). These agencies produce Highway Administration defines inva- using reed canarygrass in weavings listings, based on their jurisdictions sive species as “Non-native [sic.] flora nor of its native status (E. Ito, D. Dal- along with additional input from the and fauna that can cause significant botten, and H. Pellerin, personal National Invasive Species Information changes to ecosystems, upset the eco- communication). Deliberations within Center, the National Agricultural Pest logical balance, and cause economic and among the Minnesota Tribal Information Center, and the U.S. harm to the agricultural and recrea- NationsaswellaswithGLIFWC National Plant Board (Table 1). Mon- tional sectors” (U.S. Department of regarding the “native, invasive” status itoring agencies in the separate 50 Transportation, Federal Highway of reed canarygrass may change the U.S. States, the 10 Canadian Provin- Commission, n.d.). Additional advisory current classification of this species. ces and its three territories as well as taskforces or boards for U.S. federal Until that has been accomplished, local or municipal government listings agencies include: the Aquatic Nuisance actions by tribal land managers are in both countries were not included Species Taskforce (2011), the USDA uncertain. Likewise, future efforts for due to their inherent complexity. National Invasive Species Information potential control of reed canarygrass The included U.S. federal agen- Center (n.d.), National Agricultural would seem unlikely because this is a cies show that, for “native, non- Pest Information Center (2004), and native species with cultural heritage invasive” species, the reference point the U.S. National Plant Board (2014). ties both in Minnesota and elsewhere is for identifying and preventing entry Reed canarygrass is neither listed throughout North America. or exit of endangered or threatened on any of the U.S. federal agency list- Identification of “exotic, invasive” species, based on the Convention on ings nor by any advisory taskforce or Eurasia species growing within “native, International Trade in Endangered board listings (Table 1). Thus, the invasive” stands of reed canarygrass in Species (1975) (Table 1). Although change in designation for most, if not Tribal Nations would require extensive reed canarygrass is neither endangered all, populations of reed canarygrass in and costly SNP testing for nuclear or threatened, apparently no listings Minnesota (Noyszewski et al., 2020, DNA (Noyszewski et al., 2019, 2020, or regulations for any “native, non- 2021) from an “exotic, invasive” to a 2021) until specific genetic sequences invasive” species exist (Table 1). “native, invasive” should not presently are identified to distinguish between For the categories of “exotic, be affected by U.S. federal agencies Eurasian and North American species. noninvasive” and “exotic, invasive,” and regulations. While the chloroplast genome has been the importation or exportation of the sequenced in reed canarygrass (Xiong species may be regulated, particularly Tribal land managers et al., 2020), it has not yet been com- if they affect commercial agriculture The Great Lakes Indian Fish and pleted for the nuclear genome where and interstate commerce (Table 1). Wildlife Commission-Exotic Plants the SNP markers are located. On the
360 August 2021 31(4) other hand, ITS regions may hold pos- canarygrass as “Invasive” (MNDNR, In the State of Connecticut, reed can- sibilities for earlier detection of native 2021e). MNDOT has four classifica- arygrass is listed as “invasive” but not vs. exotic reed canarygrass accessions, tions for “exotic, invasive” species: Pro- regulated (University of Connecticut, provided our initial ITS findings (that hibited, Eradicate; Prohibited, Connecticut Invasive Plant Working separated most native, North American Control; Restricted Noxious Weeds; Group, 2018) whereas in the adjacent types in Minnesota from elsewhere in and Specially Regulated (MNDOT, Commonwealth of Massachusetts, as the U.S. and Eurasia) are accurate 2020) (Table 1). Reed canarygrass, an invasive species it is listed as (Noyszewski et al., 2021). however, is not listed herein. In coordi- “prohibited” (Massachusetts Natural nation with MISAC (2020), Minne- Resources Council, 2021). Only Public land managers (State of sota Department of Agriculture’s Washington State lists it as a “Class C Minnesota) (MNDA) Invasive Species Programs noxious weed” (Washington State fi “ For “native, noninvasive” species, provide four classi cations for exotic, Noxious Weed Control Board, ” the State of Minnesota Department of invasive species: Plant Protection, Pest 1995). If future research proves that Natural Resources (MNDNR), pro- Detection and Management Unit, additional populations of reed canary- vides vascular plant checklists in two Weed Team, Noxious Weed Law, Early grass in these and other U.S. states resources: The Native Plant Encyclo- Detection (Table 1). This program, and/or Canadian provinces are also pedia (MNDNR, 2021a) and Mn- however, lacks any species listings; native to North America, the refor- Taxa: The State of Minnesota Vascular thus, reed canarygrass is also not mulation of approaches in the State of Plant Checklist (MNDNR, 2021b). A included herein. Minnesota may have applicable value definition of interest is provided: Public land managers within the to these states. “ ” “ ” State of Minnesota are thus confronted native is exclusive of invasive. Private land managers (State of This would require modification for with myriad issues, depending on the reed canarygrass identified as “native, state agency in which they operate as Minnesota) invasive” (Noyszewski et al., 2019, well as lack of listings and/or conflict- The Nature Conservancy (2021) 2020, 2021) as well as for other spe- ing categorizations across agencies. is a large primary private land holder cies in this category, such as poison ivy These all would require modification in the State of Minnesota that protects (Toxicodendron radicans Kuntze ssp. to accommodate the new findings that native plant communities and controls negundo Gillis). The Minnesota Inva- reed canarygrass populations in Min- invasive species on its lands of 60,845 sive Species Taskforce provides cate- nesota are predominantly, if not exclu- acres (S. Edmunds, personal commu- gorical definitions in Minnesota sively, “native, invasive” instead of nication). Reed canarygrass is not Noxious Weed Risk Assessments being classified as “exotic, invasive.” listed in any category, particularly as [Minnesota Invasives Species Advisory Such changes would require legislative an “exotic, invasive” species (Table Council (MISAC), 2020] by answer- action by the Minnesota Legislature to 1). Additional smaller private land- ing the question: “Is the plant species affect changes throughout the Minne- owners, such as residential properties, or genotype nonnative?” with either sota Terrestrial Invasive Plant Resour- would be mandated to follow state “disputed, yes, and no.” ces listings (Minnesota Board of Soil regulations for invasive species control Reed canarygrass is not listed in and Water Resources, 2011) and the should they be legislated. Thus, any The Native Plant Encyclopedia but is Minnesota State Management Plan for private land manager is at liberty to found in MnTaxa with a listing as Invasive Species (MNDNR, 2009). control or not control reed canary- invasive. As such, it would require Current discussions have included a grass on their lands at their discretion. reclassification as “native, invasive” range of responses from land managers It remains to be seen whether the reed canarygrass. In contrast, the tasked with the control and eradication reclassification of reed canarygrass as a Minnesota Department of Transpor- of reed canarygrass from public lands “native, invasive” species would tation (MNDOT, 2020) has a Minne- (N.O. Anderson, unpublished data), change landowners’ perspectives on sota Noxious Weeds list, which lists a based on control methods recom- its control. “Specially Regulated” category for mended by USDA Natural Resources “native, invasive” species (e.g., poison Conservation Service (2002). Some NGOs (State of Minnesota) ivy; Table 1). However, reed canary- welcome potential changes to accom- A wide variety of NGOs operate grass is not included in this category modate the new listing of reed canary- in various capacities with differing and should be. grass as a “native, invasive” although services in the State of Minnesota For “exotic, noninvasives,” both whether this change would alter cur- (Table 1). For “native noninvasive” the MNDNR and MNDOT do not rent management plans is unclear. species, the Minnesota Native Plant recognize this category, and thus, reed These could range from no change in Society (MNPS, 2021) is dedicated to canarygrass is not listed. The “exotic, control approaches to selective control the preservation of native plant species invasive” category in the MNDNR of particularly aggressive populations within the state. The MNPS Con- provides an Invasive, Nonnative Plant that threatens water resources or rare/ servation Committee informs the list (MNDNR, 2021c), as well as an endangered native species. membership on “issues relating to Invasive Terrestrial Plant list (MND- The lack of regulation in the conservation, including, but not lim- NR, 2021d); reed canarygrass is in the State of Minnesota for reed canary- ited to, promotion of the use of native latter listing because it “outcompetes grass (MNDNR, 2021f) and its classi- plant species, preservation of native most native species in natural wetlands” fication as a “nuisance species,” differs plant species and communities, and (Table 1). MNTaxa also lists reed from at least three other U.S. states. conservation of rare and endangered
August 2021 31(4) 361 species.” Thus, invasive species con- constraints in risk assessment, and Lithuania. River Res. Appl. 34:300–309, trol would be of paramount interest in continued evolution of all species doi: 10.1002/rra.3259. this organization, which may result in regardless of their native origin or ten- Anderson, N.O., A. Noyszewski, A.G. lobbying efforts and other supportive dency to spread in either native or Smith, A. Kilian, E. Ito, D. Dalbotten, A. roles to public and private as well as nonnative ranges (Broennimann and Timm, and H. Pellerin. 2020a. Manage- tribal land managers. As part of the Guisan, 2008). Thus, a thorough and ment and control issues for native, invasive MNDNR, Restore Your Shore is an deliberative reevaluation and reassess- species: Philosophical, managerial and leg- advisory group that published an ment of reed canarygrass by all inter- islative issues. HortScience 55(9S):S191. online searchable database: Native est groups as a native, invasive species (Abstr.). Plant Encyclopedia (MNDNR, in Minnesota and elsewhere in North Anderson, N.O., A. Noyszewski, A.G. 2019). The University of Minnesota America is warranted before any party Smith, D. Dalbotten, E. Ito, A. Timm, and Andersen Horticulture Library main- can propose sound solutions for its H. Pellerin. 2020b. Reed canarygrass: tains an online searchable database, preservation and/or control. Implications for control of an invasive Plant Information Online (University species when it is native. 1 Aug. 2020. of Minnesota Libraries, 2021), listing
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