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Contributions to the flora of Colorado

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Title:
Contributions to the flora of Colorado
Creator:
Majack, Marike Laurel ( author )
Place of Publication:
Denver, CO
Publisher:
University of Colorado Denver
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Language:
English
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1 electronic file (138 pages). : ;

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Botany -- Colorado -- Denver ( lcsh )
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bibliography ( marcgt )
theses ( marcgt )
non-fiction ( marcgt )

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Review:
With increasing population and urbanization there is a pressing need to catalog biodiversity and understand the impacts of urban expansion on native ecosystems. Habitat degradation, fragmentation, and loss contribute to the extirpation of native plants, while facilitating the spread and establishment of introduced species. Floristic inventories are important tools used to document biodiversity, increase our understanding of species distributions, and inform management decisions. Traditionally, these inventories have been compiled by individuals conducting extensive fieldwork collecting vouchers over a discrete and usually brief period of time, typically not involving extensive herbarium study. Museum (herbarium) collections vouchering plant occurrences provide invaluable resources for many types of research, even more so with the recent focus on digitization of herbarium accessions. Virtual access to these collections has sparked the use of novel research methods, techniques and approaches, advancing research in many fields and adding richness to phytogeographic studies. However, the use of these new technologies does not come without risks. Here, I demonstrate the used of web-based herbarium databases for two floristic products traditionally created from a combination of fieldwork and study of physical specimens in herbaria; a checklist of vascular plant species occurring outside of cultivation in the City and County of Denver, Colorado and updates to the distribution of the pondweed family (Potamogetonaceae) in Colorado. To demonstrate a novel method of documenting a flora by rigorously mining the web-based herbarium databases, a flora of Denver was compiled using existing vouchers from 20 herbaria. After multiple queries, much manual revision, and supplementation from fieldwork, a total of 758 species collected between 1861 and 2013 were documented for the county; these represent 410 genera and 101 families. Web-based herbarium databases were also used to guide work further documenting pondweeds in Colorado by identifying gaps in previous collecting efforts. Subsequent fieldwork resulted in 62 new and noteworthy collections of pondweeds in Colorado, including a species new to the state and 34 new county records. These case studies demonstrate the utility of web-based herbaria in floristics while highlighting the limitations of these databases, which can potentially led to misleading data.
Thesis:
Thesis (M.S.)--University of Colorado Denver. Biology
Bibliography:
Includes bibliographic references.
General Note:
Department of Integrative Biology
Statement of Responsibility:
by Marika Laurel Majack.

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University of Colorado Denver
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|Auraria Library
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All applicable rights reserved by the source institution and holding location.
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904608461 ( OCLC )
ocn904608461

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CONTRIBUTIONS TO THE FLORA OF COLORADO by MARIKA LAUREL MAJACK B.S., University of Colorado, Boulder, 2005 A thesis submitted to the Faculty of the Graduate School of the University of Colorado in partial fulfillment of the requirements for the degree of Master of Science Biology 2014

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This thesis for the Master of Science degree by Marika Laurel Majack has been approved for the Department of Integrative Biology by Leo P. Bruederle, Chair Melissa B. Islam Laurel M. Hartley !"#$%&'()'*# !+",# ii

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Majack, Marika Laurel (M.S., Biology) Contributions to the Flora of Colorado Thesis directed by Associate Professor Leo P. Bruederle. ABSTRACT With increasing population and urbanization there is a pressing need to catalog biodiversity and understand the impacts of urban expansion on native ecosystems. Habitat degradation, fragmentation, and loss contribute to the extirpation of native plants, while facilitating the spread and establishment of introduced species. Floristic inventories are important tools used to document biodiversity, increase our understanding of species distributions, and inform management decisions. Traditionally, these inventories have been compiled by individuals conducting extensive fieldwork collecting vouchers over a discrete and usually brief period of time, typically not involving extensive herbarium study. Museum (herbarium) collections vouchering plant occurrences provide invaluable resources for many types of research, even more so with the recent focus on digitization of herbarium accessions. Virtual access to these collections has sparked the use of novel research methods, techniques and approaches, advancing research in many fields and adding richness to phytogeographic studies. However, the use of these new technologies does not come without risks. Here, I demonstrate the use of web-based herbarium databases for two floristic products traditionally created from a combination of fieldwork and study of physical specimens in herbaria: a checklist of vascular plant species occurring outside of cultivation in the City and County of Denver, Colorado and iii

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updates to the distribution of the pondweed family (Potamogetonaceae) in Colorado. To demonstrate a novel method of documenting a flora by rigorously mining web-based herbarium databases, a flora of Denver was compiled using existing vouchers from 20 herbaria. After multiple queries, much manual revision, and supplementation from fieldwork, a total of 758 species collected between 1861 and 2013 were documented for the county; these represent 410 genera and 101 families. Web-based herbarium databases were also used to guide work further documenting pondweeds in Colorado by identifying gaps in previous collecting efforts. Subsequent fieldwork resulted in 62 new and noteworthy collections of pondweeds in Colorado, including a species new to the state and 34 new county records. These case studies demonstrate the utility of web-based herbaria in floristics while highlighting the limitations of these databases, which can potentially lead to misleading data. The form and content of this abstract are approved. I recommend its publication. Approved: Leo P. Bruederle iv

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I dedicate this work to my grandmother Shirley Lange, who inspired my love of botany, and continues to demonstrate that a brave woman can accomplish anything she sets her mind to. v

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ACKNOWLEDGMENTS This project would not have been possible without the complete and unwavering support of Jacob and Nan Uhl, for which I am forever grateful. Additional funding was provided by the Colorado Native Plant Society, Colorado Garden Foundation, Society of Herbarium Curators, and USDA Forest Service (PO# AG-82AT-P-14-0034) I am so thankful for the kind guidance of my advisor, Leo Bruederle, whose enthusiasm and patience kept me both challenged and grounded throughout this program. I am very grateful for the invaluable advice, encouragement, and technical assistance I have received from many professional botanists, especially Melissa Islam, Pam Regensberg, and the entire Research Department at the Denver Botanic Gardens, Steve Popovich (USDA Forest Service), Zdenek Kaplan (PRA), Tim Hogan and Dina Clark (COLO), Jen Ackerfield (CS), Elizabeth Brown (CPW), Ron Wittmann, Mo Ewing, and CNHP staff, especially Denise Culver, Pam Smith, Bernadette Kuhn, and Jill Handwerk. I would also like to extend my sincere gratitude to those who made my fieldwork possible, especially Jacob Uhl for providing transportation, lodging, and donating four months of his time to assist with fieldwork in seven counties in Colorado. I am also very grateful to Nan and Genny Uhl for their week of assistance in Chaffee County. Thank you to Jackie Sanderson for kindly escorting me to many sites on Douglas County Open Space, and to Tom Bates and Trevor Richards (USDA Forest Service) for accompanying me in Arapaho National Forest. CU Denver students Chelsea Beebe, Elizabeth Pansing, Jill Pyatt, and Daniel Gee deserve special thanks for their contributions to the flora of Denver project and assistance with aquatic fieldwork. ! vi

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TABLE OF CONTENTS CHAPTER I. FLORISTICS IN THE TWENTY-FIRST CENTURY 1 II. THE VASCULAR FLORA OF DENVER, COLORADO: A CASE STUDY OF FLORISTICS IN THE TWENTY-FIRST CENTURY 4 Introduction 4 Methods 6 Results 10 Discussion 11 Tables and Figures 21 Checklist of the Vascular Flora of the City and County of Denver 30 III. NEW AND NOTEWORTHY COLLECTIONS OF PONDWEEDS (POTAMOGETONACEAE) IN COLORADO 73 Introduction 73 Methods 76 Results 78 Discussion 105 Tables and Figures 113 REFERENCES 121 APPENDIX 125 vii

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LIST OF FIGURES FIGURE II.1. Location and political boundaries of the City and County of Denver, Colorado. 21 II.2. Proportions of native and non-native vascular plant species documented in the City and County of Denver, Colorado. 24 II.3. The most species-rich vascular plant families in the City and County of Denver Colorado. 24 II.4. Proportion of cultivated species included on a checklist automatically generated by SEINet for Denver County, Colorado. 28 II.5. Data sources used to voucher the checklist of vascular plants growing outside of cultivation in the City and County of Denver, Colorado. 28 II.6. Temporal distribution of databased records (1861-2013) of vascular plant species growing outside of cultivation in the City and County of Denver, Colorado. 29 III.1. Documented distribution of pondweeds ( Potamogeton and Stuckenia ) in Colorado based on herbarium records collected between 1873 and 2012. 114 III.2. Sites surveyed for pondweeds ( Potamogeton and Stuckenia ) 2013-2014. 115 III.3. Updated distribution of Potamogeton crispus L. in Colorado. 118 III.4. Populations of pondweeds ( Potamogeton and Stuckenia ) documented in Colorado between 1873 and 2014. 119 III.5. Colorado county records of pondweeds ( Potamogeton and Stuckenia ) documented between 1873 and 2014. 120 viii

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LIST OF TABLES TABLE II.1. Herbaria providing discoverable records of vascular plants occurring outside of cultivation in the City and County of Denver, Colorado. 22 II.2. New species records documented in the City and County of Denver, Colorado in 2013. 23 II.3. Species of concern tracked by the Colorado Natural Heritage Program (CNHP) documented in the City and County of Denver, Colorado between 1861 and 2013. 25 II.4. Endemic and near-endemic species documented in the City and County of Denver, Colorado between 1886 and 2013. 26 II.5. Noxious weeds listed by the Colorado Department of Agriculture (CDA) documented in the City and County of Denver between 1861 and 2013. 27 III.1. Herbaria providing discoverable records of pondweeds ( Potamogeton and Stuckenia ) from Colorado 113 III.2. Pondweeds ( Potamogeton and Stuckenia ) documented in Colorado by existing herbarium records (1873-2012) and new records from fieldwork (2013-2014). 116 A.A. Herbaria searched for records from the City and County of Denver, Colorado, and for pondweeds (Potamogetonaceae) from Colorado. 127 ix

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LIST OF ABBREVIATIONS APG3 Angiosperm Phylogeny Group III classification ARIZ University of Arizona Herbarium ASC Northern Arizona University BLM Bureau of Land Management BONAP Biota of North America Project BRY S. L. Welsh Herbarium, Brigham Young University CNHP Colorado Natural Heritage Program CDA Colorado Department of Agriculture COCO Colorado College Herbarium COLO University of Colorado Museum Herbarium CPNWH Consortium of Pacific Northwest Herbaria CS Colorado State University Herbarium DMNS Denver Museum of Nature and Science F Field Museum of Natural History FLD Fort Lewis College Herbarium FNA Flora of North America G1 NatureServe Global Conservation Rank: Critically Imperiled G2 NatureServe Global Conservation Rank: Imperiled G3 NatureServe Global Conservation Rank: Vulnerable G4 NatureServe Global Conservation Rank: Apparently Secure G5 NatureServe Global Conservation Rank: Secure GU NatureServe Global Conservation Rank: Unrankable GH Gray Herbarium, Harvard University ID Stillinger Herbarium, University of Idaho KANU University of Kansas Herbarium KHD Kathryn Kalmbach Herbarium, Denver Botanic Gardens KSC Kansas State University Herbarium MO Missouri Botanical Gardens x

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NMC New Mexico State University NRCS Natural Resource Conservation Service NY New York Botanical Garden OKLA Oklahoma State University Herbarium PRA Institute of Botany, Academy of Sciences, Pr" honice PCA Colorado Natural Heritage Program Potential Conservation Area S1 NatureServe Subnational Conservation Rank: Critically Imperiled S2 NatureServe Subnational Conservation Rank: Imperiled S3 NatureServe Subnational Conservation Rank: Vulnerable S4 NatureServe Subnational Conservation Rank: Apparently Secure S5 NatureServe Subnational Conservation Rank: Secure SNR NatureServe Subnational Conservation Rank: Unranked SU NatureServe Subnational Conservation Rank: Unrankable SJNM San Juan College Herbarium SEINet Southwest Environmental Information Network RM Rocky Mountain Herbarium, University of Wyoming RMBL Rocky Mountain Biological Laboratory UCD University of C olorado Denver UCR University of California, Riverside US United States National Herbarium, Smithsonian Institution USDA United States Department of Agriculture UTC Intermountain Herbarium, Utah State Universit y WGS84 World Geodetic System 1984 WS Marion Ownbey Herbarium, Washington State University WSC Western State College Herbarium WTU University of Washington WWB Western Washington University xi

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CHAPTER I FLORISTICS IN THE TWENTY-FIRST CENTURY Recent advances in technology have had paradigm-shifting impacts in plant systematics. The resources traditionally used in floristics are now increasingly available in electronic form. Herbaria are being digitized and published on the Internet, offering instant access to specimen records. The Flora of North America Project is now available online (efloras.org), as is the USDA Natural Resources Conservation Service PLANTS Database (plants.usda.gov). Websites such as Google Books (books.google.com), The Biodiversity Heritage Library (biodiversitylibrary.org), and The Internet Archive (archive.org) provide free access to digitized versions of historic floras, journals and books that could be otherwise unavailable for floristic research. Additionally, novel resources are being created as websites compiling information as searchable databases benefitting from multiple collaborators and constant internal and public review. Examples of these resources include The Plant List (theplantlist.org), a collaborative database of plant species, which is an invaluable resource for determining synonymy. The closely related International Plant Names Index (ipni.org) is a website providing access to plant names and biographical information. This increased availability of information comes at an appropriate time, as there is a push internationally to document and catalog biodiversity as the current extinction event continues. Tracking changes in species distribution is increasingly important as climate change and anthropogenic impacts threaten native plant communities. It is absolutely imperative that biodiversity 1

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information be current and accurate, as successful conservation depends on frequent surveys of native flora, with early recognition of invasive exotics. Increasingly, herbaria are digitizing their collections, providing access through individual or consortia websites. Those herbaria that allow access to herbarium records through their institutional website do so through various web interfaces, with some providing dataset downloads, regional checklists, or similar resources. In contrast, consortium websites combine specimen data from multiple institutions into a single, searchable database, which vary in scope and geographic coverage. In the United States, the National Science Foundation provides a large portion of the funding for virtual herbaria through the Advancing Digitization of Biological Collections program (iDigBio 2013). Announced in 2010, the Biodiversity Information Serving Our Nation project funded by ADBC will provide access to United States natural history collection records through GBIF website (Global Biodiversity Information Facility) and iDigBio (Integrated Digitized Biocollections) (Barkworth and Murrell 2012). International organizations like GBIF provide access to all types of biodiversity data, combining records from many natural history collections. GBIF is a government-initiated and funded initiative established in 2001, which currently includes 10,004 datasets from 464 publishers (GBIF 2013). Regional databases are more specialized, in that they comprise herbarium records from a specific geographical area. For example, SEINet (Southwest Environmental Information Network), funded by NSF grants and led by Arizona State University, focuses on the flora of the Southwestern United States. SEINet currently provides access to records in 48 herbaria, and offers many additional resources and tools, such as high! 2

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resolution images of herbarium specimens, custom maps and checklists, interactive keys, games, and links to the Flora of North America project (SEINet 2013a). The digitization of herbarium holdings is far from complete, and the usefulness of the existing electronic records is further hampered by the lack of a single database. The US Virtual Herbarium project endeavors to create a single portal providing access to an electronic database of all herbarium specimens housed in the United States. Although not currently funded, the project's focus is to facilitate linking of smaller herbaria to a regional network by extending the coverage of existing consortium databases, or by creating new ones. In connection with the USVH project, Utah State University conducts an annual survey of herbaria in the United States to determine digitization progress. The first survey, conducted in 2011, showed that of the 729 herbaria in the United States registered with Index Herbariorum (Theirs 2014 ), 601 appear to be active. The survey was sent to active herbaria, and 329 responses were received as of 2012. These responses reveal that 228 herbaria maintain a database, 71 of which provide access through a regional website, with an additional 59 providing local access only. Based on responses to the 2012 survey, these herbaria house a total of 70,800,276 specimens, 3,383,947 (32%) of which have been databased (Barkworth and Murrell 2012). 3

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CHAPTER II THE VASCULAR FLORA OF DENVER, COLORADO: A CASE STUDY OF FLORISTICS IN THE TWENTY-FIRST CENTURY Introduction As digitization of herbarium collections continues (Barkworth and Murrell 2012), research opportunities afforded by electronic formats have sparked the use of novel research methods, techniques and approaches. Institutional and consortium websites facilitate access to natural history collections, advancing research in many fields (e.g., phenology, phenotypic plasticity) and adding richness to phytogeographic studies. Floristic inventories are important tools used to document biodiversity, increase our understanding of species distributions, and inform land use decisions. Traditionally, these inventories have been compiled by individuals conducting extensive fieldwork collecting vouchers over a discrete and usually brief period of time, typically not involving extensive herbarium study. The widespread digitization of herbarium collections facilitates the creation of vouchered species checklists without the time and cost of visiting individual herbaria and adds value to these projects with the addition of historic or rare records housed in out-of-state institutions. However, this does not come without risks. To demonstrate a novel method of documenting a flora by rigorously mining webbased herbarium databases, a checklist of native and naturalized vascular plants is created for the City and County of Denver, Colorado. With increasing population and urbanization there is a pressing need to catalog biodiversity and understand the impacts of urban expansion on native ecosystems. 4

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Habitat degradation, fragmentation, and loss contribute to the extirpation of native plants, while facilitating the spread and establishment of introduced species. A recent study (Dolan et al. 2011) comparing historic and current species composition for Marion County, Indiana (including Indianapolis) revealed a decrease in native species of 2.4 species per year, and an increase in introduced species of 1.4 species per year since 1940. Lososova et al. (2012) demonstrated that large cities in arid climates typically have the highest percentage of non-native plant species, suggesting that some cities in drier regions of the United States have higher rates of native plant loss and invasion by exotic species than those documented in Marion County. A recent comparison of 110 urban floras (La Sorte et al. 2014) revealed checklists have been compiled for cities in the Northeast, Midwest and Pacific West regions of the United States, yet some regions (e.g., the Mountain West) remain unrepresented. A flora of Denver, the largest city in the Mountain West, has not been compiled in over 100 years. Denver is located in the center of a large metropolitan area in the semi-arid Front Range Urban Corridor in north-central Colorado. Herbarium accessions for Denver date back to 1863, providing records of the city's flora since near its founding in 1858. The efforts of early botanists, especially Alice Eastwood, offer the unique opportunity to compile a comprehensive checklist of the flora of Denver including the species present before urbanization. In 1893, not long after Denver was founded, Eastwood published A Popular Flora of Denver, Colorado. Most of the fieldwork documenting the flora was performed by Eastwood, as evidenced by accessions dating from 1882. Her book also refers to a few collections by Thomas C. Porter, Thomas Meehan, Miss E. Eaton, Dr. 5

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George Smith, and his son, Benjamin H. Smith. In total, Eastwood listed 487 species in 265 genera and 62 families. Eastwood cited 36 species as being rare in Denver (4 of these were introduced species), and noted the presence of 48 introduced and naturalized plants, only 35 years after the first Europeans settled in the region. After the publication of her book in 1893, Eastwood continued to document the flora of Denver. She was joined by many other botanists in this effort who, over the last 120 years, have contributed enough herbarium specimens to make Denver County one of the best-documented counties in the Western United States (Taylor 2014). Here, I use these invaluable collections to create a checklist of the native and naturalized vascular flora of the City and County of Denver, demonstrating the utility of web-based herbarium databases in the creation of a regional flora. Methods Site descriptio n Denver is located in the Front Range Urban Corridor in north-central Colorado, just east of the Front Range of the Southern Rocky Mountains in the High Plains ecoregion. Founded in 1858, Denver has quickly grown to be the most populous city in Colorado. In 2010, the population of the county was 600 158 in a metro area of approximately 2.6 million (US Census 2010). The average population density in 2010 was 1500 people per km 2 The consolidated City and County of Denver occupies 401 km 2 within the extremes of 39.614461¡ to 39.913401¡ latitude, and -105.109634¡ to -104.602718¡ longitude, bordered by Jefferson, Adams, and Arapahoe Counties (Fig. 1). The elevation ranges from 1567 to 1729 m above sea level. These data, and this study, do 6

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not include Denver Mountain Parks, 22 parks and 24 conservation areas comprising 57 km 2 owned by the City and County of Denver. These Mountain Parks and properties are outside of the city limits at elevations between 1800 m and 4000 m. Not surprisingly, the flora of these areas is extremely varied, not representative of Denver County proper, and as such, not discussed here. Herbarium database mining In the creation of a checklist of all native and naturalized vascular plants reported for Denver, major herbaria worldwide, as well as smaller herbaria in Colorado and surrounding regions were searched for records from Denver (Appendix A). Web-based herbarium databases were first searched with queries for "Denver" in the county field, and then again in the locality field (leaving the county field empty). Herbaria that did not offer online databases with downloadable data (e.g., COCO, COLO) were contacted by email with a request for results from the two queries above. Records resulting from these efforts (Table 1) were compiled into a spreadsheet and specimen label data were reviewed, removing records that were not of native or non-native vascular plants occurring outside of cultivation within the current boundaries of the City and County of Denver, Colorado. As none of these specimens were annotated during this project, I accepted the assumptions that the identification and label data were correct as published to the databases, with a few exceptions (see discussion). Records with duplicate accession numbers were removed, as well as records of specimens that were incompletely identified (e.g., Aster sp.). Records based on cultivated plants, including those transplanted or collected from a greenhouse (e.g., at the Denver 7

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Botanic Gardens) were also excluded. The borders of the City and County of Denver were determined using Google Earth, and specimens collected outside of these limits were removed from the spreadsheet, including those collected from Denver Mountain Parks. Records with notes indicating collection "near Denver" were removed, unless the collection was dated prior to 1900. As some collection years were entered incorrectly into databases (e.g., 1920 changed to 2020), collection dates were corrected if determined to be inaccurate (e.g., future dates, collector not alive at time of collection). Supplemental data sources Records from alternative sources were also investigated during the creation of this checklist. Students at the University of Colorado Denver collected plants in Denver as part of coursework for Flora of Colorado during 2011 and 2012. Daniel Gee, a CU Denver student, processed and confirmed the identification of these specimens, and added a subset of these to the teaching herbarium at the University. Herbarium sheets from the Denver Museum of Nature and Science (DMNS) were digitized by Brooke Sekora, ( CU Denver student) at the Denver Botanic Gardens (KHD). The species vouchered for Denver through these efforts were compared to the checklist generated from herbarium database mining in an effort to reveal any novel species. One species, Berteroa incana (L.) DC. (Brassicaceae) was added to the checklist, vouchered for Denver by one accession: Jones 137 [DMNS]. In 2013, I surveyed aquatic habitats in Denver for pondweeds as part of research for Chapter III of this thesis. Additionally, I made collections of aquatic plants at Parkfield Lake with Denver Botanic Gardens staff members Pam Regensberg and Daniel 8

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Gee in October 2013. These collecting efforts, combined with new records resulting from the Colorado Natural Heritage Program's assessment of the wetlands in Denver County (2013) documented an 33 additional species in Denver (Table 2). Although it is not vouchered due to the small population size, Limnorchis sp. has been observed in Denver and is believed to be extant as of 2013 (Pam Smith, CNHP, personal communication). Checklist creation A species list of vascular plants occurring outside of cultivation in Denver was compiled from existing herbarium records and new records resulting from 2013 surveys. Nomenclature, synonymy, endemic, native status follows Colorado Flora: Eastern Slope (Weber and Wittmann 2012a), using species names. Where subspecific variation is used in Colorado Flora: Eastern Slope it is indicated in the checklist by CFES after the taxon name. When synonymy could not be resolved using Colorado Flora: Eastern Slope (Weber and Wittmann 2012a), I consulted the Catalog of the Colorado Flora: A Biodiversity Baseline (Weber and Wittmann 2000). The Plant List (2010) was used to adjudicate nomenclature, and the format of authorities follows the International Plant Names Index (IPNI 2014). Where synonyms were encountered, they are provided below the species name (following Weber and Wittmann 2012a) using the original nomenclature of the owner institution. The Colorado Natural Heritage Program (CNHP 2013a) tracks species of concern in the state of Colorado. These species are indicated in the checklist with their tracking status, global and state ranks, as well as Federal, BLM or USFS sensitive status, if applicable. 9

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The Colorado Department of Agriculture lists 74 noxious weeds for Colorado, with an additional 26 species on the Noxious Weeds Watch List (CDA 2014). State-listed noxious weed species are indicated on the Denver checklist with their listing rank (List A, List B, List C or Watch List ). Results The checklist of vascular plants growing outside of cultivation in the City and County of Denver contains 758 species in 410 genera and 101 families collected since 1861. Of these 758 species, 247 (33%) are non-native (Fig. 2). Five species of ferns and fern allies have been documented, in five genera and four families. One gymnosperm has been documented: Pinus ponderosa P.Lawson & C.Lawson. Asteraceae, Poaceae, Fabaceae and Brassicaceae are the most species-rich families, each comprising more than five percent of the flora. The largest family represented in the Denver flora is Asteraceae, with 134 species (35 non-native, 11 noxious) in 70 genera, followed by Poaceae, with 121 species (58 non-native, 6 noxious) in 63 genera. Fabaceae is represented by 52 species (13 non-native, 0 noxious) in 24 genera, and Brassicaceae by 40 species (20 nonnative, 3 noxious) in 22 genera (Fig. 3 ). Nineteen species tracked by the Colorado Natural Heritage Program (CNHP 2013a) have been documented in Denver (Table 3). Denver County was once, or currently is habitat for seven species ranked critically imperiled in Colorado (six S1, one S1S2), seven species ranked imperiled (S2), and five species ranked vulnerable (four S3, one S3S4). One species, Astragalus sparsiflorus A.Gray, is ranked globally imperiled (G2). Five species are ranked globally vulnerable (four G3, one G3G4). Two species are 10

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apparently secure (G4?, G4G5) and the remaining 11 species tracked by CNHP are ranked globally secure (G5). One species documented in Denver, Asclepias uncialis Greene subsp. uncialis is considered a sensitive species by the BLM and USDA Forest Service. Seven endemic (or near-endemic) species have been documented in Denver (Table 4). Endemic status was assigned following Colorado Flora, Eastern Slope (Weber and Wittmann 2012a). Two of these seven species are tracked by the Colorado Natural Heritage Program: Ambrosia linearis (Rydb.) W.W.Payne, and Physaria vitulifera Rydb. Only one of these endemic species, Ambrosia linearis (Rydb.) W.W.Payne has been collected in Denver since 1940 Thirty-nine of the 74 species on the Colorado Noxious Weeds List (CDA 2014) have been documented in Denver (Table 5), including two species on List A: Lythrum salicaria L. and Euphorbia myrsinites L., 26 List B species, and 11 List C species. Additionally, four of the 26 species on the Colorado Noxious Weeds Watch List have been documented in Denver. Discussion Biodiversity This checklist of the spontaneously occurring vascular flora of the City and County of Denver was created to update the historic book A Popular Flora of Denver, Colorado (Eastwood 1893), and demonstrate the utility of web-based herbarium databases in the creation of a regional flora. This checklist comprises 758 native and nonnative species representing 410 genera and 101 families, based on over 3000 accessions 11

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collected from 1861 to 2013. Native species (511) comprise 67% of the species collected in Denver over the last 152 years (Fig. 2), including 19 species of concern (3%), and seven endemic species (~1%). Not surprisingly for Colorado, Asteraceae (134 species or 18%) and Poaceae (121 species, 16%) are the most species-rich families in Denver, followed by Fabaceae (52 species, 7%) and Brassicaceae (40 species, 5%) (Fig. 3). These results are based on the most current manual for Colorado (Weber and Wittmann 2012a), which does not always follow the commonly accepted nomenclature used by the PLANTS database (NRCS, USDA 2013) or Flora of North America (FNA). Historic context. Alice Eastwood reported 487 species in A Popular Flora of Denver, Colorado (1893), noting incomplete treatment of Poaceae and Cyperaceae. Eastwood collected extensively in Denver, and many of her specimens voucher species records on the modern checklist. Unfortunately, a direct parallel cannot be drawn between Eastwood's work and the present study because her book was not intended to be an exhaustive species list for Denver, uses outdated nomenclature for which synonymy is difficult to determine, and is not completely vouchered by physical specimens Much of the diversity on this checklist is the result of historic vouchers from over 20 years ago. Of all accessions from Denver, only 32% (948) were collected after 1930 (Fig. 6). Many species are vouchered only by historic accessions. For example, only five of 19 species of concern tracked by the Colorado Natural Heritage Program (CNHP) have been documented in Denver in the last 90 years (Table 3). Acorus calamus L., Callitriche heterophylla Pursh and Sparganium eurycarpum Engelm. were documented by CNHP in 2013. Virgulus novae-angliae (L.) Reveal & Keener was last collected in 1984. Ambrosia 12

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linearis (Rydb.) W.W.Payne was vouchered in 2004, and is also the only endemic species (of seven) documented in Denver since 1940. Introduced species. Introduced species represent 33% (247 of 758) of all species collected in Denver since 1861, including 43 noxious weeds (Fig. 2, Table 5). Poaceae contributes the highest number of non-native species (58, or 23% of non-native species). Thirty-five non-native species are in Asteraceae (14% of non-native species), including 11 noxious weeds (Fig. 3). Eastwood reported 48 introduced species in A Popular Flora of Denver, Colorado (1893) indicating that an additional 199 non-native species have been introduced to Denver in the last 121 years (Table 5). It is likely that a future survey of all habitats in Denver County will result in additional county records of invasive weeds, as they do not appear to be well-documented in herbaria. Especially for floras of urban areas, it seems critical that many vouchers of invasive species be documented in herbaria, (especially those that share data on large consortium websites ), so that these data are available to monitor range expansions and provide material for future research. Limitations of web-based herbarium databases Digital herbaria provide detailed specimen records instantly and efficiently, but the value of these data should not be over-emphasized until evaluated through expert review. Although checklists generated automatically by consortium databases (e.g., SEINet) offer many advantages when compared to traditional methods of compiling floristic data, the checklist of the Denver flora presented here is the result of multiple queries, much manual revision, and supplementation from fieldwork. While using webbased herbarium databases to compile records for this project, I observed three general 13

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types of limitation s: the limited number of records available, lack of standardization among herbaria, and erroneous or missing label data. Dataset size. Intrinsically, the usefulness of any database is limited primarily by the number of records available, due to undercollection, institution participation, and digitization progress. With approximately 32% of records digitized in those herbaria that responded to the 2012 Utah State survey (Barkworth and Murrell 2012), the actual percentage of digitized specimens in the United States is likely much lower. Those records that have been digitized (a relatively small percent of accessions) are freely available for download from institutional and consortia websites. As these datasets are utilized by a wider audience than would be examining physical herbarium specimens (and benefitting from interactions with curatorial staff), it is misleading to present these datasets to the public without communicating their shortcomings. Considering that Denver is in the 90th percentile for herbarium specimen density in the Western United States (Taylor 2014), one would assume that all species in the county would be represented by at least one voucher. This was not the case, as 33 species were added to the checklist as a result of fieldwork, including the 2013 CNHP survey of Denver County wetlands. As Taylor (2014) illustrated, the entire flora of most of the Western United States is severely underdocumented. In geographic areas or taxonomic groups suffering from low collection density, even if all existing herbarium specimens were digitized, databases could not possibly represent biodiversity accurately. It must be made clear that the records available on any given website are only a sample of the actual 14

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collecting effort housed in herbaria nationwide, and no dataset should be interpreted as being complete. Standardization in consortia. The regional consortium database (SEINet) was found to contain records for approximately 78% of databased species records for Denver (75% of total species with new records). Individual herbarium databases contributed 156 species beyond those discoverable on SEINet (Fig. 5), although 142 of these were available on the COLO website. These results suggest that searching two websites (SEINet and COLO) give a relatively accurate representation of the known flora of Denver as 97% of databased species (94% of the species on the checklist after records from 2013 fieldwork were added). Encouraging as this is, consortia inherently contain inconsistencies arising from the lack of standardization among herbaria, threatening the reliability of the data. Generally, specimen records available online do not indicate if a specimen is native, introduced, or cultivated. Dangerous assumptions (e.g., that cultivated species are indicated as such, or not vouchered at all) may lead to an impression of false biodiversity. Even when assumptions are not made, these inconsistencies create the need for the end user to research each species to determine its native status. In the present study, raw data from SEINet returned twice as many cultivated species than native species (Fig. 4), requiring much revision to remove the many records of cultivated species Some herbaria indicate cultivation status in databases, enabling the user to easily remove these records if needed, but in this study, hundreds of records were eliminated on a case-by-case basis by reviewing label data, an extremely time-consuming process. 15

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As not all herbaria use the same resources to identify specimens, it is expected that checklists or downloads from consortium databases include many synonyms. This lack of nomenclatural standardization on consortium websites also leads to inflated species counts. Taxonomic thesauri, as on SEINet, resolve synonymy on automatically generated species lists, but problems still exist. For example, a species list for Denver County generated by SEINet using the taxonomic thesauri included both Chrysothamnus nauseosus (Pallas) Britt. and its synonym Ericameria nauseosa (Pallas ex Pursh) G.L. Nesom & Baird). Also, synonyms Potamogeton pectinatus L. and Stuckenia pectinatus (L.) Bšrner were included in addition to the currently accepted name Stuckenia pectinata (Pers.) Bšrner These two examples alone added three superfluous names to the checklist automatically generated for Denver by SEINet. Issues such as these are expected in the early stages of herbarium digitization, and are easily repaired. More difficult problems arise when duplicate accessions are identified to different species. Several accessions on the Denver checklist were found to be assigned different non-synonymous names when duplicates were deposited at different herbaria. These inconsistencies seemed to arise from the use of different manuals, the retention of outdated, non-synonymous nomenclature, or the annotation of one or more of the duplicate specimens. For example, duplicate accessions collected from the Denver airport in 1939 ( Gierisch 917 ) are identified as Abronia fragrans Nutt. ex Hook. at Colorado State (CS), and as Abronia elliptica A.Nelson at Rocky Mountain Herbarium (RM). Interestingly, this problem can also occur within an herbarium, as illustrated by duplicate accessions from City Park in Denver, from the same collector and year, where one of the 16

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accessions was identified as Oenothera pallida Lindl. var. pallida and the other Oenothera albicaulis Pursh ( Gierisch 902 [RM]). In both of these cases, I included only one of the species names ( Oenothera albicaulis and Abronia fragrans ) after Abronia elliptica and Oenothera pallida subsp. pallida were determined to be species only found on the Western Slope of Colorado. These chance observations are examples of records that were relatively easy to recognize because it did not seem likely the species occur in Denver. It is probable that additional species on this checklist are inappropriately vouchered in this way, especially with older records that frequently do not include collection numbers or detailed location descriptions. Missing or erroneous label data. The usefulness of data from web-based herbarium databases was also limited by errors or missing information in the records themselves. The most common error encountered in these electronic databases was a missing or incorrect county. These inaccuracies resulted in many records being erroneously attributed to Denver County, requiring line-by-line evaluation of raw data from databases to remove records that were obviously not collected from Denver County. Some of these errors were obvious (e.g., locality: "Steamboat Springs," "48 miles W of Denver," "Mountains near Denver," or "Along the Cog Wheel Railway to Pike's Peak") but other were more subtle (e.g., locality: "Daniels Park" [Denver Mountain Park], or "Federal Center" [in Jefferson County]). I was able to recognize these erroneous records because of my familiarity with Colorado geography, but these records would have been indistinguishable to out-of-state users of these databases. 17

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Conversely, queries for Denver as a locality returned an additional 960 records from Denver County, adding 85 species to the checklist (Fig. 6). The majority of these records (855) had an unknown county in databases, while 95 had an outdated or incorrect county. With few exceptions, all records that were not attributed to Denver County in herbarium databases were collected prior to 1930, possibly due to the changes in political boundaries early in the 19th century. Many of Alice Eastwood's collections from Denver County were excluded from earlier drafts of my checklist because the county was either recorded as Arapahoe County (Denver was the seat of Arapahoe County until 1902) or not listed at all. Searching for "Denver" in the locality field without a county uncovered these records. I suspect additional records may be retrieved by conducting similar searches for specific locations in Denver, such as "Berkeley" or "Valverde." Although not a major issue, I encountered many records with an incorrect year in databases. This appeared to be due to the automatic formatting of dates in the 1800's into those in the 1900's or those in the 1900's into the 2000's (e.g., 1878 became 1978, and 1917 became 2017). Results of research, especially that concerning climate change, historic distributions, or conservation are dramatically impacted by dates that are off by 100 years. These mistakes are only caught and corrected by those familiar with the collector or with some other indicator in the record, such as the description of a historic location. Although likely specific to Colorado, I also encountered the problem of "Co." (county) being interpreted as "CO" (Colorado) during transcription of several herbarium labels, which if left unnoticed, could attribute erroneous records to Colorado from other states. 18

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Summary. Denver presents an excellent subject for this study because its flora was documented early in the city's development when native plant communities were still intact. The flora continued to be well-collected throughout the early 1900s, then after a 50 year period with relatively few collections, approximately 800 specimens have been collected from 1970 to present today (Fig. 6), resulting in a specimen density of 7.47 specimens per km 2 very high compared to other counties in the Western United States (Taylor 2014). Even with this high density of collections, numerous problems arose during the creation of this checklist. Hundreds of hours were spent on data quality control. Records based on cultivated plants were removed, as were many records erroneously attributed to Denver. Synonymy and nomenclature presented their own suite of challenges, as taxonomic thesauri on SEINet were not 100% effective, and could not recognize the duplicate specimens that were identified differentially when sent to multiple herbaria. The creation of a regional floristic checklist using web-based herbarium databases can be very useful for some applications, such as understanding the comprehensive floristic history of an area. However, checklists created using this method do not represent the current state of biodiversity, especially for urban areas. Dolan et al. (2011), in their study of the effects of urbanization on the flora of Marion County, Indiana, observed an average decrease of 2.4 native species per year, and an average increase of 1.4 non-native plant species per year. The recently documented flora of Marion County (through surveys completed 1996-2009) has a species richness similar to Denver County (about 700 species), of which 27% (189) are introduced, an increase of 7% compared to 19

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estimates of historic diversity (Dolan et al. 2011). Although Denver is considered wellcollected overall (Taylor 2014), it is not possible to conduct a study similar to that done in Marion County until Denver's contemporary flora is documented. Currently, only 984 accessions have been made since 1930, averaging 11 per year (Fig. 6). Considering that 199 non-native species have been documented since Alice Eastwood's book (1893), a checklist compiled for Denver solely using recent collections and fieldwork would be expected to comprise fewer species overall, with a significantly higher proportion of introduced species, and fewer species of concern. Urban floras have been prepared for cities in Northeast, Midwest and Pacific West regions of the United States (La Sorte et al. 2014) but none have documented the flora of a city in the arid Western United States. Any comparison to these existing floras would be misleading, as Denver's climate and ecology is strongly dissimilar to these more mesic locations. For example, the flora for Philadelphia lists 2528 species (La Sorte et al. 2014). Additionally, there is incredible variation in study design, surveying effort, and nomenclature used. Clearly, more work remains to be done documenting the flora of the Western United States, including urban areas. This checklist compiled for Denver is intended to provide historical context for future research, and should supplement, but not replace, efforts to document the contemporary flora of Denver. ! 20

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Tables and Figures Figure II.1. Location and political boundaries of the City and County of Denver, Colorado. The study area did not include Denver Mountain Parks. ! 21 N 0 25 50 100 km Data source: ESRI

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Table II.1. Herbari a providing records of vascular plants occurring outside of cultivation in the City and County of Denver, Colorado. Herbarium a cronym Herbarium name Source ARIZ University of Arizona SEINet COLO University of Colorado Museum of Natural History Tim Hogan (email) CS Colorado State University SEINet DMNS Denver Museum of Nature and Science DMNS physical specimens GH Gray Herbarium, Harvard University GH website ID Stillinger Herbarium, University of Idaho CPNWH KHD Kathryn Kalmbach Herbarium, Denver Botanic Gardens SEINet KS C Kansas State University Herbarium KSC Online Herbarium MO Missouri Botanical Garden TROPICOS website NMC New Mexico State University SEINet NY New York Botanical Garden SEINet RM Rocky Mountain Herbarium, University of Wyoming SEINet SJNM San Juan College SEINet UMO University of Missouri TROPICOS website US United States National Herbarium, Smithsonian Institution NMNH website UTC Intermountain Herbarium, Utah State University SEINet WS Marion Ownbey Herbarium, Washington State University CPNWH WSC Western State College Herbarium SEINet WTU University of Washington CPNWH WWB Western Washington University CPNWH 22

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Table II.2. New species records documented for the City and County of Denver in 2013. Collections made during the Colorado Natural Heritage Program's assessment of wetlands in Denver County and fieldwork performed by Majack, Regensberg (KHD) and Gee (UCD/KHD). Nomenclature follows Weber and Wittmann (2012a) ! 23 Family Species Collectors [Herbarium] Acoraceae Acorus calamus L. Kuhn and Smith (CHNP) [ CS] Brassicaceae Neolepia campestris (L.) W.A.Weber Kuhn and Smith (CHNP) [ CS] Callitrichaceae Callitriche heterophylla Pursh Kuhn and Smith (CHNP) [ CS] Ceratophyllaceae Ceratophyllum demersum L. Regensberg, Majack, Gee [KHD] Chenopodiaceae Bassia hyssopifolia (Pall.) Kuntz Kuhn and Smith (CHNP) [ CS] Chenopodium simplex (Torr.) Raf. Kuhn and Smith (CHNP) [ CS] Cyperaceae Carex pellita Muhl. ex Willd. Kuhn and Smith (CHNP) [ CS] Carex siccata Dewey Kuhn and Smith (CHNP) [ CS] Carex utriculata Boott Kuhn and Smith (CHNP) [ CS] Cyperus odoratus L. Kuhn and Smith (CHNP) [ CS] Eleocharis macrostachya Britton Kuhn and Smith (CHNP) [ CS] Eleocharis rostellata (Torr.) Torr. Kuhn and Smith (CHNP) [ CS] Haloragaceae Myriophyllum spicatum L. Kuhn and Smith (CHNP) [ CS] Hydrocharitaceae Elodea canadensis Michx. Kuhn and Smith (CHNP) [ CS] Elodea bifoliata H.St.John Regensberg, Majack, Gee [KHD] Lamiaceae Lycopus uniflorus Michx Kuhn and Smith (CHNP) [ CS] Lemnaceae Lemna minuta Kunth Kuhn and Smith (CHNP) [ CS] Wolffia columbiana H.Karst. Kuhn and Smith (CHNP) [ CS] Regensberg, Majack, Gee [KHD] Najadaceae Najas guadalupensis (Spreng.) Magnus Kuhn and Smith (CHNP) [ CS] Oleaceae Forestiera pubescens Nutt. Kuhn and Smith (CHNP) [ CS] Onagraceae Epilobium ciliatum Raf. Kuhn and Smith (CHNP) [ CS] Poaceae Eragrostis lutescens Scribn. Kuhn and Smith (CHNP) [ CS] Eragrostis spectabilis (Pursh) Steud. Kuhn and Smith (CHNP) [ CS] Glyceria striata (Lam.) Hitchc. Kuhn and Smith (CHNP) [ CS] Leymus triticoides (Buckley) Pilg. Kuhn and Smith (CHNP) [ CS] Polygonaceae Persicaria amphibia (L.) Gray Kuhn and Smith (CHNP) [ CS] Polygonaceae Rumex utahensis Rech. Kuhn and Smith (CHNP) [ CS] Potamogetonaceae Potamogeton crispus L. Majack [COLO, KHD] Potamogeton nodosus Poir. Kuhn and Smith (CHNP) [ CS] Majack, Regensberg, Gee [KHD] Potamogeton pusillus L. Kuhn and Smith (CHNP) [ CS] Majack [KHD] Rhamnaceae Rhamnus frangula L. Kuhn and Smith (CHNP) [ CS] Salicaceae Salix famelica (C.R.Ball) Argus Kuhn and Smith (CHNP) [ CS] Scrophulariaceae Linaria vulgaris P.Mill. Majack [KHD]

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Figure II.2. Proportions of native and non-native vascular plant species documented in the City and County of Denver, Colorado. A total of 758 species occurring outside of cultivation have been documented in Denver between 1861 and 2013, representing 410 genera and 101 families. Figure II.3. The most species-rich vascular plant families in the City and County of Denver, Colorado. Asteraceae, Poaceae, Fabaceae, and Brassicaceae each represent more than 5% of spontaneously occurring species collected in Denver between 1886 and 2013. Poaceae contributes the most non-native species (58), although Asteraceae contributes the greatest number of noxious weeds (11). ! 24

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Table II.3. Species of concern tracked by the Colorado Natural Heritage Program (CNHP) documented in the City and County of Denver, Colorado between 1886 and 2013. Global and subnational (Colorado) conservation ranks follow NatureServe (2014). Family Species Most recent collection CNHP status NatureServe r ank Acoraceae Acorus calamus L. 2013 Fully tracked G4? S1 Alismataceae Sagittaria montevidensis Cham. & Schltdl. subsp. calycina (Engelm.) Bogin 1887 Fully tracked G5T5? S1 Asclepiadaceae Asclepias uncialis Greene subsp. uncialis 1895 Fully tracked G3G4T2T3 S2 BLM USFS Asclepias stenophylla A.Gray 1871 Fully tracked G4G5 S2 Asteraceae Ambrosia linearis (Rydb.) W.W.Payne 2004 Fully tracked G3 S3 Virgulus novae-angliae (L.) Reveal & Keener 1984 Fully tracked G5 S1 Ošnopsis engelmannii (A.Gray) Greene 1924 Fully tracked G3 S3? Liatris ligulistylis (A.Nelson) K.Schum. 1873 Fully tracked G5? S1S2 Boraginaceae Oreocarya cana A.Nelson 1895 Fully tracked G5 S2 Brassicaceae Physaria vitulifera Rydb. 1901 Fully tracked G3 S3 Callitrichaceae Callitriche heterophylla Pursh 2013 Fully tracked G5 S1 Elatinaceae Elatine triandra Schkuhr 1914 Fully tracked G5 S1 Elatine rubella Rydb. 1910 Fully tracked G5 S2 Fabaceae Astragalus sparsiflorus A.Gray 1895 Fully tracked G2 S2 Apios americana Medik. 1886 Fully tracked G5 S1 Gentianaceae Eustoma grandiflorum (Raf.) Shinners 1887 Watchlisted G5 S3S4 Grossulariaceae Ribes americanum Mill. 1911 Fully tracked G5 S2 Loasaceae Nuttallia speciosa (Osterh.) Greene 1916 Fully tracked G3 S3 Sparganiaceae Sparganium eurycarpum Engelm. 2013 Fully tracked G5 S2 25

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Table II.4. Endemic and near-endemic species documented in the City and County of Denver, Colorado between 1886 and 2013. Endemic status and nomenclature follows Weber and Wittmann (2012a). State distributions follow BONAP maps (Kartesz 2014). Family Species Most recent collection CNHP tracking status NatureServe rank State distribution Asteraceae Ambrosia linearis (Rydb.) W.W.Payne 2004 Fully tracked G3 S3 CO Grindelia subalpina Greene 1903 G4 S3 CO, NM, WY Helianthus pumilus Nutt. 1916 G4 SNR CO, ID, MT, UT, WY Boraginaceae Oreocarya virgata (Porter) Greene 1886 G4 SNR CO, UT Brassicaceae Physaria vitulifera Rydb. 1901 Fully tracked G3 S3 CO Fabaceae Astragalus parryi A.Gray 1916 G4 S4 CO, UT Scrophulariaceae Penstemon virens Pennell 1939 G3G4 SNR CO, WY 26

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Table II.5. Noxious weeds listed by the Colorado Department of Agriculture (CDA) documented in the City and County of Denver between 1861 and 2013. Nomenclature follows Weber and Wittmann 2012a. 27 CDA List Family Species First Denver record List A Euphorbiaceae Euphorbia myrsinites L. 1988 Lythraceae Lythrum salicaria L. 1978 List B Asteraceae Acosta diffusa (Lam.) Soj‡ k 2004 Acroptilon repens (L.) DC. 2004 Anthemis cotula L. 1891 Breea arvensis (L.) Less. 1978 Carduus nutans L. 2004 Cirsium vulgare (Savi) Ten. 1913 Onopordum acanthium L. 2004 Tanacetum vulgare L. ca. 191 6 Tripleurospermum inodorum (L.) Sch.Bip. 1968 Boraginaceae Cynoglossum officinale L. 2004 Brassicaceae Cardaria draba (L.) Desv. 1955 Cardaria latifolia (L.) Spach 2004 Caryophyllaceae Saponaria officinalis L. 1916 Cyperaceae Cyperus esculentus L. 1997 Dipsacaceae Dipsacus laciniatus L. 2004 Elaeagnaceae Elaeagnus angustifolia L. 1914 Euphorbiaceae Tithymalus esula Hill 2004 Haloragaceae Myriophyllum spicatum L. 2013 Malvaceae Hibiscus trionum L. 1982 Poaceae Cylindropyrum cylindricum (Host) .Lšve 2004 Elytrigia repens (L.) Nevski 1916 Ranunculaceae Clematis orientalis L. 2007 Scrophulariaceae Linaria vulgaris L. 2012 Verbascum blattaria L. 2011 Solanaceae Hyoscyamus niger L. 1984 Tamaricaceae Tamarix chinensis Lour. 2009 List C Apiaceae Conium maculatum L. 2004 Asteraceae Arctium minus (Hill) Bernh. ca. 1900 Cichorium intybus L. 1891 Convolvulaceae Convolvulus arvensis L. 1902 Geraniaceae Erodium cicutarium (L.) L'Her. 1934 Malvaceae Abutilon theophrasti Medik. 1982 Poaceae Anisantha tectorum (L.) Nevski 1924 Panicum miliaceum L. 1978 Sorghum halepense (L.) Pers. 1986 Scrophulariaceae Verbascum thapsus L. 2004 Zygophyllaceae Tribulus terrestris L. 1910 W atch List Boraginaceae Anchusa officinalis L. 1914 Brassicaceae Alliaria petiolata (M.Bieb.) Cavara & Grande 2004 Onagraceae Epilobium hirsutum L. 2009 Poaceae Phragmites australis (Cav.) Trin. 1954

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Figure II.4. Proportion of cultivated species included on a checklist automatically generated by SEINet for Denver County, Colorado (5 November 2014). Synonymy was resolved using the APG3 taxonomic filter on SEINet. The number of native and nonnative spontaneously occurring species illustrated here (758) includes records from sources beyond SEINet (see Fig. 5). Figure II.5. Data sources used to voucher the checklist of vascular plants growing outside of cultivation in the City and County of Denver, Colorado. Records were collected between 1861 and 2013. 28

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Figure II.6. Temporal distribution of databased records (1861-2013) of vascular plant species growing outside of cultivation in the City and County of Denver, Colorado. Light blue bars represent the number of records databased with an unknown (855 records) or incorrect (95 records) county. 29

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Checklist of the Vascular Flora of the City and County of Denver, Colorado The following checklist of vascular plant species spontaneously occurring in the City and County of Denver, Colorado (excluding Denver Mountain Parks) was compiled using available existing herbarium records (1861-2013) supplemented with new records resulting from fieldwork (2013-2014) by the Colorado Natural Heritage Program (CNHP), the Denver Botanic Gardens (KHD) and University of Colorado Denver students (UCD). Nomenclature follows Colorado Flora: Eastern Slope (Weber and Wittmann 2012a). Accession numbers of voucher specimens (which were not annotated by the author) are listed below the species name used by the owner institution. Herbarium codes follow Index Herbariorum (Theirs 2014). Species not native to Colorado (Weber and Wittmann 2012a)   State-listed noxious weed (CDA 2014) Species of concern tracked by CNHP (2013a) listed with NatureServe conservation rank (2014) # Endemic or near-endemic (Weber and Wittmann 2012a) listed with state distribution (Kartesz 2014) CFES 2012 Indicates subspecies listed in Weber and Wittmann (2012a) Pteridophytes Athyriaceae Gymnocarpium Gymnocarpium dryopteris (L.) Newman (RM 105349) Dryopteridaceae Dryopteris Dryopteris filix-mas (L.) Schott (KHD 419, 5261) Hypolepidaceae Pteridium Pteridium aquilinum (L.) Kuhn Pteridium aquilinum (L.) Kuhn subsp. lanuginosum (Bong.) HultŽ n (KHD 5512) Equisetaceae Equisetum Equisetum arvense L. (KHD 37834) Hippochaete Hippochaete laevigata (A.Braun) Farw. (COLO 47) Gymnosperms Pinaceae Pinus Pinus ponderosa P.Lawson & C.Lawson (US 3615904) 30

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Angiosperms Aceraceae Acer Acer glabrum Torr. (COLO 1323) Negundo Negundo aceroides Moench (COLO 1365, 1380) Negundo aceroides Moench subsp. interius (Britton & Shafer) .Lšve & D.Lšve CFES 2012 (RM 113480) Acer negundo L. (KHD 5589, 5606, UTC 951, 1297) Acoraceae Acorus *Acorus calamus L. [CNHP tracked: G4? S1] (CS: CNHP D-2013-01, -02 ) Agavaceae Yucca Yucca glauca Nutt. (COLO 27273, 27277, 27282, KHD 39531, MO 148459, RM 97111) Alismataceae Alisma Alisma triviale Pursh (COLO 1439, 1443, KHD 5816, RM 226816) Sagittaria Sagittaria cuneata E.Sheld. (MO 86916, RM 89124) Sagittaria latifolia Willd. (KHD 5255, 7860, 26153) Sagittaria montevidensis Cham. & Schltdl. [ CHNP tracked: G5T5? S1] Sagittaria montevidensis Cham. & Schtdl. subsp. calycina (Engelm.) Bogin CFES 2012 (COLO 1480) Alliaceae Allium Allium geyeri S.Watson (UTC 1626) Allium textile A Nelson & J.F.Macbr. (COLO 27519, 27528, 27664, CS 5219, RM 87994) Alsinaceae Alsine *Alsine media L. (COLO 4674, 4725, 4726) Stellaria media (L) Vill. (KHD 9591, 13174, 22593) Cerastium *Cerastium fontanum Baumg. (COLO 4323) Spergularia *Spergularia media (L.) C.Presl (COLO 539842, KHD 46923) Stellaria Stellaria graminea L. (WSC 7054) 31

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Amaranthaceae Amaranthus *Amaranthus albus L. (COLO 1509, 1512, 1513, 1514, RM 225499) Amaranthus arenicola I.M.Johnst. (COLO 20134, 20135, 106059, 404103, KHD 6690, 7953, 7954) Amaranthus blitoides S.Watson (COLO 1505, 1524, KHD 7962, 43922) Amaranthus hybridus L. (KHD 31012) Amaranthus powellii S.Watson (KHD 22749, 39820) Amaranthus retroflexus L. (COLO 1519, 1521, 1522, 1527, 170919, 170919, KHD 7970, 7971, 7972, 7974, 20289) Froelichia Froelichia gracilis (Hook.) Moq. (COLO 1549, 1551, 1553, 14757, 106060, CS 26091, KHD 43892, RM 87784) Anacardiaceae Rhus Rhus aromatica Aiton Rhus aromatica Aiton subsp. trilobata (Nutt.) W.A.Weber (COLO 1681, 103065, 518070, KHD 5553, 7330, 13783) Toxicodendron Toxicodendron rydbergii (Small ex Rydb.) Greene (CS 13348, KHD 8183) Apiaceae Berula *Berula erecta (Huds.) Coville (COLO 101440, KHD 15658, 46914) Cicuta Cicuta maculata L. (KHD 43923) Cicuta douglasii (DC.) J.M.Coult. & Rose (COLO 41375, 41382, 41963, KHD 1058) Conium  * Conium maculatum L. [Noxious List C] (COLO 518128, KHD 39489, 39490) Cymopterus Cymopterus acaulis (Pursh) Raf. (COLO 41443, 41452, 41453, CS 15259) Cymopterus montanus Torr. & A.Gray (COLO 41455, 41799, 41800, CS 15278, 15280) Heracleum Heracleum sphondylium L. Heracleum sphondylium L. subsp. montanum (Schleich.) Briq. CFES 2012 (COLO 41532, 41543) Lomatium Lomatium orientale J.M.Coult. & Rose (COLO 41647, 41653, 41657, 41658, CS 15355) Musineon Musineon divaricatum (Pursh) Raf. (COLO 41684, 41685, 41692, CS 15049, 15051, 25188, KHD 30694, RM 88112, UTC 12752) Osmorhiza Osmorhiza longistylis (Torr.) DC. (KHD 29555, 46739) 32

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Apocynaceae Apocynum *Apocynum cannabinum L. (COLO 1758, 1779, KHD 1168, 8224, 47347) Apocynum floribundum Greene (CS 16338 ) Asclepiadaceae Asclepias Asclepias engelmanniana Woodson (COLO 1924, 1926, 2046, CS 16384, KHD 8351, 8352) Asclepias incarnata L. (CS 33643, RM 8435, 45489) Asclepias pumila (A.Gray) Vail (COLO 1958, 1965, 1967, CS 16417, 16418, KHD 8705, 40085, 47308, RM 49160, 88962, UTC 4986) Asclepias speciosa Torr. (COLO 2000, 518131, KHD 8360, 23754, MO 5661677, RM 88966) Asclepias stenophylla A.Gray [ CHNP tracked: G4G5 S2] (COLO 104122) Asclepias uncialis Greene [ CHNP tracked: G3G4T2T3 S2 BLM USFS] (COLO 1907, 14746, MO 2761608 ) Asclepias viridiflora Raf. (CS 16399) Asparagaceae Asparagus *Asparagus officinalis L. (COLO 26641, 26642, CS 26051, 60711, KHD 7145) Asteraceae Achillea *Achillea millefolium L. (KHD 7875) Achillea lanulosa Nutt. (COLO 5390, 5391) Acosta  * Acosta diffusa (Lam.) Soj‡k [CO Noxious List B] (COLO 518138) Centaurea diffusa Lam. (KHD 46509) Acroptilon  * Acroptilon repens (L.) DC. [CO Noxious List B] (COLO 518127, 530324, KHD 39494, 46453) Agoseris Agoseris glauca (Pursh) Raf. (KHD 5682) Ambrosia Ambrosia acanthicarpa Hook. (COLO 9248, 9263, 9273, KHD 10294, 10295, 16563, 43275) Ambrosia artemisiifolia L. (KHD 5711, 5712, 10298) Ambrosia confertiflora DC. (KSC 72686) # Ambrosia linearis (Rydb.) W.W.Payne [Endemic CNHP tracked: G3 S3] (COLO 522247, KHD 39732) Ambrosia psilostachya DC. (KHD 452, 5708, 5709, RM 97755) Ambrosia psilostachya var coronopifolia (Torr. & A.Gray) Farw. (COLO 5648, 5663, 5667, 5668, 5675, 103025, 103036, 518148, KHD 39591) Ambrosia tomentosa Nutt. (COLO 9267, 9268, 559181, KHD 10297) Ambrosia trifida L. (COLO 5680, 5682, 5685, KHD 5717, UTC 162079, 174624) 33

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Anthemis   *Anthemis cotula L. [CO Noxious List B] (COLO 6068, 6072) Arctium   *Arctium minus (Hill) Bernh. [CO Noxious List C] (COLO 6092, KHD 47346) Arctium tomentosum Mill. (COLO 6090) Artemisia *Artemisia biennis Willd. (COLO 6705, 518136, KHD 39818, UTC 1627) Artemisia carruthii Alph.Wood ex Carruth. (COLO 6680, 425953) Artemisia frigida Willd. (COLO 6472, 6474, 6497, 6704, KHD 6984, KSC 82185) Artemisia ludoviciana Nutt. (COLO 518139, KHD 16123, 17057, UTC 1896) Artemisia ludoviciana Nutt. subsp. incompta (Nutt.) D.D.Keck CFES 2012 (COLO 6558, 6565, 6571) Artemisia ludoviciana Nutt. subsp. ludoviciana CFES 2012 (COLO 6339, 6549, 6555, 6557, 6617) Artemisia ludoviciana Nutt. subsp. typica (Nutt.) D.D.Keck (UTC 1974) Bahia Bahia dissecta (A.Gray) Britton Amauriopsis dissecta (A.Gray) Rydb. (RM 95419, 160975) Bidens *Bidens cernua L. (COLO 7445, 7449, CS 33267, KHD 9641, 9645, 20236, RM 105344) Bidens frondosa L. (COLO 7421, 7430, 7431, KHD 9642, 9643, 9644, 24413) Bidens tenuisecta A.Gray (COLO 7474) Brachyactis Brachyactis ciliata Ledeb. Brachyactis ciliata Ledeb. subsp. angusta (Lindley) A.G.Jones CFES 2012 (COLO 6734, 6735, 6736) Symphyotrichum ciliatum (Ledeb.) G.L.Nesom (KHD 8174) Brachyactis frondosa (Nutt.) A.Gray Symphyotrichum frondosum (Nutt.) G.L.Nesom (CS 20747, KHD 8632) Breea   *Breea arvensis (L.) Less. [CO Noxious List B] (COLO 559184) Cirsium arvense (L.) Scop. (KHD 443, 40222) Brickellia Brickellia eupatorioides (L.) Shinners (COLO 10298, 10315, KHD 39816) Brickellia eupatorioides var. chlorolepis (Wooton & Standl.) B.L.Turner ( CS 33257) Brickellia rosmarinifolia subsp. chlorolepis (Wooton & Standl.) W.A.Weber (COLO 10299, 10322) Carduus   *Carduus nutans L. [CO Noxious Weed List B] Carduus nutans L. subsp. macrolepis (Peterm.) Kazmi (KHD 40239) 34

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Chrysothamnus Chrysothamnus nauseosus (Pall. ex Pursh) Britton Chrysothamnus nauseosus (Pall. ex Pursh) Britton subsp. graveolens (Nutt.) Piper CFES 2012 (COLO 7517, 7520, 7526, 518144, KHD 8060, 39596) Ericameria nauseosa var. graveolens (Nutt.) Reveal & Schuyler (KHD 9808, 9809, 9810, RM 57442, 89126) Ericameria nauseosa var. nauseosa (Pall. ex Pursh) G.L.Nesom & G.I.Baird (KHD 9825, 40110, 46514, RM 87237, 89138) Chrysothamnus viscidiflorus (Hook.) Nutt. Chrysothamnus viscidiflorus (Hook.) Nutt. subsp. lanceolatus (Nutt.) H.M.Hall & Clem. CFES 2012 (COLO 101017) Cichorium   *Cichorium intybus L. [CO Noxious Weed List C] (COLO 8068, 518108, KHD 8709, 47344) Cirsium Cirsium canescens Nutt. (KHD 40111) Cirsium flodmanii ( Rydb.) Arthur (COLO 8178, 8181) Cirsium parryi (A.Gray) Petr. (COLO 8149, 8151)  * Cirsium vulgare (Savi) Ten. [CO Noxious Weed List B] (KHD 21199, RM 169268) Conyza Conyza canadensis (L.) Cronquist (COLO 8471, 8476, 8490, 8491, 518085, KHD 451, 9880, 39576, UTC 5645 ) Conyza ramosissima Cronquist (COLO 106030) Coreopsis Coreopsis tinctoria Nutt. (COLO 8238, KHD 9898) Cyclachaena Cyclachaena xanthiifolia (Nutt.) Fresen. (COLO 10285, 10286, 10289, 106065, 518143, KHD 16944, 16946, RM 49223, 97079) Iva xanthifolia Nutt. (CS 32018) Dyssodia Dyssodia aurea (A.Gray) A.Nelson Thymophylla aurea (A.Gray) Greene (COLO 8377) Dyssodia papposa (Vent.) Hitchc. (COLO 8381, 8389, 8390, 8396) Erigeron Erigeron bellidiastrum Nutt. (CS32233, KHD 43891) Erigeron canus A.Gray (COLO 8887) Erigeron compositus Pursh (COLO 8529) Erigeron divergens Torr. & A.Gray (COLO 8461, 518133, KHD 479, 9965, 87419) Erigeron flagellaris A.Gray (COLO 8699) Erigeron pumilus Nutt. (COLO 8888, 101579, 518092, 518230, KHD 4796, 6383, 6385, 6404, 10060, 10073, RM 92936, 163702) Erigeron concinnus (Hook. & Arn.) Torr. & A.Gray (UTC 13270) Euthamia Euthamia graminifolia (L.) Nutt. (COLO 11763) Euthamia occidentalis Nutt. (COLO 11761, 11764, 518231, CS 38356, KHD 17604, 39815, 40527, 47343, RM 158428) 35

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Galinsoga *Galinsoga parviflora Cav. (COLO 559076, KHD 439, 10345, 24778, 32673, 37940, MO 11554) *Galinsoga quadriradiata Ruiz & Pav. (KHD 24775) Gnaphalium Gnaphalium uliginosum L. (COLO 9403, KHD 10357, 10359) Grindelia Grindelia hirsutula Hook. & Arn. (RM 88047) Grindelia inornata Greene (COLO 9440, 9445, 9475, 9488, CS 30968) Grindelia squarrosa (Pursh) Dunal (COLO 9471, 9491, 9498, 106049, KHD 440, 10221, 10383, 10384, 39587, 46502, KSC 94568, RM 43855, 88049, 92979) !Grindelia subalpina Greene [Endemic ] (CS 30965) Gutierrezia Gutierrezia sarothrae (Pursh) Britton & Rusby (COLO 103033, 518184, KHD 47342, RM 87228) Helenium Helenium autumnale L. (UTC 9331) Helianthus Helianthus annuus L. (COLO 9758, 9759, 9768, 103022, 518115, CS 31186, KHD 16764, 39830, 47341) Helianthus maximiliani Schrad. (CS 31171, MO 5713184) Helianthus nuttallii Torr. & A.Gray (COLO 9801, 9807, 9814, RM 89130) Helianthus petiolaris Nutt. (COLO 9819, 9820, 9821, 9828, 103021, 518150, CS 21733, 89729, KHD 16775, 16776, 16779, 16780, 39588, 47340, RM 97771) # Helianthus pumilus Nutt. [Endemic ] (COLO 9853, 9854, 9856, 9860) Heterotheca Heterotheca canescens (DC.) Shinners (COLO 7833) Heterotheca villosa ( Pursh) Shinners (COLO 106035, 518149, KSC 94942 ) Heterotheca villosa (Pursh) Shinners var. nana (A.Gray) Semple (CS 30979) Heterotheca villosa (Pursh) Shinners var. villosa (RM 221464) Heterotheca foliosa (Nutt.) Shinners (KHD 40242, RM 87230, 87231) Heterotheca horrida (Rydb.) V.L.Harms (RM 83086, 87229, 92938, 171858, 221475, 221493) Hymenopappus Hymenopappus filifolius Hook. Hymenopappus filifolius Hook. var. cinereus (Rydb.) I.M.Johnston CFES 2012 (COLO 7782, RM 87275) Iva Iva axillaris Pursh (COLO 10267, 10270, 10271, RM 16936, 16937, 16938, 16942) Lactuca Lactuca ludoviciana (Nutt.) Riddell (RM 49486) Lactuca serriola L. (COLO 10329, 10341, 10380, 10394, CS 33147, KHD 16969, 16979, 22661, 40088) Lactuca tatarica (L.) C.A.Mey. Lactuca tatarica (L.) C.A.Mey. subsp. pulchella (Pursh) Stebbins CFES 2012 (COLO 10362, 10366, 10376, 10377) 36

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Lapsana *Lapsana communis L. (KHD 16989, RM 105350) Lepidotheca Lepidotheca suaveolens (Pursh) Nutt. Matricaria matricarioides (Less.) Porter (KHD 17081) Liatris Liatris ligulistylis (A.Nelson) K.Schum. [CNHP tracked: G5 S1S2] (COLO 100958 ) Liatris punctata Hook. (COLO 10490, 10490, 518185, KHD 13842, 37147, 39827, KSC 98269, 98273, RM 37501, 84645, 92910) Liatris punctata Hook. f. coloradensis Gaiser (GH 9775, 9783) Lygodesmia Lygodesmia juncea (Pursh) D.Don (COLO 10546, 10555, 10560, 10561, CS 31685, 45234, KHD 8719, 37146, 40240, RM 87771, 88952) Machaeranthera Machaeranthera annua (Rydb.) Shinners Rayjacksonia annua (Rydb.) R.L.Hartm. & M.A.Lane (COLO 9577, 9578, 9580, 9665) Machaeranthera bigelovii (A.Gray) Greene Dieteria bigelovii (A.Gray) D.R.Morgan & R.L.Hartm. (KHD 9242, 9243, 9244) Machaeranthera canescens (Pursh) A.Gray (COLO 6820, KHD 10005) Machaeranthera canescens (Pursh) A.Gray var. canescens (MO 5713185) Machaeranthera canescens (Pursh) A.Gray var. glabra A.Gray (CS 20849) Dieteria canescens (Pursh) Nutt. (KHD 46523) Dieteria canescens var. glabra (A.Gray) D.R.Morgan & R.L.Hartm. (RM 84417, 93050, 160033) Machaeranthera pinnatifida ( Hook.) Shinners (COLO 9667, 9674, 9676, 9679, 518141, KHD 16084, 16627, UTC 10379) Xanthisma spinulosum var. spinulosum (Pursh) D.R.Morgan & R.L.Hartm. (RM 49215, 92959) Machaeranthera tanacetifolia (Kunth) Nees (COLO 6837, 6838, 7211, 7225, 7226, KHD 9245) Nothocalais Nothocalais cuspidata (Pursh) Greene (COLO 5583, 100975, CS 23072, RM 29225) Oligoneuron Oligoneuron rigidum (L.) Small (COLO 11814) Solidago rigida var. humilis Porter (RM 244309) Ogliosporous Oligosporus dracunculus (L.) Poljakov Oligosporus dracunculus (L.) Poljakov subsp. dracunculinus (S.Watson) W.A.Weber CFES 2012 (COLO 6374, 6375, 6396) Oligosporus dracunculus (L.) Poljakov subsp. glaucus (Pallas) .Lšve & D.Lšve CFES 2012 (COLO 6290) Artemisia dracunculus L. (KHD 6974, RM 93043) Oligosporus filifolius (Torr.) Poljakov (COLO 6403, 6404, 6407, 6408, 6410, 104163) Artemisia filifolia Torr. (KHD 6982, RM 49221, 87280, 105347, UTC 1680, 1731, 1825) Oligosporus pacificus (Nutt.) Poljakov (COLO 6343) Artemisia campestris L. (KHD 47345) 37

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Onopordum  * Onopordum acanthium L. [CO Noxious List B] (COLO 518084, KHD 562, 44403 ) Ošnopsis Ošnopsis engelmannii (A.Gray) Greene [CNHP tracked: G3 S3?] (MO 1809826, RM 87239, 169698, UTC 13270) Packera Packera fendleri (A.Gray) W.A.Weber & .Lš ve (KHD 3588, 3591, 3596, 17699) Packera neomexicana (A.Gray) W.A.Weber & .Lšve (KHD 37240) Packera plattensis (Nutt.) W.A.Weber & .Lš ve (COLO 539974, KHD 926, 37946, RM 274794) Senecio plattensis Nutt. (MO 880546) Packera tridenticulata (Rydb.) W.A.Weber & .Lšve (COLO 11031, 518089, CS 22873, 39810, RM 93051, 93081) Senecio tridenticulatus Rydb. (KHD 30833, WSC 4206 ) Pectis Pectis angustifolia Torr. (KHD 17130) Picradeniopsis Picradeniopsis oppositifolia (Nutt.) Rydb. (COLO 7377, 7378, 7379, 7380, 7386, 103044, 106142, KHD 39590, 46518, RM 87274) Bahia oppositifolia (Nutt.) A.Gray (UTC 5306) Podospermum Podospermum laciniatum (L.) DC. Scorzonera laciniata L. (KHD 25303, 25304) Pseudognaphalium Pseudognaphalium stramineum (Kunth) W.A.Weber (COLO 9400) Psilochenia Psilochenia occidentalis (Nutt.) W.A.Weber (COLO 8298) Crepis occidentalis Nutt. (CS 23244, KHD 18524) Ratibida Ratibida columnifera (Nutt.) Wooton & Standl. (COLO 10798, 10802, 518147, 518238, KHD 39592, 39593, 47339, RM 92915) Ratibida tagetes (E.James) Barnhart (COLO 10835, 537632, 537633, 538062, KHD 17179, 46779) Rudbeckia Rudbeckia hirta L. (COLO 10940, 10942, 100970, KHD 442) Senecio Senecio flaccidus Less. Senecio flaccidus Less. var. douglasii (DC.) B.L.Turner & T.M.Barkley CFES 2012 (KHD 47338) Senecio riddellii Torr. & A.Gray (KHD 17359, 17361, RM 92752, 92975) Senecio spartioides Torr. & A.Gray (COLO 11124, 518142, KHD 17345, 17347, 17350, 17351, 19943, 19944, KSC 87737, RM 92751, 92977, UTC 87116, 116760) Senecio vulgaris L. (COLO 318409, CS 42062, KHD 17403, 17406, 17409, 26086, 26087, RM 315063, UMO 153612, UTC 120307) Seriphidium Seriphidium tridentatum ( Nutt.) W.A.Weber (COLO 6662) Shinnersoseris Shinnersoseris rostrata ( A.Gray) Tomb (COLO 10569) 38

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Solidago *Solidago altissima L. Solidago canadensis L. (COLO 11551, 518180, CS 37956, KHD 8104, 17485, 17486, 17487, 17491, 39835) Solidago lepida DC. Solidago lepida var. salebrosa (Piper) Semple (RM 84646) Solidago gigantea Aiton (KHD 450, 17522, RM 84412) Solidago missouriensis Nutt. (CS 45225, 45228, KHD 17536, 37745, 38402, 40112, 40553) Solidago mollis Bartl. (COLO 11732, 11753, 11754, CS 20271, 32217, KHD 17549, 17552) Solidago nana Nutt. (KHD 17559) Sonchus Sonchus asper (L.) Hill (COLO 11909, CS 31613, 39767, KHD 22590) Sonchus oleraceus L. (COLO 11913, KHD 610, 40115) Stenactis *Stenactis strigosa (MŸhl. ex Willd.) DC. (COLO 8976) Erigeron strigosus MŸ hl. ex Willd. (KHD 491, 18180) Stephanomeria Stephanomeria pauciflora (Torr.) A.Nelson (COLO 10571, 518075, KHD 39600) Symphyotrichum Symphyotrichum laeve (L.) .Lšve & D.Lšve Aster laevis L. (COLO 6981) Symphyotrichum lanceolatum (Willd.) G.L.Nesom Symphyotrichum lanceolatum (Willd.) G.L.Nesom var. hesperium CFES 2012 (KHD 6333, 9225, RM 43854, 92290, 93085, 95517, 170839, 312031) Aster lanceolatus Willd. subsp. hesperius (A.Gray) Semple & Chmiel. (COLO 7162) Symphyotrichum porteri (A.Gray) G.L.Nesom (CS 20781) Aster porteri A.Gray (COLO 7111, 100885) Tanacetum   *Tanacetum vulgare L. [CO Noxious Weed List B] (COLO 14868, KHD 46440) Taraxacum *Taraxacum officinale F.H.Wigg. (COLO 11964, 518072, KHD 17718, 39190) Tetraneuris Tetraneuris acaulis (Pursh) Greene (COLO 5526) Thelesperma Thelesperma filifolium (Hook.) A.Gray (RM 87273, 88950) Thelesperma filifolium (Hook.) A.Gray var. intermedium (Rydb.) Shinners (COLO 12020, 106070) Thelesperma megapotamicum (Spreng.) Kuntze (COLO 12028, 12032, KHD 22567, 40245, KSC 68955, RM 92953) Townsendia Townsendia exscapa (Richardson) Porter (COLO 12098, CS 20632) Townsendia grandiflora Nutt. (KHD 20890) 39

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Tragopogon *Tragopogon dubius Scop. (COLO 518105, KHD 17866, 27537, 39803) Tragopogon dubius Scop. subsp. major (Jacq.) Vollm. CFES 2012 (COLO 12128) Tragopogon porrifolius L. (COLO 12112, 12114, KHD 17869) Tripleurospermum   *Tripleurospermum inodorum (L.) Sch.Bip. [CO Noxious Weed List B] Matricaria perforata Merat (KHD 17079) Virgulaster Virgulaster ascendens (Lindl.) Semple (COLO 7128) Symphyotrichum ascendens (Lindl.) G.L.Nesom (KHD 15414, 47337) Virgulus Virgulus ericoides ( L.) Reveal & Keener (COLO 7027, 7036) Symphyotrichum ericoides (L.) G.L. Nesom (KHD 6346, 8616, 23999) Virgulus falcatus (Lindl.) Reveal & Keener (COLO 7028, 7030, 7031, 7035) Symphyotrichum falcatum (Lindl.) G.L.Nesom (KHD 8612) Symphyotrichum falcatum var. commutatum (Torr. & A. Gray) G. L. Nesom (RM 162547, 312668, 84408) Virgulus fendleri (A.Gray) Reveal & Keener (COLO 6895, 6898, 135528, KHD 8618) Virgulus novae-angliae (L.) Reveal & Keener [ CHNP tracked G5 S1] Symphyotrichum novae-angliae (L.) G.L. Nesom (KHD 24013) Virgulus x amethystinus (Nutt.) Reveal & Keener (COLO 7032) Xanthium *Xanthium strumarium L. (COLO 12260, 12269, 12272, 12273, 518174) Xanthium strumarium var. canadense (Mill.) Torr. & A.Gray (CS 33174) Ximenesia *Ximenesia encelioides Cav. (COLO 12154, 12158, 12175, 12177, 170920) Verbesina encelioides (Cav.) Benth. & Hook.f ex A.Gray (KHD 17892, 17893, 17894, 37832, 40086, 47336, RM 92774, 92984, 97756) Berberidaceae Berberis *Berberis vulgaris L. (CS 8726) Betulaceae Alnus Alnus incana (L.) Moench Alnus incana (L.) Moench subsp. tenuifolia (Nutt.) Breitung CFES 2012 (KHD 8466) Betula Betula occidentalis Hook. (KHD 8474, 8484) 40

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Boraginaceae Anchusa   *Anchusa officinalis L. [CO Noxious Weed Watch List] (CS 17351) Cryptantha Cryptantha crassisepala (Torr. & A.Gray) Greene (KHD 8572) Cryptantha fendleri (A.Gray) Greene (COLO 2452, 106078, RM 48372, 170837) Cryptantha minima Rydb. (COLO 2425, 2434, 518097, CS 17294, KHD 39808, RM 435978) Cynoglossum   *Cynoglossum officinale L. [CO Noxious Weed List B] (COLO 518243, KHD 39510) Heliotropium *Heliotropium curassavicum L. Heliotropium curassavicum L. subsp. oculatum (A.Heller) Thorne CFES 2012 (COLO 2755) Lappula Lappula redowskii (Hornem.) Greene (COLO 2767, 2776, 2777, 2804, 2809, 2811, 2813, 106063) Lappula occidentalis var. cupulata (A.Gray) L.C.Higgins (CS 17196, KHD 8797) Lappula occidentalis (S.Wats.) Greene var. occidentalis (CS 17174, KHD 40535) Lithospermum Lithospermum incisum Lehmann (COLO 2848, 518066, KHD 8828, 13686, RM 92991) Mertensia Mertensia lanceolata (Pursh) DC. (COLO 3076, 3091, KHD 1354, 1461) Onosmodium Onosmodium molle Michx. (KHD 8958) Onosmodium molle Michx. var. hispidissimum (Mack.) Cronquist (NY 21471, 21472) Onosmodium molle Michx. subsp. occidentale (Mack.) Cochrane CFES 2012 (COLO 3227, 3234, 3241, 22975, 106055) Oreocarya Oreocarya cana A.Nelson [CNHP tracked G5 S2] Cryptantha cana (A.Nels.) Payson (CS 17261, 17262) Oreocarya suffruticosa (Torr.) Greene (COLO 2491, 2545, 2549, 104157) Cryptantha cinerea (Greene) Cronquist (CS 17280, 17285) Cryptantha cinerea (Greene) Cronquist var. jamesii (Torr.) Cronquist (KHD 8592, RM 87733) # Oreocarya virgata (Porter) Greene [Endemic ] (COLO 2591) Plagiobothrys Plagiobothrys scouleri (Hook. & Arn.) I.M.Johnst. Plagiobothrys scopulorum (Greene) I.M.Johnst. (GH 93611) Symphytum *Symphytum officinale L. (COLO 459886, 459887) 41

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Brassicaceae Arabis Arabis pycnocarpa M.Hopkins Arabis hirsuta (L.) Scop. (CS 9526) Arabis hirsuta (L.) Scop. var. pycnocarpa (M.Hopkins) Rollins (COLO 12761) Alliaria  *Alliaria petiolata (M.Bieb.) Cavara & Grande [CO Noxious Weed Watch List] (COLO 538069, KHD 46778) Alyssum Alyssum desertorum Stapf (COLO 286270, 518068, KHD 3912, 10310, 17949, 40528) Berteroa *Berteroa incana (L.) DC. (DMNS 589) Boechera Boechera fendleri (S.Watson) W.A.Weber Arabis fendleri (S.Watson) Greene (KHD 1460) Boechera stricta (Graham) Al-Shehbaz Boechera drummondii (A.Gray) .Lšve & D.Lš ve (COLO 12735) Brassica *Brassica juncea (L.) Czern. (COLO 12899, 12903, KHD 29414, RM 89019) Brassica nigra (L.) K.Koch (COLO 12918, KHD 18090) Camelina *Camelina microcarpa Andrz. ex DC. (COLO 12932, 12943, KHD 18094, 22606, UTC 5458) Capsella *Capsella bursa-pastoris (L.) Medicus (COLO 12967, KHD 10282, 18100, 23839, 23840) Cardaria   *Cardaria draba ( L.) Desv. [CO Noxious Weed List B] (COLO 518106, KHD 18134, 18137, 18138, 39802, 40221 )  * Cardaria latifolia (L.) Spach [CO Noxious Weed List B] (COLO 518118, KHD 39550) Cardaria pubescens (C.A.Mey.) Jarm. (COLO 518203, KHD 39891) Chorispora *Chorispora tenella (Pall.) DC. (COLO 518112, KHD 18150, 18153, 39544, WSC 6976 ) Descurainia Descurainia incana (Bernh. ex Fisch. & C.A.Mey.) Dorn (KHD 5511, 9282, 18179) Descurainia incisa (Engelm.) Britton (COLO 13105) Descurainia pinnata (Walter) Britton (KHD 27974, 46924) Descurainia pinnata (Walter) Britton subsp. halictorum (Cockerell) Detling (COLO 13102) Descurainia sophia (L.) Webb ex Prantl (COLO 518126, KHD 39113, 39535) Draba Draba cuneifolia Nutt. (KHD 39112) Draba nemorosa L. (COLO 13223) Draba reptans (Lam.) Fernald (COLO 13186, 13250, KHD 45203, RM 41266, 98706) 42

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Erysimum *Erysimum repandum L. (RM 89039) Lepidium *Lepidium densiflorum Schrad. (COLO 13563, 13564, KHD 8869, 18349, 18352, 25488) Lepidium perfoliatum L. (COLO 13592) Lepidium ramosissimum A.Nels. (KHD: Wingate 9940 2014) Lepidium virginicum L. (KHD 37532) Nasturtium Nasturtium officinale R.Br. (COLO 13763, 101478, KHD 1623, RM 12972) Neolepia *Neolepia campestris (L.) W.A.Weber Lepidium campestre (L.) W.T. Aiton (CS: CNHP D-2013-78 ) Noccaea Noccaea fendleri (A.Gray) Holub Noccaea montana (L.) F.K.Mey. (KHD 1341, 1356) Physaria Physaria ludoviciana (Nutt.) O'Kane and Al-Shehbaz Lesquerella ludoviciana (Nutt.) S.Watson (COLO 13637, 13640, 13660, 106147, 518093, 530323, KHD 39806) Physaria montana (A.Gray) Greene (RM 92838) Lesquerella montana (A.Gray) S.Watson (COLO 13666, 13667, 100915) # Physaria vitulifera Rydb. [Endemic CNHP tracked: G3 S3] (COLO 13821 ) Rorippa Rorippa palustris (L.) Besser Rorippa palustris (L.) Besser subsp. hispida (Desv.) Jonsell (KHD 22398, 26048, 26712, RM 87789) Rorippa sinuata (Nutt.) Hitchc. (COLO 492650, CS 9174, KHD 18452, 32112, 40522) Rorippa sphaerocarpa (A.Gray) Britton (COLO 13809, RM 87790) Rorippa teres (Michx.) Stuckey (COLO 13770, 13771, 13773, 518121, 530325, KHD 16553) Sinapis *Sinapis arvensis L. (COLO 12896, 12897, 452416, KHD 22636) Sisymbrium *Sisymbrium altissimum L. (COLO 13901, KHD 18468, 18469, 18472, 39811, 40056) Sisymbrium loeselii L. (KHD 32116) Thlaspi *Thlaspi arvense L. (COLO 518129, KHD 8121, 31848) 43

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Cactaceae Coryphantha Coryphantha vivipara (Nutt.) Britton & Rose (COLO 3312, 3313, 518225) Opuntia Opuntia fragilis (Nutt.) Haw. (MO 2015204, US 441679, 1487504) Opuntia macrorhiza Engelm. (COLO 3324, MO 2015554, US 535665) Opuntia polyacantha Haw. (COLO 3332, 518095, KHD 9006, 39507, MO 2015623, US 535759, 3046516) Callitrich aceae Callitriche Callitriche heterophylla Pursh [CNHP tracked: G5 S1] (CS: CNHP D-2013-89 ) Campanulaceae Campanula Campanula parryi A.Gray (CS 19835, 19840) Campanula rapunculoides L. (KHD 32234, 37821) Campanula rotundifolia L. (COLO 3467, ID 1892, NY 1877) Lobelia Lobelia siphilitica L. Lobelia siphilitica L. var. ludoviciana A.DC. CFES 2012 (COLO 3556) Cannabaceae Cannabis Cannabis sativa L. (COLO 318676, KHD 1018, 1020, RM 105342) Humulus Humulus lupulus L. Humulus lupulus L. subsp. americanus (Nutt.) .Lšve & D.Lš ve (COLO 42037) Humulus lupulus var. neomexicanus A.Nelson & Cockerell CFES 2012 (CS 6320, KHD 1027) Caprifoliaceae Symphoricarpos Symphoricarpos albus (L.) S.F.Blake (KHD 41011) Symphoricarpos occidentalis Hook. (COLO 3890, 3895, 3896, 3919, 3927, 3928, 101018, 106116, 492631, KHD 37517) Caryophyllaceae Gastrolychnis Gastrolychnis drummondii (Hook.) . Lšve & D. Lšve Silene drummondii var. striata (Rydb.) Bocquet (KHD 10102) Saponaria   *Saponaria officinalis L. [CO Noxious Weed List B] (COLO 4453, KHD 26204) Silene *Silene noctiflora L. (COLO 4545) 44

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Ceratophyllaceae Ceratophyllum demersum L. (KHD 3824496) Chenopodiaceae Atriplex Atriplex argentea Nutt. (COLO 4796, 4797, 4809, KHD 10487) Atriplex canescens (Pursh) Nutt. (COLO 4816, 4836, 4846, KHD 10495, 10497, 10498, 40107, 40109) Atriplex confertifolia (Torr. & FrÂŽm.) S.Watson (RM 48838) Atriplex heterosperma Bunge (KHD 10516, 39631, 44409) Atriplex hortensis L. (COLO 4873, 4874, KHD 10507, 10512) Atriplex dioica Raf. Atriplex patula L. (COLO 4869, 4870, 4885, KHD 10514, 10515, 10517, 32794) Atriplex rosea L. (KHD 10522, 10523) Atriplex truncata (Torr.) A.Gray (COLO 4894) Bassia Bassia hyssopifolia (Pall.) Kuntz (CS: CNHP D-2013-77) *Bassia sieversiana (Pall.) W.A.Weber (COLO 5124, 5126, 5127, 5128, 5129, 5130) Kochia scoparia (L.) Schrad. (KHD 2210, 10608, 10610, 10611, 10612, 10613, 10614, 22746, 22750, 47309, RM 138231) Chenopodium *Chenopodium album L. (KHD 10553, 10554, 10556, 32808, 32809) Chenopodium berlandieri Moq. (COLO 4928, 4951, 5042, KHD 34317, 47310) Chenopodium desiccatum A.Nelson (COLO 5019, RM 225692) Chenopodium glaucum L. (COLO 5000, 5002, 5044, KHD 1285, 39553, 43424) Chenopodium incanum (S.Watson) A.Heller (COLO 4990, 4991, 5016, 5018, KHD 10578, RM 225679) Chenopodium leptophyllum (Moq.) Nutt. ex S.Watson (COLO 5020, 5028, 5031) Chenopodium pratericola Rydb. (COLO 4940) Chenopodium simplex (Torr.) Raf. (CS: CNHP D-2013-175, -176, -179 ) Corispermum Corispermum americanum (Nutt.) Nutt. (COLO 5091) Corispermum hyssopifolium L. p.p. (UTC 5652) Cycloloma Cycloloma atriplicifolium (Spreng.) Coult. (COLO 5068, 5070, 5072, 5078, 5079, KHD 10600, RM 48840, UTC 7635, 7668) Dysphania *Dysphania botrys (L.) Mosyakin & Clemants (CS 6807, KHD 10559) Teloxys botrys (L.) W.A.Weber (COLO 4962, 4966) Krascheninnikovia Krascheninnikovia lanata (Pursh) A.Meeuse & A.Smit (COLO 5097, 5099, 5100, 5105, 5113, KHD 10544, 10545, 46464) 45

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Monolepis Monolepis nuttalliana (Schult.) Greene (KHD 10619, 10620) Salsola *Salsola australis R.Br. (COLO 5141, 5146, 5154) Salsola tragus L. (CS 7097, 7102, KHD 10636, 10637, 10638, 10639, 10640, 10646, 37574) Suaeda Suaeda calceoliformis (Hook.) Moq. (COLO 5183, 5184, 5186) Suaeda nigra (Raf.) J.F.Macbr. (COLO 5194) Cleomaceae Peritoma Peritoma serrulata (Pursh) DC. Cleome serrulata Pursh (COLO 3643, 3654, 492630, CS 93789, KHD 9106, 9107, 47306, RM 92422, 92983, 97633, UTC 5508) Comandraceae Comandra Comandra umbellata (L.) Nutt. Comandra umbellata (L.) Nutt. subsp. pallida (A.DC.) Piehl (COLO 38030, 38031, 38045) Commelinaceae Commelina Commelina erecta L. (US 222848) Tradescantia Tradescantia occidentalis (Britton) Smyth (KHD 39804, 47311, US 247252, 270385, 1100525) Tradescantia occidentalis (Britton) Smyth var. occidentalis (RM 48293, 92989, 97126) Tradescantia occidentalis (Britton) Smyth var. scopulorum (Rose) E.S.Anderson & Woods. (COLO 5245, 5246, 5261, 5282) Convallariaceae Maianthemum Maianthemum stellatum (L.) Link (COLO 27083, KHD 37786) Convolvulaceae Calystegia Calystegia sepium (L.) R.Br. Calystegia sepium (L.) R.Br. subsp. angulata Brummitt (COLO 12322, 12330, KHD 42617) Convolvulus   *Convolvulus arvensis L. [CO Noxious Weed List C] (COLO 14961, CS 89727, KHD 4803, 10701, 37822, 47312) Evolvulus Evolvulus nuttallianus Roem. & Schult. (COLO 12348, 12350, 12358, 518099, CS 16568, KHD 10727, 39809) Ipomoea Ipomoea leptophylla Torr. (COLO 12367, 12370, 101516, 518186, CS 16640, 45229, KHD 10735, RM 45303, 87735, 97470) Ipomoea purpurea (L.) Lam. (COLO 459885) 46

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Cornaceae Swida Swida sericea (L.) Holub (COLO 12464) Cucurbitaceae Cucurbita Cucurbita foetidissima Kunth (COLO 518137, KHD 17420, 47304) Echinocystis Echinocystis lobata (Michx) Torr. & A.Gray (COLO 103017, KHD 10816, 47305) Cuscutaceae Grammica Grammica indecora (Choisy) W.A.Weber Grammica indecora (Choisy) W.A.Weber var. neuropetala (Engelm.) W.A.Weber (COLO 14209, 14210, 14213) Cuscuta indecora Choisy (UTC 6515) Cuscuta indecora Choisy var. neuropetala (Engelm.) Hitchc. (CS 16540) Cyperaceae Bolboschoenus Bolboschoenus maritimus (L.) Palla Bolboschoenus maritimus (L.) Palla subsp. paludosus (A.Nelson) .Lšve & D.Lšve CFES 2012 (COLO 16006, KHD 45974) Carex Carex aquatilis Wahlenb. (UTC 5469) Carex atherodes Spreng. (COLO 496430) Carex brevior (Dewey) Mack. ex Lunell (CS 4226, KHD 46925, RM 88125) Carex douglasii Boott (COLO 14480, CS 4245) Carex duriuscula C.A.Mey. (KHD 6511, 40533) Carex stenophylla Wahlenb. subsp. eleocharis (L.H.Bailey) HultŽ n (COLO 14524) Carex emoryi Dewey (COLO 286271, KHD 44404, 44405) Carex filifolia Nutt. (COLO 522244, KHD 8250, 39179) Carex hystericina Muhl. ex Willd. (MO 890032, RM 88010) Carex inops L.H.Bailey Carex pensylvanica Lam. subsp. heliophila (Mack.) W.A.Weber (KHD 6193) Carex nebrascensis Dewey (CS 4315, COLO 6211, 6222, 32631, 32688, RM 313362) Carex pellita Muhl. ex Willd. (CS: CNHP D-2013-110 ) Carex praegracilis W.Boott (COLO 15300, 93743, 518117, KHD 3934, 6461, 6462, 6465, 40077, 46922) Carex siccata Dewey (CS: CNHP D-2013-111 ) Carex utriculata Boott (CS: CNHP D-2013-107 ) 47

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Cyperus Cyperus erythrorhizos Muhl. (KHD 40523)  * Cyperus esculentus L. [CO Noxious Weed List B] (KHD 1379, 21071) Cyperus odoratus L. (CS: CNHP D-2013-101) Cyperus squarrosus L. (CS 3886, RM 48701, 95500) Cyperus aristatus Rottb. (COLO 15650, 15683) Eleocharis Eleocharis colorado‘nsis (Britton) Gilly Eleocharis parvula (Roem. & Schult.) Link ex Bluff, Nees & Schauer var. anachaeta Svenson (COLO 518083) Eleocharis macrostachya Britton (CS: CNHP D-2013-182, -183, -184, -185, -186, -187, -189, -193, -195 ) Eleocharis palustris (L.) Roem. & Schult. (COLO 15798, 15815, KHD 6602, 6609) Eleocharis rostellata (Torr.) Torr. (CS: CNHP D-2013-191, -192 ) Lipocarpha Lipocarpha aristulata (Coville) G.C. Tucker Hemicarpha micrantha (Vahl) Pax var. aristulata Coville (COLO 21204) Schoenoplectus Schoenoplectus pungens (Vahl) Palla (COLO 15979, 518122, 518156, KHD 6780, 6781, 26151, 39538, 39555, RM 93013 ) Schoenoplectus americanus (Pers.) Volkart ex Schinz & R. Keller (CS 4009, 4010, KHD 11787 ) Schoenoplectus tabernaemontani ( C.C.Gmel.) Palla (KHD 46913) Schoenoplectus acutus (Muhl. ex J.M.Bigelow ) .Lš ve & D.Love var. acutus (KHD 7604) Schoenoplectus lacustris (L.) Palla subsp. acutus (Muhl. ex J.M.Bigelow ) .Lšve & D.Lš ve (COLO 518119) Schoenoplectus validus (Vahl) Lšve & Lš ve (COLO 16060) Scirpus Scirpus microcarpus J. & K. Presl (KHD 6212, RM 105345) Scirpus pallidus (Britton) Fernald (COLO 16032, KHD 6783) Dipsacaceae Dipsacus   *Dipsacus laciniatus L. [CO Noxious Weed List B] (COLO 518183) Elaeagnaceae Elaeagnus   *Elaeagnus angustifolia L. [CO Noxious Weed List B] (COLO 16134, 16137, 518104) KHD 1624, 4903, 10867) Elatinaceae Elatine Elatine triandra Schkuhr [CNHP tracked: G5 S2] Elatine americana A.Gray (US 582405) Elatine rubella Rydb. [CNHP tracked: G5 S2, not listed in CFES2012] (COLO 16175, 16176) 48

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Euphorbiaceae Agaloma Agaloma marginata (Pursh) .Lšve & D.Lšve (COLO 16760, 16761, 16764, 16778) Euphorbia marginata Pursh (CS 27040, 60712, RM 67227) Chamaesyce Chamaesyce fendleri (Torr. & A.Gray) Small (COLO 16722) Chamaesyce glyptosperma (Engelm.) Small (COLO 16741, 16747, 16753, 106140, KHD 39560, 47332) Chamaesyce maculata (L.) Small (CS 82997, KHD 38966) Chamaesyce supina (Raf.) Moldenke (COLO 500031) Chamaesyce missurica (Raf.) Shinners (COLO 16848, 16850, 16851, 16852, 16853, 106141, 418726, KHD 21068, RM 37698, 288951) Chamaesyce prostrata (Aiton) Small (KHD 11115, 33926, 34297, 38033) Chamaesyce serpyllifolia (Pers.) Small (COLO 16870, 16872, CS 27151, KHD 11117, 11119, 24162, RM 88959, 97226) Euphorbia serpyllifolia Pers (UTC 8740) Croton Croton texensis (Klotsch) MŸll.Arg. (COLO 16658, 16662, 16663, 16664, 31457, 103026, KHD 11130, 22681, 40075, RM 113414, UTC 6455 ) Poinsettia *Poinsettia dentata (Michx.) Klotsch & Garcke (COLO 16703, 16704, 16705) Euphorbia dentata Michx. (CS 82671, 82672, KHD 25330, 40397) Tithymalus Tithymalus brachycerus (Engelm.) Small (COLO 16818, 16839, RM 92961)  * Tithymalus esula Hill [CO Noxious Weed List B] Euphorbia esula var. uralensis (Fisch. ex Link) Dorn (KHD 39188 47331)  * Tithymalus myrsinites (L.) Hill [CO Noxious Weed List A] Euphorbia myrsinites L. (KHD 25513) Tithymalus peplus (L.) Hill Euphorbia peplus L. (CS 13252) Tithymalus spathulatus (Lam.) W.A.Weber (COLO 16689) Tithymalus uralensis (Fisch. ex Link) Prokh. (COLO 16679, KHD 40246) Fabaceae Amorpha Amorpha fruticosa L. (COLO 23979, 23981, 23977) Apios Apios americana Medik. [CNHP tracked: G5 S1] (COLO 23993) Astragalus Astragalus agrestis Douglas ex G.Don (ARIZ 701, COLO 24384, KHD 12198, NY 1214991, RM 95426) Astragalus bisulcatus (Hook.) A.Gray (COLO 24066, 24518, 24519, 518109, NY 1244808, RM 87143, 87144) Astragalus canadensis L. (COLO 24082) Astragalus ceramicus E Sheld. 49

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(KHD 40530) Astragalus ceramicus E.Sheld. var. filifolius (A.Gray) F.J.Herm. CFES 2012 (COLO 24271, 24426, 24503, 52900, CS 12220, 26400, RM 87941, 167824) Astragalus cicer L. (COLO 518154, KHD 39557) Astragalus crassicarpus Nutt. (ARIZ 514, COLO 24097, KHD 11393, RM 28220, 95481) Astragalus drummondii Douglas ex Hook. (COLO 24261) Astragalus flexuosus Douglas ex G.Do n (NY 1249390) Astragalus gracilis Nutt. (COLO 24353, 24436, 24437, 518090, CS 12268, 12273, KHD 39527) Astragalus hallii A. Gray (COLO 24525) Astragalus haydenianus A. Gray Astragalus bisulcatus var. haydenianus (A. Gray) Barneby (KHD 39547) Astragalus laxmannii Lam. Astragalus adsurgens Pall. var. robustior Hook. (COLO 24005, NY 1203762) Astragalus lotiflorus Hook. (KHD 40534, 43079, NY 1261125, 1261128) Astragalus missouriensis Nutt. (COLO 24446, UTC 246737, NY 1262015) # Astragalus parryi A.Gray [Endemic ] (COLO 24473, 24633) Astragalus shortianus Nutt. (COLO 24534) Astragalus sparsiflorus A.Gray [CNHP tracked G2 S2] (COLO 24554, 24555, CS 12385) Astragalus sparsiflorus A.Gray var. majusculus A.Gray (UTC 249456) Colutea *Colutea arborescens L. (KHD 45918) Dalea Dalea candida Michx. ex Willd. (UTC 7721) Dalea candida Willd. var. oligophylla (Torr.) Shinners (COLO 25793, 25797, 25810, 103029, KHD 8069, 31819, RM 107501) Dalea cylindriceps Barneby (COLO 25805, 453007, KHD 30070, 30079, NY 1259200, 1259213) Dalea leporina (Aiton) Bullock (COLO 24775, 24776, CS 32595, NY 1270105) Dalea purpurea Vent. (COLO 25849, 25865, 101590, NY 1277504, RM 89033) Desmanthus Desmanthus illinoensis (Michx.) MacMill. (COLO 24819) Desmodium Desmodium rigidum (Elliott) DC. [not listed in CFES2012, extirpated?] (COLO 106011) Glycyrrhiza Glycyrrhiza lepidota Pursh (COLO 24907, 103030, 518110, KHD 12505, 39546) Lathyrus Lathyrus eucosmus Butters & H.St.John (COLO 24987, 25031, 25052, KHD 34918, RM 92520, 92925) Lathyrus decaphyllus Pursh (ARIZ 18) Lathyrus latifolius L. (KHD 31895) 50

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Lotus Lotus corniculatus L. [not listed in CFES2012] (KHD 1625, 29147) Lupinus Lupinus argenteus Pursh (COLO 25270, 25271, KHD 8080) Lupinus argenteus Pursh var. argenteus (RM 81544) Lupinus argenteus var. laxiflorus (Douglas ex Lindl.) Dorn (RM 92462, 93072, 165558, 165559) Lupinus plattensis S.Watson (COLO 25272, 492636) Lupinus pusillus Pursh (COLO 25412, 25413, CS 11666, KHD 8200, RM 95412) Medicago *Medicago lupulina L. (CS 32530, KHD 1832, 22902, 37835) Medicago sativa L. (COLO 25515, 518086, KHD 37169, 39812, 41720) Melilotus *Melilotus albus Medik. (COLO 25536, 25537, 518111, KHD 39545, 41718) Melilotus officinalis (L.) Pall. (COLO 518242) Onobrychis *Onobrychis viciifolia Scop. (COLO 25602) Orophaca Orophaca tridactylica (A.Gray) Rydb. Astragalus tridactylicus A.Gray (CS 12353, RM 28236, 167941) Oxytropis Oxytropis lambertii Pursh (COLO 25672, 52901, 100960, 518098, CS 12472, KHD 5164, 12915, 37173) Pediomelum Pediomelum hypogaeum (Nutt. ex Torr. & A.Gray) Rydb. (RM 164780) Psoralidium Psoralidium lanceolatum (Pursh) Rydb. (COLO 25945, 101513, 106052, 106132, KHD 13002, 32102, 40241, 47330, RM 88964) Psoralidium tenuiflorum (Pursh) Rydb. (COLO 25966, 25973, 25980, 25988, 101587, 103050, CS 11973, KHD 13012, 40238, RM 84655, 93088) Robinia *Robinia pseudoacacia L. (KHD 40108) Securigera *Securigera varia (L.) Lassen Coronilla varia L. (KHD 21840, 26762) Thermopsis Thermopsis divaricarpa A.Nelson (COLO 26164) Thermopsis rhombifolia (Nutt. ex Pursh) Nutt. ex Richardson (RM 92455, 93070, 165534) Trifolium *Trifolium fragiferum L. (KHD 5258, 16437, KSC 59234 ) Trifolium repens L. (KHD 13143) Vexibia Vexibia nuttalliana (B.L.Turner) W.A.Weber (COLO 24628, 26028, 106139) Sophora nuttalliana B.L.Turner (CS 11538, RM 92921, 92923) 51

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Vicia Vicia americana Muhl. ex Willd. (KHD 13238, 13656, 34900, WSC 4205 ) Vicia linearis (Nutt.) Greene Vicia americana var. minor Hook. (RM 89137, 95418, 168076, 225409) Fumariaceae Corydalis Corydalis aurea Willd. (COLO 100834, KHD 3469, 11421, 43408) Corydalis aurea Willd. var. occidentalis Engelm. ex A.Gray (RM 92693, 92945) Corydalis curvisiliqua Engelm. Corydalis curvisiliqua Engelm. ex A.Gray subsp. occidentalis (Engelm. ex A.Gray) W.A.Weber CFES 2012 (COLO 29934, 29935, 29941) Fumaria *Fumaria vaillantii Loisel. (COLO 539979, KHD 39491) Gentianaceae Eustoma Eustoma russellianum (Hook.) G.Don [CNHP tracked: G5 S3S4] Eustoma grandiflorum (Raf.) Shinners (COLO 17179, RM 161733) Geraniaceae Erodium   *Erodium cicutarium (L.) L'Her. [CO Noxious Weed List C] (COLO 518071, KHD 5797, 11608, 26181, 26184) Grossulariaceae Ribes Ribes americanum Mill. [CNHP tracked: G5 S2] (COLO 106031, CS 10136, RM 155349) Ribes aureum Pursh (COLO 518234, KHD 14549, 39189) Haloragaceae Myriophyllum Myriophyllum sibiricum Kom. (COLO 22419, RM 92929) Myriophyllum exalbescens Fernald (US 1100511)  * Myriophyllum spicatum L. [CO Noxious Weed List B] (CNHP D-2013-86, -87 ) Helleboraceae Delphinium Delphinium carolinianum Walter Delphinium carolinianum Walter subsp. virescens (Nutt.) M.C. Johnston CFES 2012 (COLO 33806, 103058, RM 92943, 95401, 168950) Delphinium virescens Nutt. (KHD 3773, 3774) Hydrocharitaceae Elodea Elodea bifoliata H.St.John (KHD 3824499) Elodea canadensis Michx. (CS: CNHP 5-27-2014, PS 1520 ) 52

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Hydrophyllaceae Ellisia *Ellisia nyctelea (L.) L. (COLO 22453) Phacelia Phacelia alba Rydb. (COLO 22722, 100993) Phacelia hastata Douglas ex Lehm (COLO 22582) Phacelia heterophylla Pursh (COLO 22632, 22644) Iridaceae Sisyrinchium Sisyrinchium montanum Greene (COLO 22884) Juncaceae Juncus Juncus arcticus Willd. Juncus arcticus Willd. subsp. ater (Rydb.) HultŽn CFES 2012 (COLO 22973, 507774, 518130) Juncus arcticus var. balticus (Willd.) Trautv. (KHD 395333) Juncus articulatus L. (CS 73932, 73935, 73936, 73937, 74021, KHD 3413) Juncus bufonius L. (COLO 23043, KHD 35772) Juncus compressus Jacq. (KHD 31894) Juncus dudleyi Wiegand (COLO 507779) Juncus longistylis Torr. (COLO 23136, 86914, 106029, 106103, MO 3266171, 3266177, RM 3266177) Juncus nodosus L. (COLO 23178, KHD 6956) Juncus tenuis Willd. (MO 3267191) Juncus torreyi Coville (COLO 15697, 106081, KHD 35873, MO 3266564, 3266568) Juncaginaceae Triglochin Triglochin maritima L. (KHD 7053, RM 87772) Lamiaceae Glecoma *Glechoma hederacea L. (CS 38099, KHD 26899) Hedeoma Hedeoma hispida Pursh (COLO 23298, KHD 24788) Lamium *Lamium amplexicaule L. (KHD 11866, 11893, 11894, 11895, 22670, 22671, 45507) Lycopus Lycopus americanus MŸ hl. ex W.P.C. Barton (COLO 23338) Lycopus asper Greene (KHD 11909, 23040, RM 89025) Lycopus uniflorus Michx. (CS: CNHP D-2013-73 ) 53

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Mentha Mentha arvensis L. (COLO 23379, 23380, 23397, 23399, KHD 11945) Mentha spicata L. (COLO 23435) Monarda Monarda fistulosa L. Monarda fistulosa L. var. menthifolia (R. Graham) Fernald (COLO 23474) Monarda pectinata Nutt. (COLO 23501, 23504, 104119, KHD 11987, 20886, RM 95473, 229553) Nepeta *Nepeta cataria L. (COLO 518181, KHD 12010, 23319, 39834, 47334) Prunella Prunella vulgaris L. (KHD 37825) Salvia *Salvia reflexa Hornem. (COLO 23680, 23683, 23684, 104118) Scutellaria Scutellaria brittonii Porter (COLO 23743, 23755, 106039, 518091, KHD 39528, 45888) Scutellaria epilobiifolia A.Ham. (RM 105348) Stachys Stachys palustris L. Stachys palustris L. subsp. pilosa (Nutt.) Epling (COLO 106018) Teucrium Teucrium canadense L. Teucrium canadense L. subsp. occidentale (A.Gray) D.C.McClint. & Epling CFES 2012 (COLO 23814, 23855) Lemnaceae Lemna Lemna minor L. (KHD 3824498 ) Lemna minuta Kunth (CS: CNHP D-2013-85, -94 ) Lemna turionifera Landolt (COLO 26594) Spirodela *Spirodela polyrrhiza (L.) Schleid. (COLO 26603) Wolffia Wolffia columbiana H.Karst. (KHD 3824497, CS: CNHP D-2013-88 ) Liliaceae Leucocrinum Leucocrinum montanum Nutt. ex A.Gray (COLO 26868, 26870, 100942, 100950, KHD 7238, 7239, 7240, 39191, 47329, RM 92917, 97102) Linaceae Adenolinum Adenolinum lewisii (Pursh) .Lšve & D.Lšve (COLO 27476) Linum lewisii Pursh var. lewisii (CS 27397) 54

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Loasaceae Acrolasia Acrolasia albicaulis (Douglas ex Hook.) Rydb. (COLO 27715, 492665) Mentzelia albicaulis (Douglas ex Hook.) Douglas ex Torr. & A.Gray (UTC 10423, 12286) Nuttallia Nuttallia multiflora (Nutt.) Greene [CNHP tracked: G3 S3] (COLO 27768) Nuttallia speciosa (Osterh.) Greene (COLO 27756, 27759, 27762, 27777, 27830) Nuttallia nuda (Pursh) Greene (COLO 27770, 27778, 27815, 27816, 27817, 27818, 27819, 86920, 518073, CS 89728, 96503, KHD 13349, 13352, UTC 12575,12576, RM 81534, 87156, 88965, 92361, 92368, 92957, 97283, 237756) Nuttallia rusbyi (Wooton) Rydb. (COLO 27772) Lythraceae Ammannia *Ammannia robusta Heer & Regel (COLO 27894, 27898, 27899, 27901, RM 105341, 788949) Lythrum Lythrum alatum Pursh (COLO 27902, 27911, 27914, 27915, 103031, 539647, CS 96220 )  * Lythrum salicaria L. [CO Noxious Weed List A] (COLO 318898, KHD 30397) Malvaceae Abutilon   *Abutilon theophrasti Medik. [CO Noxious Weed List C] (CS 82996, KHD 3719, 31483) Callirho‘ Callirho‘ involucrata (Torr. & A.Gray) A.Gray (CS 13915) Hibiscus   *Hibiscus trionum L. [CO Noxious Weed List B] (CS 274155, KHD 1628) Malva *Malva crispa (L.) L. (CS 27298) Malva neglecta Wallr. (COLO 27979, 498752, KHD 5003, 13469, 13470, 22743, 37830) Sphaeralcea Sphaeralcea coccinea (Nutt.) Rydb. (COLO 28146, 28148, CS 13852, 45232, KHD 13512, 13517, 13522, 37836, 40236, 47328, RM 92913, 97223, UTC 141881) Melanthiaceae Toxicoscordion Toxicoscordion venenosum (S.Watson) Rydb. (COLO 27356, 27365, 100949) Zigadenus venenosus var. gramineus (Rydb.) Walsh ex M.Peck (KHD 7552) Molluginaceae Mollugo *Mollugo verticillata L. (KHD 9634, 34316) 55

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Moraceae Maclura *Maclura pomifera (Raf.) C.K.Schneid. (COLO 28206, 28207, 28211, 43921) Najadaceae Najas *Najas guadalupensis (Spreng.) Magnus (CS: CNHP D-2013-83 ) Nyctaginaceae Abronia Abronia fragrans Nutt. ex Hook. (ARIZ 44751, COLO 28337, 28340, 28344, 28353, 101533, 518087, CS 26133, 74366, KHD 1062, 22617, 39530, KSC 86750, RM 84649, 92974, 224549, UTC 500, 947) Oxybaphus Oxybaphus linearis (Pursh) B.L.Rob. (COLO 28399, 28450) Allionia linearis Pursh (UTC 1597) Mirabilis linearis (Pursh) Heimerl. (KHD 40076, RM 49645, 87766, 92440, 92916) Oxybaphus nyctagineus (Michx.) Sweet (COLO 28465, 28474, 106056, 518152) Mirabilis nyctaginea (Michx.) Macmill. (KHD 39559, RM 87767) Tripterocalyx Tripterocalyx micranthus (Torr.) Hook. (COLO 28364, 28367, 28368, UTC 141882, 141886) Oleaceae Forestiera Forestiera pubescens Nutt. (CS: CNHP D-2103-44, -80 ) Onagraceae Calylophus Calylophus serrulatus (Nutt.) P.H.Raven (COLO 29099, 29243, KHD 5662) Chamerion *Chamerion danielsii ( D. Lš ve) Czerep. Chamerion angustifolium var. canescens (Alph.Wood) N.H.Holmgren & P.K.Holmgren (CS 14561) Epilobium Epilobium ciliatum Raf. (CS: CNHP D-2013-55, 63, -64, -70 )  * Epilobium hirsutum L. [CO Noxious Weed Watch List] (COLO 539991, KHD 46915, 46916) Gaura Gaura coccinea Nutt. ex Pursh (COLO 28965, 106045, CS 14884, KHD 1226, 1227, 1232, 1233, 1237, 34914, 39525, 40235, 47327, UTC 9001) Oenothera suffrutescens (Ser.) W.L.Wagner & Hoch (RM 92911, 97288) Gaura mollis E.James (COLO 29000, 29002, 29005, KHD 1243) Gaura parviflora Dougl. ex Lehmm. (KHD 40243, 46487) 56

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Oenothera Oenothera albicaulis Pursh (COLO 29187, 29372, 106149, CS 14698, KHD 1290, 1313, 1314, 1358, 2592, 10558, 34911, 47326, RM 92978, 518363, US 44466, 44469, 44561, 1070231, 270390, 1486866) Oenothera coronopifolia Torr. & A.Gray (COLO 29150, 29159, CS 14712, KHD 9372, 40118, RM 92912, US 44558, 270394) Oenothera flava Garrett (COLO 518155, CS 14807, KHD 1371, 39556, RM 265028) Oenothera nuttallii Sweet (COLO 29165, 29168, 29170, 29172, 29210, CS 14737, 14740, KHD 1777, RM 88967) Oenothera villosa Thunb. Oenothera villosa Thunb. subsp. strigosa (Rydb.) W.Dietr. & P.H.Raven (KHD 47325, 88963) Oenothera biennis L. (US 91802, 91803, 91805, 91806, 91807, 91808, UTC 13259) Orobanchaceae Aphyllon Aphyllon fasciculatum (Nutt.) Torr. & A.Gray (COLO 29726, 29757, 29763) Orobanche Orobanche ludoviciana Nutt. (COLO 29766) Oxalidaceae Oxalis Oxalis dillenii Jacq. (COLO 29782) Oxalis stricta L. (COLO 29798, KHD 1521, 3128, 22975, 22976, 37831) Papaveraceae Argemone Argemone hispida A.Gray (COLO 29839, 29841, CS 8835, UTC 174639) Argemone polyanthemos ( Fedde) Ownbey (COLO 29848, 29850, 29859, 43393, 518132, CS 8825, KHD 39529, 47335, RM 97492) Plantaginaceae Plantago Plantago elongata Pursh (COLO 21237, 30073) Plantago eriopoda Torr. (COLO 30107, KHD 1643) Plantago lanceolata L. (COLO 30090, 30091, KHD 1644) Plantago major L. (COLO 30101, 30105, 30106, KHD 1645, 25260, RM 485467) Plantago patagonica Jacq. (COLO 30112, 30123, CS 19323, 19327, KHD 1611, 1653, 1654, 1737) Poaceae Achnatherum Achnatherum hymenoides (Roem. & Schult.) Barkworth (COLO 20884, 106089, KHD 5166, 5172, 39305, 39327, MO 2974620, RM 49378, 93004) Oryzopsis hymenoides (Roem. & Schult.) Barkworth (UTC 13532) Achnatherum pinetorum (M.E.Jones) Barkworth (CS 1800) Achnatherum robustum (Vasey) Barkworth (COLO 106088) 57

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Agropyron *Agropyron cristatum (L.) Gaertn. (COLO 518103, KHD 4910) Agrostis *Agrostis exarata Trin. (COLO 18293, MO 775742) Agrostis gigantea Roth (COLO 18254, KHD 4814, 11785, 16390, 21811) Agrostis scabra Willd. (COLO 18324, MO 778978) Agrostis stolonifera L. (MO 775741, 928268) Alopecurus Alopecurus aequalis Sobol. (COLO 18474, 104120, MO 2872654, RM 243910) Alopecurus arundinaceus Poir. (COLO 500082, 518069, KHD 5906, 36828, 46911) Andropogon Andropogon gerardii Vitman (MO 778968) Andropogon hallii Hack. (COLO 18567, 402181, KHD 20157) Anisantha   *Anisantha tectorum (L.) Nevski [CO Noxious Weed List C] (COLO 518102 ) Bromus tectorum L. (KHD 5071, 7306, 7307, 7308, 7309, MO 1129346, NY 664579, UTC 5308) Aristida Aristida divaricata Humb. & Bonpl. ex Willd. (KHD 46917) Aristida purpurea Nutt. (COLO 18653, 18674, 106153, 518100, KHD 5073) Aristida purpurea Nutt. var. fendleriana (Steud.) Vasey (GH 267784) Aristida purpurea Nutt. var. longiseta (Steud.) Vasey (KHD 3753, 40249, MO 1041132, 2874095) Avena *Avena fatua L. (KHD 4698, 16330) Avena fatua L. var. sativa (L.) Hausskn. (COLO 18725) Beckmannia Beckmannia syzigachne (Steud.) Fernald (KHD 3747, MO 770518, 2875506) Beckmannia syzigachne (Steud.) Fernald subsp. baicalensis (Kusn.) T.Koyama & Kawano CFES 2012 (COLO 13918, 18750, 18755, 18756, 18757, 106145) Bothriochloa *Bothriochloa bladhii (Retz.) S.T.Blake (KHD 11961) Bothriochloa ischaemum (L.) Keng (KHD 20446) Bothriochloa ischaemum (L.) Keng var. songarica (Rupr.) Celarier & J.R.Harlan CFES 2012 (COLO 392518, KHD 44407, 44408) Bouteloua Bouteloua curtipendula (Michx.) Torr. (COLO 18801, 18811, 106155, KHD 39601, MO 780433, 1713334) Bromopsis *Bromopsis inermis (Leyss.) Holub (COLO 518241, KHD 4639, 14298, 33505, 49281) 58

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Bromus *Bromus briziformis Fisch. & C.A.Mey. (KHD 3494, 3509) Bromus commutatus Schrad. (KHD 28288) Bromus japonicus Thunb. (COLO 518124, KHD 3543, 3544, 3545, 3546, 39532) Bromus secalinus L. (MO 796007) Buchlo‘ Buchlo‘ dactyloides (Nutt.) Engelm. (COLO 19190, 19199, 106084, 518179, KHD 3811, 3850, 3853, 39836, MO 779317, 1207506, 2958399, 2958408, RM 87037, 150622, UTC 5309, 5454) Calamagrostis Calamagrostis canadensis (Michx.) P.Beauv. Calamagrostis canadensis (Michx.) P.Beauv. var. macouniana (Vasey) Stebbins (MO 778988) Calamagrostis purpurascens R.Br. (COLO 100927) Calamovilfa Calamovilfa longifolia (Hook.) Scribn. in Hack. (COLO 19314, 106091, 402182, 518074, KHD 5064, 5065, 20158, 39602, MO 770511 ) Cenchrus Cenchrus longispinus (Hack.) Fernald (COLO 19330, KHD 3944, 3945, 3948) Ceratochloa *Ceratochloa carinata (Hook. & Arn.) Tutin Bromus marginatus Nees ex Steud. (KHD 3551) Chloris *Chloris verticillata Nutt. (COLO 404123, 518135, CS 79384, KHD 17685, 17838, 20206, 21456, 21936, 30489) Chloris virgata Sw. (COLO 404124, CS 79380, 92629, KHD 17326, 17839, 33928) Chondrosum Chondrosum gracile Kunth (COLO 18840, 40664, 106154, 518145, MO 778999, 2875953) Bouteloua gracilis (Kunth) Lag. ex Griffiths (KHD 39594, 46494, RM 48775) Chondrosum hirsutum (Lag.) Sweet Bouteloua hirsuta Lag. var. hirsuta (KHD 20153) Chondrosum prostratum (Lag.) Sweet (COLO 418721) Bouteloua simplex Lag. (KHD 21466, 21536, 39496) Critesion *Critesion brachyantherum (Nevski) Barkworth & D.R.Dewey Hordeum brachyantherum Nevski (KHD 4607) Critesion jubatum (L.) Nevski (COLO 20403, 20446, 518123, 518229) Hordeum jubatum L. (KHD 4594, 4603, 39537, MO 778835, 2506628, 2506634, 5391750) Hordeum jubatum nothosubsp. intermedium Bowden (MO 1618063, 2506537) Critesion murinum (L.) .Lš ve Critesion glaucum (Steud.) .Lš ve (COLO 404119) Hordeum murinum L. subsp. glaucum (Steud.) Tzvelev (KHD 14300, 21817, 39833, UTC 9370) Critesion pusillum (Nutt.) .Lšve Hordeum pusillum Nutt. (KHD 10550, 40529, MO 2506414, RM 421514, UTC 9531) 59

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Cylindropyrum   *Cylindropyrum cylindricum (Host) .Lšve [CO Noxious Weed List B] (COLO 518239) Aegilops cylindrica Host (KHD 39843, 47318) Cynodon *Cynodon dactylon (L.) Pers. (CS 30989) Dactylis *Dactylis glomerata L. (COLO 19379, CS 77589, KHD 3648, 3649) Dichanthelium Dichanthelium oligosanthes (Schult.) Gould Dichanthelium oligosanthes (Schult.) Gould var. scribnerianum (Nash) Gould (COLO 21048) Digitaria *Digitaria ischaemum (Schreb.) Muhl. (COLO 403886, 426257, KHD 20197, 20218, 23425) Digitaria sanguinalis (L.) Scop. (COLO 19555, KHD 3961, 3962, 16694, 19858, 20945) Diplachne Diplachne dubia (Kunth) Scribn. Leptochloa dubia Kunth (KHD 23437, 43887) Diplachne fascicularis (Lam.) P.Beauv. (COLO 426258) Leptochloa fascicularis (Lam.) Gray (UTC 9704) Leptochloa fusca subsp. fascicularis (Lam.) N.Snow (CS 79382, KHD 21838, 23440, 44442, MO 781116) Distichlis Distichlis stricta (Torr.) Rydb. (COLO 19568, 19575, 518134, KHD 3665, 19952, 39814 ) Distichlis spicata (L.) Greene (KHD 3663, 3664, 46471) Distichlis spicata (L.) Greene var. stricta (Torr.) Scribn. (MO 778958, 2964189) Echinochloa *Echinochloa crus-galli (L.) P.Beauvois (COLO 21016, 21033, 106086, 518080, KHD 3975, 3976, 21836) Echinochloa muricata (P.Beauvois) Fernald (MO 778964, 1607601, RM 9821) Eleusine *Eleusine indica (L.) Gaertn. (COLO 392516, 392527, 392562, KHD 20321, 20477, 21837, 35721) Elymus Elymus canadensis L. (COLO 19635, 19649, 518079, KHD 1817, 5449) Elymus brachystachys Scribn. & C.R. Ball (MO 928264) Elymus elymoides (Raf.) Swezey (COLO 518096, CS 3844, KHD 4659, 4795, 40244, 47317, 49086, RM 427516, UTC 7722) Elymus sitanion Schult. (MO 4153866, 4153882) Elymus trachycaulus (Link) Gould ex Shinners (COLO 17992, 22989, KHD 2273, 5386, 5438, MO 768567, 768990, 1057784, 1810691) Elymus virginicus L. Elymus virginicus L. var. submuticus Hook. (MO 2965232) Elytrigia   *Elytrigia repens (L.) Nevski [CO Noxious Weed List B] Elymus repens (L.) Gould (MO 928267) 60

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Eragrostis *Eragrostis barrelieri Daveau (COLO 419825) Eragrostis cilianensis (All.) Vignolo ex Janch. (COLO 19797, 19802, KHD 3667, 3668, 3669, 16085, 16696, 46472) *Eragrostis curvula (Schrad.) Nees (COLO 425547) Eragrostis lutescens Scribn. (CS: CNHP D-2013-139 ) Eragrostis minor Host (COLO 398280, KHD 20447) Eragrostis pilosa (L.) P.Beauv. (KHD 4326, MO 2966273) Eragrostis pectinacea (Michx.) Nees (COLO 19819) Eragrostis spectabilis (Pursh) Steud. (CS: CNHP D-2013-141 ) Festuca Festuca arizonica Vasey (KHD 25135) Festuca baffinensis Polunin (COLO 106156) Festuca brachyphylla Schultes Festuca ovina L. (MO 771795) Festuca rubra L. (KHD 25137) Festuca saximontana Rydb. (COLO 106158) Festuca trachyphylla (Hack.) Krajina (KHD 40554) Glyceria Glyceria grandis S.Watson (COLO 20240, KHD 4501, 45920, UTC 9003, 9084) Glyceria striata (Lam.) Hitchc. (CS: CNHP D-2013-133, -134, -160 ) Hesperostipa Hesperostipa comata (Trin. & Rupr.) Barkworth (COLO 22080, 22987, 539748, KHD 4915, 5305, 5309, 40248, RM 49380, 92920, UTC 9332) Stipa comata Trin. & Rupr. (MO 2489718, 2489721, 778870) Hordeum *Hordeum vulgare L. (COLO 20454, KHD 38459) Koeleria Koeleria macrantha (Ledeb.) Schult. (COLO 20551, 106151, 518224, KHD 4765, 4797, MO 2971389) Leymus Leymus triticoides (Buckley) Pilg. (CS: CNHP D-2013-174 ) Lolium *Lolium perenne L. Lolium multiflorum Lam. (KHD 3928, 4630) Munroa Munroa squarrosa (Nutt.) Torr. (COLO 20662, 106085, 402190, KHD 20152, 47316, MO 778998, 2974386, 2974389, RM 159883, 180735) 61

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Muhlenbergia Muhlenbergia asperifolia (Nees & Meyen) Parodi (COLO 20689, 20690, 20692, 22990, KHD 2363, MO 778976, 795477, 2973547, RM 48803, UTC 12688,12707) Muhlenbergia montana (Nutt.) Hitchc. (UMO 18878) Muhlenbergia racemosa (Michx.) Britton, Sterns, & Poggenb. (COLO 20780, 106162, MO 778843) Muhlenbergia schreberi J.F.Gmel. (COLO 392528, 398298, CS 1894, KHD 3911, 11972, 17598, 20323, 20751, 46293) Muhlenbergia torreyi (Kunth) Hitchc. ex Bush (COLO 20831, MO 928199) Nasella Nassella viridula (Trin.) Barkworth (COLO 22171, 518101, KHD 5070, 5874, 5875, 5882, 5899, 40237, MO 778866, 1057019, RM 107525) Panicum *Panicum capillare L. (COLO 20964, 518176, 518226, KHD 4018, 39857, 47315, MO 778969, 780719, 2974899) Panicum dichotomiflorum Michx. (COLO 416175, 459786, 468603, KHD 9830, 21543, 31308)  * Panicum miliaceum L. [CO Noxious Weed List C] (COLO 398279, KHD 4055, 17837, 20154, 20205) Panicum obtusum Kunth (KHD 39495) Panicum virgatum L. (COLO 21088, 21089, 21090, 21091, 21095, 21096, 518116, KHD 4093, 47314, MO 773168, 775829, 2975581, 2975585, UTC 13888) Pascopyrum Pascopyrum smithii (Rydb.) .Lšve (COLO 18111, 518125, KHD 5387, 5388, 8602, 39536, 46292, 46507) Elytrigia smithii (Rydb.) .Lšve (MO 2555095, 2553858, 2554763, 2555672, 2555760, 2555591, 2553201, 2553608, 2554099) Paspalum Paspalum setaceum Michx. Paspalum setaceum Michx. var. stramineum (Nash) D.J.Banks (COLO 402183, KHD 20325) Phalaroides *Phalaroides arundinacea (L.) Rauschert (COLO 518236, KHD 19827, 39548) Phleum *Phleum pratense L. (KHD 4442, 5190) Phragmites   *Phragmites australis (Cav.) Trin. [CO Noxious Weed Watch List] (COLO 518182, 518228, CS 77588, KHD 46457, 46486) Poa *Poa annua L. (COLO 21268, 21313, 518233, KHD 556, 39192) Poa arida Vasey (KHD 3698) Poa compressa L. (COLO 21349, KHD 3707, 3708, 23334) Poa fendleriana (Steud.) Vasey (CS 3085, KHD 5974) Poa nemoralis L. Poa nemoralis L. subsp. interior (Rydb.) W.A.Weber (COLO 21475) Poa pratensis L. (KHD 5923) Poa secunda J.Presl. Poa sandbergii Vasey (MO 928192) 62

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Polypogon *Polypogon interruptus Kunth (KHD 40136) Polypogon monspeliensis (L.) Desf. (COLO 21801, 518078, 518237, KHD 5212, 5213, 23333, 39551, 39585) Puccinellia Puccinellia nuttalliana (Schult.) Hitchc. Puccinellia airoides S.Watson & J.M.Coult. (COLO 21745, 21815, 21816) Puccinellia distans (L.) Parl. (KHD 6017) Schedonnardus Schedonnardus paniculatus (Nutt.) Branner & Coville (COLO 518077, KHD 3896, 26345, 39598, MO 768953, 778852) Schedonorus *Schedonorus arundinacea (Schreb.) Dumort. Festuca arundinacea Schreb. (COLO 518153, KHD 21069, 39558) Schedonorus pratensis (Huds.) P.Beav. Festuca pratensis Huds. (KHD 4354, 4355, 4356, MO 1129367) Secale *Secale cereale L. (KHD 39523, 40169) Setaria *Setaria pumila (Poir.) Roem. & Schult. (KHD 20005, 20156, 20338, 23584) Setaria glauca (L.) P.Beauv. (COLO 21867) Setaria verticillata (L.) P.Beauv. (CS 29576, KHD 16695, 20065, 37828, 38809, 38812) Setaria viridis (L.) P.Beauv. (COLO 21888, 21890, KHD 4097, 4098, 4099, 4100, 19877, 37829, MO 778946) Sorghum   *Sorghum halepense (L.) Pers. [CO Noxious Weed List C] (KHD 23439) Spartina *Spartina gracilis Trin. (COLO 21948) Spartina pectinata Bosc. ex Link (COLO 21938, 106152) Sporobolus Sporobolus airoides (Torr.) Torr. (COLO 518076, KHD 39599, 46492) Sporobolus cryptandrus (Torr.) A.Gray (COLO 22008, 106161, 404118, CS 2085, KHD 1807, 5298, 15238, 20159, 46513, MO 781094, 795518, 3055109, 3055113) Thinopyrum *Thinopyrum intermedium (Host) Barkworth & D.R.Dewey (MO 770208) Thinopyrum ponticum (Podp.) Barkworth & D.R.Dewey (COLO 518146, KHD 39597) Trisetum Trisetum spicatum (L.) K.Richt. Trisetum spicatum (L.) K.Richt. subsp. congdonii (Scribn. & Merr.) HultŽn CFES 2012 (COLO 106090) Triticum *Triticum aestivum L. (KHD 4674) Vulpia Vulpia octoflora (Walter) Rydb. (COLO 106157, KHD 4390, 39079, MO 771839, 820383, RM 93066) 63

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Polemoniaceae Collomia Collomia linearis Nutt. (COLO 30186) Gilia Gilia ophthalmoides Brand (COLO 30429) Gilia pinnatifida Nutt. Aliciella pinnatifida (Nutt. ex A.Gray) J.M.Porter (COLO 30353, 30354, 30369, 30370, 425954, KHD 1818, 39520) Ipomopsis Ipomopsis aggregata (Pursh) V.E.Grant Ipomopsis aggregata (Pursh) V.E.Grant subsp. collina (Greene) Wilken & Allard (COLO 30328, 30338, CS 26880) Ipomopsis laxiflora (Coulter) V.E.Grant (COLO 30433, 30434, 30437, 30453, 32514, 518094, CS 16813, 26949, 60638, 60639, KHD 1822, 39805) Leptodactylon Leptodactylon pungens (Torr.) Rydb. (COLO 30476, 30477, CS 16650, 60605) Linanthus pungens (Torr.) J.M.Porter & L.A.Johnson (RM 92973, 97172) Microsteris Microsteris gracilis ( Hook.) Greene Microsteris gracilis ( Douglas ex Hook.) Greene subsp. humilis (Greene) V.E.Grant (COLO 30649, CS 16888, 16891, RM 29322, 164287) Polygonaceae Acetosella *Acetosella vulgaris Fourr. Rumex acetosella L. (KHD 2375) Eriogonum Eriogonum annuum Nutt. (COLO 31081, 31087, 31089, 103020, CS 26093, UTC 7965) Eriogonum cernuum Nutt. (COLO 31107, SJNM 12727, UTC 8002) Eriogonum effusum Nutt. (COLO 31140, 31141, 31142, 31155, 31161, 31251, KHD 2174, 37744, 39595, 46515, KSC 87320, 87321, UTC 8385) Eriogonum microthecum var. effusum (Nutt.) Torr. & A.Gray (RM 84656, 144360) Eriogonum umbellatum Torr. (COLO 254429) Fallopia *Fallopia baldshuanica (Regel) Holub Fallopia aubertii (L.Henry) Holub (COLO 287963) Fallopia convolvulus (L.) .Lšve (COLO 31629, 31636, 31645) Fallopia japonica (Houtt.) Ronse Decr. (KHD 37468) 64

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Persicaria Persicaria amphibia (L.) Gray (CS: CNHP D-2013-142 ) Persicaria bicornis (Raf.) Nieuwl. (COLO 518082) Persicaria coccinea (Muhl.) Greene Persicaria wardii Greene (US 47280, 133945) Persicaria hydropiper (L.) Opiz (KHD 44448) Persicaria lapathifolia (L.) Gray (COLO 518175, RM 89002) Persicaria maculata (L.) Gray (COLO 31672, 31673, 31714, 31727, 31744, 518213) Persicaria maculosa Gray Polygonum persicaria L. (KHD 2353, 25589) Persicaria pensylvanica (L.) M.G—mez (COLO 31675, 31677, 31678) Persicaria punctata (Elliott) Small (COLO 31514, 31754, 31786) Polygonum *Polygonum argyrocoleon Steud. (COLO 518204, KHD 40113) Polygonum aviculare L. (KHD 2326, 2327, 2328, 25321, 25322) Polygonum arenastrum Boreau (COLO 31531, 31533, 31760, 100909) Polygonum douglasii Greene (KHD 2346, 25320, RM 2561) Polygonum erectum L. (KHD 2347, 2348, 28072, UTC 22637) Polygonum ramosissimum Michx. (COLO 31535, 31713, 31757, 31759, 31761, CS 6715, KHD 4964, KSC 89936, RM 92417, 92971, 124060, 302904) Rumex *Rumex crispus L. (COLO 31945, 32038, KHD 2378, 2379, 40116) Rumex hymenosepalus Torr. [not in CFES2012. West Slope only] (KSC 98098) Rumex triangulivalvis (Danser) Rech.f. (COLO 31934, 31992) Rumex utahensis Rech. (CS: CNHP D-2013-143 ) Rumex venosus Pursh (COLO 32018, 32020, 32021, KHD 2394, 47313, RM 48954, 95449) Pontederiaceae Heteranthera Heteranthera limosa (Sw.) Willd. (COLO 32062, 32063, 32064, 43813, CS 4911, RM 89026) Portulacaceae Phemeranthus Phemeranthus parviflorus (Nutt.) Kiger Phemeranthus confertiflorus (Greene) Hershkovitz (UTC 13889) Talinum parviflorum Nutt. (COLO 32180, 32189, 32194, 32196) Portulaca *Portulaca oleracea L. (KHD 25329, 40074, UTC 22650) 65

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Potamogetonaceae Potamogeton Potamogeton crispus L. (COLO, KHD: Majack 13130, 13131 ) Potamogeton foliosus Raf. (COLO 28291, RM 48860) Potamogeton nodosus Poir. (KHD: Majack 13155 CS: CNHP D-2013-92, -96 ) Potamogeton pusillus L. (COLO, KHD: Majack 13155 CS: CNHP D-2013-91 ) Stuckenia Stuckenia filiformis (Pers.) Bšrner (KHD 7561, 7565) Stuckenia pectinata (L.) Bšrner (COLO 28265, CS 952, RM 93026) Primulaceae Anagallis Anagallis arvensis L. (KHD 46929) Androsace Androsace occidentalis Pursh (COLO 32226, KHD 45202) Glaux Glaux maritima L. (CS 15998, 16007, RM 88119, 144485) Glaux maritima L. var. angustifolia B.Boivin (COLO 8684, 32498, 32502) Lysimachia Lysimachia ciliata L. (COLO 32516) Ranunculaceae Anemonidium Anemonidium canadense (L.) .Lšve & D.Lšve (COLO 32755, CS 26052) Batrachium Batrachium aquatile (L.) Dumort. Batrachium trichophyllum (Chaix) Bosch (COLO 106067) Ranunculus aquatilis L. (KSC 74439) Ranunculus aquatilis L. var. diffusus With. (KHD 2809, RM 87120) Ceratocephala *Ceratocephala orthoceras DC. (COLO 451876) Clematis Clematis ligusticifolia Nutt. (COLO 33317, 33336, 33342, 33358, 106041, RM 97439)  * Clematis orientalis L. [CO Noxious Weed List B] (KHD 43894 ) Halerpestes Halerpestes cymbalaria (Pursh) Greene Halerpestes cymbalaria (Pursh) Greene subsp. saximontana (Fernald) Moldenke CFES 2012 (COLO 34338, 34339, 34343) Ranunculus cymbalaria Pursh (KHD 4624, KSC 74489) 66

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Hecatonia *Hecatonia scelerata (L.) Fourr. (COLO 34250, 34252) Ranunculus sceleratus L. (KHD 45919) Ranunculus sceleratus L. var. multifidus Nutt. (CS 26042) Myosurus Myosurus minimus L (COLO 33882) Ranunculus Ranunculus acriformis A.Gray (KHD 25647) Ranunculus gmelinii DC. (KHD 43080) Ranunculus macounii Britton (COLO 34137, 34189) Ranunculus pensylvanicus L. (KSC 74662) Ranunculus repens L. (COLO 34241, 34282, 54405, KHD 25316, 43384, RM 87977, 169225) Resedaceae Reseda *Reseda lutea L. (COLO, 459787, KHD 9844) Reseda luteola L. (COLO 34585) Rhamnaceae Rhamnus Rhamnus frangula L. (CS: CNHP D-2013-60 ) Rosaceae Agrimonia Agrimonia striata Michx. (COLO 34783) Amelanchier Amelanchier alnifolia ( Nutt.) Nutt. ex M.Roem. (KHD 3087) Argentina Argentina anserina (L.) Rydb. (COLO 35553, 35561) Cerasus Cerasus pumila (L.) Michx. Prunus pumila L. var. besseyi (L.H.Bailey) Gleason (KHD 13736, 13741, RM 87770, 87961, 89010) Cercocarpus Cercocarpus montanus Raf. (KSC 68269) Fragaria Fragaria virginiana P.Miller Fragaria virginiana P.Miller subsp. glauca (S.Watson) Staudt CFES 2012 (COLO 100889) Oreobatus Oreobatus deliciosus (Torr.) Rydb. Rubus deliciosus Torr. (KHD 8880) Padus Padus virginiana (L.) P.Miller Padus virginiana (L.) P.Miller subsp. melanocarpa (A.Nelson) W.A.Weber CFES 2012 (COLO 36201, 106164) 67

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Potentilla *Potentilla norvegica L. (COLO 35905, KHD 25848) Potentilla monspeliensis L. (UTC 7635) Potentilla rivalis Nutt. (COLO 518235, KHD 26713, 39539, 40252) Potentilla supina L. Potentilla supina L. subsp. paradoxa (Nutt. ex Torr. & A.Gray) Soj‡ k (COLO 36018) Potentilla paradoxa Nutt. (UTC 22657) Prunus Prunus americana Marshall (COLO 36117, 36124, KHD 2476, 5523) Rosa Rosa arkansana Porter (COLO 493019, RM 113681, UTC 22754, 22772) Rosa woodsii Lindl. (COLO 36353, 36355, 36392) Sanguisorba *Sanguisorba minor Scop. (COLO 36719) Rubiaceae Galium Galium aparine L. (CS 28558) Galium odoratum (L.) Scop. (COLO 52902) Galium spurium L. (COLO 36883, 106114, 493053, KHD 40532) Galium verum L. (COLO 353882, KHD 14895, 30396) Salicaceae Populus Populus acuminata Rydb. (COLO 37161, 37162, 101015, 103067, KHD 5349,14068) Populus angustifolia E.James (KHD 33164, 33174) Populus deltoides W.Bartram ex Marshall (KHD 5178, 5318) Populus deltoides Marshall subsp. monilifera (Aiton) Ecken. CFES 2012 (COLO 103066, 518114, 518232) Salix Salix amygdaloides Andersson (COLO 37276, 37277, 37302, KHD 2850, 14099, RM 92926, 165984, 718082) Salix exigua Nutt. (COLO 37518, 37519, 37520, 37525, 37526, 37528, 37529, 37611, 37612, 518240, CS 44897, KHD 4232, 19962, 39549, RM 718080) Salix famelica (C.R.Ball) Argus (CS: CNHP D-2013-61 ) Salix x fragilis L. (COLO 37573, 37574, KHD 14132, RM 718081) Salix irrorata Andersson (COLO 37622) Salix lasiandra Benth. Salix lasiandra Benth. subsp. caudata (Nutt.) A.E.Murray CFES 2012 (COLO 37477, 37479, 37481, 37482) Salix lucida var. caudata (Nutt.) Cronquist (KHD 3201) 68

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Salix ligulifolia C.R.Ball ex C.K.Schneid. (COLO 37303 ) Salix purpurea L. (COLO 37831, CS 44382) Saururaceae Anemopsis *Anemopsis californica (Nutt.) Hook. & Arn. (COLO 518081, KHD 1730, 46519, 47321, 47322) Scrophulariaceae Agalinis Agalinis tenuifolia Raf. (COLO 39411, 39422) Bacopa Bacopa rotundifolia Wettst. (COLO 39007, CS 18759, RM 88960) Collinsia Collinsia parviflora Lindl. (COLO 39335, 39342, KHD 14871, RM 313309) Limosella Limosella aquatica L. (COLO 39436, 39438, 39441, CS 18764, RM 88945) Linaria   *Linaria vulgaris P.Mill. [CO Noxious Weed List B] (UCD) Lindernia Lindernia dubia (L.) Pennell Lindernia dubia (L.) Pennell var. anagallidea (Michx.) Cooperr. CFES 2012 (COLO 39435, CS 18770, 18771, RM 88961) Mimulus Mimulus glabratus Kunth (COLO 493050, KHD 14946) Mimulus ringens L. [Not listed in CFES2012. Extirpated?] (COLO 496405) Penstemon Penstemon albidus Nutt. (COLO 39966, 39967, 39971, KHD 4372, 15093, 39524, NY 541823, 541827, 541835, 541836, RM 56985, 93020, 170996) Penstemon angustifolius Nutt. (COLO 40020, 40022, 40036, 40041, 518067, KHD 13682, 47320, NY 569878, 569879, RM 93053) Penstemon glaber Pursh Penstemon glaber var. alpinus (Torr.) A.Gray (RM 84687, 88070, 97448) Penstemon glaber Pursh var. glaber (COLO 39988, 39996, NY 600270, 600271, 600272, 600273) Penstemon secundiflorus Benth. (COLO 40348, 40350, 40353, 40364, 40365, NY 651873, 651874, 651891, 651909) # Penstemon virens Pennell [Endemic ] (KHD 14800, NY 763271) Penstemon virgatus A.Gray Penstemon virgatus A.Gray subsp. asa-grayi Crosswhite CFES 2012 (COLO 40477, NY 763345, RM 95434) Pocilla *Pocilla polita (E.Fries) Fourr. (COLO 104151, 200531, 419818) Veronica persica Poir. (KHD 602, 11625, 21818, 32314, 43418) Scrophularia Scrophularia lanceolata Pursh (COLO 40646) 69

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Verbascum   *Verbascum blattaria L. [CO Noxious Weed List B] (KHD 43395)  * Verbascum thapsus L. [CO Noxious Weed List C] (COLO 518178, 518227, KHD 39839) Veronica Veronica americana Schwein. ex Benth. (COLO 40761, 40763) Veronica catenata Pennell (COLO 518157, KHD 3414, 15437, 39552) Veronica anagallis-aquatica L. (CS 95180) Veronica peregrina L. Veronica peregrina L. subsp. xalapensis (Kunth) Pennell (COLO 40803, KHD 832, 15444, 20470, 46921) Simaroubaceae Ailanthus *Ailanthus altissima (Mill.) Swingle (COLO 541412, KHD 48497) Solanaceae Datura Datura wrightii Regel (KHD 23583) Hyoscyamus   *Hyoscyamus niger L. [CO Noxious Weed List B] (KHD 20144) Lycium *Lycium barbarum L. (KHD 23034, 23035) Physalis Physalis hederifolia A.Gray Physalis hederifolia A.Gray var. comata (Rydb.) Waterf. (COLO 41031, CS 28586, 45182) Physalis heterophylla Nees (COLO 40984, 40987, 40990) Physalis pumila Nutt. (CS 18129, 18146, 18149) Physalis pumila Nutt. subsp. hispida (Waterfall) Hinton (COLO 41000) Physalis virginiana P.Miller (COLO 40999, 41015, 41034, 101432, KHD 39817) Quincula Quincula lobata (Torr.) Raf. (COLO 41003, 41006, 41013, CS 18135, RM 88954) Solanum Solanum americanum P.Miller (COLO 41064, 41066, 492647, KHD 22413, 43047) Solanum nigrum L. (NY 759944) Solanum dulcamara L. (COLO 443793, KHD 28066) Solanum physalifolium Rusby Solanum physalifolium Rusby var. nitidibaccatum (Bitter) Edmonds (COLO 41109, KHD 22395, 42834) Solanum rostratum Dunal (COLO 41076, 41079, 41080, 103051, 518177, KHD 15511, 15517, 16789, 39856, NY 820930, 820933, 820934, RM 97399) Solanum triflorum Nutt. (COLO 41086, 41098, CS 96499, KHD 3138, 15521, 15521) 70

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Sparganiaceae Sparganium Sparganium eurycarpum Engelm. [CNHP tracked: G5 S2] (COLO 41147, CS: CNHP D-2013-04 ) Tamaricaceae Tamarix   *Tamarix chinensis Lour. [CO Noxious Weed List B] Tamarix ramosissima Ledeb. (KHD 40187) Typhaceae Typha Typha angustifolia L (KHD 40247, RM 87773) Typha latifolia L. (KHD 46912) Ulmaceae Ulmus *Ulmus pumila L. (COLO 518107, KHD 39522) Urticaceae Urtica Urtica gracilis Aiton (COLO 42074) Verbenaceae Glandularia Glandularia bipinnatifida (Nutt.) Nutt. (COLO 42239, 42240, 42241, 42246, CS 17521) Phyla Phyla cuneifolia (Torr.) Greene (COLO 42220, 42223, 42231, CS 17578) Verbena *Verbena bracteata Lag. & Rodr. (COLO 42253, 42254, 42256, 42260, 101510, 518088, CS 17512, KHD 39512) Verbena hastata L. (COLO 42319, 42336, CS 17528, KHD 23166) Verbena stricta Vent. (COLO 42334, 104117) Violaceae Viola Viola nuttallii Pursh (COLO 42530, 42543, 42545, 42550, KHD 39518, RM 92918, WSC 7066 ) Viola odorata L. (KHD 33197) Viola sororia Willd. Viola sororia Willd. var. affinis (Le Conte) L.E. McKinney (COLO 42520) Viscaceae Arceuthobium Arceuthobium vaginatum (Willd.) K.Presl Arceuthobium vaginatum (Willd.) K.Presl subsp. cryptopodum (Engelm.) Hawksworth & Wiens (US 3615904) 71

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Vitaceae Parthenocissus Parthenocissus vitacea (Knerr) A.S.Hitchcock (KHD 40117) Zannichelliaceae Zannichellia Zannichellia palustris L. (COLO 493046) Zygophyllaceae Tribulus   *Tribulus terrestris L. [CO Noxious Weed List C] (COLO 42785, 42791, 518151, KHD 16566, 16557, 37819, 39561, 47319) 72

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CHAPTER III NEW AND NOTEWORTHY COLLECTIONS OF PONDWEEDS (POTAMOGETONACEAE) IN COLORADO Introduction Potamogetonaceae (pondweed family) are one of the most ecologically important plant families in aquatic ecosystems (Preston 1995; Haynes and Hellquist 2000), comprising approximately 100 species worldwide of submerged and floating-leaved aquatic vascular plants in three genera; Potamogeton L., Stuckenia Bšrner, and Groenlandia J.Gay (Haynes & Hellquist 2000). The family is taxonomically complex, with some species (e.g., Potamogeton pusillus L.) having as many as 51 synonyms (Wiegleb and Kaplan 1998). Pondweeds are notoriously difficult to identify, due to microscopic diagnostic characters and a propensity for phenotypic plasticity and hybridization (Wiegleb and Kaplan 1998) Collection biases have resulted in a paucity of pondweed accessions throughout much of the family's range, and existing collections do not fully represent the range of plasticity within species. It is not uncommon for herbarium specimens to be misidentified, even confused with members of other aquatic families (e.g., Zannichelliaceae, Ruppiaceae). Even expert determinations using morphological characters alone cannot reliably identify some species of linear-leaved pondweeds, such as Potamogeton clystocarpus Fernald (Whittall et al. 2004) and Potamogeton foliosus Raf. subsp. fibrillosus (Fernald) Haynes & C.B. Hellquist (Hellquist et al. 2014), for which expert determinations were refuted by molecular techniques. In their treatment of 73

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Potamogetonaceae in Flora of North America (FNA) Haynes and Hellquist (2000) suggest that only fruiting specimens should be collected. This practice of avoiding vegetative specimens has not contributed to a better understanding of pondweed diversity in North America, as the majority of hybrids do not produce fruit (Preston 1995). For example, 33 pondweed hybrids are recognized in the USDA PLANTS database (USDA, NRCS 2014), and additional hybrids continue to be described with the increased use of molecular techniques (Kaplan et al. 2009; Les et al. 2009). Documenting plant distributions through herbarium specimens is the first step toward accurate assessment and subsequent conservation of biodiversity. Aquatic ecosystems are especially susceptible during the current extinction crisis because of the numerous human impacts that have lead to the widespread degradation of waterbodies. Not surprisingly, it has been shown that freshwater fauna experience extinction rates five time higher than those of terrestrial animals (Riccardi and Rasmussen 1999). Despite the critical importance of aquatic plants in freshwater ecosystems, they are often neglected during botanical surveys, resulting in gaps in knowledge about their distribution, taxonomy, and ecology (Shelth et al. 2012). As demonstrated by recent survey of aquatic habitats in Yellowstone National Park (Hellquist et al. 2014), focused fieldwork can result in many new and noteworthy collections, even in areas where the flora is considered well-documented. As expected, pondweeds are undercollected in Colorado, thereby contributing to a poor understanding of their diversity and distribution in the state. Prior to my fieldwork (2013 and 2014), herbarium vouchers documented 13 species of Potamogeton and three 74

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species of Stuckenia, after nomenclature was standardized following FNA (Haynes and Hellquist 2000). Only one species, Potamogeton diversifolius Raf., has been assigned a subnational conservation status in Colorado (S1, critically imperiled). The remaining native pondweed species in Colorado are not ranked (SNR) due to insufficient data (NatureServe 2014). The first and only rigorous survey of aquatic plants in Colorado was completed 50 years ago, culminating in the publication of The True Aquatic Plants of Colorado (Harrington and Matsumura 1955). Collections made by Harrington and Matsumura in the 1950s frequently remain the only specimen(s) vouchering the occurrence of a species in a county. Current distribution maps for pondweeds, such as those included in the Field Guide to Colorado's Wetland Plants (Culver and Lemly 2013), were compiled using existing herbarium records. Due to the small number of current and historic collections of pondweeds in Colorado, these maps likely reflect patterns in collecting efforts rather than actual distributions. In fact, the authors noted that several pondweed species are uncommon and likely undercollected: Potamogeton am pl i f ol i us Tuck., Potamogeton epihydrus Raf., Potamogeton illinoensis Morong, Potamogeton natans L. In the most recent treatment of the flora of Colorado, Weber and Wittmann (2012a, 2012b) preface the key to Potamogetonaceae by stating that the habitat and distribution of the family cannot be determined from the inadequate number of available specimens. Kelso et al. (2008) also brought attention to the paucity of Colorado pondweed specimens in their treatment of the Pikes Peak flora, and discussed the likelihood that additional species occur in the area. 75

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These uncertainties in the diversity and distribution of pondweed species in Colorado prompted my investigation into the known distribution of the family by using web-based herbarium databases to identify undercollected areas of the state. Here I report the results of my analysis of previous collecting efforts of pondweeds in Colorado, and noteworthy collections from my subsequent fieldwork (2013 and 2014) focusing on filling in these knowledge gaps. Methods Herbarium database mining and site selection To identify gaps in previous collecting efforts of pondweeds in Colorado, I compiled all available records of Potamogeton and Stuckenia from Colorado into a single database. In addition to searching Colorado herbaria, efforts were made to include herbaria outside Colorado that may house accessions not duplicated within the state (e.g., F, KSC, OKLA, MO, NY, US). Specimen records were located by querying web-based herbarium databases (CPNWH, F, KAVU, KSC, MO, SEINet, US), and by contacting herbaria directly (COCO, COLO, GREE, OKL, OKLA). Twenty-six of 82 herbaria (Appendix A) provided specimen label data through online databases (ARIZ, ASC, BRY, CS, F, FLD, FLFO, ID, KAVU, KHD, KSC, MESA, MO, NMC, NY, RM, RMBL, SJNM, UCR, UMO, UTC, WSC) or contributed records via email (COCO, COLO, OKLA). In total, these 26 herbaria house a total of 717 Colorado pondweed specimens collected between 1873 and 2013 (Table 1). I compiled these records into a single database and standardized nomenclature using FNA (Haynes and Hellquist 2000). I made efforts to identify duplicate collections by matching collector numbers, revealing that 76

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more than 200 of the accessions (about 30%) were duplicates. Further analysis revealed that only 462 populations of pondweed have been documented in Colorado since 1873, after repeat collections from the same site were removed. Specimen label data were used to map the approximate location of all sites in Colorado from which pondweeds had been collected (Fig. 1). Records lacking latitude and longitude were georeferenced manually with Google Earth using collection locality descriptions from label data. The resulting map (Fig. 1) revealed eight counties that contained significant aquatic habitat from which pondweeds had not been previously collected: Chaffee, Delta, Denver, Douglas, Grand, Mesa, Pitkin, and Summit counties. Study sites within each county were selected from a variety of aquatic habitats on National Forest or other public lands (Denver and Douglas counties). Efforts were made to include sites from multiple water body types (streams, rivers, ponds, lakes, reservoirs) exhibiting a variety of physical and chemical characters, as well as varying degrees of human modification and disturbance. Field collections and identification In 2013, I surveyed a total of 122 sites in Colorado (Fig. 2) for pondweeds from June through September, focusing on Chaffee, Delta, Denver, Douglas, Mesa, Pitkin, and Summit counties. Because Potamogeton crispus L., the only introduced pondweed in North America, reaches maximum growth and fruits earlier in the season than the native pondweeds, sites were visited early in the season (June to July) and again late in the season (August to September) when possible. In 2014, I surveyed an additional 30 sites in Boulder, Grand, Jackson, Lake, and Larimer counties between August and September 77

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coincident with an aquatic plant survey for Arapaho and Roosevelt National Forests (PO# AG-82AT-P-14-0034). In October 2014, I visited one site on Boulder County Open Space. Collection permits or permissions were obtained from the USDA Forest Service, Boulder County, City of Carbondale, City and County of Denver, and Douglas County Open Space. Forest Service permits did not allow collection of plant material from Wilderness Areas (a significant percent of land in Pitkin and Summit counties), so waterbodies in these areas were not visited. Site surveys were conducted from the shoreline with binoculars or in the water using waders or a kayak. Specimens were collected by hand or with an avalanche probe fitted with an improvised grappling hook. Herbarium vouchers were created from representative specimens of each pondweed population encountered and deposited at the Kathryn Kalmbach Herbarium at the Denver Botanic Gardens (KHD) and the University of Colorado Museum of Natural History (COLO). In addition, I annotated select pondweed specimens at COLO, CS, KHD and RM. Species determinations were made following FNA (Haynes and Hellquist 2000) Results Summary Here I report 62 new and noteworthy collections of pondweeds in Colorado, including a new state record and 34 new county records. Potamogeton robbinsii Oakes was documented in Colorado for the first time. I also documented a new population of Potamogeton amplifolius a species thought to be extirpated in Colorado. I discovered the first known population of Potamogeton diversifolius (critically imperiled) in Douglas 78

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County, as well as documenting the second known population in Boulder County. In total, I documented 29 new county records of native pondweeds, including several rare species with limited distributions in Colorado: Potamogeton epihydrus Potamogeton nodosus Poir., Potamogeton richardsonii (A.Benn) Rydb., and Potamogeton praelongus Wulfen. Additionally, my fieldwork significantly increased the known range of the invasive Potamogeton crispus from three to eight countie s in Colorado. Of the 153 aquatic habitats I surveyed in 2013 and 2014 (Fig. 2), at least one species of pondweed was present at 58 sites (38%). Fifteen species of Potamogeton and three species of Stuckenia were confirmed in Colorado through herbarium database mining and fieldwork in 2013 and 2014 (Table 2). Existing herbarium specimen records were used to provide distributional context for new collections, but the majority of these accessions were not annotated as part of this study. Known distributions (e.g., Culver and Lemly 2013) are based on very few vouchers, and many counties are vouchered by a single, historic accession. As such, one misidentified specimen has the potential to significantly affect the known distribution of a species. All distributions presented here should be considered provisional pending additional fieldwork to fill in knowledge gaps from undercollected areas and the annotation of all existing Colorado pondweed specimens to reflect current taxonomic concepts. A portion of Colorado pondweed specimens were annotated by Robert Haynes or C. Barre Hellquist during the preparation of their treatment for FNA (2000), and I annotated others following their treatment. My limited annotations resulted in a county being removed from the known distribution of five species. Other discrepancies between known distributions presented here and 79

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recently published distribution maps (e.g., Culver and Lemly 2013) are due to the inclusion here of only those counties for which a voucher could be located. Global and subnational conservation ranks are given for each species following NatureServe (2014). Latitude and longitude are reported in WGS84 datum. New and noteworthy collections Potamogeton alpinus Balb. Herbarium accessions collected between 1878 and 2010 document 39 known populations of Potamogeton alpinus in Colorado. The species has been previously documented at 39 sites in 18 of 64 Colorado counties, 11 of these sites documented in eight counties in the last 20 years: Conejos, Grand, Hinsdale, Jackson, Larimer, Rio Grande, San Juan, and Teller counties. Potamogeton alpinus is globally secure (G5), and although it has not been assigned a conservation rank for Colorado (CO SNR), it is ranked imperiled/vulnerable in Wyoming (WY S2S3). I documented seven new populations of P. alpinus in 2013, and collected specimens from one historic site. The first known occurrences of P. alpinus in Chaffee County were documented from Alpine Lake, Chalk Lake, and Cottonwood Lake. Potamogeton alpinus was also vouchered for the first time for Lake County, collected from a beaver pond 3.3 km north of the town of Twin Lakes. Two previously unknown populations were documented in Summit County from Cataract Creek and the Curtain Ponds near Copper Mountain Ski Resort. Although P. alpinus has been vouchered previously for Summit County, only two historic collections exist, one from a pond seven miles southwest of Dillon in 1952 ( Harrington 7132 [CS]; Matsumura 1387 [CS]), and 80

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from the Blue River valley in 1980 ( Wittmann 1211 [RM]). I also documented a previously unknown population in Larimer County north of Chambers Lake, representing the first collection in the county since 1996. I collected from the population in Red Rock Lake in Boulder County collected previously documented by Francis Ramaley in 1907, documenting the persistence of the population in the lake after 107 years. I found that characters of P. alpinus in Colorado diverge slightly from the FNA treatment, as our plants have up to 11 veins in submerged leaves. FNA gives a range of 5-9 veins for submerged leaves (Haynes and Hellquist 2000), whereas Wiegleb and Kaplan list a range of (7-)9-15 veins in their monograph of Potamogeton (1998). Potamogeton alpinus material from Lake County (which exhibited 11 veins in submerged leaves) was verified by Dr. Zdenek Kaplan (Head of Department of Taxonomy and Biosystematics, Institute of Botany, Pr" honice). I also note that although I agree that P. alpinus takes on a reddish tinge in fresh and dry material, it should not be considered diagnostic for the species as implied by Haynes and Hellquist (2000), as I have observed this character in other species (e.g., Potamogeton amplifolius Potamogeton crispus ). USA. COLORADO. ROOSEVELT NATIONAL FOREST. Boulder County : Red Rock Lake, elev. 3099 m, 40.08189, -105.54041, 11 August 2014, M. Majack, C. Beebe, and E. Pansing 14178, 14179 Larimer County : Twin Lakes north of Chambers Lake, elev. 2898 m, 40.631160, -105.834330, 16 August 2014, M. Majack and J. Uhl 14183 SAN ISABEL NATIONAL FOREST. Chaffee County : Cottonwood Lake, elev. 2918 m, 38.78091, -106.27825, 20 July 2013, M. Majack and J. Uhl 13166, 13167 Alpine Lake inlet, elev. 2847 m, 38.71330, -106.29733, 22 July 2013, M. Majack and J. Uhl 13178, 81

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13179, 13183-13185 Chalk Lake, elev. 2658 m, 38.711754, -106.234292, 22 June 2013, M. Majack and J. Uhl 13192 Lake County : beaver pond 3.3 km north of Twin Lakes, elev. 3175 m, 39.111498, -106.388708, 7 September 2014, M. Majack and J. Uhl 14189 WHITE RIVER NATIONAL FOREST. Summit County : Cataract Creek, elev. 2622 m, 39.83682, -106.31123, 25 August 2013, M. Majack and J. Uhl 13270 Curtain Ponds near Gore Range Trailhead, elev. 2950 m, 39.513586, -106.145816, 1 September 2013, M. Majack and J. Uhl 13287-13289 Potamogeton amplifolius Tuck. Prior to my fieldwork, Potamogeton amplifolius was vouchered for Colorado in 1952 by a single accession from Lower Twin Lake in Lake County ( Harrington and Matsumura 7146 [CS]). Lower Twin Lake was impounded by Twin Lakes Dam in 1980 as part of the US Bureau of Reclamation's Fryingpan-Arkansas Project. It is unlikely that P. amplifolius persists in Lower Twin Lake or in the surrounding area, as it has not been collected since 1952, nor was it seen during my surveys in June 2013 or September 2014. Potamogeton amplifolius is not reported for Colorado in FNA (Haynes and Hellquist 2000). It is ranked globally secure (G5), and although its subnational conservation status has not been determined for Colorado (SNR), it is ranked critically imperiled (S1) in the surrounding states of Kansas, Nebraska, and Wyoming. Potamogeton amplifolius was considered extirpated in Colorado until August 2014, when the species was discovered in Strawberry Lake in Grand County. This site represents the only known extant population in Colorado. 82

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USA. COLORADO. ARAPAHO NATIONAL FOREST. Grand County : Strawberry Bench, Strawberry Lake, elev. 2805 m, 40.109988, -105.773083, 21 August 2014, M. Majack, T. Bates, and T. Roberts 14184, 14185 Potamogeton crispus L. Potamogeton crispus is the only non-native pondweed in North America. Herbarium records first document the occurrence of this species in Colorado in 1952. Although it has not yet been assigned legal noxious status in Colorado, it is considered a serious threat in much of the United States, where it has had negative impacts on water quality, native plants and animals, power generation, recreation, and property values (Tamayo and Olden 2014). The invasive qualities of the species are compounded by its ability to begin spring growth in colder temperatures than native species and to reproduce vegetatively involving extensive fragmentation. Existing herbarium records document six populations of P. crispus in Arapahoe, Jefferson, Larimer, and Moffat counties (Fig. 3). The most recent published distribution of P. crispus (Culver and Lemly 2013) indicates that the species is also present in Jackson County. This is likely due to a transcription error of specimen label data from a Jefferson County collection (RM 236913) that was uploaded onto SEINet. I documented six new populations of P. crispus in 2013, adding five counties to the species known distribution in Colorado (Fig. 3). Potamogeton crispus was found in Greys Creek in Chaffee County, an unnamed pond in Bear Creek Greenbelt in Denver County, the South Platte River in Douglas County (Pike National Forest), and the Blue River in Summit County. Of special significance are the first vouchers from Grand Lake 83

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and Shadow Mountain Reservoir in Grand County (Roosevelt National Forest), as these waterbodies are the primary source of water for the Colorado-Big Thompson Project, which diverts water from Colorado's Western Slope to many waterbodies on the Front Range. Based on my limited observations of P. crispus populations, the species appears to create the most significant threat to native species in small waterbodies with slow moving waters (creeks and ponds), where it was observed creating monocultures of dense growth. Although the presence of P. crispus in the Blue and South Platte rivers is disturbing from a dispersal perspective, the plants were vegetative, smaller in size, and growing intermixed with native species (e.g., Stuckenia pectinata (L.) Borner, Zannichellia palustris L., Ranunculus aquatilis L.). Potamogeton crispus almost certainly occurs at additional sites in Colorado; the documentation of which should be a priority. USA. COLORADO. Denver County : Bear Creek Park, pond 17 m north of West Kenyon Blvd., elev. 1635 m, 39.650164, -105.044289, 26 June 2013, M. Majack and E. Pansing 13130, 13131 ARAPAHO NATIONAL FOREST. Grand County : Grand Lake, elev. 2555 m, 40.245490, -105.825680, 15 June 2013, M. Majack 13109 Shadow Mountain Lake, elev. 2555 m, 40.242187, -105.829187, 15 June 2013, M. Majack 13110-13112 PIKE NATIONAL FOREST. Douglas County : South Platte River, elev. 1902 m, 39.342430, -105.179500, 8 September 2013, M. Majack and L. Bruederle 13298, 13299 SAN ISABEL NATIONAL FOREST. Chaffee County : Greys Creek, elev. 2646 m, 38.420448, -106.128926, 19 July 2013, M. Majack and J. Uhl 13156-13160 WHITE RIVER NATIONAL FOREST. Summit County : Blue River, elev. 2575 m, 39.727223, -106.132666, 25 August 2013, M. Majack and J. Uhl 13271, 13272 84

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Potamogeton diversifolius Raf. Potamogeton diversifolius is ranked critically imperiled in Colorado (G5 CO S1) and is the only pondweed in the state assigned a subnational conservation rank. The species and the Potamogeton diversifolius Herbaceous Vegetation community type (G1? CO S1?) are fully tracked by the Colorado Natural Heritage Program (CNHP 2013a, 2013b). The USDA Forest Service Rocky Mountain Region evaluated P. diversifolius for Sensitive Species status but determined that there was insufficient information available to make a recommendation (Burkhart and Crook 2002). Prior to 2013, herbarium records documented P. diversifolius at seven sites in five counties in Colorado, exclusively along the Front Range. One accession collected from a pond near Horsetooth Reservoir in 1962 vouchers Larimer County ( Norton s.n. [CS]) but no populations appear to have been collected from the county in over 50 years. Two populations have been documented in El Paso County, in a pond between Peyton and Falcon in 1957 ( Penland 4940 [COCO]), and in 2000 at the intersection of Squirrel and Walker roads ( Leininger 18 [CS]). In Las Animas County, P. diversifolius has been documented in a side canyon of the Purgatoire River in 1985 ( Galatowitsch 938 [CS]) and in 2010 at J.E. Canyon Ranch ( Ackerfield and Islam 3516 [CS]). One population is known from Weld County, collected from Pawnee National Grasslands in 1995 ( Hazlett 9311 [COLO, RM]). Potamogeton diversifolius is also reported for Baca County (Culver and Lemly 2013) but does not appear to be vouchered. Yuma County was removed from the known distribution of P. diversifolius as the only known voucher for the county ( Clark 1836 [COLO]) was determined to be Potamogeton foliosus Raf. (det. Majack 2014). 85

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My fieldwork documented two previously unknown populations of P. diversifolius in Colorado, bringing the number of known sites to nine total. I documented the first known occurrence of the species in Douglas County at Prairie Canyon Ranch (Douglas County Open Space). The plants were growing in the typical habitat of the species in Colorado, in plunge pools in sedimentary rock (here, Castle Rock Conglomerate). When plants were collected in June of 2013, the creek bed was dry, with the exception of three small pools, all of which contained P. diversifolius in fruit and flower. In 2014, I documented a second population of P. diversifolius at Heil Valley Ranch (Boulder County Open Space). Potamogeton diversifolius was documented previously in Boulder County in 1990 from Silver Lake Ditch ( Weber and Dahnke 18097 [BRY, COLO, RM, UCR, UTC]). It is probable that the population persists, as Silver Lake Ditch is on City of Boulder Open Space, but no specimens have been collected since it was originally documented in 1990. The population I documented in Boulder County at Heil Valley Ranch occurs in an abandoned sandstone quarry (Lyons Sandstone). This population represents the highest known elevation for P. diversifolius in Colorado, at 2075 meters. Although P. diversifolius has been documented from sites not associated with sandstone pools, my observations from Boulder and Douglas counties suggest that the species has an affinity for this habitat type. At Prairie Canyon Ranch in Douglas County, four adjacent waterbodies were surveyed (two ponds, one permanent creek and one intermittent creek, with a maximum of 900 meters separation), but P. diversifolius was only present in plunge pools in the intermittent creek. Similarly, at Heil Valley Ranch in Boulder County, P. diversifolius was absent from the only other open 86

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water in the area, a pond less than a kilometer from the quarry. These observations, albeit limited to two sites, imply that further fieldwork on the Front Range focused on areas of exposed sandstone could likely result in discoveries of additional populations of P. diversifolius. USA. COLORADO. Boulder County : Boulder County Open Space: Heil Valley Ranch, abandoned sandstone quarry, elev. 2075 m, 40.176244, -105.305088, 21 October 2014, M. Majack, S. Hauptli, and C. LeFevre 14186, 14187 Douglas County : Douglas County Open Space: Prairie Canyon Ranch, plunge pools in unnamed intermittent creek, elev. 2028 m, 39.31215, -104.71412, 21 June 2013, M. Majack and J. Sanderson 13116-13123 Potamogeton epihydrus Raf. Potamogeton epihydrus has not been assigned a conservation rank in Colorado (CO SNR), but it is ranked critically imperiled (S1, Utah and Wyoming) or imperiled (S2, Nebraska) in surrounding states where it occurs. Existing herbarium records document 11 populations in Colorado, widely dispersed throughout the state in eight of 64 counties. Only four of these populations were documented in the last 20 years, in Jackson, Rio Grande, Summit and Weld counties. Potamogeton epihydrus is also reported for Routt County (Culver and Lemly 2013) but does not appear to be vouchered. In 2013, I documented two previously unknown populations of P. epihydrus representing the first records of P. epihydrus in Delta and Pitkin counties. The Delta County population was collected from Round Lake on Grand Mesa, a small subalpine lake along a hiking trail at 3170 m elevation. The Pitkin County population was found in Sellar Lake at a similar elevation (3218 m) in the subalpine along a rough jeep trail. 87

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USA. COLORADO. GRAND MESA NATIONAL FOREST. Delta County : Round Lake, elev. 3170 m, 39.069030, -107.833345, 15 August 2013, M. Majack and J. Uhl 13237-13240 WHITE RIVER NATIONAL FOREST. Pitkin County : Sellar Lake, elev. 3218 m, 39.069320, -106.583869, 6 August 2013, M. Majack and J. Uhl 13198-13202 Potamogeton foliosus Raf. Potamogeton foliosus is a relatively common species of pondweed in the United States, occurring in all 50 states (Kartesz 2014). Existing herbarium records document P. foliosus at 61 sites in 31 of 64 counties in Colorado. In the last 20 years, collections have been made from 12 sites in nine counties: Arapahoe, El Paso, Gunnison, Jefferson, Larimer, Lincoln, Weld, and Yuma. Despite its apparent ubiquity in the United States, I documented many new populations of P. foliosus in Colorado in 2013 and 2014. Four counties were added to the known distribution of P. foliosus in Colorado (Chaffee, Delta, Douglas and Mesa), and vouchers were collected from three counties for the first time this decade (Denver, Jackson and Lake). The number of new records made for this common species further highlights the gaps in collecting effort for aquatic plants in Colorado. As with most linear-leaved pondweeds, the species is difficult to identify with absolute certainty, especially when collected in a vegetative state, and the above records should be considered provisional in the absence of fruiting voucher specimens. Resources currently used in the identification of Colorado pondweeds (Culver and Lemly 2013, Weber and Wittmann 2012a, 2012b) use the absence of nodal glands as the main character to separate Potamogeton foliosus from Potamogeton pusillus L., the other 88

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common linear-leaved pondweed known in Colorado. FNA (Haynes and Hellquist 2000) and Weigleb and Kaplan (1998) acknowledge the occasional to frequent occurrence of nodal glands on P. foliosus. Further complicating identification is the subtle nature of glands of P. pusillus subsp. pusillus as they are small and the same color as the stem and often occur at only a few nodes (Haynes and Hellquist 2000). I agree that the presence of nodal glands is not a reliable character for the separation of the linear-leaved pondweeds in Colorado, as I encountered many populations of P. foliosus with nodal glands whose identification was confirmed by the presence of diagnostic keeled fruits. Instead, I emphasize the importance of using leaf, stipule, and turion characters. In FNA, Haynes and Hellquist (2000) list two subspecies: Potamogeton foliosus Raf. subsp. foliosus and P. foliosus Raf. subsp. fibrillosus (Fernald) R.R.Haynes & Hellq. Recently, the validity of the latter has been questioned, as plants identified as P. foliosus subsp. fibrillosus in the field were determined by Dr. Zdenek Kaplan to be P. pusillus P. foliosus or hybrids using DNA fingerprinting (Hellquist et al. 2014). Although I collected plants showing characteristics P. foliosus subsp. fibrillosus (nodal glands and stipules disintegrating into fibers) during 2013 and 2014, I will not recognize subspecies here due to the morphological variation I observed between and within populations that did not align with FNA descriptions. Clearly, this group requires additional study, and it seems highly likely that cryptic hybrids and/or species state records have already been collected in Colorado. Potamogeton foliosus has not been assigned a conservation rank in Colorado (G5 CO SNR), although the Potamogeton foliosus Herbaceous Vegetation community type 89

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(G3? CO S3) is fully tracked by the Colorado Natural Heritage Program (CNHP 2013b), and the species is ranked vulnerable (S3) in Wyoming. In 2013 and 2014, I documented many new occurrences of P. foliosus in eight counties, four of which are new county records (Chaffee, Delta, Douglas and Mesa counties). Additionally, new sites for P. foliosus were documented in three counties vouchered previously by single historic collections (Denver, Lake, and Jackson counties). My collections from Bear Creek Park and Rocky Mountain Lake in Denver County represent the first vouchers for the county in 136 years, since P. foliosus was documented in the South Platte River in 1898 ( Jones 600 [COLO, RM, UTC]). In Lake County, also previously vouchered by a single collection in 1878 ( Jones s.n. [COLO]), I documented P. foliosus in a beaver pond north of Twin Lakes Reservoir. My collections of P. foliosus from the east branch of Willow Creek in Jackson County were the first in the county in 116 years, as Jackson County was vouchered previously by a single collection from the North Platte River in 1898 ( Shear 3997 [RM]). I also documented one new population of P. foliosus in Larimer County in a beaver pond north of Chambers Lake. Larimer County was vouchered previously by five collections, most recently in 1992. In 2013, I documented P. foliosus in four new counties: Chaffee Delta, Douglas, and Mesa. Potamogeton foliosus was documented for the first time in Mesa County from Atkinson Creek on Grand Mesa. Also on Grand Mesa, the first occurrence of P. foliosus in Delta County was documented from Weir and Johnson Reservoir, found growing with Potamogeton pusillus, and possibly a hybrid taxon showing characters of the two species. Chaffee County is vouchered from Cottonwood Lake, and Douglas County is vouchered 90

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from Cherry Creek and East Plum Creek, both on Douglas County Open Space. These collections from Chaffee and Douglas counties were collected without fruit and may represent cryptic hybrids. USA. COLORADO. Denver County : Bear Creek Park, pond 17 m north of West Kenyon Blvd., elev. 1635 m, 39.650164, -105.044289, 26 June 2013, M. Majack and E. Pansing 13134-13137 Rocky Mountain Lake, elev. 1636 m, 39.782688, -105.027248, 10 June 2013, M. Majack 13152-13154 Douglas County : Columbine Open Space, East Plum Creek, elev. 1988 m, 39.269375, -104.895154, 27 June 2013, M. Majack and J. Sanderson 13138-13140 Prairie Canyon Ranch, Cherry Creek, elev. 2010 m, 39.307499, -104.722313, 21 June 2013, M. Majack and J. Sanderson 13124, 13125 GRAND MESA NATIONAL FOREST. Delta County : Weir and Johnson Reservoir, elev. 3195 m, 39.066565, -107.830292, 15 August 2013, M. Majack and J. Uhl 13241-13247 Mesa County : Atkinson Creek, elev. 3006 m, 39.103142, -107.897537, 21 August 2013, M. Majack and J. Uhl 13256-13259 ROOSEVELT NATIONAL FOREST. Larimer County : beaver pond adjacent to Twin Lakes, north of Chambers Lake, elev. 2898 m, 40.631160, -105.834330, 16 August 2014, M. Majack and J. Uhl 14188 ROUTT NATIONAL FOREST. Jackson County : East Branch Willow Creek, elev. 2757 m, 40.392442, -106.128642, 22 August 2014, M. Majack and J. Uhl 14190, 14191 SAN ISABEL NATIONAL FOREST. Chaffee County : Cottonwood Lake, elev. 2918 m, 39.83547, -106.31767, 20 July 2013, M. Majack and J. Uhl 13164, 13165 Lake County : beaver pond north of Twin Lakes, elev. 2808 m, 39.085155, -106.372674, 21 July 2013, M. Majack and J. Uhl 13168-13171 91

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Potamogeton gramineus L. Potamogeton gramineus has been documented previously from 47 (-49) sites in 20 (-22) of 64 counties in Colorado. Only nine of these populations have been documented in the last 20 years, in eight counties: Delta, Gunnison, Hinsdale, Jefferson, La Plata, Larimer, Montezuma, and San Juan. Potamogeton gramineus is sometimes difficult to differentiate from Potamogeton illinoensis and as a result, two counties (Montrose and Rio Blanco) are vouchered by collections identified as P. gramineus at one herbarium, and P. illinoensis at another. Although it has not been assigned a conservation rank in Colorado (CO SNR), P. gramineus is ranked vulnerable (S3) in Wyoming. In 2013, I documented two new populations of P. gramineus resulting in one county record (Summit County). Potamogeton gramineus was vouchered previously in Delta County by a collection in 1997 ( Lyon 8725 [FLD]). I documented the second known population in the county in 2013 from an unnamed pond south of Youngs Creek Reservoirs. Additionally, I added Summit County to the known distribution of P. gramineus, vouchered by my collections from Dillon Reservoir. USA. COLORADO. GRAND MESA NATIONAL FOREST. Delta County : unnamed pond south of Youngs Creek Reservoirs, elev. 3118 m, 39.033142, -107.911284, 20 August 2013, M. Majack and J. Uhl 13248-13255 WHITE RIVER NATIONAL FOREST. Summit County : Dillon Reservoir, elev. 2753 m, 39.581186, -106.056775, 1 September 2013, M. Majack and J. Uhl 13273-13275 92

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Potamogeton nodosus Poir. Potamogeton nodosus has been documented previously at 41 sites in Colorado, those accessions vouchering 17 of 64 Colorado counties. Fourteen of those populations were collected in the last 20 years, from eight counties: Archuleta, Huerfano, La Plata,, Pueblo, Saguache, San Juan, Weld, and Yuma counties. Las Animas County was previously included in the distribution of P. nodosus, but vouchers for the county ( Clark 2797 [COLO, KHD]) are likely a large form of Potamogeton diversifolius (or even possibly Potamogeton spirillus Tuck). Potamogeton nodosus has not been assigned a conservation rank in Colorado (G5, CO SNR), but it is ranked critically imperiled (S1) in Wyoming. In 2013, I documented P. nodosus in Denver County from Parkfield Lake, a eutrophic urban lake surrounded by densely populated suburbs. USA. COLORADO. Denver County : Parkfield Lake Park, Parkfield Lake, elev. 1624 m, 39.79119, -104.804008, 2 October 2013, M. Majack, P. Regensberg, and D. Gee 13312 [KHD]. Potamogeton praelongus Wulfen Potamogeton praelongus has been documented previously at 15 sites in 11 of 64 counties in Colorado. Five of these populations were documented in the last 20 years, from four counties: Clear Creek, Jackson, Las Animas, and San Juan. Potamogeton praelongus has not been assigned a conservation rank in Colorado (G5, CO SNR) but is ranked critically imperiled (S1) in Nebraska and Wyoming (S1S2). The USDA Forest Service Rocky Mountain Region has evaluated P. praelongus for Sensitive Species status, 93

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but it was determined that there was insufficient information available to make a recommendation (Houston and Sidle 2002). In 2013, I documented P. praelongus in Chaffee and Summit counties for the first time. Although P. praelongus has been previously reported for Chaffee County (Culver and Lemly 2013), my collections from Cottonwood Lake appear to represent the first vouchered occurrence of the species for the county. Potamogeton praelongus was also documented in Summit County for the first time, from Lower Cataract Lake in Arapaho National Forest, at the edge of Eagles Nest Wilderness. In 2014, I visited the only known population of P. praelongus in Clear Creek County (Echo Lake, Denver Mountain Parks). The population was healthy and filled most of the lake (no specimens were collected). Hellquist et al. reported atypical collections of P. praelongus from their recent survey of the aquatic plants of Yellowstone National Park (2014). These unusual plants did not have keeled leaf apices (a diagnostic character for P. praelongus ), suggesting that the plants may be of hybrid origin. Although plants collected from both Cottonwood Lake and Lower Cataract Lake in Colorado were not in flower or fruit, which may indicate hybrid taxa, the leaf apices split when pressed and were typical in all other characters as described in FNA (Haynes and Hellquist 2000). USA. COLORADO. SAN ISABEL NATIONAL FOREST. Chaffee County : Cottonwood Lake, elev. 2918 m, 39.83547, -106.31767, 20 July 2013, M. Majack and J. Uhl 13161, 13162 WHITE RIVER NATIONAL FOREST. Summit County : Lower Cataract Lake, elev. 2633 m, 39.835388, -106.317769, 25 August 2013, M. Majack and J. Uhl 13265-13269 94

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Potamogeton pusillus L. Potamogeton pusillus is an apparently common species in the United States, occurring in all 50 states (Kartesz 2014). Available herbarium records voucher 56 populations of P. pusillus in 24 of 64 Colorado counties. Of these, only 11 populations have been documented in the last 20 years, in seven counties: Dolores, El Paso, Jackson, La Plata, Larimer, Park, and Teller. My fieldwork in 2013 added six new counties to the known distribution of P. pusillus in Colorado: Chaffee, Delta, Denver, Douglas, Mesa, and Pitkin. The species has not been assigned a conservation rank in Colorado (G5, CO SNR ) or other surrounding states, but it is ranked vulnerable (S3) in Wyoming. Potamogeton pusillus is taxonomically problematic, as is common with the linearleaved pondweeds, with 51 synonyms listed in Wiegleb and Kaplan (1998). Because of this confusing history, it has even been proposed that the name be rejected (Brummitt 1986). FNA recognizes two subspecies of P. pusillus in Colorado: Potamogeton pusillus L. subsp. pusillus and P. pusillus L. subsp. tenuissimus (Mert. & W.D.J.Koch) K.Richt. (Haynes and Hellquist 2000). Potamogeton pusillus subsp. tenuissimus is a synonym for Potamogeton berchtoldii Fieber, a name frequently encountered in European literature and in some North American herbaria. DNA sequence data (Les et al. 2009) support the separation of P. berchtoldii (and other taxa) from P. pusillus and confirm the occurrence of hybrid taxa in subsection Pusilli Les et al. (2009) call attention to the difficulty of reliable identification of species in this group using morphology alone, as well as the presence of cryptic hybrids. Because Colorado manuals (Culver and Lemly 2013; Weber and Wittmann 2012a, 2012b) use the absence of nodal glands as the main 95

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character to separate P. foliosus from P. pusillus it is likely that taxa are commonly confused in herbaria, as some morphotypes of P. foliosus exhibit nodal glands (Wiegleb and Kaplan 1998; Haynes and Hellquist 2000). Currently, most counties in Colorado are vouchered by one to few pondweed accessions, which result in misidentifications having a significant effect on county-level distributions. As P. foliosus and P. pusillus appear to be relatively common pondweeds in Colorado, I speculate that both taxa occur in most, if not all counties, and most discrepancies in distributions resulting from misidentification of P. pusillus will resolve themselves as more pondweed specimens are vouchered from all areas of the state. In 2013, I added six counties to the known distribution of P. pusillus in Colorado through collections in Chaffee, Delta, Denver, Mesa, and Pitkin counties. In Denver County, the species was documented in Rocky Mountain Lake, a highly eutrophic urban lake. The first occurrence of P. pusillus in Chaffee County was documented from a beaver pond at the inlet of Alpine Lake near the town of St. Elmo. In Delta County, a population was documented on Grand Mesa at Weir and Johnson Reservoir, where the species appears to grow with P. foliosus and/or possibly a hybrid of the two. Also on Grand Mesa, P. pusillus was documented for the first time in Mesa County from Jumbo Reservoir and an unnamed pond near Ball Reservoir. Pitkin County was also added to the known distribution of P. pusillus in 2013, documented from the outlet of Chapman Dam. Although P. pusillus has been previously documented in Grand County in 1961 and 1988, two of my collections from the county were of special interest. Plants with coriaceous expanded leaves below each inflorescence were collected from a muddy 96

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depression near Lost Lake in Grand County. Harrington and Matsumura (1955) recognized this morphotype as Potamogeton lateralis Morong., citing two records from Larimer County, also noting that these plants may be an unusual variant of P. pusillus The name Potamogeton lateralis Morong. has since been rejected, as it was described based on both P. pusillus and Potamogeton vaseyi J.W. Robbins (Hellquist et al. 1988). Although Haynes and Hellquist (2000) do not mention expanded leaves in FNA, these plants are identifed as P. pusillus following their treatment. Another notable collection of P. pusillus from Grand County was made from a kettle pond on Strawberry Bench, where plants exhibited fruit and leaf apex characters similar to Potamogeton obtusifolius Mert. & W.D.J. Koch Both of these populations from Grand County would be excellent additions to future molecular work in the group. USA. COLORADO. Denver County : Rocky Mountain Lake, elev. 1636 m, 39.782688, -105.027248, 10 June 2013, M. Majack 13155 ARAPAHO NATIONAL FOREST. Grand County : depression along Lost Lake Trail, elev. 2902 m, 40.303353, -105.960231, 3 August 2014, M. Majack and J. Uhl 14130, 14131 Strawberry Bench, kettle pond 170 meters southeast of Strawberry Lake, elev. 2805 m, 40.105608, -105.767499, 21 August 2014, M. Majack, T. Bates, and T. Roberts 14189 GRAND MESA NATIONAL FOREST. Delta County : Weir and Johnson Reservoir, elev. 3195 m, 39.066565, -107.830292, 15 August 2013, M. Majack and J. Uhl 13241-13247 Mesa County : unnamed pond near Ball Reservoir, elev. 3025 m, 39.060115, -108.085166, 12 August 2013, M. Majack and J. Uhl 13223-13226 Jumbo Reservoir, elev. 2985 m, 39.053868, -108.091553, 12 August 2013, M. Majack and J. Uhl 13227-13231 SAN 97

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ISABEL NATIONAL FOREST. Chaffee County : Alpine Lake inlet, elev. 2847 m, 38.71330, -106.29733, 22 July 2013, M. Majack and J. Uhl 13178, 13179, 13183-13185 WHITE RIVER NATIONAL FOREST. Pitkin County : outlet of Chapman Lake below dam, elev. 2603 m, 39.316513, -106.641213, 6 August 2013, M. Majack and J. Uhl 13194-13197 Summit County : Curtain Ponds near Gore Range Trailhead. elev. 2950 m, 39.513586, -106.145816, 1 September 2013, M. Majack and J. Uhl 13287-13289 Potamogeton richardsonii (A.Benn.) Rydb. Existing herbarium specimens document P. richardsonii at 22 sites in 15 of 64 counties in Colorado, although only five of these accessions are from the last 20 years, from Archuleta, Custer, Jackson, Larimer, and Routt counties. Potamogeton richardsonii has not been assigned a conservation rank in Colorado (G5, CO SNR), but it is ranked vulnerable (S3) in Wyoming. In 2013, I documented P. richardsonii for the first time in Delta and Summit counties. The Summit County voucher was collected from Dillon Reservoir, directly below a boat slip at the Frisco Marina. It may be of interest that the collection site was above the water level a few months prior, according to a marina employee. In Delta County, I collected P. richardsonii from Island Lake on Grand Mesa, where it was growing with Potamogeton robbinsii a species new to the state. Although both collections of P. richardsonii were made late in the season (21 August and 1 September), flowering or fruiting plants were not found at either site. USA. COLORADO. GRAND MESA NATIONAL FOREST. Delta County : Island Lake, elev. 3119 m, 39.032590, -108.007630, 21 August 2013, M. Majack and J. Uhl 13260-13261 WHITE RIVER NATIONAL FOREST. Summit County : Dillon 98

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Reservoir, elev. 2752 m, 39.578558, -106.088548, 1 September 2013, M. Majack and J. Uhl 13276-13277 Potamogeton robbinsii Oakes In 2014, I documented the first known occurrences of P. robbinsii in Colorado. Potamogeton robbinsii is a strictly North American species, and is considered rare where present in all but the northernmost United States (Kartesz 2014). The species is known from a few sites in Wyoming and Utah, where it is critically imperiled (S1). Potamogeton robbinsii has not been documented in other states adjacent to Colorado: Arizona, Kansas, Nebraska, or New Mexico (Kartesz 2014). The USDA Forest Service Rocky Mountain Region evaluated P. robbinsii for Sensitive Species status, but it was determined that insufficient information was available to make a recommendation (Warren 2003). In 2013, I documented the first known occurrences of P. robbinsii in Colorado from Alexander Lake and Island Lake in Delta County. These clear, subalpine lakes are located on Grand Mesa, a large basalt-topped mesa in Grand Mesa National Forest. Interestingly, the elevation and volcanic geology of the Colorado sites is similar to the only other known population in the Southern Rocky Mountain region (Fish Lake, Sevier County, Utah). These sites appear to represent the southern limit of P. robbinsii in the Rocky Mountains, and are among the southernmost documented populations throughout the range of the species (Kartesz 2014). In Wyoming, P. robbinsii is known from the Wind River Range and the Yellowstone Plateau (USDA 2014). In their recent survey of aquatic plants of Yellowstone National Park, Hellquist et al. (2014) reported the occurrence of P. robbinsii in four lakes, two of which were only recently documented in 99

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2010. Collectively, these data suggest that further fieldwork in Colorado may result in the discovery of new populations of P. robbinsii in subalpine lakes associated with volcanic rock. USA. COLORADO. GRAND MESA NATIONAL FOREST. Delta County : Alexander Lake, elev. 3093 m, 39.04107, -107.96419, 11 August 2013, M. Majack and J. Uhl 13213 Island Lake, elev. 3123 m, 39.03305, -108.008752, 21 August 2013, M. Majack and J. Uhl 13262-13264 Stuckenia filiformis (Pers.) Bšrner Stuckenia filiformis is one of three Stuckenia species documented in Colorado. Although the genus is relatively easy to distinguish from Potamogeton (having longer leaf sheaths and different leaf and peduncle structure), Stuckenia is notoriously difficult to identify to species, as plants exhibit extensive morphologic variation (Kaplan 2008). The manuals currently used for Colorado plants (Haynes and Hellquist 2000; Weber and Wittmann 2012a, 2012b; Culver and Lemly 2013) separate Stuckenia species using the shape of the leaf apex as the primary diagnostic character. This character has since been shown to be unreliable, leading to the incorrect identification of many herbarium specimens (Kaplan 2008). Additional characters used in the keys, such as overall plant size, branching pattern, or leaf width have been found to be similarly plastic. Therefore, Kaplan (2008) recommends the use of leaf sheath and fruit characters for identification. Although not mentioned in FNA (Haynes and Hellquist 2000), S. filiformis is the only species in the key that has connate (closed) leaf sheaths (Kaplan 2008). FNA (Haynes and Hellquist 2000) recognizes two subspecies of S. filiformis; S. filiformis (Pers.) Bšrner 100

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subsp. alpina (Blytt) R.R.Haynes, Les & M.Kr‡l and S. filiformis (Pers.) Bšrner subsp. occidentalis (J.W.Robbins) R.R.Haynes, Les & M.Kr‡l, both of which have been documented for Colorado. It has since been shown that the delimiting characters for these subspecies (e.g., overall plant size, leaf width) are not genetically fixed and vary in response to environmental conditions (Kaplan 2008). Based on available records in herbarium databases, Stuckenia pectinata (L.) Bšrner and S. filiformis are relatively common and widespread in Colorado (Table 2). Six records of Stuckenia vaginata (Turcz.) Holub are present in herbaria, but four of these have duplicate specimens identified as S. filiformis or S. pectinata (e.g., N.H. Russell 10466a [CS] is identified as S. pectinata but N.H. Russell 10466b [CS] was annotated recently to S. vaginata ). Kaplan (2008) points out that S. pectinata is considered the most plastic of the pondweeds, and is sometimes mistaken for S. vaginata when collected in certain habitats (e.g., deep water), or life stages (e.g., overwintering). Duplicate collections of Stuckenia from 11 Colorado sites are identified as both S. filiformis and S. pectinata (e.g., C.L. Porter 3686 [US] is identified as S. filiformis subsp. alpina and C.L. Porter 3686 [RM] is identified as S. pectinata ). These inconsistencies are likely due to differences in the quality of specimens (e.g., presence of mature fruit, visibility of diagnostic characters), the use of different resources for identification, or the annotation of only one of two or more specimens. Existing herbarium records report S. filiformis at 68 sites in 25 of 64 counties in Colorado. Of these, nine sites in seven counties have been documented in the last 20 years: Archuleta, Eagle, El Paso, Gunnison, Larimer, La Plata, Park, and San Miguel 101

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counties. Hinsdale, Moffat, and Weld counties are vouchered only by accessions with duplicates identified as S. pectinata Given that these specimens were likely identified using the unreliable character of leaf apex shape, and contradicting identification of duplicates, current distributions should be considered temporary pending the annotation of all Stuckenia accessions from Colorado. Although it is not ranked in Colorado (G5, CO SNR) S. filiformis is considered vulnerable (S3) in Wyoming. The Stuckenia filiformis Herbaceous Vegetation community type (GU CO SU) is fully tracked by the Colorado Natural Heritage Program (CNHP 2013b). In 2013, I documented three previously unknown populations of S. filiformis representing the first known occurrences of the species in Summit and Mesa counties. In Mesa County, S. filiformis was documented on Grand Mesa in South Mesa Lake. In Summit County, the species was documented at two sites within meters of Interstate 70, in Officers Gulch Pond and the Curtain Ponds north of Copper Mountain Ski Resort. Fresh material from the Curtain Ponds was sent to Dr. Zdenek Kaplan (PRA) in 2014. All accessions listed below are of fruiting plants. USA. COLORADO. GRAND MESA NATIONAL FOREST. Mesa County : South Mesa Lake, elev. 3034 m, 39.041335, -108.090248, 14 August 2013, M. Majack and J. Uhl 13234-13236 WHITE RIVER NATIONAL FOREST. Summit County : Curtain Ponds, elev. 2950 m, 39.513586, -106.145816, 21 September 2013, M. Majack and J. Uhl 13290-13293, 13303-13306 Officers Gulch Pond, elev. 2885 m, 39.540577, -106.144849, 20 September 2013, M. Majack and J. Uhl 13282-13283 102

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Stuckenia pectinata (L.) Bšrner Stuckenia pectinata is apparently the most common pondweed in Colorado, but considering the difficulty of identification of this genus (see discussion under S. filiformis ), these distributional data should be considered temporary pending annotation of all Colorado Stuckenia accessions. Herbarium databases report S. pectinata at 75 sites in 35 of 64 counties in Colorado. In the last 20 years, vouchers have been collected in 17 of 64 counties: Archuleta, Baca, Dolores, Eagle, Grand, Hinsdale, Jefferson, Kiowa, La Plata, Larimer, Lincoln, Moffat, Montezuma, Park, Pueblo, San Juan, and San Miguel. Huerfano, Lake and San Juan counties are included in the distribution vouchered only by accessions with duplicates identified as S. filiformis Yuma County was previously included the distribution of S. pectinata but its inclusion appears to have been based on a single misidentified voucher ( Clark 2735 [KHD]). Stuckenia pectinata has not been assigned a conservation rank in Colorado (G5, CO SNR), but is considered vulnerable (S3) in Wyoming. Despite being one of the best-collected pondweeds in Colorado, I documented seven new populations of S. pectinata in 2013, resulting in four new county records. In Summit County, I documented S. pectinata from Dillon Reservoir in Summit County, from Dinkle Lake in Pitkin County, Alpine Lake in Chaffee County, and from the South Platte River in Douglas County. Although my collections were not the first records of S.pectinata in Denver County, the new populations I documented in Parkfield Lake and Rocky Mountain Lake represent the first collections from the county in almost 100 years, the most recent collection from Denver dated 1919 ( Bethel and Payson s.n. [COLO]). 103

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Similarly, my collections of S. pectinata in Garfield County do not represent a new county record, as the species was documented from Sweetwater Lake in 1991 ( Vanderhorst 4050 [COLO, RM]), but I include the record here because the collection site was in an interesting habitat: a roadside ditch on Main Street in the City of Carbondale. USA. COLORADO. Denver County : Parkfield Lake Park, Parkfield Lake, elev. 1624 m, 39.79113, -104.801954, 2 October 2013, M. Majack, P. Regensberg, and D. Gee 13310, 13311 Rocky Mountain Lake, elev. 1637 m, 39.782713, -105.027214, 10 June 2013, M. Majack 13149-13151 Garfield County : Town of Carbondale, ditch at 6th and Main streets, elev. 1885 m, 39.400920, -107.214110, 8 August 2013, M. Majack and J. Uhl 13205-13208 PIKE NATIONAL FOREST. Douglas County : South Platte River, elev. 1924 m, 39.297454, -105.207448, 8 September 2013, M. Majack and L. Bruederle 13294, 13295 SAN ISABEL NATIONAL FOREST. Chaffee County : Alpine Lake inlet, elev. 2847 m, 38.71330, -106.29733, 22 July 2013, M. Majack and J. Uhl 13189, 13190 WHITE RIVER NATIONAL FOREST. Pitkin County : Dinkle Lake, elev. 2597 m, 39.303670, -107.117770, 8 August 2013, M. Majack and J. Uhl 13203, 13204 Summit County : Dillon Reservoir, elev. 2754 m, 39.619220, -106.034930, 31 August 2013, M. Majack and J. Uhl 13280-13281 104

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Discussion My fieldwork in 2013 and 2014 resulted in the documentation of 62 previously unknown populations of pondweeds in Colorado (Table 2, Fig. 4), leading to a new state record and 34 new county records (Table 2, Fig. 5). This work demonstrated that our current understanding of the distribution of pondweeds in Colorado, and likely most of North America, is far from complete. Prior to this study, 16 pondweed species were documented from 462 populations in Colorado, based on herbarium accessions collected between 1873 and 2013 (Fig. 1). My analysis of previous collecting efforts of pondweeds revealed that of these 462 populations, only 114 of them have been documented in the last 20 years. Many existing county records are vouchered by single or few historic accessions, leading to significant changes to known county-level distributions from misidentified or erroneously attributed specimens (Table 2). Although I made limited annotations in 2014, they resulted in a county being removed from the distributions of five species (Jackson County for Potamogeton crispus Yuma County for Potamogeton diversifolius and Stuckenia pectinata and Las Animas County for Potamogeton nodosus and Potamogeton pusillus ). Historic records are also very likely to represent populations that no longer exist, given the dynamic nature of aquatic ecosystems and recent anthropogenic influences such as changes in hydrology, land use, eutrophication, pollution, and the introduction of invasive species. Considering the small number of sites documented for pondweeds in Colorado and the high probability of extirpated historic populations, coupled with the difficult nature of pondweed identification, distributions 105

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created using herbarium accessions are often misleading due to the significant impact of one inaccurate or outdated accession. Due to the paucity of information about the distribution of pondweeds in Colorado, only one species, Potamogeton diversifolius has been assigned a subnational conservation rank (S1, critically imperiled). The remaining native pondweed species in Colorado are not ranked (SNR) due to insufficient data (NatureServe 2014), although the many of these species are ranked critically imperiled (S1) in Wyoming (Table 2). The minimum conservation ranking given to our native pondweed species in Wyoming is vulnerable (S3). Similarly, the USDA Forest Service Rocky Mountain Region has evaluated three pondweed species ( P. diversifolius, P. praelongus, and P. robbinsii ) for Sensitive Species status but recommendations were not made due to insufficient information (Burkhart and Crook 2002; Houston and Sidle 2002; Warren 2003). An important first step in the conservation of pondweeds in Colorado is to conduct extensive fieldwork to document their diversity and distribution in the state. In the present study, the known distributions of 14 pondweed species were expanded (Figs. 4 and 5), including the documentation of P. robbinsii in Colorado for the first time. Potamogeton amplifolius, assumed to be extirpated in the state, was documented at a new site in Colorado. Including the new state record for P. robbinsii 14 Potamogeton and three Stuckenia species are documented for Colorado (Table 2), following the species concepts in FNA (Haynes and Hellquist 2000). In 2013 and 2014, I documented previously unknown sites for all of these species (Table 2, Fig. 4), with the exception of Potamogeton illinoensis, Potamogeton natans and Stuckenia vaginata. 106

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Potamogeton illinoensis is known from seven sites in Colorado, historically from Gunnison, Montrose, and San Miguel counties. Recent collections have been made from Jackson County in 2001, and Routt County in 2000. Databased herbarium records indicate P. illinoensis is vouchered for Rio Blanco County as well, but a duplicate of this accession is identified as P. gramineus Potamogeton illinoensis is ranked critically imperiled (S1) in Kansas, Utah, and Wyoming, but remains unranked in Colorado. Potamogeton natans has been documented from 16 sites in ten of 64 counties in Colorado. Four of these sites have been documented in the last 20 years, in El Paso, Jackson, and Larimer counties. The species has not been assigned a conservation rank in Colorado (G5, CO SNR), although the Potamogeton natans Herbaceous Vegetation community type is fully tracked by the Colorado Natural Heritage Program (CNHP 2013b) and is ranked critically imperiled in Colorado (G5? CO S1). Furthermore, the species is ranked critically imperiled or imperiled in states surrounding Colorado: Kansas (S1), Oklahoma (S1), Utah (S1), and Wyoming (S2). Stuckenia vaginata has been documented at six sites in Colorado, historically in Park and Gunnison counties, and more recently in Jackson County (1996) and Larimer County (1998). Four of these six accessions have a duplicate specimen that has a conflicting identification of Stuckenia filiformis or Stuckenia pectinata introducing doubt into the true distribution of S. vaginata in the state. Stuckenia vaginata is unranked in Colorado (CO SNR), but is considered critically imperiled (S1) in Utah, and imperiled (S2) in Wyoming. 107

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Continued fieldwork is needed to document new populations of all species of pondweeds in Colorado, and to visit historic sites to document the continued presence of those populations. Further work in the field and herbarium will certainly reveal additional state and county records. For example, Potamogeton friesii Rupr. and Stuckenia striata (Ruiz & Pav.) Holub are reported for Colorado by FNA (Haynes and Hellquist 2000) but do not appear to have been vouchered by physical specimens. Additionally, Potamogeton zosteriformis Fernald is known from surrounding states (Utah, Wyoming, Kansas, and Nebraska), but not yet from Colorado (Haynes and Hellquist 2000) even though similar habitat occurs in the state. In addition to the documentation of new populations of native pondweeds in Colorado, special attention should be given to the documentation of the range expansion of Potamogeton crispus so that new populations can be detected early and management can be put into place if necessary. In 2013, I documented 6 new populations of P. crispus adding five counties to the known distribution of the species in Colorado (Fig. 3). Based on my observations in 2013 and 2014, P. crispus is most easily detected from late May to early July, as the plants appear to reach their maximum size and fruit earlier than the native pondweeds. If surveying for P. crispus later in the season, during August and September, the plants can usually be found sprouting from turions or fragments in protected sites at the bottom of waterbodies. My experience from the casual monitoring of P. crispus in a pond at Bear Creek Park in Denver County, plants could be collected in vegetative, but clearly identifiable condition even at the beginning of March. Although P. crispus appears to die back early enough in the year to compete with native pondweeds in 108

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Colorado, the species is receiving increased attention in other states where populations are spreading (Tamayo and Olden 2014). In the present study, I identified uncollected areas of the state by identifying gaps in past collecting efforts (Fig. 1). I focused on the Chaffee, Delta, Grand, Mesa, Pitkin and Summit counties on the Western Slope of Colorado, and Denver and Douglas counties on the Eastern Slope (Fig. 2). Despite this collecting effort, most counties in Colorado remain underdocumented, or even unsurveyed. The plains east of the Front Range are largely undocumented, as much of this land is privately owned. Further fieldwork on the plains is clearly needed, and will likely be most efficiently completed through surveys of State Wildlife Areas. State Parks, land owned by local governments or by initiatives such as Great Outdoors Colorado offer other alternatives to gaining access to private land. Many areas of the Rocky Mountains remain unsurveyed, with Wilderness Areas offering the most potential for discovery of new populations of rare pondweeds that are intolerant of disturbance. The Flat Tops Wilderness in Routt and White River National Forests and surrounding areas should be a priority in the future, as should further study of Grand Mesa National Forest, Rio Grande National Forest and San Juan National Forest, due to the abundance of suitable aquatic habitat associated with few existing records of pondweeds. The deserts of the Western Slope of Colorado do not have extensive aquatic habitat, but as they have not been well surveyed for aquatics, fieldwork may result in interesting discoveries. My method of targeting previously unsurveyed areas was quite effective, as it resulted in the documentation of 62 new populations of pondweeds (Fig. 4), resulting in 109

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34 new county records (Fig. 5). Despite the number of new populations I documented in 2013 and 2014, pondweeds were encountered at only 38% of sites surveyed. I recommend that anyone surveying for aquatic taxa in Colorado take advantage of the species lists compiled by the Colorado Natural Heritage Program for Potential Conservation Areas (PCAs). The purpose of these broad-scale surveys is to identify sites of high biodiversity significance and as such, often identify aquatics to genus only. During this study, I visited one PCA that included Potamogeton sp. on the species list (Strawberry Lake, Grand County). The investigation of this report resulted in the documentation of Potamogeton amplifolius currently the only known extant population in Colorado, and the documentation of an interesting morphotype of Potamogeton pusillus During my surveys for pondweeds, I also documented three other aquatics previously unknown from the site, two of which were rare, and represented the first collections from Grand County. In addition to surveys for new pondweed populations, I also recommend that historic populations of pondweeds be confirmed by the collection of voucher specimens. Although my focus in this study was on the expansion of known distributions, I visited a few sites from which pondweeds were documented previously. In the case of P. amplifolius attempts to locate the species at Lower Twin Lake were unsuccessful, as the construction of Twin Lakes Reservoir appears to have caused the extirpation of the species in the area. Other sites, such as Echo Lake in Clear Creek County, have supported a population of P. praelongus that has been documented since 1952. I also collected P. alpinus from Red Rock Lake in Boulder County, previously collected by Francis 110

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Ramaley in 1907. Accessions such as this, over a hundred years later, offer unique opportunities for future research in microevolution, population dynamics, phenology, or the study of plant responses to changes in hydrology, climate, invasive species, or increased recreation. The continued monitoring of established sites can also provide insight into pondweed dispersal vectors and allow for the identification and mitigation of detrimental impacts before populations are locally extirpated. The pondweed specimens collected during this study represent the first steps toward the creation of a baseline to better understand diversity, distribution and ecological affinities of Potamogetonaceae in Colorado. There remains much work to be done in the field and herbarium. It is crucial that existing pondweed specimens from Colorado be annotated to reflect the most current taxonomic concepts. Without annotation, these collections are of limited value, in some cases perpetuating erroneous distributions or misleading efforts to identify new collections. Due to high rates of misidentification in herbaria, the use of distributional data downloaded from web-based herbarium databases is not recommended for pondweeds at this time. The unreliability of these data is illustrated by the changes in county-level distributions resulting from the select annotations I made, as well as by the significant number of duplicate specimens with conflicting identifications (i.e., two plants collected from a single population are identified as one species at one herbarium, and as a different species at another). I recommend that future fieldwork focus primarily on (1) documenting pondweeds in Colorado to levels acceptable to inform conservation rankings for known native species and inform management of P. crispus (2) conducting surveys rigorous 111

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enough to document expected state records (if present), and (3) investigate the hybrid taxa present in Colorado. Once these baselines are established, further studies should aim to determine the ecological affinities of individual species to create habitat models and to predict possible responses to changes in hydrologic regimes, water quality, and/or spread of invasive species. 112

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Tables and Figures Table III.1. Herbaria providing discoverable records of pondweeds ( Potamogeton and Stuckenia ) from Colorado 113 Herbarium a cronym Herbarium name Source Number of records ARIZ University of Arizona SEINet 2 ASC Northern Arizona University SEINet 1 BRY S. L. Welsh Herbarium, Brigham Young University SEINet 16 COCO Colorado College Tass Kelso (email) 2 3 COLO University of Colorado Museum of Natural History Tim Hogan (email) 168 CS Colorado State University SEINet 19 6 F Field Museum of Natural History FMNH website 1 FLD Fort Lewis College SEINet 7 FLFO Florissant Fossil Beds National Monument SEINet 3 ID Stillinger Herbarium, University of Idaho CPNWH 5 KANU University of Kansas Herbarium KANU website 7 KHD Kathryn Kalmbach Herbarium, Denver Botanic Gardens SEINet 57 KS C Kansas State University Herbarium KSC Online Herbarium 4 MESA Walter A. Kelly Herbarium, Colorado Mesa University SEINet 1 MO Missouri Botanical Garden TROPICOS website 2 NMC New Mexico State University SEINet 5 NY New York Botanical Garden SEINet 1 OKLA Oklahoma State University Mark Fishbein (email) 3 RM Rocky Mountain Herbarium, University of Wyoming SEINet 134 RMBL Rocky Mountain Biological Laboratory SEINet 4 SJNM San Juan College SEINet 44 UCR University of California, Riverside SEINet 4 UMO University of Missouri TROPICOS website 5 US United States National Herbarium, Smithsonian Institution NMNH website 11 UTC Intermountain Herbarium, Utah State University SEINet 11 WSC Western State College Herbarium SEINet 5 Totals 26 herbaria 10 databases 717

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Figure III.1. Documented distribution of pondweeds ( Potamogeton and Stuckenia ) in Colorado based on herbarium records collected between 1873 and 2012. Available records compiled from 26 herbaria (Table 1) document 462 populations of pondweeds collected in Colorado in the last 141 years. 114 Existing pondweed records 1873-1964 (documented >50 years ago) 1965-1994 (documented 20 to 50 years ago) 1995-2012 (documented recently: <20 years old) P ! ! P P P ( P ( ( P ( ( ! P ( ( P ! ( ( P ( ( ( ( ( ( ( ( ( ( P ( ( ( ( ! ( P P P ! P ( ( ( ( ( ( ( ( ( ( P ! ! ( ( ( ( ( ( ( P ( P P P ! ! ! P ! ! ( ( ! ( ( ( P P P P ! ( ( ! ! ( ! P ( P P ! P ( ! ! ! ! ( ! ( ! ! P P P ( ( ( P P ! ( ( ! P ! ( ( P P ( P P P ( ( ( ( ( ( ! ( ( ( ! P ( ( ( ! ( ( P ( ( ( ( ( P P P ! P P P ! ( ( ( ! ( ( ( ! ( P ( ! P P P P ! P ! P ( ( ( ( P P P ( ( ( ! ! ! ( ( P ( ( ( ( P ! P ( ( P P ( P ( ( ( ( ( ! ! ( ! ! P P ! ! P ( ! ( ( ( ( ( ! ( ! ( ( P ( ( P ! P P ! ! P P P ( ( ( P P ( ! ! ( ( ( ( ( ( P ( ! ( ! ( P P ! ! ( ( ( ! P ! ( ( ( ( ( ( ( ! ( ( ( ( ! ! ! ( ! ( ( ( ( ( ( ( ( ! ( ( ( ( ( P P ! P ( ( ( ( ( ! ! ( P ( ( ( ! ( ( ( ! ( ( ! ( ( ! ! ! P ( ( ( ( ( ( ! ! ( P P ( ( ( ( ( ( P P ( ( ! ( ( ( ( P ! ! P P ! P P P ! ! ( ( ( ! ( ( ! P ( ( ( ( P ( ( ( ( ! ( P ( P ! P ( ( ! ( P ! ! ! ! P ( ( P ! P ! ( P ( ! ( ( ( ( ! ( ! ( ! P ! ! P P ! ( P ! ( ! ! P ( ( ! P P P P P ( ( P P ! P ! ( ( ( P P ( ( ! ( ! P ( P ( ( P P ! P P P P P P P P ( P ( ( ( ( P W e l d M o f f a t M e s a B a c a P a r k L a s A n i m a s R o u t t Y u m a G a r f i e l d P u e b l o G u n n i s o n S a g u a c h e L i n c o l n B e n t L a r i m e r R i o B l a n c o E a g l e E l b e r t L o g a n E l P a s o G r a n d K i o w a M o n t r o s e O t e r o W a s h i n g t o n D e l t a J a c k s o n L a P l a t a P r o w e r s K i t C a r s o n C h e y e n n e F r e m o n t M o n t e z u m a A d a m s H u e r f a n o M o r g a n C o n e j o s C o s t i l l a P i t k i n D o l o r e s A r c h u l e t a C h a f f e e H i n s d a l e S a n M i g u e l M i n e r a l C u s t e r D o u g l a s C r o w l e y T e l l e r B o u l d e r P h i l l i p s J e f f e r s o n A l a m o s a O u r a y A r a p a h o e R i o G r a n d e S u m m i t L a k e S e d g w i c k S a n J u a n C l e a r C r e e k G i l p i n D e n v e r B r o o m f i e l d Data sources: herbarium databases COCO, COLO, CPNWH, F, KANU, KSC, OKLA, MO, SEINet, US (Table 2), ESRI 0 25 50 100 km N

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Figure III.2. Sites surveyed for pondweeds ( Potamogeton and Stuckenia ) in 2013 and 2014. Fieldwork performed June to September 2013 focused on seven Colorado counties: Denver, Delta, Douglas, Chaffee, Pitkin, Mesa and Summit. Surveys from August to October 2014 focused on Grand County, with additional sites visited in Boulder, Jackson and Lake counties. 115 Sites surveyed 2013-2014 Pondweeds present Pondweeds not present ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( ( W e l d M o f f a t M e s a B a c a P a r k L a s A n i m a s R o u t t Y u m a G a r f i e l d P u e b l o G u n n i s o n S a g u a c h e L i n c o l n B e n t L a r i m e r R i o B l a n c o E a g l e E l b e r t L o g a n E l P a s o G r a n d K i o w a M o n t r o s e O t e r o W a s h i n g t o n D e l t a J a c k s o n L a P l a t a P r o w e r s K i t C a r s o n C h e y e n n e F r e m o n t M o n t e z u m a A d a m s H u e r f a n o M o r g a n C o n e j o s C o s t i l l a P i t k i n D o l o r e s A r c h u l e t a C h a f f e e H i n s d a l e S a n M i g u e l M i n e r a l C u s t e r D o u g l a s C r o w l e y T e l l e r B o u l d e r P h i l l i p s J e f f e r s o n A l a m o s a O u r a y A r a p a h o e R i o G r a n d e S u m m i t L a k e S e d g w i c k S a n J u a n C l e a r C r e e k G i l p i n D e n v e r B r o o m f i e l d N 0 25 50 100 km Data sources: Majack field data 2013-2014, ESRI

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Table III.2. Pondweeds ( Potamogeton and Stuckenia ) documented in Colorado by existing herbarium records (1873-2012) and new records from fieldwork (2013-2014). Species NatureServe conservation rank USFS Sensitive Species Status Populations in CO (estimate : previous records unverified) Known distribution (estimate: previous records unverified) Updates to known county distribution Potamogeton alpinus Balb. G5 CO SNR (WY S2S3) 46 populations 7 new (Majack) 39 previous 20 of 64 counties 2 new (Majack) 18 previous New : Chaffee Lake Potamogeton amplifolius Tuck. G5 CO SNR (KS S1 NE S1 WY S1) 1 population 1 new (Majack) 1 previous 1 of 64 counties 1 new (Majack) 1 previous (Lake 1952) New : Grand Removed : Lake (extirpated) Potamogeton crispus L. introduced 12 populations 6 new (Majack) 6 previous 9 of 64 counties 5 new (Majack) 4 previous New : Chaffee Denver Douglas Grand Summit Removed : Jackson Potamogeton diversifolius Raf. G5 CO S1 (Community G1? CO S1?) (NE S2, OK S1, WY S1) USFS insufficient data (2002) 9 populations 2 new (Majack) 7 previous 6 of 64 counties 1 new (Majack) 5 previous New : Douglas Removed : Yuma Potamogeton epihydrus Raf. G5 CO SNR (NE S2 UT SI WY S1) 13 populations 2 new (Majack) 11 previous 10 of 64 counties 2 new (Majack) 8 previous New : Delta Pitkin Potamogeton foliosus Raf. G5 CO SNR (Community G3? CO S3 ) (WY S3) 71+ populations 10+ new (Majack) 61 previous 35 of 64 counties 4 new (Majack) 31 previous New : Chaffee Delta Douglas Mesa Potamogeton gramineus L. G5 CO SNR (WY S3) 49-51* populations 2 new (Majack) 47-49* previous 21-23* of 64 counties 1 new (Majack) 20-22* previous New : Summit Provisional : Montrose* Rio Blanco* Potamogeton illinoensis Morong G5 CO SNR (KS S1 UT S1 WY S1 ) 6-8* populations 0 new (Majack) 6-8* previous 5-6* of 64 counties 0 new (Majack) 5-6* previous no new records Provisional : Rio Blanco Potamogeton natans L. G5 CO SNR (Community G5? CO S1) (KS S1 OK S1 WY S2) 15 populations 0 new (Majack) 15 previous 10 of 64 counties 0 new (Majack) 10 previous no new records Potamogeton nodosus Poir. G5 CO SNR (WY S1) 42-43* populations 1 new (Majack) 41-42* previous 18 of 64 counties 1 new (Majack) 17 previous New : Denver Removed : Las Animas Potamogeton praelongus Wulfen G5 CO SNR (NE S1, WY S1S2) USFS insufficient data (2002) 17 populations 2 new (Majack) 15 previous 13 of 64 counties 2 new (Majack) 11 previous New : Chaffee Summit 116

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* Voucher specimen with a duplicate accession identified as a different species Potamogeton pusillus L. G5 CO SNR (WY S3) 65+ populations 9+ new (Majack) 56-60 previous 29-30 of 64 counties 6 new (Majack) 23-24* previous New : Chaffee Delta Denver Douglas Mesa Pitkin Removed : Las Animas Provisional : Huerfano Potamogeton richardsonii (A.Benn) Rydb. G5 CO SNR (WY S3) 24 populations 2 new (Majack) 22 previous 17 of 64 counties 2 new (Majack) 15 previous New : Delta Summit Potamogeton robbinsii Oakes G5 CO SNR (UT S1 WY S1) USFS insufficient data (2003) 2 populations 2 new (Majack) 0 previous 1 of 64 counties 1 new (Majack) 0 previous New to Colorado New : Delta Stuckenia filiformis (Pers.) Bšrner G5 CO SNR (Community GU CO SU) (WY S3) 59-71* populations 3 new (Majack) 56-68* previous 24-27* of 64 counties 2 new (Majack) 22-25* previous New : Mesa Summit Provisional : Hinsdale Moffat Weld Stuckenia pectinata (L.) Bšrner G5 CO SNR (WY S3) 69-83* populations 7 new (Majack) 62-76* previous 34-39* of 64 counties 4 new (Majack) 32-35* previous New : Chaffee Douglas Pitkin Summit Removed : Yuma Provisional : Huerfano* Lake* San Juan* Stuckenia vaginata (Turcz.) Holub G5 CO SNR (UT S1 WY S1) 2-6* populations 0 new (Majack) 2-6* previous 1-4* of 64 counties 0 new (Majack) 1-4* previous no new records Provisional : Park* Gunnison* Jackson* Totals 17 Potamogeton and Stuckenia species 16 native species 1 introduced species 524 populations 62 new (Majack) 462 previous 348 historic 114 recent 34 new county records total (Majack ) 29 for native species 5 for introduced P. crispus Chaffee: 6 Delta: 5 Denver: 3 Douglas: 5 Grand: 2 Lake: 1 Mesa: 3 Pitkin: 3 Summit: 6 Species NatureServe conservation rank USFS Sensitive Species Status Populations in CO (estimate : previous records unverified) Known distribution (estimate: previous records unverified) Updates to known county distribution 117

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Figure III.3. Updated distribution of Potamogeton crispus L. in Colorado. Six new populations of P. crispus were documented in 2013 (Majack), adding five new county records (Chaffee, Denver, Douglas, Grand and Summit counties). One county (Jackson) was found to have been added to the distribution erroneously. 118 P P P ! P ! P ( ( ! ! ! ! W e l d M o f f a t M e s a B a c a P a r k R o u t t L a s A n i m a s Y u m a G a r f i e l d P u e b l o L i n c o l n G u n n i s o n S a g u a c h e B e n t L a r i m e r E a g l e R i o B l a n c o E l b e r t L o g a n E l P a s o G r a n d K i o w a M o n t r o s e O t e r o D e l t a W a s h i n g t o n J a c k s o n L a P l a t a P r o w e r s K i t C a r s o n C h e y e n n e F r e m o n t M o n t e z u m a A d a m s H u e r f a n o M o r g a n C o s t i l l a C o n e j o s P i t k i n D o l o r e s A r c h u l e t a C h a f f e e H i n s d a l e S a n M i g u e l C u s t e r M i n e r a l D o u g l a s T e l l e r C r o w l e y B o u l d e r P h i l l i p s J e f f e r s o n O u r a y A l a m o s a A r a p a h o e S u m m i t R i o G r a n d e L a k e S e d g w i c k S a n J u a n C l e a r C r e e k G i l p i n D e n v e r B r o o m f i e l d Documented locations of P. crispus in Colorado 1952-1953 (collected 50+ years ago) 1978-1986 (collected 20 to 50 years ago) 2010-2011 (recent collections, in the last 20 years) 2013 (Majack) County-level distribution of P. crispus in Colorado Previously vouchered (1 952-2011) New county record (Majack, 2013) County removed (database error) N 0 25 50 100 km Data sources: Herbarium databases (COLO, CS, KHD, RM, UTC), Majack 2013 collections, ESRI

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Figure III.4. Populations of pondweeds ( Potamogeton and Stuckenia ) documented in Colorado between 1873 and 2014. A total of 524 populations of pondweeds have been documented in Colorado, including 62 new sites documented in 2013 and 2014 (Majack) and 462 existing herbarium records (114 in the last 20 years, 348 historic). 119 Species documented in Colorado Potamogeton alpinus Potamogeton amplifolius Potamogeton crispus Potamogeton diversifolius Potamogeton epihydrus Potamogeton foliosus Potamogeton gramineus Potamogeton illinoensis Potamogeton natans Potamogeton nodosus Potamogeton praelongus Potamogeton pusillus Potamogeton richardsonii Potamogeton robbinsii Stuckenia filiformis Stuckenia pectinata Stuckenia vaginata Number of documented populations in Colorado 0 10 20 30 40 50 60 70 80 90 100 Site unreliable (duplicate specimens identified differently) Historic sites (most recent collection > 20 years ago) Recently documented (collected in the last 20 years) New site documented by 2013-2014 fieldwork (Majack)

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Figure III.5. Colorado county records of pondweeds ( Potamogeton and Stuckenia ) documented between 1873 and 2014. Thirty-four new county records were documented during 2013 and 2014 (Majack). Select annotations revealed five erroneously vouchered counties (four misidentified vouchers, one database error). Thirteen additional counties are vouchered by single accessions with a duplicate specimen identified as a different species.. ! 120 Species documented in Colorado Potamogeton alpinus Potamogeton amplifolius Potamogeton crispus Potamogeton diversifolius Potamogeton epihydrus Potamogeton foliosus Potamogeton gramineus Potamogeton illinoensis Potamogeton natans Potamogeton nodosus Potamogeton praelongus Potamogeton pusillus Potamogeton richardsonii Potamogeton robbinsii Stuckenia filiformis Stuckenia pectinata Stuckenia vaginata Colorado county records -8 4 16 28 40 52 64 Provisional county records (inconsistent identification of only voucher between herbaria) Erroneous county records (e.g., misidentified voucher) Historic counties (species not collected since 1994) Current county distribution (existing records from 1995-2012) New county records (Majack 2013-2014)

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REFERENCES Barkworth, M. E., and Z. E. Murrell. 2012. The U.S. Virtual Herbarium: working with individual herbaria to build a national resource. ZooKeys 209: 55-73. Brummitt, R. K. 1986. Report of the committee for Spermatophyta: 30. Taxon 35: 556-563. Burkhart, B., and R. Crook. 2002. R2 individual species recommendation for Potamogeton diverisifolius [online]. USDA Forest Service, Rocky Mountain Region Species Conservation Program. Website: http://www.fs.fed.us/r2/projects/scp/ evalrationale/index.shtml [accessed 30 September 2014]. CNHP. 2013a (updated September 2013). CNHP Tracked Vascular Plant Species [online]. Colorado Natural Heritage Program, Colorado State University, Fort Collins, Colorado, USA. Website http://www.cnhp.colostate.edu/download/list/vascular.asp [accessed 2 September 2014]. CNHP. 2013b. (updated September 2013). CNHP Tracked Natural Plant Communities [online]. Colorado Natural Heritage Program, Colorado State University, Fort Collins, Colorado, USA. Website http://www.cnhp.colostate.edu/download/list/ communities.asp [accessed 2 September 2014]. Culver, D. R., and J. M. Lemly. 2013. Field guide to Colorado's wetland plants: identification, ecology and conservation. Colorado Natural Heritage Program, Colorado State University, Fort Collins, Colorado, USA. Dolan, R.W., Moore, M. E., and J. D. Stephens. 2011. Documenting effects of urbanization on flora using herbarium records. Journal of Ecology 99: 1055-1062. Eastwood, A. 1893. A popular flora of Denver, Colorado. Zoe Publishing Company, San Francisco, C alifornia, USA. Environmental Systems Research Institute, Inc. (ESRI). 2013. ArcGIS, version 10.1. Redlands, California, USA. GBIF. 2013 (continuously updated). Global Biodiversity Information Facility [online]. Copenhagen, Denmark. Website http://www.gbif.org/ [accessed 9 April 2013 onward]. Google. 2013. Google Earth, version 7.1.2.2041 for Mac OSX. Mountain View, California USA. 121

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Harrington, H. D., and Y. Matsumura. 1955. The true aquatic vascular plants of Colorado. Colorado Agricultural Experiment Station Technical Bulletin, vol. 57, Colorado Agricultural College, Fort Collins, C olorado, USA. Haynes, R. R. and C. B. Hellquist. 2000. Potamogetonaceae Dumortier. In Flora of North America Editorial Committee [eds.], Flora of North America north of Mexico, vol. 22, Magnoliophyta: Alismatidae, Arecidae, Commelinidae (in part), and Zingiberidae, 4774. Oxford University Press, New York, New York, USA. Hellquist, C. B., C. T. Philbrick, and R. L. Hilton. 1988. The taxonomic status of Potamogeton lateralis Morong (Potamogetonaceae). Rhodora 90: 1520. Hellquist, C. E., C. B. Hellquist, and J. J. Whipple. 2014. New records for rare and undercollected aquatic vascular plants of Yellowstone National Park. Madro–o 61:159-176. Houston and Sidle. 2002. R2 individual species recommendation for Potamogeton praelongus [online]. USDA Forest Service, Rocky Mountain Region Species Conservation Program. Website http://www.fs.fed.us/r2/projects/scp/evalrationale/ index.shtml [accessed 30 September 2014]. IPNI. 2013 (continuously updated). The International Plant Names Index [online]. Website http://www.ipni.org [accessed 2013 onward]. Kaplan, Z. 2008. A taxonomic revision of Stuckenia (Potamogetonaceae) in Asia, with notes on the diversity and variation of the genus on a worldwide scale. Folia Geobotanica 43: 159-234. Kartesz, J. T. 2014. The Biota of North America Program (BONAP) [online]. North American Plant Atlas. Chapel Hill, North Carolina, USA. Website http://bonap.net/ napa [accessed 5 November 2014]. Kelso, T., E. Hilpman, B. Hockaday, and G. Maentz. 2008 Flora of the Pikes Peak Region, vol. 8. Colorado College, Colorado Springs, Colorado, USA. La Sorte, F. A., M. F. J. Aronson, N. S. G. Williams, L. Celesti-Grapow, S. Cilliers, et al. 2014. Beta diversity of urban floras among European and non-European cities. Global Ecology and Biogeography 23: 769779. Lososova, Z., M. Chytry, L. Tichy, J. Danihelka, K. Fajmon, O. Hajek, K. Kintrova, I. Kuhn, D. Lanikova, Z. Otypkova and V. Rehorek. 2012. Native and alien floras in urban habitats: a comparison across 32 cities of central Europe. Global Ecology and Biogeography 21: 545555 122

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Kaplan, Z., J. Fehrer, and C.B. Hellquist. 2009. New hybrid combinations revealed by molecular analysis: the unknown side of North American pondweed diversity ( Potamogeton ). Systematic Botany 34: 625-642. Les, D. H., N. M. Murray, and N. P. Tippery. 2009. Systematics of two imperiled pondweeds ( Potamogeton vaseyi, P. gemmiparus ) and taxonomic ramifications for subsection Pusilli (Potamogetonaceae). Systematic Botany 34: 643-651. NatureServe. 2014. NatureServe Explorer: An online encyclopedia of life, version 7 [online]. Arlington, Virginia, USA. Website http://www.natureserve.org/explorer [accessed March 2013 onward]. The Plant List. 2010. The Plant List, version 1 [online]. Website http:// www.theplantlist.org/ [accessed May 2012 onward]. Preston, C. D. 1995. Pondweeds of Great Britain and Ireland. BSBI handbooks for field identification. Botanical Society of the British Isles, London. Riccardi, A., and J. B. Rasmussen. 1999. Extinction rates of North American freshwater fauna. Conservation Biology 13: 1220-1222. SEINet. 2012 onward (continuously updated). Southwest Environmental Information Network database [online]. Website http://swbiodiversity.org/seinet/ [accessed May 2012 onward]. Shelth, S. N., L. G. Lohmann, T. Distler, and I. Jimenez. 2012. Understanding bias in geographic range size estimates. Global Ecology and Biogeography 21: 732-742. Tamayo, M., and D. O. Julian. 2014. Forecasting the vulnerability of lakes to aquatic plant invasions. Invasive Plant Science and Management 7: 32-45. Taylor, D. W. 2014. Large inequalities in herbarium specimen density in the Western United States. Phytoneuron 53: 18. Thiers, B. 1998 onward (continuously updated). Index Herbariorum: A global directory of public herbaria and associated staff [online]. New York Botanical Garden. Website http://sweetgum.nybg.org/ih/ [accessed March 2013 onward]. USDA Forest Service. 2014. Rocky Mountain Region Species Conservation Program: Species Evaluation IndexMonocots [online]. USDA Forest Service Rocky Mountain Region, Renewable Resources, Golden, Colorado, USA. Website http:// www.fs.fed.us/r2/projects/scp/evalrationale/evaluations/monocots/ [accessed 21 September 2014]. 123

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US Census Bureau. 2011 (last revised 27 October 2011). American Fact Finder: Denver County, Colorado [online]. Website http://www.census.gov [accessed 13 December 2011]. USDA, NRCS. 2014 (continuously updated). The PLANTS Database [online]. National Plant Data Team, Greensboro, North Carolina, USA. Website http://plants.usda.gov/ [accessed 2012 onward]. Warren, N. 2003. R2 individual species recommendation for Potamogeton robbinsii [online]. USDA Forest Service, Rocky Mountain Region Species Conservation Program. Website: http://www.fs.fed.us/r2/projects/scp/evalrationale/index.shtml [accessed 30 September 2014]. Weber, W. A. and R. C. Wittmann. 2000. A catalog of Colorado flora. University Press of Colorado, Boulder, Colorado, USA. Weber, W. A. and R. C. Wittmann. 2012a. Colorado flora: eastern slope, a field guide to the vascular plants, 4th ed. University Press of Colorado, Boulder, Colorado, USA. Weber, W. A. and R. C. Wittmann. 2012b. Colorado flora: western slope, a field guide to the vascular plants, 4th ed. University Press of Colorado, Boulder, Colorado, USA. Whittall, J. B., C. B. Hellquist, E. Schneider, and S. A. Hodges. 2004. Cryptic species in an endangered pondweed community ( Potamogeton Potamogetonaceae) revealed by AFLP markers. American Journal of Botany 91: 2022-2029. Wiegleb, G., and Z. Kaplan. 1998. An account of the species of Potamogeton L. (Potamogetonaceae). Folia Geobotanica 33: 241-316. 124

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APPENDIX Appendix A. Herbaria searched for records from the City and County of Denver, Colorado, and for pondweeds (Potamogetonaceae) from Colorado. Herbarium a cronym Herbarium name Source ALA Museum of the North, University of Alaska, Fairbanks CPNWH ARIZ University of Arizona SEINet ASC Northern Arizona University SEINet BABY B.A. Bennet Herbarium, Yukon Government CPNWH BNF Bitterroot National Forest herbarium SEINet BRY S. L. Welsh Herbarium, Brigham Young University SEINet CCH Cochise County Herbarium SEINet CIC Harold M. Tucker Herbarium, College of Idaho CPNWH COCO Colorado College Tass Kelso (email) COLO University of Colorado Museum of Natural History Tim Hogan (email) CONABIO Comisi—n Nacional para el Conocimiento y Uso de la Biodiversidad SEINet CS Colorado State University SEINet DAV University of California, Davis DAV website DES Desert Botanical Garden Herbarium Collection SEINet DMNS Denver Museum of Nature and Science DMNS physical specimens ENLC Eastern Nevada Landscape Coalition SEINet ENMU ENMU Natural History Collection Herbarium SEINet EPHR Snow College Herbarium SEINet EVE The Evergreen College CPNWH F Field Museum of Natural History FMNH website FLD Fort Lewis College SEINet FLFO Florissant Fossil Beds National Monument SEINet GCNP Grand Canyon National Park SEINet GH Gray Herbarium, Harvard University GH website GILA Gila National Forest Herbarium SEINet GREE University of Northern Colorado (email) HJAEF H.J. Andrews Experimental Forest CPNWH HNT Huntington Botanical Gardens Herbarium SEINet HSC Humboldt State University CPNWH ID Stillinger Herbarium, University of Idaho CPNWH IDS Idaho State University CPNWH KANU University of Kansas Herbarium KANU website 125

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KEW Kew Botanic Gardens KEW website KHD Kathryn Kalmbach Herbarium, Denver Botanic Gardens SEINet KS C Kansas State University Herbarium KSU Online Herbarium LEA University of Lethbridge CPNWH MABA Madrean Archipelago Biodiversity Assessment Observations SEINet MESA Walter A. Kelly Herbarium, Colorado Mesa University SEINet MEVE Mesa Verde National Park SEINet MNA Museum of Northern Arizona SEINet MO Missouri Botanical Garden TROPICOS website MONT Montana State University CPNWH MONTU University of Montana CPNWH NAVA Navajo Nation Herbarium SEINet NMC New Mexico State University SEINet NMCR NMSU Center for Natural History Collections Range Science SEINet NY New York Botanical Garden SEINet OKLA Oklahoma State University Mark Fishbein (email) OKL University of Oklahoma (email) OSU Oregon State University CPNWH PLU Pacific Lutheran University CPNWH PRI College of Eastern Utah SEINet PSM University of Puget Sound CPNWH PUA Pacific Union College Hebarium SEINet PUSC Colorado State University Pueblo SEINet REED REED College CPNWH RM Rocky Mountain Herbarium, University of Wyoming SEINet RMBL Rocky Mountain Biological Laboratory SEINet SJNM San Juan College SEINet SNM Gila Center for Natural History, Western New Mexico University SEINet SOC Southern Oregon University CPNWH SRP Snake River Plains Herbarium, Boise State University CPNWH SUU Southern Utah University SEINet SWANER Swaner Preserve and EcoCenter SEINet SWRS Southwestern Research Station SEINet TES TEUI Herbarium, US Forest Service Southwestern Region SEINet TSJC Trinidad State Junior College SEINet Herbarium a cronym Herbarium name Source 126

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UBC University of British Columbia CPNWH UC University of California, Berkeley Andrew Doran (email) UCR University of California, Riverside SEINet UMO University of Missouri TROPICOS website UNM University of New Mexico SEINet US United States National Herbarium, Smithsonian National Museum of Natural History NMNH website USON Herbario de la Universidad de Sonora (DICTUS) SEINet USUUB Utah State University Uintah Basin SEINet UTC Intermountain Herbarium, Utah State University SEINet UVSC Utah Valley University Herbarium SEINet V Royal British Columbia Museum CPNWH WCW Whitman College CPNWH WS Marion Ownbey Herbarium, Washington State University CPNWH WSC Western State College Herbarium SEINet WTU University of Washington CPNWH WWB Western Washington University CPNWH Herbarium a cronym Herbarium name Source 127