Front Range, Eastern Slope, Rocky Mountains Checklist Flora of Native and Naturalized Vascular Plants of Golden and Vicinity, Jefferson County, Colorado (Continued)  

Tom Schweich  

Home Page
Topics in this Article:
History of Botanic Exploration
Useful Publications
 Golden, Colorado sits in a valley formed by erosion along the Golden fault, the geotectonic boundary between the North American Cordillera and the Great Plains. Somewhat like Mono Lake, for which I have also prepared a checklist flora, it sits at a boundary, or perhaps ecotone. Things are always more interesting at the boundaries. I started this project when I realized no such list had been prepared for my newly adopted city. I hope you find this checklist flora helpful. Please write to me if you have questions or comments.





Literature Cited:
- Colwell, Robert K., 2008.
- Savard, Jean-Pierre L., Philippe Clergeau, and Gwenaelle Mennechez, 2000.  

What is biodiversity?





1 — Introduction

2 — Geography

3 — History of Botanic Exploration

4 — Useful Publications

  • 411 — Floras
    • 4112017w — "411" & publication year & author.
  • 412 — Botanical but not a flora.

5 — Methods

6 — Results

  • 61 — Collections Found
  • 62 — Collections Made

63 — Discussion

  • 631 — Rare Plants
  • 632 — Notable Native Plants
    • 6321 -- Ferns, if any.
    • 6322 -- Gymnosperms, if any.
    • 6323 -- Dicots
      • 6323280Erinaugra -- "6323" & family & genus & species & subspecies or variety
    • 6324 -- Monocots
  • 633 — Notable Non-Native Plants
  • 6330 — Noxious Weeds
    • 63402 — Non-Native Grasses in the Golden Landscape.
      • 63404020Agrcri -- "63404" & family & genus & species & sequence number, if needed.
    • 6342 — Non-Native Mustards in the Golden Landscape
      • 6342106Alyaly -- "6342" & family & genus & species & sequence number, if needed.
    • 6343 — Non-Native Sunflowers in the Golden Landscape
    • 6349 — Other Families

7 — Conclusion

8 — Acknowledgements

  • 90 — Ecological Systems of Colorado
  • 91 — GIS Resources to Accompany the Checklist Flora
  • 92 — How the Flora is Built
  • 93 — Source Data
      • 9305 — Names and Abbreviations of Historical Reports
    • 931 — Types from the Golden Area
    • 932 — Namesakes of the Golden Area
    • 933 — General Information, and Mysteries Regarding Other Non-Types or Non-Namesakes
    • 934 — Studies of Prairie or Foothill Ecosystems
    • 935 — Restoration, and Effects of Attempts Thereof
    • 936 — Recreation in Foothill and Prairie Ecosystems
  • 94 — Keys
  • 95 — Schemas
  • 96 — Vegetation Descriptions
  • 97 — Reminders
  • 98 — Mystery Locations



Ecological Systems of Colorado

  The Colorado Natural Heritage Program page on Ecological Systems of Colorado is found at: .

Literature Cited:
- Faber-Langendoen, Don, Ralph H. Crawford, and David L. Tart, 2009.
- Federal Geographic Data Committee, 2008.
- Jennings, Michael D., Don Faber-Langendoen, Orie L. Loucks, Robert K. Peet,m and David Roberts, 2009.  

  Comparison of published vegetation types.
CNHP, 2005O'Shea-Stone, 2002Kilburn & White, 1992Zeise, 1976
    Lichen-rock type. Lichen stand types. Areas of bare rock from steep lava cliffs to the conical peaks on the mesa surface.
  • Short-grass grassland. Bouteloua gracilis, Bromus tectorum, with Alyssum parviflorum, and Opuntia sp., Echinocereus viridiflorus, Coryphantha missouriensis, Coryphantha vivipara var. vivipara. Also Hesperostipa comata (Syn: Stipa c.), and Yucca glauca. Some short shrubs of Chrysothamnus nauseosus ssp. graveolens, Prunus virginiana (Syn: Padus v.), Rhus aromatica ssp. trilobata. Celtis reticulata at edge of mesa. Traditionally dominated by Bouteloua gracilis and Buchloe dactyloides, but now dominated by Bromus tectorum.
Grassland type. Bromus tectorum and Alyssum simplex (Syn: A. minus. Occasional Achnatherum scribneri (Syn: Stipa s.) and Andropogon gerardii. Mixed-grass stand types. Dominated by Bromus tectorum and Agropyron sp. (Elymus sp. ?), with Buchloe dactyloides, and Alyssum alyssoides, Eriogonum umbellatum Torr., Heterotheca villosa, Opuntia compressa, Yucca glauca, and Ericameria nauseosa (Syn: Chrysothamnus nauseosus). West, south, and east exposures.
  Mixed-grass grassland. Stipa comata, Pascopyrum smithii, Bouteloua gracilis, Bromus tectorum, with Andropogon gerardii, Bouteloua curtipendula, Aristida purpurea, and Nassella viridula, with a large number of forbs. Mesa slopes and toe areas of STM.
RM Aspen Forest and Woodland - - -
RM Cliff, Canyon and Massive Bedrock - - -
RM Dry-Mesic and Mesic Montane Mixed Conifer Forest and Woodland - - -
SRM Pinyon-Juniper Woodland - - -
SRM Ponderosa Pine Woodland - - -
Rocky Mountain Lower Montane - Foothill Shrubland.
  • Cercocarpus montanus Shrubland Alliance
    • Series determination requires more data collection.
Upland shrubland. Cercoparpus montanus, with sparse cover of Bromus tectorum intermixed with Hesperostipa comata (Syn: Stipa c.), Yucca glauca, and many cacti.

Ravine shrubland. Skunkbush, chokecherry and Prunus americana, in dense thickets. Few plains cottonwoods and Salix amygdaloides

Shrubland type

  • Mixed shrub community. Symphoricarpos occidentalis, Cercocarpus montanus, Rhus [aromatica] ssp. trilobata, Ribes cereum, Prunus americana, and Prunus virginiana. Understory of Poa pratensis, Bromus tectorum, Elymus trachycaulus (Syn: Agropyron trachycaulum), Eriogonum umbellatum, Alyssum alyssoides, etc.
Mixed shrub stand types. Rhus [aromatica] ssp. trilobata, Ribes cereum Dougl., Symphoricarpos occidentalis, Cercocarpus montanus, Prunus virginiana L., Prunus americana Marsh. Acer glabrum in dense patches. Mostly north exposures.
    Shrubland type.

  • Mountain mahogany community. Cercocarpus montanus with an understory of Alyssum alysoides, Bromus tectorum, Agropyron cristatum (Syn: A. desertorum), Eriogonum umbellatum, etc.
Pure shrub stand types. Cercocarpus montanus, with Bromus tectorum, Alyssum alyssoides, and Eriogonum umbellatum.
    Grassland-shrub type. Common foothills species: Symphoricarpos occidentalis, Prunus americana, Rhus trilobata, and Ribes cereum. Grasses are Poa pratensis, Bromus tectorum, and Elymus trachycaulus (Syn: Agropyron trachycaulum). Also Cercoparpus montanus, Symphoricarpos rotundifolius (Syn: S. oreophilus), Prunus virginiana melanocarpa, Rosa arkansana, Physocarpus monogynus, and Ribes aureum. Shrub cover within grassland matrix is significant, but less than 50%. Shrub-grass stand types. Shrubs of Crataegus succulenta (Syn: C. erythropoda), Rosa sp., Rhus trilobata, Prunus virginiana, Prunus americana, Celtus reticulata, and Ribes cereum, with Agropyron sp. (Syn: Elymus sp. ?), Bromus tectorum, Achnatherum hymenoides (Syn: Oryzopsis h., and Alyssum alyssoides. Patches of shrubs in mixed grass-forb areas.
NA Arid West Emergent Marsh - - -
  Wetlands. Cottonwoods and willows, with Carex spp., and Juncus spp., and a variety of grasses and forbs. Patches of Typha spp.. Hydrology alteration.   Riparian (streamside) stand types. Salix exigua, Populus sargentii, Eleocharis macrostachya, Scirpus lacustris L., and Mentha spicata L.
    Woodland type.

  • Mountain maple community. Dense community of small Acer glabrum just below cliffs or in ravines with a dense understory of mixed shrub. North and east slopes.
  • Cottonwood woodland community. Scattered cottonwoods (Populus deltoides ssp. monilifera (Syn: P. sargentii), and P. angustifolia) along permanent and intermittent streams. In Big Ravine, Acer negundo, Salix exigua, and S. amygdaloides also occur. Other shrubs also form a dense understory.
  • Juniper Savannah community. Juniperus scopulorum with an understory of typical grassland.
Woodland stand types. Juniperus scopulorum, with Bromus tectorum.
WGP (Western Great Plains) Cliff, Outcrop, and Shale Barrens - - -
WGP (Western Great Plains) Closed Depression Wetland - - -
WGP (Western Great Plains) Foothill and Piedmont Grassland - - -
WGP (Western Great Plains) Riparian Woodland, Shrubland and Herbaceous - - -
WGP (Western Great Plains) Shortgrass Prairie - - -


  1. [R3C3] Stipa comata grassland of 30-60 acres near western rim of mountain, may be partly due to effects of the 1988 fire.


Recognized Ecological Systems



Ecological Systems Recognized by the Colorado Natural Heritage System


Literature Cited:
- Colorado Natural Heritage Program, 2005.  

Ecological systems are dynamic assemblages or complexes of plant and/or animal communities that 1) occur together on the landscape; 2) are tied together by similar ecological processes, underlying abiotic environmental factors or gradients; and 3) form a readily identifiable unit on the ground. These systems provide a coarser level unit than plant associations and alliances as defined under the International Vegetation Classification standard, and are more easily identified on the ground.

The descriptions and summarized viability guidelines presented here are intended to serve as a tool for conservation and management planning by providing a context for conservation and management (i.e., what systems do we have in Colorado), and by providing easy access to ranking and evaluation criteria for key ecological attributes of each system (i.e., what is the condition of our systems).

System descriptions and viability guidelines are based on materials compiled by NatureServe or developed by the Colorado Natural Heritage Program. Funding for the development of these documents was provided in part by the Bureau of Land Management, The Nature Conservancy, and the USDA Forest Service (CNHP, 2005).

Literature Cited:
- Colorado Natural Heritage Program, 2005.  



Southwest Regional Gap Analysis Project (SWReGAP)


Literature Cited:
- Rondeau, R., K. Decker, J. Handwerk, J. Siemers, L. Grunau, and C. Pague, 2011.  

The Southwest Regional Gap Analysis Project (SWReGAP) was a mapping and assessment of biodiversity for the five-state region encompassing Arizona, Colorado, Nevada, New Mexico, and Utah. The area comprises approximately 150 million hectares (560,000 square miles) representing 1/5 the coterminous United States. The primary objective of the project was to use a coordinated approach to create detailed, seamless maps of the land cover, habitat for native terrestrial vertebrate species, land stewardship, and management status for the Southwest region. This information was analyzed to identify animal species habitats and natural land cover types that are underrepresented on land managed for their long term conservation. SWReGAP was a multi-institutional effort with scientists based in all five southwest states.


USNVC -- United States National Vegetation Classification
  The U.S. National Vegetation Classification is supported by a formal partnership between the federal agencies, the Ecological Society of America (ESA), and NatureServe, working through the Federal Geographic Data Committee (FGDC) Vegetation Subcommittee. Primary signators include the U.S. Forest Service (which chairs the subcommittee), ESA, NatureServe, and the U.S. Geological Survey Core Science Systems (USGS/CSS). Together we are committed to supporting the implementation and maintenance of the National Vegetation Classification (NVC) Standard (FGDC 2008).
  The overall objective of the Vegetation and Information Standards is to support the use of a consistent national vegetation classification system (NVCS) to produce uniform statistics in vegetation resources from vegetation cover data at the national level. It is important that, as agencies map or inventory vegetated Earth cover, they collect enough data accurately and precisely to translate it for national reporting, aggregation, and comparisons. Adoption of the Vegetation Classification and Information Standards in subsequent development and application of vegetation mapping schemes will facilitate the compilation of regional and national summaries. In turn, the consistent collection of such information will eventually support the detailed, quantitative, geo-referenced basis for vegetation cover modeling, mapping, and analysis at the field level.




Literature Cited:
- Faber-Langendoen, Dom, Todd Keeler-Wolf, Del Meidinger, Dave Tart, Bruce Hoagland, Carmen Josse, Gonzalo Navarro, Serguei Ponomarenko, Jean-Peirre Saucier, Alan Weakley, and Patrick Comer, 2014.



GIS Resources to Accompany the Checklist Flora




How the Flora is Built




Source Data




Literature Cited:
- Colbry, Vera Lyola, 1957.  





Literature Cited:
- Hufford, Larry, Michelle M. McMahon, Anna M. Sherwood, Gail Reeves, and Mark W, Chase, 2003.  

Names recognized by Harrington (1954) Names recognized by Snow (2009) Names recognized by Weber and Wittmann (2012) Names recognized by Ackerfield (2015) Names recognized by FNANM Mentzelia (s.l.) represented by collections in Jefferson County, Colorado
Section Bartonia
Mentzelia multiflora (Nutt.) Gray
(Syn: M. speciosa Osterhout., Nuttallia multiflora (Nutt.) Greene, N. speciosa (Osterh.) Greene, N. sinuata Rydb.)
Mentzelia multiflora (Nutt.) A. Gray var. multiflora Nuttallia multiflora (Nuttall) Greene
(Incl: N. sinuata, N. speciosa)
Mentzelia multiflora (Nutt.) Gray Mentzelia multiflora (Nutt.) Gray M. multiflora (Nutt.) Gray
Mentzelia sinuata (Rydb.) R. J. Hill Mentzelia speciosa Osterh. Mentzelia speciosa Osterh. Mentzelia sinuata (Rydb.) R. J. Hill
Mentzelia speciosa Osterh. Mentzelia speciosa Osterhout
Mentzelia nuda (Pursh) T. & G.
(Syn: Nuttallia nuda (Pursh) Greene)
Mentzelia nuda (Pursh) Torr. & A. Gray Nuttallia nuda (Pursh) Greene Mentzelia nuda (Pursh) Torr. & A. Gray Mentzelia nuda (Pursh) Torr. & A. Gray M. nuda (Pursh) Torr. & Gray
Section Trachyphytum
Mentzelia albicaulis Dougl ex Hook. Mentzelia albicaulis (Douglas ex Hook.) Douglas ex Torr. & A. Gray Acrolasia albicaulis (Douglas) Rydberg Mentzelia albicaulis (Douglas ex Hook.) Douglas ex Torr. & A. Gray
(Syn: M. montana (Davidson) Davidson)
M. albicaulis (Dougl. ex Hook.) Dougl. ex Torr. & Gray M. albicaulis (Dougl. ex Hook.) Dougl. ex Torr. & Gray
Not recognized Mentzelia montana (Davids.) Davids. Not recognized M. montana (Davidson) Davidson M. montana (Davidson) Davidsona
Mentzelia dispersa Wats. Mentzelia dispersa S. Watsonb Acrolasia dispersa (S. Watson) Davidson Mentzelia dispersa S. Watson M. dispersa S. Wats. M. dispersa S. Wats.
Harrington, H. D. 1954. Manual of the plants of Colorado. Denver, CO.: Sage Books., 1954. Snow, Neil. 2009. Checklist of Vascular Plants of the Southern Rocky Mountain Region (Version 3). 316 p. Weber, William A., and Ronald C. Wittmann. 2012. Colorado Flora: Eastern Slope. 4th Edition. Boulder, Colorado: University Press of Colorado, 2012. Ackerfield, Jennifer. 2015. The Flora of Colorado. Fort Worth, TX. Botanical Research Institute of Texas. Flora of North America Editorial Committee, eds. 1993+. Flora of North America North of Mexico. 16+. New York and Oxford. Published on the Internet; (accessed 2013 through 2019) Source: Southwest Environmental Information Network, SEINet. 2014. http// Accessed on September 04, 2014.


  1. bMentzelia dispersa S. Watson. Snow (2009) follows Dorn (2001) in not recognizing varieties in our region.
  2. aMentzelia montana (Davidson) Davidson is represented by two collections made by George E. Osterhout with Ira W. Clokey, Osterhout's #3095 (RM95508) and #5741 (RM162001). The collections were made 22 June 1918 and the locality is Morrison, Jefferson County, Colorado.



Literature Cited:
- Bell, Charles D., 2010.  

Authors for Symphoricarpos in FNANM are: Applequist, Wendy L./wendy.applequist at and Bell, Charles D./valerianaceae1969 at Caprifoliaceae will be contained in Volume 18, which, as of this date, 3 August 2014, is under production.

Literature Cited:
- Bell, Charles D., 2010.  

“Towards a Species Level Phylogeny of Symphoricarpos (Caprifoliaceae), Based on Nuclear and Chloroplast DNA”

Names of Historical Reports

  The following is an intentionally empty table …
Abbreviation Full Title Availability Online
  • …abbreviation…
… full name of publication … URL: … if available …

Literature Cited:
- Gmelin, Johan Georg, 1747-1769.  

Abbreviation Full Title Availability Online
  • Gmel. Sibir.
Gmelin, Johan Georg. 1747-1769. Flora Sibirica sive Historia Plantarum Siberiae. St. Petersburg: Imperial Academy of Sciences. URL:

Literature Cited:
- Pursh, Frederick, 1814.  

Pursh (1814) is an often cited reference for taxa found in Golden s.l.
Abbreviation Full Title Availability Online
  • Pursh Fl. Sept. Americ.
Pursh, Frederick. 1814. Flora Americae Septentrionalis; or, A Systematic Arrangement and Description of the Plants of North America. 2 vols. London: White, Cochrane, and Co., 1814. URL:

There are forty-four taxa named by Pursh that are known to occur in Golden s.l..

  • Juniperus communis var. depressa Pursh. Common juniper
  • Musineon divaricatum (Pursh) Raf.. Leafy Wild Parsley
  • Agoseris glauca (Pursh) Raf.. Pale Goat-Chicory
  • Arnica fulgens Pursh. Shining Leopardbane
  • Balsamorhiza sagittata (Pursh) Nutt.. Arrow-Leaf Balsamroot
  • Dieteria canescens (Pursh) Nutt.. Hoar False Tansy-Aster
  • Erigeron compositus Pursh. Dwarf Mountain Fleabane
  • Gaillardia aristata Pursh. Great Blanket-Flower
  • Grindelia squarrosa (Pursh) Dunal. Curly-Cup Gumweed
  • Gutierrezia sarothrae (Pursh) Britt. & Rusby. Kindlingweed
  • Heterotheca villosa (Pursh) Shinners. Hairy False Golden-Aster
  • Lygodesmia juncea (Pursh) D. Don ex Hook.. Rush Skeleton-Plant
  • Nothocalais cuspidata (Pursh) Greene. Wavy-Leaf Prairie-Dandelion
  • Xanthisma spinulosum (Pursh) D.R. Morgan & R.L. Hartman. Lacy Sleepy Daisy
  • Mertensia lanceolata (Pursh) DC.. Prairie Bluebells
  • Phacelia heterophylla Pursh. Variable-Leaf Scorpion-Weed
  • Cleome serrulata Pursh. Rocky Mountain Beeplant
  • Euphorbia marginata Pursh. Snow-on-the-Mountain
  • Amorpha canescens Pursh. Leadplant ???
  • Glycyrrhiza lepidota Pursh. American Licorice
  • Lupinus argenteus Pursh. Silver-Stem Lupine
  • Oxytropis lambertii Pursh. Stemless Locoweed
  • Pediomelum argophyllum (Pursh) J. Grimes. Silver-Leaf Indian-Breadroot
  • Psoralidium lanceolatum (Pursh) Rydb.. Lemon scurfpea
  • Psoralidium tenuiflorum (Pursh) Rydb.. Slimflower scurfpea
  • Ribes aureum Pursh. Golden Currant
  • Linum lewisii Pursh. Prairie Flax
  • Mentzelia decapetala (Pursh ex Sims) Urb. & Gilg ex Gilg. Gumbo-Lily
  • Mentzelia nuda (Pursh) Torr. & A. Gray. Goodmother
  • Claytonia lanceolata Pursh. Lance-Leaf Springbeauty
  • Lewisia rediviva var. rediviva Pursh. Bitter root
  • Mirabilis linearis (Pursh) Heimerl. Narrow-Leaf Four-O'clock
  • Oenothera albicaulis Pursh. White-Stem Evening-Primrose
  • Castilleja sessiliflora Pursh. Great Plains Indian-Paintbrush
  • Mimulus lewisii Pursh. Great Purple Monkey-Flower
  • Ipomopsis aggregata (Pursh) V. Grant. Scarlet Skyrocket
  • Rumex venosus Pursh. Veiny Dock
  • Androsace occidentalis Pursh. Western Rock-Jasmine
  • Clematis hirsutissima Pursh. Sugarbowls
  • Ranunculus cymbalaria Pursh. Alkali Buttercup
  • Geum triflorum Pursh. Old-Man's-Whiskers
  • Scrophularia lanceolata Pursh. Lance-Leaf Figwort
  • Viola nuttallii Pursh. Yellow Prairie Violet
  • Pseudoroegneria spicata (Pursh) A. Löve. Bluebunch-Wheat Grass

Literature Cited:
- Nuttall, Thomas, 1818.

Other articles:
• Golden Checklist Flora:  Bouteloua gracilis, Nuttall, 1818;  

Abbreviation Full Title Availability Online

Nuttall, Thomas. 1818. The Genera of North American Plants and a catalogue of the species to the year 1817. 2 Vols.. Philadelphia: 1818.

Types published by Nuttall in this volume that are found in Golden s.l.

  • Agoseris parviflora (Nutt.) D. Dietr.
  • Ambrosia tomentosa Nutt.
  • Artemisia ludoviciana Nutt. Silver Wormwood.
  • Cirsium undulatum (Nutt.) Spreng. Wavy Leaved Thistle.
  • Coreopsis tinctoria Nutt. Golden Tickseed.
  • Crepis occidentalis Nutt. Largeflower Hawksbeard.
  • Erigeron pumilis Nutt. Shaggy Fleabane.
  • Helianthus pumilus Nutt. Little Sunflower.
  • Heliomeris multiflora Nutt. Showy Golden Eye
  • Packera plattensis (Nutt.) W. A. Weber & Á. Löve. Prairie Groundsel.
  • Ratibida columnifera (Nutt.) Woot. & Standl. (Syn: Rudbeckia columnifera Nutt. ) Upright Prairie Coneflower.
  • Senecio integerrimus Nutt. Columbia Ragwort.
  • Solidago missouriensis Nutt. Missouri Goldenrod.
  • Solidago nana Nutt. Baby Goldenrod.
  • Solidago speciosa Nutt. Showy Goldenrod.
  • Leucocrinum montanum Nutt. Ex A. Gray. Star Lily.
  • Aristida purpurea Nutt. Purple Threeawn.
  • Buchloe dactyloides (Nutt.) Engelm. Buffalo Grass
  • Hordeum pusillum Nutt. Little Barley.
  • Muhlenbergia montana (Nutt.) Hitchc. Mountain Muhly.
  • Munroa squarrosa (Nutt.) Torr. False Buffalograss.
  • Schedonnardus paniculatus (Nutt.) Trel. Tumblegrass.

Literature Cited:
- Emory, William H., 1848.  

Abbreviation Full Title Availability Online

Emory's Rep.
Emory, Notes milit. Reconn. 1848
Emory, William H. 1848. Notes of a military reconnaissance, from Fort Leavenworth, in Missouri, to San Diego, in California, including parts of the Arkansas, Del Norte, and Gila rivers. Washington: Wendell and Van Benthuysen, printers, 1848.

Literature Cited:
- Marcy, Randolph B., 1854.  

Abbreviation Full Title Availability Online

Explor. Red River Louisiana
Marcy, Randolph B., 1854. Exploration of the Red River of Louisiana. Exploration of the Red River of Louisiana. Washington: A. O. P. Nicholson, Public Printer, 1854. URL:


Types from the Golden Area



Namesakes of the Golden Area



General Information, and Mysteries Regarding Other Non-Types or Non-Namesakes


Populus angustifolia E. James

James provided a validating diagnosis: "The long leaved cotton-wood … is found intermixed with the common cotton-wood, resembling in size and general aspect. Its leaves are long and narrow, its trunk smoother, and its branches more slender and flexile than those of Populus anuglata. Some of its fruits was fortunately still remaining …"


Other articles:
• Golden Checklist Flora:  Quercus gambelii;  

Quercus gambelii Nutt. “Gambel Oak”


Literature Cited:
- Jercinovic, Gene, n.d..
- Nuttall, Thomas, 1848.  

William Gambel (June 1823 – December 13, 1849) was an American naturalist, ornithologist, and botanist from Philadelphia. As a young man he worked closely with the renowned naturalist Thomas Nuttall. At the age of eighteen he traveled overland to California, becoming the first botanist to collect specimens in Santa Fe, New Mexico and parts of California.

In March 1841, at the age of eighteen, Gambel set off on his own for California to collect plants and other specimens for Nuttall. He planned to take a more southerly route than that taken in 1834 by Nuttall and John Kirk Townsend. Upon reaching Independence, Missouri he joined a group of traders and headed for Santa Fe following the Santa Fe Trail. Gambel reached Santa Fe in June and spent the next couple months collecting plants. In September Gambel joined a party heading to California and accompanied them along the Old Spanish Trail, arriving in Mexican Alta California in early November, 1841, becoming the first botanist to enter California overland from the east.

Nuttall (1848) described an oak collected on the Rio Grande by William Gambel.

Original Text Comments and Interpretation
Q. Gambelii. Leaves obovate, shortly petiolate, narrowed below, sinuately lobed, dilated and somewhat 3-lobed at the summit, beneath pubescent, the lobes rather obtuse, the upper ones subdentate ; fruit sessile, small, the cup hemispherical, scales ovate-acute ; the glande ovate and acute, about half immersed in the cup ; the conic summit short.
HAB. On the banks of the Rio del Norte, but not abundant. With the aspect of our northern oaks, but very distinct ; in the leaf approaching a little to L. obtusiloba, but without any near affinity. [The Rio del Norte is now called the Rio Grande. – Ed.]


Eriogonum umbellatum Torr.


Literature Cited:
- Torrey, John G., 1827.  

Published in 1827 by John Torrey.




Literature Cited:
- Ray, John, 1686.
Full Size ImageRay's (1686) description of Aquilegia.  

Latin Automated Translation to English My Interpreted English
Aquilegia recens vocabulum est, quasi Aquilina ob florum mucrones aduncos ut sunt Aquiline ungues. Anglis Columbine dicitur, quia florum calcaria recurva columbarum cervicem cum capite & rostro tum figura tum colore referunt. An veteribus Graecis vel Latinis cognita, aut quo nomine dicta nondum constat. Aguilegia recent designation is, as Aquilina of flowers blades that are AQUILINE hooked claws. English Columbine called for bits of flowers and beaks and curved shape and color the dove's neck and head. Is it known to the ancient Greeks, or to the Latins, the name of which it was said, or not yet been agreed. … flower petals that are like an eagle's hooked claws …
Notae illius genericae sunt folia Umbelliferarum modo divisa, flores penduli cum pluribus corniculis recurvis, femina nigra lucida. Note that the generic Umbelliferarum newspapers are now divided, with many little flowers hanging bend, female black cloud.
Aquilegiam stellatam, quae caret illis corniculis recurvis in flore, pro specie degenere habeo. Aquileia the Star, which has no unto them, little curves that bend in the flower, here standing for species degenerates I have.


  1. “aquiline” like an eagle
    • (of a person's nose) hooked or curved like an eagle's beak.

Literature Cited:
- Al-Shehbaz, Ihsan A., and Steve L. O'Kane, 2002.

Other articles:
• Glossary:  cytology;  


The taxonomic world knew for a long time that keeping Physaria and Lesquerella separate was very tenuous. More recently molecular data clearly showed that Physaria is nested within Lesquerella. Also, the data showed that Physaria evolved more than once from Lesquerella. Regardless of how many times Physaria evolved from Lesquerella, Physaria is polyphyletic and Lesquerella definitely paraphyletic. The two genera could no longer be maintained separately on either molecular, morphological, cytological, biogeographic, or ecological grounds. Therefore the two genera should be combined. Physaria is the older of the two names and thus has priority, but there are far more species in Lesquerella. It might make more sense to conserve Lesquerella and transfer the Physaria thereto. An proposal to conserve the name Lesquerella against Physaria was made to the Committee for Spermatophyta (2000). However, the proposal was denied. Therefore, names in Physaria were proposed for most of the former Lesquerella (Al-Shehbaz & O'Kane, 2004).

One of the taxa transferred to Physaria was P. montana a bladderpod found in Jefferson County, including Golden.

Seven taxa of Physaria have been reported for Jefferson County. Four of the taxa are unlikely to occur here, as the collections are misidentified or otherwise quesitonable data. The three confirmed Jefferson County taxa are:

  • P. bellii
  • P. montana
  • P. vitulifera
The four spurious reports of Physaria in Jefferson County are:
  • P. acutifolia — there may be two collections of P. acutifolia from the Evergreen - Kittridge - Parmalee Gulch area. They are old records and a bit sketchy. Otherwise this is an western slope taxon.
  • P. didymocarpa -- misidentified, two vouchers at YU, duplicate at CS9266 determined P. vitulifera.
  • P. floribunda — western slope taxon. Data point from NY showing P. floribunda in Golden is misidentified.
  • P. rollinsii — generally known only from the Gunnison River basin, except for Rollins No. 5151, July 6, 1951, 2 miles west of Deckers, GH 01691383, which has been annotated P. rollinsii, date and name of annotator unknown.

The following key is abstracted from Ackerfield (2015).

(1b.) Fruit (and ovary) pubescent with stellate hairs ... 4
(4b.) Plants not present in the alpine, found below 11,000 ft in elevation ... 6
(6b.) Inflorescence usually obviously exceeding the leaves, or if included then the plants otherwise unlike the above; plants sometimes mound-forming, but the mounds usually not small and button-like ... 8
(8b) Mature fruiting pedicels S-curved (sigmoid), ascending, or sometimes horizontal ... 12
(12a) Plants of the eastern slope ... 13

(13a) Fruit entire at the apex, not or only slightly inflated; ovules 4-20 per ovary ... 14
(14a) Basal leaves suborbicular, obovate, or elliptic, mostly over 4 mm wide (rarely narrowly elliptic and 2-3 mm wide), usually with a well-defined petiole; stem leaves usually secund; ovules (8) 12-24 per ovary ... P. montana

(13b) Fruit notched at the apex (didymous), usually inflated; ovules 4 per ovary ... 16
(16b) Plants of the foothills, absent from the northeastern plains; fruit not conspicuously broader at the apex or appearing flared from the base ... 17

(17a) At least some basal leaf margins deeply and broadly incised, rarely almost entire ... P. vitulifera

(17b) Basal leaf margins shallowly dentate or entire ... 18
(18a) Basal leaves gradually tapering to an ill-defined petiole; fruit 2-8 mm wide ... P. bellii


Other articles:
• Golden Checklist Flora:  Saskatoon Serviceberry;  

Amelanchier alnifolia (Nutt.) Nutt. Ex M. Roem. “Saskatoon Serviceberry”


Literature Cited:
- Moulton, Gary E., 1999.  

Collected by Lewis & Clark on April 15, 1806 at The Dalles of the Columbia River. However, it was not published by Pursh.

Literature Cited:
- Nuttall, Thomas, 1818.
Full Size ImageNuttall (1818) description of Aronia alnifolia.  

Order IV. — Pentagynia

340. Aronia. Persoon. Mespilus. L.

Calix 5-toothed. Petals 5. Berry inferior 5 to 10-celled; cells 1 or 2-seeded. Seeds cartilaginous.

Shrubs without spines, having alternate undivided leaves, and flowers which are corymbose or racemose, generally white; fruit a small black purple or scarlet pomois berry, containing seeds similar to those of apples.

Species. 1. … 6. * Alnifolia. Smooth: leaves roundish, upperpart toothed, pinnately nerved, under side somewhat glaucous; raceme simple, elongated; fruit black and sweet. Habitat. In ravines and on the elevated margins of small streams from Fort Mandam to the Northern Andes. Observation. A shrub 4 or 5 feet high; leaves roundish and retuse, somewhat attenuated at the base, toothed towards the summit; fruit dark purple, somewhat pruinose, very agreeable and saccharine; ripening about July and August.

Literature Cited:
- Nuttall, Thomas, 1834a.
Full Size ImageNuttall (1834) use of the name Amelanchier alnifolia  

In 1834, Nuttall referred to Amelanchier alnifolia in his review of plants collected by Nathaniel Wyeth:

37. Amelanchier alnifolia. Source of the Missouri.

Unfortunately this possible publication of a new name for the taxon is invalid because there is no reference (either direct or indirect) to the intended basionym Aronia alnifolia Nutt. (1818).

Literature Cited:
- Roemer, M. J., 1847.
Full Size ImageRoemer (1847) description of A. alnifolia  

Roemer validly published Amelanchier alnifolia in 1847, referring to Nuttall's 1834 use of the name.
3) prope apicem tantum serrata, subrotunda v. late elliptica, utrinque obtusissima v. retusa ; racemi densiflori ; petala lineari-oblonga, calyce 3–4plo longiora; stamina brevissima.

18. A. alnifolia Nutt. In Joun. Acad. Philad. VII. 22

A. ovalis β semiintegrifolia Hook. L. c. 201. – G.Don l. c.

A. canadensis δ alnifolia Torr. & A.Gr. l. c.

Aronia alnifolia Nutt. gen. I. 306

Pyrus alnifolia Ser. l. c. 637. 39.G.Don l. c. 649. 56.

Ad flum. Columbia pr. Fort Vancouver et ad “the grand Rapids,” in editoribus ad flumen Multnomah. 5 5.

Literature Cited:
- Jones, George Neville, 1946.  


Lupinus argenteus Pursh

  Harrington (1954) and Ackerfield (2015) neither use keel decoration as a key character nor do they describe the decoration of the keel. California floras, e.g., Munz (1965) and Baldwin (2012), use a ciliate keel, as a key character to identify L. argenteus. Welch, et al. (1993) note that the keel can be glabrous or variously ciliate.


Other articles:
• Golden Checklist Flora:   Rhus trilobata;
Full Size ImageRhus trilobata at the southern end of the Survey Field.  

Rhus trilobata Nutt. Squawbush.

Rhus L. is a Linnean name published in his Species plantarum in 1753, although the name had been previously used in six other works that Linnaeus cites. The genus is native to temperate and tropical Asia, Europe, Africa, the Pacific region, and North and South America. There are ten other names that are synonyms for Rhus including Lobadium Raf. that is now treated as a group within Rhus and contains our R. trilobata.

Literature Cited:
- Aiton, William, 1789.

Other articles:
• Glossary:  ♄;
Full Size ImageAiton (1789) description of Rhus aromatica  

Since Rhus trilobata is sometimes reduced to the rank of variety or subspecies under Rhus aromatica Ait. we need to understand the source of that name and its distribution. R. aromatica was described from a plant growing in the Kew Gardens, in 1789. The plant was provided to the garden in 1772 by John Bartram. The source of the plant was “Carolina,” which we now separate into North and South Carolina. Aiton's description is:
Mixed Latin and English My Interpretation of the Mixed Latin and English
11. R. foliis ternatis : foliolis sessilibus ovato-rhombeis inciso-serratis pilosiusculis.
Aromatic Sumach.
Nat. of Carolina. Mr. John Bartram.
Introd. 1772.
Fl. May.     H. ♄.
11. R[hus] leaves in three parts : leaflets sessile, ovate-rhomboid, serrate, puberulent(?).
Aromatic Sumac.
Native to Carolina. [Collected by] Mr. John Bartram.
Introduced (to Kew Garden) in 1772.
Flowers in May.       Hardy. Shrubby.
As was common at the time, Aiton used the astrological sign for Saturn to indicate shrubby plants.
Full Size Image
Abbreviations used by Aiton (1789)

Literature Cited:
- Graustein, Jeannette E., 1967.
- Torrey, John, and Asa Gray, 1838-1843.
Full Size ImageNuttall's description of Rhus trilobata in Torrey & Gray (1838).  

Rhus trilobata was published by Torrey & Gray (1838-1843) from a manuscript written by Nuttall. From the location “Rocky Mountains” and the date of publication — 1838 — it is likely Nuttall made his collection on the Nathaniel Wyeth expedition of 1834-1836 (Graustein, 1967, pp. 277-304).
§ 3. Flowers dioecious or polygamous : disk glandular, deeply 5-lobed (lobes opposite the petals) : drupe globose, villous : nut smooth, compressed: flowers in short aments, preceding the leaves. Leaves 3-foliolate. — Lobadium, Raf.
8. R. aromatica (Ait.) …
9. R. trilobata (Nutt ! Mss.) : “leaves glabrous, small ; lateral leaflets obovate, obtuse, 3-lobed at the apex or nearly entire ; terminal leaflet cuneiform, 3-lobed at the summit, the middle lobe sometimes 3-toothed.
“In the central chain of the Rocky Mountains. — A low leafy shrub ; the leaves much smaller than in R. aromatica : terminal leaflet broad, ½–1  inch in length ; lateral ones smaller. Drupes scarlet, acid ; the nut flat, scarcely striate.” Nutt.

Graustein (1967), Nuttall's biographer, does not mention Rhus trilobata or Nuttall's collection of it.

Literature Cited:
- Watson, Sereno, 1871.

Other articles:
• Golden Checklist Flora:  Rhus trilobata, Barkley, 1937;

Locations: Raft River Mountains. Salt Lake Valley.  

The Geological Exploration of the Fortieth Parallel was a geological survey made by order of the Secretary of War under the direction of Brig. and Bvt. Major General A. A. Humphreys, Chief of Engineers, by Clarence King, U. S. geologist. More commonly known as the Fortieth Parallel Survey, the survey conducted field work from 1867 to 1872, exploring the area along the fortieth parallel north from northeastern California, through Nevada, to eastern Wyoming.

Volume 5, Botany, was written by Sereno Watson (1871) with the assistance of Drs. Gray and Torrey and others. Asa Gray reduced R. trilobata to the rank of variety, which was published in this volume by Watson.

Rhus aromatica, Ait., Var. trilobata, Gray. (R. trilobata, Nutt.) Leaves small, rarely exceeding 1' in length, usually glabrous ; leaflets lobed, the divisions entire or sparingly crenate. — Growing in dense leafy clumps, 3-6° high, and having a heavy disagreeable odor, It is the prevalent western form, extending from Western Texas to Southern California, and throughout the Rocky Mountains to the Upper Missouri. Found on the foot-hills around Salt Lake Valley and at the north base of the Raft River Mountains, Utah. (219.)

The Raft River Mountains are just northwest of the Great Salt Lake. The number (219) is the collection number that Watson applied when he distributed the specimens.

Full Size Image
Gray's description of Rhus aromatica var. trilobata in Watson (1871).

Literature Cited:
- Barkley, Fred Alexander, 1937.

Other articles:
• Golden Checklist Flora:  Rhus aromatica var. trilobata, Watson, 1871;  

Barkley (1937) treats both Rhus aromatica and R. trilobata at the species rank. R. aromatica var. trilobata Gray (in Watson, 1871) is placed in synonomy with R. trilobata. No comment is made regarding the treatment of either species, except that R. trilobata is an
... extremely variable assemblage of plants probably best treated taxonomically as a single polymorphis species running into several more of less consistent geographic variations…
Barkley also recognizes eight varieties of R. trilobata.

Literature Cited:
- Harrington, H. D., 1964, 2nd ed..  

Harrington (1964, 2nd ed.) accepts Rhus trilobata Nutt. Ex T. & G. and one variety, var. simplicifolia (Greene) Barkley.

Literature Cited:
- Weber, W.A., B.C. Johnston, and R. Whittmann., 1981.

Other articles:
• Golden Checklist Flora:  Rhus aromatica ssp. trilobata, Weber, 1989;  

Weber, et al. (1981) published Rhus aromatica Ait. subsp. pilosissima (Engelm.) W. A. Weber, without the similar new combination for R. trilobata. This will be corrected in Weber (1989).

Literature Cited:
- Weber, William A., 1989.

Other articles:
• Golden Checklist Flora:  Rhus aromatica, Weber, et al., 1981;  

Rhus aromatica Ait. Subsp. trilobata (Nutt.) W.A. Weber was published in Weber (1989) with the following explanation:
This is a combination inadvertantly omitted from an earlier treatment of Rhus (Weber, et al. 1981).
Neither publication gives any hint of why Weber reduces R. trilobata to a variety of Rhus aromatica

Literature Cited:
- Welsh, Stanley L., N. Duane Atwood, Sherel Goodrich, and Larry C. Higgins, 1993.  

Welsh et al. (1993) accept R. aromatica var. trilobata, and one variety, var. simplicifolia (Greene) Conq.

Literature Cited:
- Miller, Allison J., David A. Young, and Jun Wen, 2001.  

Miller, et al. (2001) paper about the phylogeny and biogeography of Rhus based on ITS sequence data is hidden behind a paywall.
Abstract. Rhus L. (sensu lato) has been considered the largest and most widespread genus in the Anacardiaceae. Controversy has surrounded the delimitation of the genus. Historically, seven segregate genera have been recognized: Actinocheita, Cotinus, Malosma, Melanococca, Metopium, Searsia, and Toxicodendron. These genera, together with Rhus s.str., are commonly referred to as the Rhus complex. Rhus s.str. Includes two subgenera, Lobadium (ca. 25 spp.) and Rhus (ca. 10 spp.). Sequences of the internal transcribed spacer (ITS) region of the nuclear ribosomal DNA were employed to examine the monophyly of Rhus s.str. And to provide insight into the phylogenetic and biogeographic history of the genus. The ITS data set indicates that Rhus s.str. Is monophyletic. Actinocheita, Cotinus, Malosma, Searsia, and Toxicodendron are distinct from Rhus s.str., although the relationships among these genera of the Rhus complex are not well resolved. Rhus subgenus Rhus is paraphyletic; the monophyletic subgenus Lobadium is nested within it. The ITS data set indicates that, for Rhus, the Madro-Tertiary floristic element (subgenus Lobadium) had a single origin within the Arcto-Tertiary floristic element (subgenus Rhus).

Literature Cited:
- Yi Tingshuang, Allison J. Miller, and Jun Wen, 2004.
- Yi, Tingshuang, Allison J. Miller, and Jun Wen, 2004.  

Phylogenetic and biogeographic diversification of Rhus (Anacardiaceae) in the Northern Hemisphere

Literature Cited:
- Yi, Tingshuang, Allison J. Miller, and Jun Wen, 2007.  

Yi, et al. (2007) published a similar paper on phylogeny of Rhus based on sequences of nuclear and chloroplast genetics found that

(1)Species of Rhus form a monophyletic group … (9) R. aromatica and R. trilobata are sister taxa.

With two notable exceptions … species of subgen. Lobadium were resolved into two clades: … and (2) R. aromatica–R. trilobata and R. integrifolia–R. ovata.

The present distributions of R. microphylla, R. aromatica–R. trilobata, R. lanceolata, and R. copallina indicate that the opportunity for hybridization between some or all of these species exist (and likely existed) in the southwestern U. S. and northern Mexico.

This is one of several phylogeny of Rhus papers that treat both R. aromatica and R. trilobata at the rank of species, although the relative rank of those two entities is not central to the purpose of those papers.

Literature Cited:
- Andres-Hernandez, A. R., and T. Terrazas, 2009.  

Andres-Hernandez (2009) published their analysis of leaf architecture of Rhus. This was part of their PhD thesis, and will be integral to a later paper (2014) on phylogeny of Rhus using structural and molecular data.

Literature Cited:
- Weber, William A., and Ronald C. Wittmann, 2012.  

Weber & Wittmann (2012) accept R. aromatica Aiton ssp. trilobata (Nuttall) W. A. Weber, which Weber published in 1989, and one subspecies, ssp. pilosissima (Engelmann) W. A. Weber, which Weber published in 1981.

Literature Cited:
- Andres-Hernandez, Agustina Rosa, Teresa Terrazas, Gerardo Salazar, and Helga Ochoterena, 2014.  

For R. aromatica, the leaf texture is described as membraneous and the aureole is described as imperfect. For R. trilobata, the leaf texture is described as chartaceous and the aureole is described as incomplete.

This is another of several phylogeny of Rhus papers that treat both R. aromatica and R. trilobata at the rank of species, although the relative rank of those two entities is not central to the purpose of those papers.

Literature Cited:
- Ackerfield, Jennifer, 2015.  

Ackerfield (2015) accepts Rhus trilobata Nutt., placing R. aromatica Aiton in synonomy, and accepts three varieties: var. simplicifolia (Greene) Barkl., var. pilosissima Engelm., and var. trilobata.


Other articles:
• Golden Checklist Flora:  Mentzelia multiflora;  

Mentzelia multiflora (Nutt.) A. Gray


Other articles:
• Golden Checklist Flora:  Evolvulus nuttallianus;
• Field Notes:  Coll. No. 2319, 6 Jun 2020;
Full Size ImageFlower of Coll. No. 2319, Evolvulus nuttallianus
Full Size ImageColl. No. 2319, Evolvulus nuttallianus  

Evolvulus nuttallianus Shaggy Dwarf Morning Glory.

Evolvulus nuttallianus is native to Colorado and is found from Montana and North Dakota south to Texas and Arizona. It is usually described as occuring on sandy and rocky prairies and plains, chaparral, pinyon-juniper and oak woodlands. It has also been found on limestone glades and bald knobs in the Ozark region of Missouri. It is a non-vining morning glory-like perennial that grows as a densely hairy subshrub to 20” tall with prostrate to decumbent stems that root at the nodes as they go. It has very small, bell-shaped, lavender to blue morning glories (to 1/2” across) from spring to mid-summer on stems with oblanceolate, hairy, silvery-green leaves (to 1” long). Flowers usually close up at night and on cloudy days. There is some confusion in the taxonomy of the genus Evolvulus. This species is sometimes listed as synonymous with E. pilosus.

The genus name comes from the Latin word “evolvo” meaning to untwist or unravel because this genus does not twine like many species in the Convolvulus family.

The specific epithet honors British botanist Thomas Nuttall (1786-1839) who traveled extensively in the U.S., as discussed below.

For gardeners, E. nuttallianus is winter hardy to USDA Zone 4. It is easily grown in organically rich, consistently moist, well-drained soils in full sun to part shade. It also performs well in sandy soils with good drainage. But they may be difficult to find in commerce.

Literature Cited:
- Graustein, Jeannette E., 1967.
- Pursh, Frederick, 1814.

Other articles:
• Glossary:  v. s.;  

Thomas Nuttall traveled up the Missouri in 1810, partially duplicating the trip of Lewis and Clark, spending the summer in and around Fort Mandan (Graustein, 1967). I have been unable to determine where Nuttall's collection of E. nuttallianus is located. Graustein notes that many of Nuttall's collections are in the British Museum, and few of their holdings are available online. Regardless, Pursh (1814) published the first description in his Flora of North America. Pursh arrived in London in November 1811. Nuttall and Pursh met in London in the spring of 1812. Nuttall must have had his collections with him. Pursh's (1814) described Nuttall's morning-glory as E. argenteus.
Latin My Interpretation
230. EVOLVULUS. Gen Pl. 524.   230. EVOLVULUS. Gen Pl. 524.  
1. E. erectus ; foliis oblongis utrinque sericeo-tomentosis, pedunculis unifloribus brevibus. argenteus. 1. Evolvulus that is erect, leaves oblong silky-felted both sides, peduncles short with a single flower. argenteus.
On the banks of the Missouri. Nuttall. v. s. Flowers yellow.   On the banks of the Missouri. Nuttall. In have seen it in the dried condition. Flowers yellow.  

Full Size Image
Pursh desription of Evolvulus argenteus
The name E. argenteus was unavailable because is was previously used by Robert Brown, 1810. Pursh notes that he saw the collection “v. s.” but it was not in the Lambert herbarium. Pursh charaterizes the flowers as yellow, but since he worked from dried material, the blue to purple color may have faded.

Literature Cited:
- Nuttall, Thomas, 1818.
Full Size ImageNuttall (1818) description of Evolvulus nuttallianus.  

Nuttall (1818) published his own description of E. nuttallianus in his Genera of North American Plants. He writes both Pursh's name argenteus and his own name of pilosus.

Calix 5-parted. Corolla rotate-campanulate; lobes subemarginate. Styles 2, deeply bifid; segments capillary and divergent. Stigma simple. Capsule 2-celled, 4-valved, 2 to 4 seeded.

Stem creeping, procumbent, or erect. Leaves alternate entire; flowers pedunculate, small; peduncles solitary, bibracteate, capsule perfecting 1, 2, or 4 seeds.

Species. …

3. argenteus, Ph. (pilosus.) Perennial; stems simple, erect, and low, many from the same root: the whole plant densely hairy and shining: leaves cuneate-oblong, acute, crowded; peduncles l-flowered, subsessile; bibracteate, bractes sessile in the axill; segments of the calix linear. — Obs. Allied to E. Commersoni. Stems 4 to 6 inches high; leaves 6 to 8 lines long, 2 to 4 wide, extremely hairy as well as the stem; flowers solitary, appearing sessile, purple, edge of the plaits hairy, capsule 4-valved, often perfecting only a single seed. — Hab. On arid gravelly hills near the confluence of Rapid river and the Missouri; flowering in May. This genus, with the above exceptions, exists exclusively within the tropical regions of India, Australia, and America.

By current standards of publication, one would think that Nuttall was putting his pilosus in synonomy with Pursh's argenteus.

Literature Cited:
- Ro¨mer, Johann Jacob, Joseph August Schultes, Julius Hermann Schultes, Jurt Polycarp Joachim Sprengel, and J. G. Cotta, 1820.
Full Size ImageRoemer & Schultes description of Evolvulus nuttallianus  

In the meantime, Roemer and Schultes (1820) published the first valid name for Nuttall's collection.
LatinMy Interpretation
18. E. Nuttallianus; erectus, foliis onlongis utrinque sericeo-tomentosis, pedunculis unifloris brevibus. E. argenteus Pursh Fl. Sept. Americ. I. p. 187.
Flores flavi. Ad ripas Missouri. Nuttall.
Obs. Nomen mutandum orat, cum sit antiquior argenteus R. Brown.
18. Evolvulus nuttallianus; erect, leaves oblong silky-felted, single flowers on short pedicels. (E. argenteus Pursh Fl. Sept. Americ. I. p. 187.)
Flowers yellow. On the banks of the Missouri River. Nuttall.
The name was changed, because argenteus was used earlier by R. Brown.

This was just a name change because Pursh's proposed E. argenteus was previously used by R. Brown (1810).

Literature Cited:
- Nuttall, Thomas, 1834.
Full Size ImageNuttall (1834) description of E. pilosus  

Then, in 1834, Nuttall first publishes his name of E. pilosus in his descriptions of plants of Arkansas territory.
2. E. pilosus. Erectus, foliis lineari-oblongis utrinque sericeo-pilosis, pedunculis uniflorus brevibus. Nuttall's Gen. Am. Pl. 1, p. 174 (E. Nuttallianus, erectus, foliis oblongis utrinque sericeo-tomentosis, pedunculis unifloris brevibus. Schultes, Syst. Veg. vol. 6, p. 198; E. argenteus, Pursh, 1, 187). — Obs. Flowers purplish, coming out in the middle of the stem; peduncle shorter than the calix; calix segments partly linear and acuminate. — Hab. On the high hills of Red river near Kiamesha,

This name is, of course, superfluous and therefore illegitimate because Roemer and Schultes previously validly published nuttallianus.


Other articles:
• Golden Checklist Flora:  Monarda pectinata;  

Monarda pectinata Nutt.


Scutellaria brittonii Porter

  Isotype: NY415671, T. C. Porter, s.n., Colorado, Clear Creek Canyon, 9000 ft, June 15, 1873. The 9000 ft elevation in Clear Creek Canyon is just below Silver Plume.

Other articles:
• Field Notes:  Coll. No. 1143, 15 Jun 2015;   Coll. No. 1379, 29 May 2016;   Coll. No. 1614, 15 May 2017;  Coll. No. 1865, 23 May 2018;  

Castilleja integra Gray

In 1849, he joined an army expedition (with Gray's help) through Texas, botanising from Galveston to San Antonio and then on to El Paso. But he had to walk most of the 673 miles, (which took over 104 days effort). He collected seeds of Penstemon baccharifolius (Hook.), between Texas and El Paso, which were later given to William Hooker. Also, Castilleja lanata (found near the Rio Grande) and Castilleja integra (found in the Organ Mountains, near El Paso) (Pennell, 1935). In the spring of 1851, he joined the United States and Mexican Boundary Survey (also with Gray's help). (Wikipedia)

Literature Cited:
- Gray, Asa, 1849.  

Castilleja integra is not listed in Gray (1849) Plantae Fendlerianae Novi-Mexicanae. Although, a good part of the report is "… to be continued."

Literature Cited:
- Gray, Asa, 1852.  

Gray (1952) does not mention any Wright collections in the Scrophulariaceae

Literature Cited:
- Torrey John, 1859.  

Emory, William H., 1859. Report on the United States and Mexican Boundary Survey. Volume II. Torrey, John, 1859. Part I. Botany of the Boundary Washington, 1859. p. 119


Castilleja integra (sp. nov.): perennis; caule stricto tomentoso; foliis linearibus integerrimis subtus tomentulosis, floralibus oblongis obocatisque integerrimis coloratis (paniceis);; spica conferta; calyce aequaliter vel postice profundius bifido, lobis bifidis lanceolatis obtusiusculis labium inferius galea multoties brevius adaequantibus. — Organ mountains, east of El Paso; Wright, (undistributed,) Bigelow. Guadaloupe cañon, Sonora; Capt. E. K. Smith. Also gathered in the Rocky Mountains further north by Dr. Kreuzfeldt, in Gunnison's expedition. Stem one or two feet high, mostly simple, rigid; leaves 1½ to 3 inches long, 2 to 3 lines wide, entire; most of the floral ones almost wholly petaloid, ample, shorted than the fully developed flowers. Calyx 8 or 12 lines long, red or reddish; “corolla reddish green;” glaea 6 to 8 lines long; the lower lip very short. Apparently a well marked new species of the section Callichroma. It is No. 584 of Fendler's New Mexican collection; and Dr. Bigelow gathered specimens in Whipple's expedition on the Llano Estacado.

  Pennell, Francis W. [Curator of Botany, Academy of Natural Sciences of Philadelphia] The Scrophulariaceae of Eastern Temperate North America The Academy of Natural Sciences Monogtaphs, Number 1 Philadelphia, 1935. p. 533

9. Castilleja integra Gray

Castilleja integra A. Gray, in Torrey, Bot. Mex. Bound. 119. 1859. “Organ mountains, east of El Paso; Wright ..., Bigelow. Guadaloupe canon, Conora; Capt. E. K. Smith."

Bracts red. Flowering from May to August.

Gravelly soil, plains and hills, Colorado to Texas, Chihuahua and Arizona. Known in our territory from a single record along the Rio grande.

Texas. Valverde: bluffs of Devils R., Havard (U).


Literature Cited:
- Herrando-Moraira, Sonia, et al., 2019.  

Tribe Cardueae

“Thistles” in the broad sense are in the tribe Cardueae Cassini, J. Phys. Chim. Hist. Nat. Arts. 88: 155. 1819 . Tribe Cynareae Lam. & DC. would seem to have precedence over Cardueae by nearly a decade. The North American representatives of Tribue Cardueae accepted by FNANM are as follows. Taxa found in Colorado are shown in bold.
  1. Leaves spiny
    1. Carduus L. Plumeless Thistle.
    2. Carlina L. Carline Thistle.
    3. Carthamus L. Distaff Thistle.
    4. Cirsium Mill. Thistle. All the native thistles in Colorado ae in the genus Cirsium Mill. In addition there are two non-native thistles that are also noxious weeds: C. arvense and C. vulgare.
    5. Echinops L. Globe Thistle.
    6. Onopordum L. Cotton Thistle.
    7. Silybum Adans. Milkthistle. Not listed for Colorado in Weber & Wittmann (2012) or Ackerfield (2015). There are two collections of Silybum marianum (L.) Gaertn. “Blessed Milk-thistle”, one from Larimer County and one from Saguache County.
  2. Leaves not spiny
    1. Acroptilon Cass. Hardheads.
    2. Amberboa (Pers.) Less. Sweet Sultan.
    3. Arctium L. Burdock.
    4. Centaurea L. Knapweed. Eight species of Centaurea are known from Colorado, all of which are non-native.
    5. Crupina (Pers.) DC. Crupina.
    6. Cynara L. Cynara. C. scolymus L. is the Globe Artichoke.
    7. Mantisalca Cass. Mantisalca.
    8. Plectocephalus D. Don in R. Sweet. Basketflower.
    9. Saussurea. Saw Wort.
    10. Volutaria Cass.

There is a more recent paper (Herrando-Moraira, Sonia, et al. 2019) describing the phylogeny of Cardueae using Hyb-Seq data. It is, however, behind a paywall until August 1, 2020, and we eagerly await its unveiling.


Balsamorhiza sagittata

Other articles:
• Tin Cup Ridge (social trail):   at Coll. 1109;
• Field Notes:   Coll. No. 1109, 14 May 2015;
Full Size ImageColl. No. 1109, Balsamorhiza sagittata  

How the heck did it get to Tin Cup Ridge?


Literature Cited:
- Ackerfield, Jennifer, 2015.  

Cirsium Mill. Thistle.

Twenty species of Cirsium “Thistle” are found in Colorado (Ackerfield, 2015). Eighteen of them are native, while two are introduced: C. arvense and C. vulgare. There are two endemic thistles: C. osterhoutii and C. perplexans.


Literature Cited:
- Flora of North America Editorial Committee, eds., 1993+.

Other articles:
• Golden Checklist Flora:  Cirsium undulatum;
Full Size ImageDistribution of collections of Cirsium undulatum  

Cirsium undulatum (Nutt.) Spreng. Wavy Leaved Thistle.

Cirsium undulatum is widely distributed in the wstern half of North America from the dry plains and plateaus of the Pacific Northwest eastward across the Great Plains to Manitoba and the Dakotas and south to Texas, New Mexico, and northwestern Mexico. It occurs in scattered localities in the Rocky Mountains and northeastern Great Basin region. At least some of the few widely scattered records from the eastern United States are probably introductions. Cirsium undulatum is both widespread and variable. Plants of the Great Plains region tend to be low-growing with a few large heads and elongate corollas. Plants of the Pacific Northwest are usually taller and produce smaller, more numerous heads with shorter corollas. A detailed study of this species might reveal races worthy of recognition as infraspecific taxa.

Wavyleaf thistle is listed by California as a noxious weed. However, most reports of Cirsium undulatum in California are based upon misidentifications of C. canescens. Cirsium undulatum is known to hybridize with C. flodmanii, C. hookerianum, and C. scariosum var. coloradense. J. T. Howell (1960b) reported that C. undulatum was suspected to hybridize with C. brevifolium in the Pacific Northwest.

— David J. Keil in Flora of North America (vol. 19, 20 and 21)

Literature Cited:
- Miller, Philip, 1754.  

The genus Cirsium was first proposed by Miller (1754) in his Gardener's Dictionary.

Literature Cited:
- Nuttall, Thomas, 1818.

Other articles:
• Glossary:  ventricose;  

Nuttall (1818) published his Cnicus undulatus in his Genera of North American Plants. He saw the plant on Lake Huron and in the upper Louisiana Territory. I have not found a type specimen, but saw a note that the material might be in the Gray Herbarium.
538. CARDUUS. L. (Thistle.) Calix ventricose, imbricate, scales spiny. Receptacle villous. Pappus pilose or plumose, deciduous.
§ II. CNICUS. Pappus plimose
11. * undulatus. Stem low and few-flowered; leaves amplexicaule, pinnatifidly sinuate, and plicately undulated, on both sides tomentose, but beneath white, lobes bifid and spiny; calix subglobose, scales lanceolate, eredt and mucronate. HAB. On the calcareous islands of lake Huron, and on the plains of Upper Louisiana. — Leaves almost like some species of Cynara, but not remarkably large, moreslenderly tomentose on the upper side; stem often 1,2, or few-flowered, and 1 to 2 feet high. Flowers large, reddish purple.

Literature Cited:
- Sprengel, Curt Polycarp Joachim, 1826.  

undulatum * 45. C. caule humili paucifloro, foliis amplexicaulibus finuato-pinnatifidis undulato-plicatus utrinque tomentofis, laciniis 2fidis fpinofis, fquamis anthodii lanceolatis erectis mucronatis. In infulis lacus Huronum et Louifiana fuper. (Cnicus Nutt.)

Literature Cited:
- Frankton, C., and R. J. Moore, 1961.  

Frankton and Moore (1961) compare and contrast of Cirsium undulatum and C. flodmanii, both of which are known to occur in Colorado is behind a paywall, so I have reproduced the abstract here.
Abstract. The morphology and specific differences of Cirsium undulatum (Nutt.) Spreng. And of C. flodmanii (Rydb.) Arthur are described and their Canadian distributions are reported in detail. The chromosome numbers are C. undulatum f. undulatum and f. album Farwell, 2n = 26; C. flodmanii f. flodmanii and f. albiflorum D. Löve, 2n = 22. The origin of four North American species of Cirsium that do not follow the world-wide base number 17 is discussed; it is postulated that reduction in number has occurred by translocations. The chromosomes of species with reduced numbers are larger than those of the unreduced species but the total length of the chromosomes of both groups is approximately the same.

Literature Cited:
- Häffner, Eva, and Frank H. Hellwig, 1999.  

Haffner and Hellwig (1999) investigated relationships within tribe Cardueae using ITS sequence data. The sole North American Cirsium in their study was C. texanum which does not occur in Colorado.

Literature Cited:
- Garcia-Jacas, Nuria, Teresa Garnatje, Alfoonso Susanna, and Rosier Vilatersana, 2002.  

Garcia-Jacas, et al. (2002) worked on tribal and subtribal delimitation of the Cardueae, unfortunately without any North American specimens.

Literature Cited:
- Kelch, Dean G., and Bruce G. Baldwin, 2003.  

Kelch and Baldwin (2003) studied historical biogeography and ecology of true thistles (Cirsium, Cardueae, Compositae) in the New World. Unfortunately, their sample of thistles did not include Cirsium undulatum. However, three other native thistles, C. muticum, C. discolor, and C. rhaphilepis, that commonly cluster with C. undulatum and did in fact cluster together in this study.

Literature Cited:
- Bodo Slotto, Tracey A., David P. Horvath, and Michael E. Foley, 2012.  

Abstract. Weedy invasive Cirsium spp. are widespread in temperate regions of North America and some of their biological control agents have attacked native Cirsium spp. A phylogenetic tree was developed from DNA sequences for the internal transcribed spacer and external transcribed spacer regions from native and non-native Great Plains Cirsium spp. And other thistles to determine if host specificity follows phylogeny. The monophyly of Cirsium spp. And Carduus within the tribe Cardinae was confirmed with native North American and European lineages of the Cirsium spp. Examined. We did not detect interspecific hybridization between the introduced invasive and the native North American Cirsium spp. Selected host-biological control agent interactions were mapped onto the phylogenic tree derived by maximum likelihood analysis to examine the co-occurrence of known hosts with biological control agents. Within Cirsium-Cardueae, the insect biological control agents do not associate with host phylogenetic lines. Thus, more comprehensive testing of species in host-specificity trials, rather than relying on a single representative of a given clade may be necessary; because the assumption that host-specificity follows phylogeny does not necessarily hold. Since the assumption does not always hold, it will also be important to evaluate ecological factors to provide better cues for host specificity.


The Erigerons that are known from collections in Golden s.l. are:
  • Erigeron compositus Pursh. (Syn: Erigeron compositus Pursh var. discoideus A. Gray, Erigeron compositus Pursh var. glabratus Macoun) Cutleaf Daisy.
  • Erigeron divergens Torr. & A. Gray. Spreading Fleabane.
  • Erigeron flagellaris A. Gray. Trailing Fleabane.
  • Erigeron pumilus Nutt. Shaggy Fleabane.
  • Erigeron strigosus Muhl. ex Willd. (Syn: Stenactis strigosa (Muhl. ex Willd.) DC.) Prairie Fleabane.
  • Erigeron tracyi Greene. (Syn: Erigeron cinereus A. Gray, Erigeron colomexicanus A. Nelson, Erigeron divergens Torrey & A. Gray var. cinereus A. Gray) Running Fleabane.
  • Erigeron vetensis Rydb. Early Bluetop Fleabane.
Erigeron compositus Pursh. Cutleaf Daisy.

Literature Cited:
- Moulton, Gary E., 1999.
- Pursh, Frederick, 1814.

Locations: Kooskoosky River.  

Of the western Erigerons found in Golden s.l. the first to be described was E. compositus Pursh. The holotype was collected by Meriwether Lewis on the Kooskoosky (Clearwater) River, date unknown, and the voucher is now at ANS.

Pursh (1814) described it as follows:

Syngenesia Superflua. Erigeron 535
14. E. pilosum, subacaule; foliis radicalibus longe petiolatis triplicato-3-partitis, laciniis linearibus divaricatis, caulinis linearibus plerumque indivisibus, caule superne nudo unifloro. compositum
  On the banks of the Kooskoosky. M. Lewis. ♃. July, Aug. v. s.; v. v. cultum. Not above a span high; flowers resembling a daisy exceedingly; they change during their flowering, from white to a lively pale red.  

My collection of this taxon was made as Little Scraggy Peak, (Buffalo Creek Recreation Area), Jefferson County, Colorado.

Full Size Image
Coll. No. 1901, Erigeron compositus
  A. Gray (1862) proposed variety discoideus. The name is now treated as a synonym of the species.
  Macoun (1884) proposed variety glabratus, but the name is not now recognized. The name is now treated as a synonym of the species.
Erigeron divergens Torr. & A. Gray. Spreading Fleabane.

Literature Cited:
- Nuttall, Thomas, 1840-1841.

Other articles:
• Glossary:   fastigiate;
Full Size ImageColl. No. 2092, Erigeron divergens  

The first publication of the entity we now call Erigeron divergens was by Nuttall (1840).
Erigeron *divaricatum; ☉ hirsute, stem branching from base; branchlets one-flowered, rather naked, fastigiate; radical leaves spathulate, the rest linear, sessile, acute, attenuated below, all entire; inner pappus of about eight setæ rays nery numerous, narrow, white.
Hab. In the Rocky Mountains and the plains of Oregon. About one foot high, at length very much branched, the leaves an inch or more long, about a line wide. Pappus double in ray and disk, the inner of remarkably few rays, very deciduous.

It might be helpful to remember that the plains of Oregon would include a good part of southwest Wyoming.

The name E. divergens had been previously used by Michaux (1803) and was not available for use be Nuttall.

Literature Cited:
- Torrey, John, and Asa Gray, 1838-1843.

Other articles:
• Field Notes:  Torrey & Gray, 1841;  

Torrey & Gray (1841, p. 175) in Flora of North America proposed Erigeron divergens for the small fleabane that was previously illegimately published by Nuttall as E. divaricatuum.
26. E. divergens: somewhat hoary with a minute hirsute pubescence, diffusely branched from the base; leaves small, entire, acute; the radical somewhat spatulate, narrowed into a short petiole; the cauline scattered, sessile, linear, narrowed at the base; heads (small) mostly solitary terminating the naked branchlets or pecuncles; rays very narrow and numerous, twice the length of the hirsute involucre; inner pappus of few (8-12) very slender and deciduous bristles. — Erigeron (Oligotrichum) divaricatum, Nutt. ! In trans. Amer. Phil. Soc. (n. ser.) 7. p. 311, not of Michx.
Rocky Mountains, and plains of the interior of Oregon, Nuttall ! — ① Stems about a foot high, diffuse, ascending, slender, at length much branched. Leaves half an inch to an inch long, 1 to 2 lines wide, mucronate-acute. Heads rather smaller than in E. tenue; the rays (white, Nutt.) nearly similar; the exterior pappus shorter.
Erigeron flagellaris A. Gray. Trailing Fleabane.

Literature Cited:
- Gray, Asa, 1849.

Other articles:
• Field Notes:  Gray, A., 1849;  

334. E. flagellare (sp. nov.): bienne? striguloso-puberulum, pumilum; caulibus gracillimis e basi ramosis, floriferis seu primariis simplicibus superne aphyllis monocephalis, sterilibus patentibus flagelliformibus; foliis spathulatis mucronulatis inferioribus in petiolum attenuatis integris seu radicalibus parce inciso-lobatus, ramealibus parvulis sublinearibus sessilibus; ligulis numerosis gracilibus (albis purpureo tinctis) involucrum hirsutum duplo superantibus; pappo radii et discii conformiduplici, exteriore coroni-forme-squamellato conspicuo, interiore e setis sub-20 fragilibus. — Low, moist places, along Santa Fe Creek; May, June. (381.) — Root slender. Flowering stems 5 to 7 inches high, very slender, few-leaved below the middle, naked, and pedunculiform above; the head rather than the preceding species; the involucre, &c., similar. Lower leaves one to two inches long, including the slender petiole; those of the runner-like sterile branches decreasing to 2 or 3 lines in length. This species should rank next to the foregoing.*

The “preceding species” and “foregoing” was Erigeron cinereum (sp. Nov.), now treated as a synonym of E. tracyi Greene.

The head was described as “… as large as those on Bellis perennis …” and the involucre was described as “… hirsutum duplo superantibus …” [… coarse erect or ascending hairs doubly … ??? ]

The asterisk “*” refers to some collections sent by Mr. Spaulding from Oregon, which I don't think are particularly relevant to E. flagellaris.

Other articles:
• Plainview Road:   near coll loc;
• Field Notes:  Coll. No. 1116, 28 May 2015;   Coll. No. 1829, 16 May 2018;
Full Size ImageColl. No. 1116, Erigeron flagellaris
Full Size ImageColl. No. 1829, Erigeron flagellaris  

Erigeron pumilus Nutt. Shaggy Fleabane.

Literature Cited:
- Nuttall, Thomas, 1818.

Other articles:
• Glossary:  nutant;  

Syngenesia. Superflua. 147
557. ERIGERON. L. (Flea-bane.)
Calix imbricated, subhemispherical, in fruit often reflected. Rays of the corolla linear, very narrow, and numerous. Receptacle naked. Pappus double, exterior minute, interior pilose, of few rays. (12 to 25?)
Herbaceous; stems simple or branched, flowers solitary, axillary and terminal, subcorymbose or fastigiately paniculate; radii white, or purplish, rarely destiture of pappus. Seed very small, rather smooth, oblong, and compressed; pappus simply pilose, deciduous, not much longer than the seed, exterior pappus minute and paleaceous.
§ I. Stem simple
SPECIES. 1. E. alpinum. 2. * pumilum. Hirsute; stems aggregated, 1-flowered, leaves oblong-linear, entire and sessile; flower large and hemispherical, before flowering nutant; calyx very hirsute. HAB. On the plains of the Missouri. Flowering in May. E. hirsutum. Ph. 2. Suppl. p. 742. but this name has been previously employed for another species. OBS. Stems several from the same root, often, indeed, connected at the base, 4 or 5 inches high, 1-flowered. Leaves 2 inches, more or less, nearly linear, attenuated downwards, scarcely 2 lines wide. Flower naked, (or pedunculate) white, and large as a Daisy; rays as long as the calix, narrow and numerous. Pappus double, internal short, about 12-rayed.
“nutant”, adj., drooping or nodding.
Erigeron strigosus Muhl. Ex Willd. Prairie Fleabane.

Literature Cited:
- Willdenow, Carl L., 1803.  

The first of the Erigeron found in the Golden, Colorado area was described from a collection made in Pensylvania. Willdenow (1803) published the name from a description written by Mühlenberg.
11. ERIGERON ʃtrigoʃum.
E. foliis lanceolatis utrinque attenuatis ſubdentatis ſtrigoſo-piloſis, floribus corymboſo-paniculatis. W.
Erigeron ſtrigoſum. Mühlenb. In litt.
Behaartes Berufungskraut. W.
Habitat in Penſylvania. (v. ʃ.)
Caulis erectus ʃtriatus, pilis ʃparʃis albis obʃitus. Folia alterna lanceolata baʃ et apice attenuata, utrinque oilis copioʃis adpresʃis obʃita, integerrima, vel medio utrunque ʃerraturis 2. ʃ. 3 inʃtructa. Flores corymboʃo-paniculati. Radius copioʃus filiformis albus. W.

Literature Cited:
- DeCandolle, Augustus Pyramus, 1836.  

DeCandolle (1836) proposed Stenactis strigosa DC.

5. S. strigosa, caule erecto sparse et patentim piloso, foliis lanceolatis utrinque stteniatis dentatis strigoso-hispodid, capitulis laxe corymboso-paniculatis, invol. Glaberrimo, ligulis numerosis filiformibus. ② ? In Pensylvania (Muhl.), Noveboraco (Torrey!). Erigeron strigosum Muhl. In Willd. Sp. 3. p. 1956. Erig. Australe Horn. Ex Spreng. An Erig. Strigosun Ell. Sketch 2. p. 394 et Doronicum ramosum Walt. Fl. Car. 206 in Carolina cresc. Ad hanc speciem referenda? (v. s. comm. A cl. Torr.)

Literature Cited:
- Nesom, Guy L., 2008.  

Stenactis is currently treated as a section of Erigeron (Nesom, 2008).
Erigeron tracyi Greene. Running Fleabane.
  Some of the current synonyms for E. tracyi are:
  • Erigeron cinereus A. Gray
  • Erigeron cinereus var. cinereus
  • Erigeron cinereus var. aridus M. E. Jones
  • Erigeron colomexicanus A. Nelson
  • Erigeron commixtus Greene
  • Erigeron dicladus Greene
  • Erigeron divergens Torrey & A. Gray var. cinereus A. Gray
  • Erigeron tephroides Greene
  • Erigeron williamsii Phil. The Plant List and the Global Composite Checklist each give E. williamsii Phil as a synonym of E. tracyi. does not. I suspect that E. williamsii is a valid name for a South American species, and does not apply to E. tracyi.

Literature Cited:
- Gray, Asa, 1849.  

The first known description of the taxon we now call E. tracyi was by A. Gray (1849) in Plantae Fendlerianae Novi-Mexicanae, a description of plants collected by Augustus Fendler in New Mexico in 1841
333. E. cinereum (sp. Nov.): bienne? Undique molliter cinereo-pilosum: caule e basi ramoso; ramis adsurgentibus apice longe nudis monocephalis; foliis spathulatis vel lineari-oblongis basi attenuatis integerrimis seu radicalibus paucidentatis incisisve; ligulus numerosissimis gracilibus (albis nunc purpureo tinctis) involucrum hirsutum duplo superantibus; pappo radii et disci conformi duplici, exteriore coroniformi-squamellato, interiore e setis sub-20 fragilibus deciduis. — Var. a. Is a dwarf, vernal form, only a span high, quite hoary, the primary flowering stems erect and almost scapiform (no. 374 of the distribution). Dry, exposed places around Santa Fe; May. Var. ß has taller and more diffuse stems (10 inches high), the leaves almost lanceolate, entire, the lower tapering into slender petioles. Low, sandy banks of the Rio del Norte and of Santa Fe Creek; May to June. (380.) Var. ? is a larger, coarser, and much more leafy state; from the valley of Santa Fe Creek, near irrigating ditches; May to July. (385.) — The heads are as large as those of Bellis perennis, solitary on peduncles, or the naked summit of the stems, of from 2 to 4 inches in length. The species belongs to the first division of the section Phalacroloma, Torr. & Gray, l. c. † Some forms of this, or of an allied species (possibly E. affine, DC.), with rather less numerous and white rays, and either entire or incised leaves, were gathered at Buena Vista and Encantada by Dr. Gregg.

Unfortunately, the name was unavailable, having been previously used by Hooker and Arnold for their E. cinereus, a native of Chile.

Literature Cited:
- Gray, Asa, and Charles Wright, 1852.  

Gray (1852, pt. 1, p. 91) places his E. cinereum as a variety of E. divergens in Plantae Wrightinae. He make no comments that describe his thinking.

268. E. divergens, var. cinereum. E. cinereum, Gray, Pl. Fendl. p. 68. New Mexico; the locality not recorded.

Gray (1852, pt. 2 p. 77) also places his E. cinereum as a variety of E. divergens in Plantae Wrightinae, part 2.

E. divergens, var. cinereum, Gray, Pl. Wright, l. c. Moutains, around the copper mines; Oct: a late, much-branched state. (1169) Hills near El Paso; March, April: the early normal form (1398.)

This placement is still accepted by Harrington one hundred years later (Harrington, 1954, 1964 2nd ed.).

Literature Cited:
- Buckley, S. B., 1861.  

These plants were collected by the author while engaged in the State Geological Survey of Texas, during 1860 and '61. Specimens of them are in the herbarium of the Academy of Natural Sciences at Philadelphia, and also in the herbarium of Elias Durand, Esq.

Erigeron (Eurigera) nudiflorum, s. n. — Hirsutum pumilum, ramosum; foliis lineari-oblongis, integris, sentis, confertissimis; ramis apice longe nudis, monocephalis; ligulis albis, plurimis, subuniseriatis, involucrum duplo excidentibus, acheniis oblongis glabris vel parum pilosis; pappo radii et disci conformi, duplici; exteriore breve setacea.

Northern Texas. May.
Very much branched from the root. Stems erect, 4-6 inches high; leaves numerous near the root and upwards on the stems about 3 inches; the upper stems, two inches below the flowers, naked; whole plant strigose hirsute, the white hairs not appressed; flowers about 1/4 of an inch in diameter; leaves 1/2 - 1 inch long and a little more than a line wide.


Literature Cited:
- Greene, Edward Lee, 1902.

Other articles:
• Field Notes:   Greene, E. L., 1902;  

E. L. Greene (1902) then published two names for what we now treat as a single species. The names he published were: Erigeron commixtus and E. tracyi.

Erigeron commixtus. With the habit of E. flagellaris, smaller, less stoloniferous, the leaves relativelt broader, some entire, others with one or more conspicuous lobes at base of the blade, the whole plant almost hoary with stiffly hirsute pubescence, this spreading on the leaves and petioles, retrorse on the stems and peduncles: heads, rays, achenes, etc., much as in E. flagellaris.
Cañon of the Limpia, mountains of western Texas, 26 April 1902, S. M. Tracy and F. S. Earle; also collected by the present writer, in the same general region, namely in the mountains near Silver City, New Mexico, 18 May 1880, and distributed for E. flagellaris. The species last named has a rather obscure, fine closely appressed hairiness. That of the new one is so extremely different, that were the plants the size of a Sunflower or Goldenrod, no botanist would confuse them as one species, were the pubescence the only character. I may remark that true E. flagellaris reaches the mountains of even southern New Mexico, where, however, it occurs only in a more elevated biological zone

Literature Cited:
- Greene, Edward Lee, 1902.
- Nesom, Guy L., 2004.  

Erigeron Tracyi. Allied to the last (Erigeron commixtus — Ed.) but dwarf, only 2 or 3 inches high, densely leafy at base and with no stolons (at least at flowering time): petioles of the spatulate-lanceolate entire leaves shorter than the blade or obsolete; the whole herbage silvery-hoary with a fine dense strigulose pubescence, or this more sparse and spreading on the solitary scapiform peduncle: involucre hispidulous: outer pappus very conspicuous though consisting of only shorter and setiform hairs, the inner of a few very delicate capillary ones.
Davis Mountains, western Texas, Tracy and Earle, 28 April 1902.

E. commixtus was published before E. tracyi, quite literally a matter of a few inches. It would therefore seem to have priority over E. tracyi.

Literature Cited:
- Coulter, John M., and Aven Nelson, 1909.

Other articles:
• Glossary:  cinereous;  

33. Erigeron colo-mexicanus A. Nels. Stems few to several from s slender annual taproot, ascending, naked pedunculiform above the middle, very leafy, flowering when very short, the later heads on stems 7-15 cm. long: leaves mostly linear-spatulate to linear, the basal sometimes oblanceolate and 3-lobed at apex, all softly cinereous-pubescent: heads musc as in E. divergens. (E cinereus Gray, Pl. Fendl. 68. 1848; not E. cinereus H. & A. Comp. Bot. Mag. 2: 50. 1836) — Southern Colorado and New Mexico.

It is interesting to me that A. Nelson does not mention Greene's (1902) publication of Erigeron tracyi, whereas other names of Greene are acknowledged.

Literature Cited:
- Rydberg, Per Axel, 1910.  

Erigeron commixtus Greene, Pittonia 5: 58. 1902
E. cinereus A. Gray, Mem. Am. Acad. 4: 68. 1849. Not E.cinereus H. & A. 1836.
E. colo-mexicanus A. Nels,; Coult. & Nels. New Man. Cent. Rocky Mts. 529. 1909.
Another specific name proposed by Professor Nelson, Erigeron colo-mexicanus, to replace the untenable E. cinereus A. Gray, is in my opinion rather distasteful. Fortunately I do not need to use the name, as the same species has been described by Dr. Greene under the name E. commixtus.

Literature Cited:
- Tidestrom, Ivar, 1925.  

Tidestrom (1925) recognizes Buckley's E. nudiflorus.

7. Erigeron nudiflorus Buckley, Proc. Acad. Phil. 1861. 456. 1862.
Erigeron cinereus A. Gray, Mem. Amer. Acad. n. ser. 4: 68. 1849. Not E. cinereus Hook. & Arn. 1836
Erigeron divergens cinereus A. Gray, Pl. Wright. 1: 91. 1852.
E. commixtus Greene, Pittonia 5: 58.1902.
Erigeron colo-mexicanus A. Nels. in Coulter, New Man. Rocky Mount. 529. 1909.
Artemisia, pinyon and yellow pine belts; Utah and Colorado to Mexico.

The author also recognizes E. divergens Torr. & Gray, and E. flagellaris Gray.

Literature Cited:
- Tidestrom, Ivar, and Sister Teresita Kittel, 1941.  

Tidestrom and Kittell (1941) recognize Buckley's E. nudiflorus.

9. Erigeron nudiflorus Buckley, Proc. Acad. Phil. 1861. 456. 1862.
E. commixtus Greene, Pittonia 5: 58.1902.
Artemisia, Pinyon and Yellow Pine belts; Utah and Colorado southward to Mexico.

The authors also recognize E. divergens Torr. & Gray, and E. flagellaris Gray.

Literature Cited:
- Cronquist, Arthur, 1947.
- Gray, Asa, and Charles Wright, 1852.  

Cronquist (1947) accepts Grays (1852) treatment of E. tracyi as a variety of E. divergens.

Along the way his key uses arrangement of hairs on stems as as a character to distinguish between E. divergens and E. flagelaris:

6. Some or all of the hairs of the stem appressed or closely ascending, or the stem glabrous.
which leads to E. flagellaris. The other half of the couplet:
6. Hairs of the stem all spreading.
leads to E. divergens where E. tracyi is treated as a variety.

Key to the varieties of Erigeron divergens
1. Earliest heads on leafy peduncles; plant without long stoloniform branches. A. var. typicus.
1. Earliest heads on long naked peduncles; plant later producing long leafy stolons or stoloniform branches. B. var. cinereus.

112B. Erigeron divergens var. cinereus A. Gray, Pl. Wright. 1: 91. 1852.
E. cinereus A. Gray, Mem. Am. Acad. II 4:68. 1849. Not H. & A. 1836.
E. nudiflorus Buckl. Prec. Acad. Phil. 1861: 456. 1862.
E. commixtus Greene, Pittonia 5:58. 1902.
E. tracyi Greene. Pittonia 5: 59. 1902.
E. divergens nudiflorus A. Nels. Man. Bot. Rocky Mts. 529. 1909.
E. colo-mexicanus A. Nels. Loc. Cit.
? E. dicladus Greene. Leafl. 2: 214. 1912.
Earliest heads on long naked peduncles; plant later producing long loafy stolons or stoliform branches; basal leaves more often persistent than in var. typicus. Southern Nevada and Utah, through Arizona and New Mexico to northern Mexico, central Texas, western Oklahome, and southwestern Kansas; reported from Colorado and western Arkansas.
Type: Fendler 374, near Santa Fe, New Mexico, 1847 (NY).
… Texas: Tracy & Earle 279, Limpia Canyon, April 26, 1902 (type of E. commixtus Greene) (MI, NY); Tracy & Earle 320, Davis Mountains, April 28, 1902 (type of E. tracyi Greene) (MI, NY); … Colorado: Rydberg & Vreeland 5438, 5 miles southwest of La Veta, Huerfano Counto, May 22, 1900 (NY); …
The variety cinereus has usually been treated as a distinct species under the name E. nudiflorus Buckl., but there are two many intermediates for it to stand. There are occasional intermediates with E. modestus, having the pubescence of the latter, and habit of E. divergens var. cinereus. These plants are more likely to have conspicuously lobed basal leaves than is typical var. cinereus.

The variety cinereus is what we are calling E. tracyi today.

Literature Cited:
- Harrington, H. D., 1954.
- Harrington, H. D., 1964, 2nd ed..  

Harrington (2nd ed., 1964, p. 565) placed E. tracyi as a variety of E. divergens. This repeats his treatment of the taxon in his first edition (Harrington, 1954).

8B. Erigeron divergens cinereus A. Gray, (var.) Pl. Wright. 1:91. 1852.
E. cinereus A. Gray; E. nudiflorus Buckl.; E. commixtus Greene; E. colo-mexicanus A. Nels.; E. divergens nudiflorus (Buckl.) A. Nels. -- Earliest heads on long naked or nearly naked peduncles; plants producing long leafy stolons later in the season. --- Dry often sandy soil. Kansas ro Nevada, south to Texas and Arizona. Our records scattered over Colorado, mostly in the southern part, at 4000-8000 feet.

Of note is Harrington's acceptance of E. commixtus but not E. tracyi.

Literature Cited:
- Nesom, Guy L., 1989.  

Nesom (1989c) accepted E. tracyi as having priority over E. colomexicanus.
Erigeron tracyi an earlier name for Erigeron colomexicanus
I have been using Erigeron colomexicanus as the name for this species, but both E. tracyi and E. commixtus were published seven years earlier, simultaneously (immediate succession in the same paper: Greene 1902). Plants of the type collections of E. tracyi, as well as those of E. commixtus, are early season forms (essentially a basal rosette with a single, subscapiform, monocephalous stem) that had not yet produced runners characteristic of the species.
Erigeron tracyi Greene, Pittonia 5:59. 1902. Type: U.S.A. Texas. [Jeff Davis Co.:] Davis Mts., 28 Apr 1902, S. M. Tracy and F. S. Earle 320 (Holotype: US!; Isotypes: GH!, NY!, OC!).
Erigeron commixtus Greene, Pittonia 5:58. 1902. Type: U.S.A. Texas. [Jeff Davis Co.:] Cañon of the Limpia, Mountains of west Texas, 26 Apr 1902, S. M. Tracy and F. S. Earle 279 (Holotype: US!; isotypes: GH!, NY!, TAES!, TEX!).
Erigeron cinereus A, Gray, Mem. Amer. Acad. Arts n.s., 4 [Pl. Fendler.]; 68. 1849 (not Hook. & Arn. 1836). Erigeron divergens Torrey & A. Gray var. cinereus (A. Gray) A. Gray, Smithsonian Contr. Knowl. 3, Art. 5 [Pl. Wright.]:91. 1852. Erigeron colomexicanus A. Nels. [nom. Nov.], Man. Bot. Rocky Mts., 529. 1909. Type: U. S. A. NEW MEXICO. [Santa Fe Co.:] near Santa Fe, 1847, A. Fendler 374 (Holotype: GH!; isotypes: GH, NY! UC-2 sheets!, US!).

Literature Cited:
- Flora of North America Editorial Committee, eds., 1993+.  

Volume 20 of Flora of North America North of Mexico was published in 2006. The treatment of Erigeron was written by Guy L. Nesom.

160. Erigeron tracyi Greene, Pittonia. 5: 59. 1902.
Running fleabane
Erigeron cinereus A. Gray 1849, not Hooker & Arnott 1836; E. colomexicanus A. Nelson; E. commixtus Greene; E. divergens Torrey & A. Gray var. cinereus A. Gray
Annuals, biennials, or short-lived perennials, 2.5–8(–12, 18) cm; usually taprooted, sometimes fibrous-rooted, caudices simple or branched. Stems first erect (greenish proximally), then producing herbaceous, leafy, prostrate runners (stoloniform branches, sometimes with rooting plantlets at tips), densely hirsutulous (hairs spreading-deflexed, of relatively even lengths and orientations), sparsely minutely glandular. Leaves mostly basal (persistent in early season); blades oblanceolate to spatulate (obovate-elliptic laminae), 10–30(–60) × 3–6(–12) mm, cauline abruptly reduced distally, margins entire, dentate, or lobed, faces densely hirsute, eglandular. Heads 1(–3 rarely, from midstem or proximal branches). Involucres 3.5–4.5(–6) × 6–9(–12) mm. Phyllaries in 3–4 series, sparsely to moderately hirsute, minutely glandular. Ray florets 60–130; corollas white, often purplish abaxially, sometimes with an abaxial midstripe, 5–9 mm, laminae not coiling or reflexing. Disc corollas 2–3 mm (throats indurate and slightly inflated). Cypselae 0.7–1.3 mm, 2-nerved, faces sparsely strigose; pappi: outer of setae, inner of 12–16 bristles. 2n = 27.
Flowering Mar–Oct. Desert scrub, grassy slopes, oak chaparral, pinyon-juniper woodlands, Douglas fir-ponderosa pine; 700–2300(–2400) m; Ariz., Colo., Kans., Nev., N.Mex., Okla., Tex., Utah; Mexico (Baja California, Chihuahua, Coahuila, Durango, Sonora, Zacatecas).
In March through June, plants of Erigeron tracyi produce leaves in a basal rosette usually with a single, erect, monocephalous, stem. Stoloniform branches are soon formed (often recognized on pressed specimens by the leaves mostly on one side of the branches), and by the end of the season (August through October), prostrate runners are usually evident, sometimes forming terminal, rooting plantlets.
Erigeron tracyi is similar in habit to E. flagellaris, particularly in the herbaceous stolons or stoloniform branches; the stem pubescence of E. tracyi is different, the stolons much less commonly produce rooting plantlets at the tips, and the plants tend to be perennial with woody or lignescent caudices, although they are variable both in habit and duration. Apparent hybrids with E. modestus and E. flagellaris are occasionally encountered, and the most common form of E. tracyi is perhaps (speculative) a stabilized, apomictic hybrid between the latter and E. divergens. All chromosome counts thus far have shown E. tracyi to be triploid and asynaptic.

Literature Cited:
- Snow, Neil, 2009.  

Erigeron tracyi Greene - FNA
[Erigeron cinereus A. Gray 1849 {not Hook. & Arnott 1836} - FNA
[Erigeron colomexicanus A. Nelson] - KTZ, E
[Erigeron colo-mexicanus A. Nelson] - SFE, W&W
[Erigeron commixtus Greene] - FNA
[Erigeron divergens Torr. & A. Gray var. cinereus A. Gray] - HDH, UTF
[Erigeron divergens Torr. & A. Gray var. cinereus (A. Gray) A. Gray] - IMF
[Erigeron modestus A. Gray] - GPF

Literature Cited:
- Weber, William A., and Ronald C. Wittmann, 2012.  

11a. [10] Heads solitary on elongate, mostly leafless stems from the basal leaf clusters; plants developing long, nonrooting, stolon-like, spreading shoots. E. tracyi Greene [for Samuel Mills Tracy, 1847-1920]. Dry, gravelly floodplains and meadows, mimicking E. flagellaris but with spreading stem hairs. Flowering very early in the spring. The FNA suggests that this is a triploid apomictic hybrid between E. divergens and E. flagellaris. (E. colomexicanus is a later name.)

Literature Cited:
- Ackerfield, Jennifer, 2015.  

Ackerfield's (2015) description of E. tracyi does not describe the stem hairs, or the direction in which they point. Also, I think the stated length of the stoloniferous branches is quite short. Perhaps, the length should be expressed in decimeters, such as “... stoloniferous branches, 1-3(6) dm. Long.” Finally, the only synonym identified is E. colomexicanus perhaps because it has long been used in Colorado and New Mexico, but E. commixtus and E. nudiflorus are not noted.

Erigeron tracyi Greene, Running Daisy. [E. colomexicanus A. Nelson]. Annuals, biennials, or short-lived perennials, 1-4 dm.; leaves oblanceolate to linear, entire or few-toothed, the stems forming stoloniferous branches, 1-3(6) cm. long; involucre 3-5 mm. hih, hirsute with curved hairs; disk flowers 2-2.5 mm. long; ray flowers 4-6 mm. long, white or pink; pappus double. Common on the eastern slope in open meadows and on dry slopes, with scattered occurrences on the estern slope, 4000-9000 ft. April-July. E/W

Couplet 24 of Ackerfield's key to Erigeron addresses the question of stem hairs:

24a Stems with spreading or tangled (pointing in all directions) hairs or mostly glandular … 25
24b. Stems with appressed or ascending hairs (at above the middle), not glandular … 34

Couplet 25 eventually ends up at Couplet 29 which differentiates between E. tracyi and E. divergens.

Couplet 34 eventually leads to E. flagellaris.

Erigeron vetensis Rydb. Early Bluetop Fleabane.

Literature Cited:
- Rydberg, Per Axel, 1905.  

Erigeron vetensis sp. nov.
Densely cespitose-puvinate perennial ; stems 5-8 cm. high, hirsute, few-leaved ; leaves linear or linear-oblanceolate, hirsute, 2-4 cm. long; heads solitary, about 7 mm, high ; bracts hnear, acuminate, hirsute as w^ell as slightly glandular-puberulent ; rays purple, 8-10 mm. long, over i mm. wide; achenes strigose; pappus more or less double.
In dry places on high mountains of southern Colorado at an altitude of 2400-3000 m. It is intermediate between E. radicatus and E. glandulosus, resembling the former most in pubescence, and the latter in habit.
Colorado: Mountains near Veta Pass, 1900, Rydberg & Vreeland 5427 (type); Veta Mountain and Ojo, 5421, 5422; West Spanish Peak, 5424.

Other articles:
• Ericameria nauseosa var. graveolens:  Introduction;  

Ericameria nauseosa (Pall. Ex Pursh) G.L.Nesom & G.I.Baird var. graveolens (Nutt.) Reveal & Schuyler

  • How did we get to the name of Ericameria nauseosa var. graveolens?
    • What is the history of the genus name Chrysothamnus?
    • What is the history of the name Ericameria nauseosa?
    • What is the history of the name ____ graveolens? And how did it become a variety of Ericameria nauseosa?

See my page about Ericameria nauseosa var. graveolens or “How did rubber rabbitbrush get that long scientific name?”

Other articles:
• Water Tank Road:   above the curve;  

Ratibida columnifera

Other articles:
• Field Notes:  Coll. No. 1198, 12 Jul 2015;
Full Size ImageColl. No. 1198, Ratibida columnifera  

So, is the cypsela ciliate on the abaxial side, or the adaxial side?

Symphyotrichum porteri (A. Gray) G. L. Nesom

Literature Cited:
- Porter, Thomas C., and John M. Coulter, 1874.  

Publication of Aster ericoides L. var strictus Porter in Porter and Coulter (1874).
Aster ericoides, L., var strictus, Porter. Low ¾ °-1° high, glabrous, except the scabrous margins and ciliate bases of the leaves, erect, slender, paniculately branched above, branches short; scales of involucre narrowly linear, lax, outer ones very acute, often entirely green, inner ones scarious with a central green line; radical leaves narrowly oblanceolate. -- “In the mountains at middle elevations,” Hall & Harbour, 254. Near Denver, Coulter. Foot-hills west of Denver, Porter; Meehan; Hoopes.

Literature Cited:
- Gray, Asa, 1880.  

basionym:Asteraceae Aster porteri A.Gray Proc. Amer. Acad. Arts xvi. (1880) 99.

Among the true Asters are several forms which have to be named, such as A. Porteri for A. ericoides, var. strictus, Porter & Coult. Fl. Colorado. 56, and A. pringlei, from the northern end of Lake Champlain …


Bouteloua Grama Grass

Genus name honors two Spanish brothers Claudius (d. 1842) and Exteban Boutelou (d. 1813) professors of botany and agriculture respectively.

Literature Cited:
- Gould, Frank W., 1979.  

Abstract. Bouteloua was established in 1805 by Mariano Lagasca. The type species is B. curtipendula (Michaux) Torrey, originally named B. racemosa by Lagasca. In the present treatment, 39 species are recognized, 29 of these restricted to North America and Central America, 2 species are endemic to the Antilles, 2 species occur in the Antilles, as well as at other locations, 5 species are distributed in both North and South America, and 1 species, B. megapotamica, is represented only in southern South America. Bouteloua is a characteristic member of the tribe Chlorideae of the subfamily Eragrostoideae (Chloridoideae). The species all are C4 plants with typical Kranz leaf anatomy and starch storage features. Chromosome numbers have been reported for 29 species with most species being diploid (2n = 20) or tetraploid (2n = 40). Aneuploid records or series of counts have been reported for 9 species.

All species of Bouteloua are characterized by features of the Kranz Syndrome. They are C4 in their photosynthesis and have a characteristic chloridoid leaf anatomy. Starch storage is in specialed plastids of the leaf sheath bundles, and the arrangement of cells in the leaf blade is typically Kranz. The 39 recognized species are all variously stapted to shortgrass prairies, desert grasslands, and xeric sites along desert shrub areas, and sandy shores.

The genus Bouteloua was described in 1805 by Mariano Lagasca. Lagasca proposed five species, B. racemosa, B. hirsuta, B. barbata, B. simplex, and B. prostrata, but did not designate a type species. Griffiths (1912), Hitchcock (1920), and Hitchcock et al. (1939) all have accepted the first species, B. racemosa, as the type. As plants of the type species had been named Chloris curtipendula by Michaux in 1803, the legitimate name for this taxon is Bouteloua curtipendula.

Bouteloua was named in honor of two Spanish gardeners, the Boutelou brothers. Lagasca's original spelling of the genus name was Botelus. In a later publication (1816), he corrected this to Bouteloua.

Two subgenera, Bouteloua and Chrondrosium (Desvaux) Gould. Weber & Wittmann (2012) retain Chrondrosium at the rank of genus, though spelling it Chrondrosum Desvaux, suggesting that we see Clayton, W. D., 1986, Genera Graminum: Grasses of the World. Kew Bull. Addit. Ser. XIII. 389 p. Peterson, et al. (2015) treat Chrondrosum as a section of Bouteloua containing B. gracilis and one other known in Colorado, B. simplex.


Other articles:
• Golden Checklist Flora:  Bouteloua curtipendula;  

Bouteloua curtipendula (Michx.) Torr. in Marcy. “Side-Oats Grama”


Literature Cited:
- Michaux, Andre, 1803.  

The basionym of B. curtipendula is Chloris curtipendula Michx.
Original Text My Interpretation
CHLORIS. Sw. CHLORIS. O. P. Swartz (1788) Nova Genera & Species Plantarum seu Prodromus
Spicae unilateriflorae, spiculis subsessiliter biseriatis : quarum gluma communis 2-valvis, 2-6-flora : floribus dissimilibus ; valva altera ( saltem nonnullorum ) aristata : uno hermaphrodito , fertili ; caeteris inperfectis , masculis neutrisve ; ultimo pedicellato. Spikes unilateral flowered, darts subsessiliter biseriatis, of which the husk common 2-doors, 2-6-flowered, flowers are different; second valve (at least some) anstata one hermaphrodito, fertile; the rest of the imperfect, or the males neutrisve; last pedicellatae.
curtipendula. C. racemo erecto longo ; e spicis plurimis , distiche alternis , e basí emittens rudimentum secundi floris inane , promisse aristatum ; intra quod bina alia rudimenta quasi in sola arista consistentia. curtipendula. Chloris, even on a long raceme; very many out of the ears of corn, distiche every other day, And I will send the initial stage of the second base of the flower out of the void, Retz promised; the rudiments of other things, and within it two of every sort as the only grain condition.
Plantae cultae statura major; spicae 6-12-glumes. If cultivated plants are taller; 6-12-spike glumes.
Hab. in aridis regionis Illinoensis ad Wabast et in rupibus ad prairie du rocher. ♃ Habitat. In arid regions of Illinois to Wabash River and watercourses of Prairie de Rocher (a town in southwest Illinois). Perennial.

Prairie du Rocher is one of the oldest communities in the 21st century United States having been founded in 1722 by French colonists, mostly migrants from Canada. About four miles to the west, closer to the Mississippi River, is Fort de Chartres, site of a French military fortification and colonial headquarters established in 1720.

Literature Cited:
- Emory, William H., 1848.  

Emory (1848) hinted as the existence of Bouteloua curtipendula as a name but this would be invalid because he really did not place B. racemosa in synonomy. I assume that Emory had heard from Torrey that B. curtipendula was the correct name but perhaps did not know that Torrey had not published the name.

____teloua racemosa, Lagasca. ? Culm erect, simple; spikes nu-
____s (20-40,) reflexed, 3-flowers; lower glume linear subulate;
____ one linear-lanceolate, scabrous, entire, nearly as long as the
____ts; lower palea of the perfect flower unequally tricuspidate,
____ent; abortive flower reduced to a slender awn which is nearly
____g as the perfect flower, furnished at the base with 2 short
____onspicuous bristles. Valley of the Gila, rare. This plant
____pretty well with Kunth's description of B. (Eutriana,) race-
____xcept in the pubescent lower palea, and the minute bristles
____base of the neiter flower. Whether it be the plant of La-
____r not is very difficult to determine from his brief character.
It certainly is very different from B. racemosa of the United States,
which has a large 3-awed neuter flower, and if distinct from La-
gasca's, must receive another name. That of B. curtipendula
would be appropriate.

Literature Cited:
- Marcy, Randolph B., 1854.  

IPNI (2020) indicates that Bouteloua curtipendula was credit to Torrey in Marcy (1854), whereas Marcy seems to credit Torrey in Emory's report. IPNI also suggest that new combination was invalid, “Torrey (in Emory, Notes milit. Reconn. 154. 1848) mentioned this comb. Nov. (invalid)”
Bouteloua racemosa, Lag. Var. Cienc. (1805) p. 141 ; Torr. In Emory's Rep., p. 154 ; not of Torr. Fl. N. York. Dinebra curtipendula, DC.? Kunth, Syn. Pl. Eq. 1, p. 281 ; excl. syn. Michx. Eutriana curtipendula, Trin. Fund. P. 161 (in part); Kunth, Enum. 1, p. 280, and Suppl. P. 233 ; excl. syn. Michx. And Willd. Main Fork of Red River; July. The detailed description of this species by Kunth, l. c., (drawn from a Mexican specimen collected by Humboldt) shows that the Chloris curtipendula of Michaux (Bouteloua curtipendula, Torr.) is a distinct species, as indicated in Emory's report, l. c.


Other articles:
• Golden Checklist Flora:  Bouteloua gracilis;  

Bouteloua gracilis (Kunth) Lag. ex Griffiths. Blue Grama Grass.


Literature Cited:
- Nuttall, Thomas, 1818.

Other articles:
• Golden Checklist Flora:  Nuttall, 1818, publication details;  

110. ATHEROPOGON. Muhlenberg.
... [Description of A. apludoides, syn: Chloris curtipendula, Mich. = Bouteloua curtipendula (Michx.) Torrey – Ed.]
2. A. * oligostachyum. Spikes 2 or 3, nearly terminal, many flowered; calix and corolla pilose; outer valve of the corolla distinctly 3-awned, the 2 lateral awns shorter, arising near the middle of the valve; neutral valve 3-awned.
On the plains of the Missouri with the above. [Bouteloua curtipendula – Ed.] Common.
Culm round, filiform, nearly naked, or with a single leaf, 8 to 12 inches high, smooth and erect. Leaves very short, smooth, and subulate, stipule and base of the spikes shortly bearded. Spikes 1, 2, or 3, about an inch long, usually curved backwards, unilateral, compressed, and pectinate, the second spikes bibracteate, rachis semiterete. Glumes in a double row, opposite; each 2-flowered; calix bluish-purple, exterior valve lanceolate, mucronate, with a single nerve; the nerve beset with a few scattered hairs arising from so many tubercles; inner valves shorter, very narrow. Corolla, outer valve lanceolate, carinate, 3-awned, pilose along the margins of the nerves, and at the base; inner valve smooth, shortly bi-cuspidate. Neutral flower 1-valved, obtuse, with 3 awns, and pubescent at the base.
This species, though certainly a congener of the preceding, is considerably allied to Chloris, appearing to unite that genus and Sesleria, agreeing partly with the latter in the structure of the flowers, and with the former in its habitus. (A North American genus)


Other articles:
• Golden Checklist Flora:  Buchloe dactyloides;
• Field Notes:  Coll. No. 1673, 16 Jun 2017;   Coll. No. 1674, 19 Jun 2017;  Coll. No. 1690, 29 Jun 2017;  

Buchloë dactyloides (Nutt.) Engelm. Buffalo Grass.


Literature Cited:
- Nuttall, Thomas, 1818.

Other articles:
• Glossary:  v. v.;  

95. SESLERIA. L. (Moor-grass.)
Calix 2 to 5-flowered. Corolla 2-valved, valvea toothed at the point. Stigmata somewhat glandulous. — Flowers spiked, often purplish, base of the spike breacteate, or involucrate. Early flowering subalpine grasses, growing in calcareous mountains.
Species. 1. S. Dactlyoides. Culm setaceous, leafy; leaves short, flat, subulate, and somewhat hairy; stipules bearded; spikes 2 or 3, few-flowered; flowers in 2 rows, disposed upon an unilateral rachis, calix mostly 2-flowered, and with the corolla acuminate and entire.
Hab. On the open grassy plains of the Missouri; abundant. Flowers in May and Junes. v. v. Root after flowering resembling a bulb.
Culm smooth and round, furnished with 2 or 3 leaves, about 4 or 5 inches high. Leaves flat, subulate, and somewhat hairy, 1 to 2 inhes in length, and about 2 lines wide; sheathes shorter than the internodes, very hairy around the stipules. Spikes 2 or 3, somewhat ovalm subtended by a single leaf, with which they are at first sheathed; rachis compressed, margined,spikelets 6 to 8, by pairs, inclined to one side. Calix 2-valved, 2 or 3-flowered, vales very unequal, each with a single nerve and carinate, the larger oblong-ovate, mucronulate. Outer valve of the corolla oblong-lanceolate, entire, 3-nerved, smooth, and menbranaceous, longer than the calix; inner 2-nerved, nearly the length of the outer. anthers linear, entire, fulvous, exserted. Styles filiform, pubescept.
This species appears on the one hand, alloed to Atheropogon, and on the other to Dactylis. Though rather a Sesleria than any other genus, it recedes from it in having the valves of the corolla entire at the apex, and thus it approaches Dactylis at least, the D. glomerata.
With the exception of the present species, the genus Sesleria is confined to the alpine regions of Northern Europe.

Literature Cited:
- Rafinesque, C. S., 1819.

Other articles:
• Glossary:  nomen nudum;  

Rafinesque (1819) published a genus name for Nuttall's grass, recognizing that the grass must be distinct from the Old World Sesleria.

18. Sesleria dactyloides must form a peculiar genus by Mr. N's. own account, it may be called Bulbilis.

The question will become whether this is a validly published name, or whether it is a nomen nudum.

Literature Cited:
- Torrey, John, and Asa Gray, 1838-1843.  

I don't see any grasses in Torrey & Gray's (1838-1843) Flora of North America.

Literature Cited:
- Marcy, Randolph B., 1854.  

Sesleria dactyloides, Nutt. Gen, 1, p. 65; Kunth, Enum, 1, p. 323; Torr, in Emory's report, p. 323, t. 10. Upper tributaries of the Red River; July. This is the well known Buffalo-grass of the western prairies. It is remarkable that neither the grain nor the fertile flowers of this grass are known.

Literature Cited:
- Engelmann, George, M.D., 1859.  

Original Text Comments and Interpretation
By George Engelmann, M.D.  
The grasses, though usually hermaphrodite, show a tendency to a separation of the sexes, and polygamous flowers are not rare among them. About 25 to 28 genera, one-twlefth of the whole number known, comprising only 75 to 80 species, about one seventy-fifth of all species, * are described as having monoecious and mostly heteromorphous flowers.  
Only two genera of dioecious grasses are known to the books; of these, Spinifex, Lin., with 6 species from the East Indies and Australia, bearing on some plants staminate and on others complete flowers, is only incompletely dioecious; the other genus is Gynerium, H. B. K., five South American species. Some other dioecious species of genera, generally hermaphrodite, are notived; such as Calamagrostis dioica, Lour., and Guadua dioca, Steud.  
The unisexual grasses mostly belong to Oryzeae, Phalarideae, Paniceae, and Rottboellieae; none have been known among the tribes of Stipeae, Agrostideae, Chlorideae, Avenaceae, Festiceae, and Hordeeae.  
They were unknown in the northern temperate zone, with the exception of Zizania and Tripsacum of North America and the cultivated Zea, all with heteromorphous staminate and pistillate flowers on the same plant. The dioecious grasses of our Flora are both species of Brizopyrum;† Eragrostis reptans is also frequently or mostly dioecious, and other species of this genus seem to be imperfectly so.  
In the following pages, two new dioecious North American grasses are described, both types of new and very distinct genera, and both, it is believed, belonging to Chloridae.  

* In the latest work on Grasses, Steudel's Glumaceae, published in 1855, about 6,000 species of Grasses are described, very unequally distributed in about 300 genera, many general containing only a single species, while Panicum alone comprises 864, Andropogon 461, Eragrostis 247, and Festuca 239 numers.  
Brizopyrum spicatum, Hook. is from the eastern seacoast, and B. strictum from the saline soils of the Missouri region and of Utah. The flowers of both sexes are conform, but the staminate plants are readily distinguished from the pistillate ones by their more slender growth, the spikes overtopping the leaves; while in the pistillate plants the latter are longer than the spikes. = Distichlis spicata (L.) Greene
  Engelmann's description of the new genus:
Original Text Comments and Interpretation
BUCHLOË, Nov. Gen. BUCHLOË, New Genus
Gramen plantitierum Americae Septentrionalis aridarum Missouriensium, Texensium, Mexicanarumque gregarium, perenne, stoloniferum, humile, sparse pilosum vel glabriusculum; ligulis barbatis. — Buchloe pro nimis longo Bubalochloe nomen vernaculum “Buffalograss,” graece reddit. Grass of arid North America, Missouri, Texas, Mexico, perennial stoloniferous, small, sparsely hairy or glabriusculus; ligules bearded. — Buchloe from a long form, Bubalochloe (Χορτ ο βου β αλ ου) The name of the breed “ Buffalograss” in Greek.

Literature Cited:
- Steudel, Ernst Gottlieb, 1855.  

Description of the new species:
Buchloe dactyloides.
Syn. Plantae masculae: Sesleria dactyloides, Nuttall, Gen. I. p. 64. Sesleria (?) dactyloides, Torrey, in Emory's Rep. 1848, p. 153, Pl. X; id. In Whipple's Rep. Pacif, R.R. Expl., IV., p. 157. Calanthera dactyloides, Kenth (?) in Hooker's account of Geyer's Rocky Mountain plants, in Kew Journ. Bot., VIII., p. 18. Triodiae spec., Bentham, in Pl. Hartweg, nro. 250, p. 28. Lasiostega humilis, Rupprecht (ined) in Benth, Pl. Hartw. Corrig., P. 347. — Drummond Tex., Ill., nro. 378. Lindheimer, Pl. Tex. Exsicc. 569. Fendler N. Mex., 940. Berlandier, nro. 1612 and 1614. Hartw, 250 (fide Gray).
Syn. Plantae faemineae: Antephora axilliflora, Steudel, Glum. I. . 111 — Drummond Tex., II., 359. Wright, 1849, 785; 1851-1852, 2079 (fide Torrey).
This remarkable plant is found in our western pariries from the British possessions throughout the Mossouri Territory, Nebraska, Kansas, and New Mexico, down to Texas and Northern Mexico, and is, under the name of “Biffalo-grass,” well known to hunters and trappers as one of the most nitricious grasses, on which, for a poart of the year, subsist and fatten immense herds of buffalo and the cattle of the hunter and emigrant. Since the time of Nuttall, who published an account of it, in his “Genera,” as early as 1818, the male plant has been collected by almost every botanist traversing those regions. The female plant had escaped the observers until it was described by Steudel, in the year 1855, from Drummond's Texan specimens, as a totally different plant and belonging even to a different tribe. Though Prof. Torrey had already, in Emory's Report, 1848, suggested the probability of the Buffalo-grass being a dioecious plant, the possibility that Nuttall's Sesleria dactyloides and Steudel's Antephora axilliflora could be the male and female of the same species was not even suspected, till finding both together in a collection sent by my brother, Henry Engelmann, who, as a geologist, accompanied the topographical corps attached to the army of Utah, I was struck with their similarity. My surmise, much doubted at first, became a certainty, when I discovered among some male plants, collected by A. Fendler, about Fort Kearny on the Platte River, a monoecious specimen, showing both male and female flowers on different stalks from the same rhizoma. A figure of this important specimen is given on Pl. XII., fig. 3.
That our plant is distinct from Sesleria has already been stated by Torrey (l. c. p. 154), and indeed by Nuttall himself (l. c. p. 65), and both have pointed to its affinity to Atheropogon or Chindrosium. The description now given fully confirms both positions. It also leaves no doubt that is is not an Antephora, nor at all paniceous. A new generic name, therefore, had to be given, and I have preferred to propose an abbreviated translation of the popular and widely known name of “Buffalo-grass,” retaining of course Nuttall's original specific appellation. The synonyms of the male plant, supplied through the kindness of Prof. Gray, are uncertain, Kunth never having published such a name as Calanthera, which, moreover, is quite unmeaning; nor can I learn that a genus Lasiostega has ever been described.
The Buffalo-grass grows in dense tufts, sending out stolons. These, in most herbarium specimens, are only a few inches long, with internodes of 1/2-2 incles in length; Lindheimer, however, sends specimens from New Braunfels, Texas, with stolons 1-2 feet long, the internodes often measuring over 3 and even as much as 5 inches. The male plant seems to throw out more numerous runners than the female, and may often overspread and kill it out. Which would account for the much greater scarcity of the latter.
Leaves 2-4 inches long, 1/2-1 1/2 lines wide, sparsely hairy or ciliate or glabrous; sheaths striate, glabrous, strongly bearded at the throat.
  Engelmann (1859) describes the male plant:
The flowering stems of the male plant are 4-6 or rarely 8 inches high, mostly glabrous or vey sparsely hairy, generally longer than the leaves, and bear 2 or 3 alternate oblique one-soded spikes. These spikes are 3-6 lines long, and bear on the lower, outer, side of the flattened, dentate, pubescent rhachis 6-15 minutely puberulent spikelets, alternate in two rows; the uppermost spkelet is usually abortive, and is represented by a bristle. The spikelets are 2 or rarely 3 lines long, with 2 or sometimes 3 subterete flowers; they are, as usual, alonf the chlorideous grasses, somewhat obliquely distorted; the glumes are broader on their lower side, but turning upwards, towards the upper end of the spike, they cover the upper edge of the flowers, leaving the lower edge free. The smaller lower glume is, as in this whole tribe of grasses, inside of the spike, and the larger upper one outside and much more conspicuous.
Lower glume ovate-lanceolate, with a scarious margin, convex, scarcely carinate, one nerved, obtuse or acutish or mucronate, one-third of one-fourth the length of the flower; on the uppermost spikelet of each spike, it is much larger, and almost equal to the upper glume.
Upper glume twice as long as the lower one, much wider ovate, obtusish, with a strong middle nerve which sometimes runs out into a point or a short awn between two membranaceous teeth; in a specimen from Fort Kearny, In find on the lower side a second nerve running out into a lateral tooth; a third nerve on the upper side is very indistinct; other spikelets of the same specimen show the ordinary structure.
Lower palea convex, obtusish, 3-nerved, middle nerve in the lower flower running out into a mucro, in the upper one mostly even with the membranaceous margin.
Upper palea as long as, or a little exceeding, the lower one, which partly envelopes it, 2-nerved, 2-carinate, obtuse and scarious at tip.
Scales, lodiculae, 2 at the margin and inside of the lower palea, minute triangular-truncate, undulate or emarginate.
Stamens 3, scarcely exceedin the paleae; anthers linear, bifid at both ends, 1 line long; filaments much shorter. Pollen grains of the dry anther, immersed in water, globular, smoothish, 0.017 line in diameter.
  Engelmann (1859) describes the feminine plant:
The stalk of the female plant is much shorter than the leaves, usually 1½-2 and very rarely 3-4 inches high; it is leafy to the top, the broad, ventricose, many-nerved sheaths of the 2 or 3 uppermost leaves serving as involucres for the flower heads. These heads, 3-3½ lines long, are usually 2 in number, one almost sessile between between both top leaves and on the side of the lower one, the other on a flattened pedicel on the side of the upper leaf; sometimes a third head os noticed still higher, and on the side of the lowest one. The pedicels of these heads are nothing but the common peduncle or rhachis terminating the stem, and the heads themselves correspond to an many lateral spikes of the male plant, in position as well as instructure; they are only more contracted, consists of fewer, one-flowered, spikelets, and turn obliquely upwards, while the male spikelets turn obliquely downwards. Where the number of spikelets does not exceed 3, the head is ventricose-cylindric, or, in fruit, subglobose; but when the number is larger, it is compressed and laterally elongated, showing distinctly enough the biserial arrangement. The pistillate spikelets are like the staminate ones arranged so that the lower glume is inside and the upper one outside in the head.
The thick rhachis of these heads is firmly united with the upper glumes, which in fruit become ligneous; They have a pale whitish or straw color, and smooth and shining surface, with a few scattered hairs; at maturity they separate entire from the common peduncle, at the insertion of which a tuft of short hair is noticed.
The lower glume is a small triangulat or ovate, obtuse, acute or acuminate, 1-nerved, carinate scale, completely hidden in the interior of the head. The lower glume of the lowest pikelet only is larger with 2 or 3 foliaceous points, turned half outside and with its lower edge united to the back of the corresponding upper glume.
Upper glumes much larger, the largest organ of the spikelets, convex outside, concave inside, broadly ovate, narrowed at base, and separated there from one another by a deep and rounded sinus; terminating into 3 herbareous, nerved, lanceolate-linear lobes, a larger central and 2 smaller lateral ones.
Lower palea, of course, opposite the upper glume and therefore turned towards the centre of the head, much smaller, also 3-pointed, with two further lateral membranaceous teeth, enveloping the upper palea.
Upper palea still smaller, 2-nerved, membranaceously 2-pointed.
Scales similar to those of the staminate flowers.
Three minute redimentary stamens at the base of the short-stipitate, compressed lenticular ovary, which at the apex bears 2 rather short erect styles and elongated stigmata, protruding from the apex of the flower. Hair of the stigma simple, rather short and scarcely dentate.
Mature head ventricose, thick, extremely hard, enclosing the loose grain. Grain about 1 line long, orbicular, ovate, flat on the outer (turned towards the lower palea and inside of the head) and convex on the inner face, 2-pointed at the apex by the persistent bases of the styles. Embryo on the flat outer side of the grain and almost as long as it.
Each spike or head has, with very rare exceptions, at the upper end and incomplete rudimentary spikelet, consisting of a single 2-3-pointed upper glume. This with the similar looking lower glume of the lowest spikelet, seems at first glance to form an involucrum for the head; or the upper glumes together have been taken for an involucrum; the lower glumes were overlooked, and the plant placed with the paniceous genus Anthephora, Schreb., which is closely allied to Cenchrus. From the analysis given above, it will be seen that this view is based on a very superficial examination of our plant; and that the structure of the head is entirely analagous to that of the staminate spike of this and the spike of the chlorideous grasses in general, from which the Buffalo-grass can in no manner be separated.
The Engelmann article then continues with a description of Monanthochloë, and M. littoralis Engelm. Also known as shore-grass.

Literature Cited:
- Plank, E. N., 1892.

Other articles:
• Golden Checklist Flora:  Buchloë dactyloides, Hitchcock, 1895;  

E. N. Plank (1892) argued that the plant was actually monoecious.
Buchloe dactyloides, Englm., not a Dioecious Grass.
The common buffalo grass of the “plains” is well known to botanists. It begins to appear near the ninety-fourth meridian, and extends westward to the Rocky Mountains, northward into the British possessions, and southward to the sands of the Gulf coast. It is a grass of short stature, seldom more than six inches tall, though I have specimens collected in this State more than a foot high. Our spcies was formerly of great repute among stockmen in the central and northern parts of its range as a winter forage plant. Much of the credit given to it belogs, however, to species of Bouteloua, especially to B. hirsuta and B. oligostachya (=B. gracilis – Ed.). It usually neighbors closely with those grasses, which are regarded by many ranchmen as merely forms of it. While doubtless a very nutritious grass, its manner of life adds very much to its valus. It has a way of sending its roots down ten or fifteen feet, which, combined with its stoloniferous character and its numerous curling linear leaves, enables the plant to live and thrive under most unfavorable climatic conditions. Naturally almost hay, it readily cures under the dry, hot winds into a valuable fodder for the wild deer and the wild buffalo, as well as for their successors, the hardly less wild steers of the cattle men, Further east and south, it largely yields place to more luxuriant grasses.
The history of our subject is interesting from the fact that, for a long time after its discovery, what has hitherto been regarded as its tue character, was unknown to botanists. Nearly seventy five years ago, Nuttall discovered the staminate form and named it Sesleria dactyloides. In 1855 Steudel names the pistillate form, collected by Drummond, Antephora axilliflora — no botanists at that time recognizing the relationship of the two forms. In 1859 the late Dr. George Engelmann, of precious memory to botanists, received what he considered a mere sprout of the plant, a form bearing both staminate and pistillate flowers, but borne on distinct stalks proceeding from different parts of the same rootstock. The practiced eye and judgement of Dr. Engelmann saw at once that Sesleria dactyloides and Antephora axilliflora were forms of the same plant. Supposing that he had before him a dioecious grass so distinct from all others as to possess generic characters, he founded a new genus and named it Buchloe, a contractions of Bubalo, Greek for the Old World buffalo, and chloa, grass.
Great and important facts and principles in science and philosophy often lie around loosely, and cuffed and are kicked about for years, sometimes for centuries, because great men, strongly intent upon another and remote object, overlook that which is nearby and underfoot. The sheep-like world follows the leader until a shance observation reveals the proximate fact, and then every one wonders why every one had not seen it before. From my first introduction to buffalo, about twelve years ago, and without personal knowledge of the plant in its original monoecious form, but knowing that such a form existed, I was led to doubt the correctness of Dr. Engelmann's conclusion that it is really a dioecious grass. The presumption of doubt strengthened when I found an individual plant bearing both staminate and pistillate flowers. Further experience confirmed my strong notion into what may be considered a fact.
During one of my botanical rambles in Kansas, while walking over soil newly moved by a freshet, I noticed the peculiar appearance of the individual plants of buffalo grass growing upon it. There were scores of them, if not hundreds. All of them appeared to be seedlings, having not yet sent out stolons. All of these plants were monoecious. That seems to be sexually the original character of the species. The fact of the unisexual flowering stems, proceeding from different parts of the plants, with its stoloniferous character generally increasing and spreading in that way, will fully account for its dioecious habit.
The facts of the rapid disappearance of the species with the advent of civilization and agriculture, the paucity of pistillate plants, and that the pistillate form, like the staminate one, is seldom, if ever, found growing singly, can now be easily explained. The staminate stolons are much stronger than the pistilate ones, so that they easily outgrow and overrum the others, becoming sometimes two to five feet lone. There are probably a thousand individuals of the staminate form to one of the pistillate. The few large seeds are eagerly sought by animals. They are often, too, destroyed by a beetle. Seed-bearing individuals being this easily destroyed, few seeds remain for reproduction, and much of that which remains only aids in giving the stronger form more chance in the race of life. So when either form is destroyed its destruction is complete.
If carefully conducted experiments in growing plants of buffalo grass from its seed should show that the plants before they send out stolons are always, or generally, dioecious, which cannot be, as these are too monoecious plants, then Dr. Engelmann's theory of the sexual character may be correct. In the absence of such proof, if we are true philosophers, we are obliged to accept the nearest-by theory that explains the phenomena that the life, character and life-work of our plant presents ; and pronounce it to be originally, as it comes from the seed, a monoecious grass, bearing unisexual flowers upon stalks arising from different parts increasing itself mainly by stolons proceeding from sexually different parts of the plant, each reproducing its own form.
E. N. Plank.

Literature Cited:
- Hitchcock, A. S., 1895.

Other articles:
• Golden Checklist Flora:  Buchloë dactyloides, Plank, 1892;  

Hitchcock (1895) refuted Plank's (1892) observation that Buffalo Grass was monoecious.
Note on buffalo grass. — I read with interest an article by Mr. Plank on “Buchloe dactyloides Englm., not a dioecious grass.” He asserts that the grass in question is not dioecious, as usually described, but monoecious, and in support records observations made in Kansas.
Wishing to satisfy myself experimentally as to the correctness of this assertion, a few seeds were germinated in the greenhouse in the spring of 1893. A single seedling was transferred to an outdoor plat. This grew vigorously through the season, sending out stolons and forming a compact mat. During 1894 the mat became larger and denser, but no flowers appeared.
However, the plant flowered this season (1895). Both staminate and pistillate flowers were present, the former preponderating. The flowers arose mostly from nodes that had taken root and thus become essentially independent plants. In no case did I find the two kinds of flowers from the same node, but from the interwoven state of the stolons I was unable to determine whether the two kinds of flowers were borne upon independent stolons.
The plant was first described by Nuttall (Gen, 1: 65. 1818) from a staminate specimen, and named Sesleria dactyloides. He is evidently doubtful about the plant belonging to the genus Sesleria.
Rafinesque having occasion to review Nuttall's Genera (Am. Monthly Mag. 2: 190. 1819) makes a note regarding this plant: “18. Sesleria dactyloides must form a peculiar genus by Mr. N.'s own account. It may be called Bulbilis.” It is upon this basis that Dr. Otto Kuntze establishes Bulbilis dactyloides (Nutt.) Raf. (Rev. Gen. Pl. 763).
Nuttall remarks in his description: “Root, after flowering, resembling a bulb,” from which, doubtless, Rafinesque derives Bulbilis. Upon the margin of the copy of the American Monthly Magazine above quoted (in the library of the Missouri Botanical Garden), someone has suggested another derivation, “bull's bile!” — A. S. Hitchcock, Kansas Agricultural College, Manhattan.

Literature Cited:
- Lamson-Scribner, F., 1900.  

Uses Rafinesque's name of Bulbilis dactyloides.

Literature Cited:
- Schaffner, John H., 1920.  

Schaffner (1920) published an article on the dioecious nature of Buffalo Grass. He combined field observations and greenhouse experiments to support his contention that Buffalo Grass is strictly dioecious.

Literature Cited:
- Hitchcock, A. S., 1927.  

Hitchcock (1927) proposed conservation of Buchloë against Bulbilis and others because Engelmann's Buchloë was the first name under which both staminate an pistillate plants were described.
Buchloe Engelm. (1859) is conserved against Bulbilis Raf. (1819), Calanthera “Nutt.&rdquo: (1856), and Casiostega Rupr. (1857). Bulbilis was proposed by Rafinesque in a review of Nuttall's Genera as follows: “Sesleria dactyloides must for a peculiar genus by Mr. N's own account, it may be called Bulbilis.” Calanthera was mentioned by Hooker in a list of Geyer's plants from the Upper Missouri. “Calanthera dactyloides Kth.–Nutt. Sesleria.” Casiostega (of the Nomina Conservanda, a lisprint for Lasiostega Rpur.; Benth, Pl. Hartw. 347. 1857) is a nomen nudum. Nuttall's description of Sesleria dactyloides was based on the staminate plant only and until Engelmann gave his full description of both sexes of the buffalo grass there had been no mention of pistillate spikelets. Though in Bulbilis and Calanthera the stype species is indicated and a previously published description is referred to, that description is so inadequate, compared to the detailed study published by Engelmann, that they may well be rejected in favor of Buchloe.

Other articles:
• Golden Checklist Flora:  Buchloë dactyloides, Hitchcock, 1971, 2nd ed.;  

See Hitchcock, 1971, below.

Literature Cited:
- Anderson, Kling, and A. E. Aldous, 1937.  

An article on monoecious Buffalo Grass by Anderson & Aldous (1937) is behind a paywall I have been unable to penetrate.

Literature Cited:
- Gernert, W. B., 1937.  

An article by Gernert (1937) appears to address the height of pistillate spikes relative to harvesting them, but is behind a paywall I have not penetrated.

Literature Cited:
- Hensel, R. L., 1938.  

Hensel (1938) wrote about perfect-flowered buffalo grass which might be interesting to read were it not, alas, behind a paywall.

Literature Cited:
- Spurlock, Clay, 1940.  

Spurlock (1940) surveys monoecious and dioecious grasses in America, finding 17 genera and 54 species of monoecious, and 13 genera and 23 species of dioecious grasses of the Western Hemisphere, of which Buffalo Grass is one of the latter. Note is made of the controversy about the degree of dioecism seen in Buffalo Grass.

Literature Cited:
- Burr, Richard D., 1951.  

Burr (1951) wrote about his observations in variations of sex along stolons of

Literature Cited:
- Rickett, H. W., and F. A. Stafleu, 1959.  

In a review of conserved names, Rickett and Stafleu (1959) note that conservation of the name “Buchloë” was superfluous.
† 308. Buchloë Engelmann, Trans. Acad. Sci. St. Louis 1 432. 1859 quadrim. 1. T.: B. dactyloides (Nuttall) Engelmann (Sesleria dactyloides) Nuttall).  
Note: Conservation superfluous: Calanthera Nuttall ex W. J. Hooker, Journ, Bot. Kew Misc. 8: 18 (1856), Bulbilis Rafinesque, Am. Mon. Mag. Crit. Rev. 4: 190. (1819), and Lasiostegia Ruprecht ex Betham, Pl. Hartw. 347 (1857), are all nomina nuda.

The meaning of the dagger (†) is unclear. All names with daggers were subjects of a superfluous conservation. However, not all with superfluous conservation were marked with a dagger.

Literature Cited:
- Hitchcock, A. S., 1971.

Other articles:
• Golden Checklist Flora:  Buchloë dactyloides, Hitchcock, 1935;  

115. BÚCHLOË Engelm. (Bulbilis Raf.)
Plants dioecious or monecious. … Type species, Buchloë dactyloides. Name contracted from Greek boubalos, buffalo, and chloë, grass, a Greek rendering of the common name, “buffalo grass.”
1. Buchloë dactyloides (Nutt.) Engelm. Buffalo Grass. … The sod houses of the early settlers were made mostly from the sod of this grass. In 1941 it was planted at Boyce Thompson Institute, Yonkers, N. Y., and is proving to be an excellent cover for exposed dry banks.

Literature Cited:
- Bai, T. J., 1990.  

Abstract : Seventy-five stands with different compositions of blue grama (B. gracilis) and buffalograss (B. dactyloides) were sampled in 1987. Multiple regression and principal component analysis techniques were used to investigate the relationship between blue grama and buffalograss composition and soil characteristics. Blue grama was often more abundant on sandy soil, while buffalograss was found on clay soil. Blue grama was also found on clay soils when lime content was high. The essential factor underlying sand, clay and lime content of soil was interpreted as water stress. Relative crowding coeff. calculated for blue grama and buffalograss showing that both species had greater height and produced more vegetation when found intermingled than when growing in monoculture.

Literature Cited:
- Quinn, James A., 1991.  

Quinn (1991) examined multiple hypotheses for dioecy in Buffalo Grass and found support only for &lqquo;out-crossing” as an advantage afforded by dioecy.
Abstract. Buchloe dactyloides is a perennial dioecious grass in which male and female inflorescences are so strikingly dimorphic that they were originally assigned to different genera. The objective of this paper is to present the results of tests for sex-specific vegetative characters, ecological differences, and sexual niche-partitioning, combining them with prior information on the reproductive biology of Buchloe for an evaluation of the key factors leading to the evolution of dioecy and sexual dimorphism. Field and greenhouse data were collected from Oklahoma and Kansas populations on vegetative characters, allocation to reproduction, and relative growth and competitive success along resource gradients. Except for greater susceptibility to leaf rust by males, there were no significant differences between males and females in vegetative characters, total biomass, or reproductive effort. Field studies of spatial distributions of males and females failed to show any relation to soil, topography, or soil moisture. In a 45-month greenhouse experiment starting at the seedling stage, the relative growth and competitive success of randomly paired individuals showed no evidence for differential competitive success or for niche-partitioning of males and females. The "outcrossing advantage" and subsequent sexual specialization of the female inflorescence appear to be the major factors underlying this dimorphic system.

Literature Cited:
- Huff, David R., and Lin Wu, 1992.  

Abstract. Variations of sex inconstancy were examined for vegetative and seed samples from eight natural populations of buffalograss located along two east-west transects crossing the shortgrass prairies of Oklahoma, New Mexico, and Texas. Each of the eight populations was found to contain inconstant (monoecious) sex forms. Sex form distributions ranged from the Guymon vegetative sample, having no inconstant sex forms, to the Chillicothe seed sample in which the frequency of inconstant sex forms was nearly 70%. Frequencies of inconstant sex forms were generally higher for seed samples than for vegetative samples. Male to female sex ratio of constant (dioecious) sex forms generally did not differ from 1:1 expectations. Inconstant sex forms were more common among peripheral populations where buffalograss vegetation coverage was sparse than for more central populations having a higher concentration of buffalograss vegetation. Quantitative measures of sex inconstancy from artificial crosses were significantly (P < 0.001) correlated with the additive linear model of general combining ability, suggesting that sex determination in buffalograss has high heritability. The possible selection forces affecting the frequency of monoecious sex forms among natural populations are discussed.

Literature Cited:
- Columbus, J. Travis, 1999.  

An expanded circumscription of Bouteloua (Graminae: Chloridoideae): New Combinations and Names

Literature Cited:
- Columbus, J. Travis, Michael S. Kinney, Maria Elena Siqueiros Delgado, and J. Mark Porter, 2000.  

Phylogenetics of Bouteloua and Relatives (Granineae: Chloridoideae): Cladistic Parsimony Analysis of Internal Transcribed Spacer (nrDNA) and trnL-F (cpDNA) Sequences.

Literature Cited:
- Peterson, Paul M., and Konstatin Romaschenko, 2015.  

Phylogeny and subgeneric classification of Bouteloua with a new species, B. herrera-arrietae (Poaceae: Chloridoideae: Cynodonteae: Boutelouinae) … Buchloe is reduced to a section of Bouteloua.


Other articles:
• Golden Checklist Flora:  Hordeum pusillum;
Full Size ImageColl. No. 1453, Hordeum pusillum  

Hordeum pusillum Nutt. Little Barley.


Literature Cited:
- Nuttall, Thomas, 1818.  

Nuttall's Text My interpretation
119. HORDEUM. L. (Barley)  
Calyces lateral, 2-valved, mostly 1-flowered, aggregted by threes, so as to resemble a setaceous 6-leaved involucrum; the central flower sessile, the lateral ones stipitate, usually sterile. Corolla 2-valved, acute; exterior valve awned.  
Very nearly allied both by habit and character to the preceding genus. Flowers spiked, imbricated mostly in 2 rows; calycine involucrum setaceous, 6 leaved, division approaching by pairs. In the H. hexastichon, the flowers are imbricated in 6 ranks, because all the flowers are hermaphrodite; probably a mere effect of cultivation. [The preceding genus was ElymusEd.]
Species. 1. H. vulgare. Cultivated … 2. * pusillum Lateral masculine or nertral flowers awnless, acute; four internal calicina glumes, coriaceous and dilated, those of the hermaphrodite sublanceolate; internal valve of the lateral masculine flower, subsemi-ovate. [The description of H. vulgare is skipped. – Ed.]
Culm 4 to 6 inches, decumbent, or somewhat genoculate at the base. Leaves rather glaucous, a little pubescent on the under surface, striate, about one and a half inches long, and almost obtuse; uppermost sheath tumid and very smooth, embracing the spike. Spike linear; about one and a half inches long. Glumes by threes, distichally imbricated. Lateral imperfect flowers awnless, acute; central sessile flower awned, the awn almost exactly the length of that of the subtending calix; awns scabrous. Calix smooth, nerveless, exterior valve in the outer flowers setaceous from its base, the inner valves obliguely dilated, and rigidly coriaceous, all awned, the inner divisions of the lateral flowers, appearing nearly semi-ovate, the central ones sublanceolate. Corolla nerveless, the inner valve furnished with a short awn, arising from its base. Nearly allied, apparently, the the H. maritimum.  
On the arid and saline plains of the Missouri.  
3. jubatum. On the calcareous islands of Lake Huron and Michigan, also on the banks of the Missouri.  
The genus Hordeum exists chiefly in Europe, extending into Northern Africa, and Tartary in Asia. The 2 species above described are natives of North America, and the F. jubatum is also common to Smyrna. [One would assume that Nuttall was referring to Smyrna in what we now call Turkey. However, there are towns named Smyrna in Georgia, Tennessee, North Carolina, Delaware, Michigan, and Maine, and Nuttall may have been in or near some of those towns in his travels. – Ed.]

“Tall Fescue” -- Lolium arundinaceum (Schreb.) Darbysh.


Literature Cited:
- Gmelin, Johan Georg, 1747-1769.
Full Size ImageGmelin's (1747) description of Tall Fescue.  


Literature Cited:
- Gmelin, Johan Georg, 1747-1769.
- Scheuzer, Johann Jacob, 1719.
- Schreber, Johann Christian Daniel von, 1771.  

There seems to be general agreement (by extensive Google search) that what is called “Tall Fescue” is the grass known by its basionym of Festuca arundinacea Schreb.

Original TextInterpretation and Comments
1000. FESTVCA (arundinacea) panicula ſpicata ſtricta, ſpiculis oblongis erectis paucifloris, ariſtatis, calycibus anguſtatis. Gmel. Sibir. 1. p. 111. 1000. Festuca arundinacea panicle spike strict, spikes oblong erect awn, calyx angular. Gmelin, Flora Sibirica sive Historia Plantarum Siberiae. 1. p. 111.
Gramen arundinaceum, locuſtis viridi-ſpadiceis loliaceis brevius ariſtatis. Scheuchz. Agr. p. 266. t. 5. f. 18. Grass reed-like, … Scheuzer, Agrostographia sive Graminum, … p. 266. t. 5. f. 18.
In prato acclivi hinter dem Biniz, loco humido. In the steep field beyond the Biniz [East Germany], in a damp location.

  • “arundinacea” would mean cane-like from arundo (“cane”) +? -aceus.
  • “loliaceis” could mean chaffy, but Lolium itself is a name given by Virgil to a troublesome weed.
  There is less agreement as to its current accepted name, whether it should be Festuca arundinacea Schreb., Lolium arundinaceum (Schreb.) Darbysh., or Schedonorus arundinaceus (Schreb.) Dumort.

Literature Cited:
- Ackerfield, Jennifer, 2015.
- Baldwin, Bruce G., Douglas H. Goldman, David J. Keil, Robert Patterson, and Thomas J. Rosatti, 2012.
- Harrington, H. D., 1954.
- Shaw, Robert B., 2008.
- Weber, William A., and Ronald C. Wittmann, 2012.
- Wingate, Janet L., 1994.  

Starting close to home, most Colorado authors use Schedonorus arundinaceus (Schreb.) Dumort., including Ackerfield (2015), Weber & Wittman (2015), and Shaw (2008).

Wingate (1994) uses Festuca arundinaceus Schreb., although it is a older reference. Harrington (1964, 2nd ed.) used Festuca elatior var. arundinaceae (Screb.) Celak..

By way of comparison, the Jepson Manual of California (Baldwin, et al., 2012) retains Festuca arundinacea Schreb.

  On a regional level, the Southwest Biodiversity Network (SEINet) taxon tree has kind of a split personality with regard to “Tall Fescue.” The current (3 October 2019) taxon tree accepts both Lolium arundinaceum (Schreb.) S.J. Darbyshire and Schedonorus asundinaceus (Schreb.) Dumort.

               [Lolium x aschersoniana]
                 Lolium arundinaceum  
                       [Festuca arundinacea]
                       [Festuca elatior subsp. Arundinacea]
                       [Festuca elatior var. arundinacea]
                 Schedonorus arundinaceus 
                       [Avena secunda]
                       [Bromus arundinaceus]
                       [Bromus elatior]
                       [Festuca elatior f. aristata]
                       [Festuca elatior f. elatior]
                       [Festuca fenas]
                       [Festuca mediterranea]
                       [Festuca orientalis]
                       [Festuca phoenix]
                       [Festuca uechtritziana]
                       [Poa elatior]
                       [Poa kunthii]
                       [Poa phoenix]
                       [Poa uliginosa]
                       [Schedonorus elatior]
                       [Schedonorus phoenix]
                       [Tragus elatior]

One consequence is that both names have to be entered into a query when searching for collections of “tall fescue.”

Full Size ImageCollections of “tall fescue” entered into SEINet  
The map at left (3 Oct 2019) shows the locations of Colorado collections of “tall fescue” that have coordinates entered into SEINet. Within Jefferson County, collections have primarily been made at Chatfield Farms, Hildebrand Park and Deer Creek Canyon Park. There are no collections of this grass from Golden s.l.

Literature Cited:
- Henson, James F., 2001.  

USDA Plants accepts Schedonorus arundinaceus (Schreb.) Dumort. as a conserved name. Festuca arundinacea Schreb and Lolium arundinaceum (Schreb.) S.J. Darbyshire are treated as synonyms. Perhaps somewhat ironic is the USDA NRCS plant guide for “Tall Fescue” (Henson, 2001) which accepts the name of Lolium arundinaceum (Schreb.) S.J. Darbyshire.

Literature Cited:
- Soreng, Robert J., Edward E. Terrell, John Wiersma, & S. J. Darbyshore, 2001.  

Conservation of Schedonorus arundinaceus (Schreb.) Dumort. was proposed by Soreng, et al. (2001). S. arundinaceus Roem. & Schult. Syst. Veg. 2: 700. 1817 was rejected because it is not the basionym of any currently accepted taxon. Rather is it a synonym of Scolochloa festucacea (Willd.) Link.

Literature Cited:
- Fribourg, H. A., D. B. Hannaway, and C. P. West, 2009.  

Tall Fescue Online monograph. (

id="tblhistlitorg"> Hexaploid tall fescue [Lolium arundinaceum (Schreb.) Darbysh. = Schedonorus arundinaceus (Schreb.) Dumort., formerly Festuca arundinacea Schreb. Var. arundinacea] is an agronomically vital member of the grass family that has been characterized based on morphological characteristics, interfertility relationships, and, more recently, genetic criteria such as gene sequences. Efforts to improve tall fescue for forage or turf and to gain insight into processes of grass evolution hinge on an accurate depiction of the interrelationships that exist between this species and other Lolium and Festuca grasses. The evolution of hexaploid tall fescue, and a majority of grasses, has involved interspecific hybridization that obscures species boundaries but, more importantly, serves as a rapid means of combining distinct genomes into novel progeny with enhanced evolutionary potential. We discuss here the systematics of the Festuca-Lolium complex within the context of such processes and highlight the dynamic and often confounding evolutionary history that characterizes tall fescue and its relatives (Fribourg, et al., 2009).

Festuca arundinacea Schreb. Tall fescue (

Forage Identification: Tall fescue. Department of Plant Sciences. Tall fescue (Festuca arundinacea (Schreb.) Darbysh) (

Phylogeny of tall fescue and related species using RFLPs. Theor Appl Genet. 1994 Aug;88(6-7):685-90. doi: 10.1007/BF01253971. (

Melanie L Hand, Noel OI Cogan, Alan V Stewart & John W Forster Evolutionary history of tall fescue morphotypes inferred from molecular phylogenetics of the Lolium-Festuca species complex BMC Evolutionary Biologyvolume 10, Article number: 303 (2010) (


Other articles:
• Golden Checklist Flora:  Muhlenbergia montana;  

Muhlenbergia montana (Nutt.) Hitchc. Mountain Muhly.


Literature Cited:
- Nuttall, Thomas, 1848.  

Calycodon montanus Nutt., J. Acad. Nat. Sci. Philadelphia Ser. II, i. 186 (1847).

Nuttall's Text Comments and Interpretation
Spikelets one-flowered, the flower sessile, bearded at the base. Glumes two, unequal, shorter than the flower, membranaceous, the lower truncate, acutely three-toothed; the lower smaller, one-toothed. Paleae two, the lower sublanceolate, carinate, terminaing in a longish scabrous awn; at length indurated, with a silky pilose margin; the upper palea lanceolate, one-nerved, indurated and involute. Anthers three. Stigmas two, plumose. — A scabrous leaved grass, with a simple inarticulated culm, terminated by a loose, narrow, somewhat spiked panicle. So called in allusion to the remarkable toothing of the calyx.
C. *montanum. Leaves short and narrow, somewhat scabrous; ligules membranaceous, elongated; panicles four or five inches long, narrow, with the branches appressed; flowers clustered on the branches, three or four together, some nearly sessile and others pedicellate; glumes variable, membranaceous and eroded at the summit, the lower, three-nerved, with three wither short, or rather long and acute teeth, sometimes with a fourth membranous tooth; the upper glume also eroded, and ending in a single tooth from the nerve; the lower palea lanceolate, carinate, scabrous, and indurated, terminated by a long, slender, scabrous awn; the inner margin silky, with soft shining hair, of which there are two tufts at the base of the paleae; the inner paleae also indurated and herbaceous in the centre, involving the germ and the stamens.
A perennial grass, with a simple, unnjointed culm, about eighteen inches high. Somewhat allied to Muhlenbergia, (when restrained to its proper limits,) but perfectly distinct by its very remarkable glumes. The ripe seed we have not seen.
Hab. In the Rocky Mountains, near Santa Fe, Mexico. Flowering in August.

Literature Cited:
- Hitchcock, A. S., 1920.  



Other articles:
• Golden Checklist Flora:  Munroa squarrosa;  

Munroa squarrosa (Nutt.) Torr. False Buffalograss.


Literature Cited:
- Nuttall, Thomas, 1818.

Other articles:
• Glossary:  squarrose;  

75. CRYPSIS. Lamarck. (Thorn-Grass.)
Calix 2-valved, oblong, 1-flowered, Corolla 2-valved, longer than the calix. Stamina 2 or 3. (Spike surrounded at the base by the sheath of the leaf; or the flowers collected into a leafy capitulum.)
Culm decumbent or procumbent, extremely branched; leaves rigid and pungent; flowers collected in squarrose heads, or short and dense irregularly involucrate, lobed spikes.
SPECIES. 1. C. * squarrosa. Stem decumbent, much branched; leaves short, all rigid, and sharply pungent; capituli squarrose, few flowered; dorsal valve of the corolla coriaceous, somewhat cleft at the point, with a shortish subulate central cusp.
On arid plains near the “Grand Detour” of the Missouri, almost exclusively covering thousands of acres, and as pungent as thorns. ☉ Not more than 3 or 4 inches high; the flowers not collected into heads, as in the European species, but merely in squarrose terminal fascicles; the outer glume of the corolla is likewise cleft so as to present 3 short coriaceous subulate points.


Literature Cited:
- Ornduff, Robert, and Marion S. Cave, .  

Leucocrinum montanum Star Lily

Sand-lily, Leucocrinum montanum Nutt., is a showy perennial that occurs widely in arid regions of the western United States. Recently, the second author reported chromosome counts of n = 11, 13, and 14 for this species (Cave, 1970). Plants from the Rocky Mountain region have n = 14 (see also Löve et al., 1971), those from several localities in California, western Nevada, and Oregon have n = 13, and one population from Nevada has n = 11. In addition, Cave noted that in some populations pollen is shed in tetrads and in others it is shed singly. In this paper we further discuss the variation in chromosome number and in the condition of pollen at the time of shedding (Ornduff and Cave, 1975).


Studies of Prairie or Foothill Ecosystems



Restoration and Effects of Attempts Thereof


Literature Cited:
- Wilson, Scott D., 1989.  

Wilson, Scott D., 1989. The suppression of native prairie by alien species introduced for revegetation. Landscape and Urban Planning. 17(2), april 1989, pp. 113-119.


Natural prairie has become uncommon in North America, making the revegetation of disturbed areas by native species a desirable goal. Alien species are often introduced for revegetation because of their abilities to stabilize and nitrify soil. The objective of this study was to test whether these attributes of introduced species would allow them to promote the recovery of native vegetation. Seven treatments (six commercially-available mixtures of introduced species and an unseeded control) were applied to a randomized field experiment in disturbed mixed-grass prairie in south-west Manitoba, Canada. Sampling eight years later revealed that introduced species suppressed native vegetation. Introduced species did not aid revegetation: plots seeded with introduced species did not produce significantly higher standing crop or below-ground biomass than did unseeded plots. Unseeded plots had the lowest frequency of bare ground. Allowing prairie to revegetate without sowing introduced species produced both the highest cover of bare ground and the greatest abundance of native species.

Literature Cited:
- Wilson, Scott D., and Joyce W. Belcher, 1989.  

Wilson, Scott D., and Joyce W. Belcher, 1989. Plant and Bird Communities of Native Prairie and Intoduced Eurasian Vegetation in Manitoba, Canada. Conservation Biology. 3(1), pp. 39-44, March 1989.

“… introduction of Eurasian plant species to North American prairie not only replaces the native plant community, but also produces significant changes in the species composition of a higher trophic level … ”

Abstract: Large areas of North American prairie are dominated by Eurasian plant species introduced either for range improvement or accidentally as weeds. We examined the impact of introduced plants on both native vegetation and bird communities in a mosaic of North American mixed-grass prairie and Eurasian vegetation. We established ten transects five in areas of native prairie and five in areas dominated by introduced plant species. Each transect comprised five sampling stations separated by 100 m. Vegetation was sampled in four 0.5m2 quadrats at each station. The cooers of eight of the ten most common plant species varied significantly (p < 0.05) between native and introduced vegetation. One common native plans Andropogon scoparius, was absent in introduced vegetation Singing birds were identified to species at each station on three occasions during the breeding season All bird species found were native to prairie. The total number of birds did not vary between vegetation types Two out of eight bird species, upland sandpiper and Sprague's pipit were signifcantly more abundant in native prairie than in introduced vegetation No bird species were significantly more common in introduced vegetation. A correlation matrix calculated for all bird species and the ten most abundant plant species divided the bird community into two groups. The first group (western meadow lark, upland sandpiper, Sprague's pipit, Baird's sparrow and savannah sparrow) was positively correlated with native plant species and negatively with introduced plants, while the second (vesper sparrow, clay-colored sparrow, and grasshopper sparrow) was negatively correlated with native species and positively correlated with introduced Discriminant analysis separated transects from native and Eurasian vegetation on the basis of their respective bird communities. The results illustrate that the introduction of Eurasian plant species to North American prairie not only replaces the native plant community, but also produces significant changes in the species composition of a higher trophic level.

Literature Cited:
- Heidinga, Lawrence, and Scott D. Wilson, 2002.  

Heidinga, Lawrence, and Scott D. Wilson, 2002. The Impact of an Invading Alien Grass (Agropyron cristatum) on Species Turnover in Native Prairie. Diversity and Distributions. 8(5), September 2002, pp 249-258.


Alien invasions typically reduce species richness of habitats, but few studies have examined their effects on species turnover, the difference in species composition between localities. Agropyron cristatum (L.) Gaertn. (crested wheat grass) has been planted on 6-10 million ha of North American prairie, and is invading native prairie. We studied the invasion of A. cristatum into native prairie by measuring species composition along a gradient from maximum to minimum A. cristatum abundance. As A. cristatum increased, the abundance of most common native species decreased, but one appeared to be unaffected (Bouteloua gracilis (H.B.K.) Lag.), and another (Poa sandbergii Vasey) increased. The effect of A. cristatum on species turnover was investigated by examining species-area curves for areas from 0.5 m2 to 8.0 m2. Species diversity was reduced by 35% at high A. cristatum abundances at all areas. A. cristatum reduced the intercept of the species-area curve, but not the slope, suggesting that A. cristatum affected species turnover proportionally in all areas and habitats. This unusual result may indicate a homogeneous environment where species are distributed randomly. A. cristatum produced almost twice as many seeds as all native grasses combined. The number of seeds collected of native grasses and A. cristatum was highly correlated with the number of seed heads immediately nearby, but not with transect position. This suggests most seeds were dispersed over distances less than 5 m. In sum, the invasion of native prairie by A. cristatum might be related to high rates of seed production, and has the effect of decreasing species turnover by reducing the intercept of the species-area curve.

Literature Cited:
- Salesman, Jessica Bolwahn, and Meredith Thomsen, 2011.  

Salesman, Jessica Bolwahn, and Meredith Thomsen, 2011. Smooth Brome (Bromus inermis) in Tallgrass Prairies: A Review of Control Methods and Future Research Directions. Ecological Restoration. 29(4), December 2011, pp. 374-381.

Abstract. Smooth brome (Bromus inermis) is a cool-season rhizomatous grass of Eurasian origin that has escaped intentional plantings and spread widely in natural areas. A large body of work exists regarding smooth brome's biology and response to conservation management strategies designed to reduce its competitive effects on native prairie species, particularly for the tallgrass prairies of North America. Here we summarize that literature to improve restoration practice. In tallgrass prairie, smooth brome benefits from the early start of its growing season and its rhizomatous growth form, making it a strong competitor against native warm-season grasses. Late-spring burns timed to target smooth brome when root reserves are at their lowest have shown promise as a control strategy. Uncertainty remains, however, about the relative efficacy of fire, herbicide, mowing, or grazing to accomplish late-spring defoliation, the effect of repeated treatments, and the potential benefits of treatment combinations. The responses of resident or seeded natives to brome control treatments and/or the resulting decreases in brome cover also remain largely unexamined. Research focused on the questions we highlight would reduce costs associated with the control of smooth brome and increase confidence in the outcomes of restoration efforts.

Literature Cited:
- Grant, Todd A., Bridgette Flanders-Wanner, Terry L. Shaffer, Robert K. Murphy, and Gregg A. Knutsen, 2009.  

Grant, Todd A., Bridgette Flanders-Wanner, Terry L. Shaffer, Robert K. Murphy, and Gregg A. Knutsen, 2009. An Emerging Crisis across Northern Prairie Refuges: Prevalence of Invasive Plants and a Plan for Adaptive Management. Ecological Restoration. 27(1), March 2009, pp. 58-65.


In the northern Great Plains, native prairies managed by the U.S. Fish and Wildlife Service (Service) can be pivotal in conservation of North America's biological diversity. From 2002 to 2006, we surveyed 7,338 belt transects to assess the general composition of mixed-grass and tallgrass prairie vegetation across five "complexes" (i.e., administrative groupings) of national wildlife refuges managed by the Service in North Dakota and South Dakota. Native grasses and forbs were common (mean frequency of occurrence 47%-54%) on two complexes but uncommon (4%-13%) on two others. Conversely, an introduced species of grass, smooth brome (Bromus inermis), accounted for 45% to 49% of vegetation on two complexes and another species, Kentucky bluegrass (Poa pratensis) accounted for 27% to 36% of the vegetation on three of the complexes. Our data confirm prior suspicions of widespread invasion by introduced species of plants on Service-owned tracts of native prairie, changes that likely stem in part from a common management history of little or no disturbance (e.g., defoliation by grazing or fire). However, variability in the degree and type of invasion among prairie tracts suggests that knowledge of underlying causes (e.g., edaphic or climatic factors, management histories) could help managers more effectively restore prairies. We describe an adaptive management approach to acquire such knowledge while progressing with restoration. More specifically, we propose to use data from inventories of plant communities on Service-owned prairies to design and implement, as experiments, optimal restoration strategies. We will then monitor these experiments and use the results to refine future strategies. This comprehensive, process-oriented approach should yield reliable and robust recommendations for restoration and maintenance of native prairies in the northern Great Plains.


Recreation in Foothill and Prairie Ecosystems












Vegetation Descriptions






Dates To Do Things


Vouchers to Examine



Letters: Wednesday, November 5, 2014.  

UTC10764: was determined Polemonium caeruleum whereas UTC19762, also bearing MEJ's #299 is determined Polemonium foliosissimum A. Gray. However, per Mary Barkworth, 11/25/2014, both are P. foliosissimum.



Mystery Locations


Locations: Gray Hill.  

Gray Hill

Harbouria trachypleura (A. Gray) J.M. Coult. & Rose. Whiskbroom Parsley. Mountain slope. Near Golden: Gray Hill. J. H. Ehlers 6848. 6/2/1938 ( RM184550 )


West Cliff

Quincula lobata (Torr.) Raf. (Syn: Physalis lobata Torr. ) Chinese Lantern. Golden, Road to West Cliff, Golden. Earl L. Johnston, with G. G. Hedgcock 813. 6/23/1917 ( RM101941 ).

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Date and time this article was prepared: 8/3/2020 9:01:59 AM