Table Of Content

'r 'Sr 'Sr FLORA OF OKLAHOMA KEYS AND DESCRIPTIONS 'Sr 2 Edition °d w 'Sr r ..r rr 'Sr `r v r.► r.r Sr Sr Sr Sr Sr 'Sr Sr Sr Sr Sr Sr Sr Sr ..r Sr Sr Sr Sr 11~. I IN~HEsjlI~j' j'~'~!'~1M~~i~il~~~~2{11~~►~i{1~~~~13{►~~~~~1~►1j~1~41' ~j~{'~'~1~5{~jl~~`l_{TjI~~-j6~ T ' ' 5 {I11 11lIi~1111~Illl~ll~iil!►if{Illf{lll~li[111i~I~Til~j11~11 TIl~lili~il~f~1118 1~?itii~Tliltli~1111111(rIii l 1fi~ll~llii V V r ~r FLORA OF OKLAHOMA KEYS AND DESCRIPTIONS V 2"d Edition v V V w by V Adam K. Ryburn, Coordinator v Susan C. Barber V Paul Buckt V V Gloria M. Caddell V Wayne J. Elisens V James R. Estes V Mark Fishbein b Patricia Folley' V Lawrence K. Magratht V Abigail J. Moore Constance L. Murray V Bruce A. Smith V Constance E.S. Taylor V Ronald J. Tyrl V Rahmona A. Thompson Id Jay B. Walker' r r Linda E. Watson * Emeritus/a tDeceased r v y with illustrations by Bellamy Parks Jansen S Cover illustration of Oklahoma Penstemon (Penstemon okiahomensis) by Katy Levings r r - © August 2018 by Flora Oklahoma Incorporated all rights reserved 3 Oklahoma City, Oklahoma r r S a a A A a A a a a a a a A A A A A A A A A A PREFACE The flora and vegetation of Oklahoma are strongly influenced by the state's location in the southern plains of the United States. The geology of this region is dominated by extensive beds of Paleozoic sedimentary rocks, mostly overlain by deep alluvial and aeolian deposits and tilted along an east-west axis. The interior location creates a strongly seasonal temperate climate influenced by the Gulf of Mexico to the southeast and the rain shadow of the Rockies to the west. Together these factors create diverse vegetation that includes temperate deciduous forests, grasslands, and semi-arid woodlands and shrublands. Climate—Oklahoma is located at the southern edge of the temperate, continental climate zone, bordering on the sub- tropical zone. Seasonal and daily temperature fluctuations are great, with long, hot summers and short mild winters over most of the state, with occasional, brief periods of extreme cold. The annual mean temperature across the state is 15.5'C (60'F). The average growing season ranges from only 171 days in the far northwest to 215 days in the southeast. Because of the combined effects of latitude and elevation (the state is tilted with the highest elevations at the western end of the Panhandle and the lowest in valleys along the eastern border), there is a temperature gradient with highest mean temperatures in the southeast and lowest in the northwest. The most conspicuous and dramatic aspect of the climate is the strong precipitation gradient along the state's long east-west axis, with the greatest precipitation found in the southeast. The gradient is driven by the combined effects of the proximity of the southeastern comer of the state to the Gulf of Mexico, the source of Gulf Stream moisture, and the gradually diminishing effect of the rain shadow of the Rocky Mountains from west to east. Mean annual rainfall ranges from 43 cm (17 in) in the western panhandle to 142 cm (56 in) in the southeast. However, annual variation in precipitation is dramatic, characterized by multi-year cycles of extreme drought and extreme rainfall. Snowfall is rare across most of the state, except for the Panhandle. Geology—Much of Oklahoma consists of rolling plains with relatively few mountainous areas. The state is situated largely in the southern Great Plains, but includes the western edge of the Ozark Plateau, the northwestern edge of the Gulf Coastal Plain, and an extreme eastern outpost of the foothills of the Rocky Mountains. There are numerous rivers, but few with large and continuous flows. The sources of the Arkansas, Cimarron, and Canadian Rivers are in the Rocky Mountains or their foothills, whereas the Washita and Red Rivers originate on the high plains. The largest of these is the Arkansas River, which enters the eastern part of the state from Kansas and exits into Arkansas between the Ozark and Ouachita Mountains. There are no large, natural lakes, but numerous reservoirs. Elevation ranges from a low of 88 m (289 ft), where the state line crosses the Little River in the southeast, to 1516 m (4974 ft) atop Black Mesa near the state line, at the western end ofthe Panhandle. The major mountain ranges are the Ozarks and Ouachitas, which extend into the eastern part of the state from Arkansas and Missouri, and the Arbuckles and Wichitas, which were uplifted as major fault blocks and are contained entirely in the south-central and southwestern parts of the state, respectively. By far the most common rock type in Oklahoma is sedimentary, with relatively limited occurrences of volcanic and metamorphic rocks. The rolling plains that make up most of the western two-thirds of the state are underlain by Permian sedimentary rocks, most commonly red shales and sandstones. Gypsum beds are common, capping hills and mesas where the bedrock is exposed. Limestone is less common. Much of this area is overlain by Neogene and Quaternary sedimentary and aeolian deposits along rivers. Sedimentary rocks are also the primary constituent of the uplifted Ozark and Ouachita Mountains of the eastern one-fourth of the state. However, these are older, Carboniferous-aged limestones, sandstones, and shales. Together, the Permian and Carboniferous sedimentary rocks and their overlying sediments cover the great majority of the land. A sliver of the Gulf Coastal Plain is present in southeastern Oklahoma north of the Red River, covered by Cretaceous sediments. Two isolated mountain ranges contain older formations. The Arbuckle Mountains consist largely of Silurian and Devonian limestones and the Wichita Mountains are composed of even older Cambrian granite, the largest outcropping of ancient igneous rocks in the state. Extrusive igneous rocks are uncommon, with the only large exposure forming the Neogene-aged Black Mesa, which just enters the Oklahoma Panhandle from New Mexico. Vegetation—Precipitation, geology, and fire are the strongest influences on the vegetation of Oklahoma. This is reflected in the classification systems of Oklahoma's ecoregions and vegetation types. The US EPA recognizes 15 major ecoregions in North America, and two of these are present in Oklahoma. The northwestern three-fourths of the state are included in the Great Plains ecoregion and the southeastern one-fourth in the Eastern Temperate Forest. a a a a Similarly, the most common natural vegetation types in the state are grasslands, followed by forests and woodlands. Much of the Great Plains ecoregion is covered by mixed-grass prairies, with smaller areas of short-grass prairie in the I `~a Panhandle and tall-grass prairie in northeastern portion of the region. However, the Great Plains ecoregion also includes the broad irregular transition to forest across much of the central portion of the state. This transition zone consists of a patchwork of mixed-grass and tall-grass prairie vegetation with a short-statured, species-poor oak-hickory a woodland widely known as the Cross Timbers. The Great Plains also consists of conifer woodlands on the slopes of Black Mesa and on surrounding hills that are outposts of the pinyon-juniper woodlands of the foothills of the Rocky Mountains. Other restricted vegetation types in the Great Plains ecoregion include sage-dominated and oak-dominated shrublands on sandy soils, mesquite-dominated woodlands, and halophytic vegetation in the Salt Plains and on playas. The Cross Timbers gradually grade into the Eastern Temperate Forest ecoregion along an amorphous line running a roughly from Durant to Miami. Continuous extents of diverse forest occur primarily in the Ozark and Ouachita Mountains. These are largely oak-pine-hickory forests with short-leaf pine the only naturally occurring pine throughout much of the region. However, loblolly pine occurs naturally in southern slopes of the Ouachita Mountains, has been extensively planted by the forest industry, and now occurs across the entire region. Another common conifer species in these forests is eastern red-cedar. Eastern red-cedar becomes increasingly conspicuous in the Cross S Timbers, where its abundance has dramatically increased in response to fire suppression. The increase in red-cedar highlights the important role of fire in controlling the ecotone between forests and grasslands, especially in the Cross Timbers transition that runs through a broad swath of Oklahoma. In the absence of fire, grasslands become invaded A by woody species (mostly natives in our state) and over time will become forests, pushing the ecotone to the west and filling in the prairie openings of the Cross Timbers. Flora—The documented flora of Oklahoma consists of 3,737 native and naturalized taxa, including subspecies and varieties. The most species-rich families in the flora are the same as those of most other floras from the temperate region of the Northern Hemisphere. These are Asteraceae, Poaceae, Fabaceae, Cyperaceae, Rosaceae, Brassicaceae, and Lamiaceae. The largest genera include Carex, Cyperus, and Eleocharis (Cyperaceae), Oenothera (Onagraceae), A Era grostis (Poaceae), Solidago and Symphyotrichum (Asteraceae), Quercus (Fagaceae), Juncus (Juncaceae), 1~ Euphorbia (Euphorbiaceae), and Asclepias (Apocynaceae). The only federally listed Threatened or Endangered plant species in Oklahoma are Great Plains white fringed orchid (Platanthera praeclara) and piedmont mock bishopweed (Harperella nodosa, or Ptilimnium nodosum by some authors). There are only a few species, subspecies, or varieties that have been considered endemic to the state, and these have been questioned taxonomically. Knowledge of the flora is uneven across the state. Several counties and numerous state parks, conservation areas, and wildlife management areas have been surveyed for plant occurrences. The areas of the state that have received the least attention from plant collectors are the eastern Panhandle and adjacent counties, and other counties of western a Oklahoma, particularly the southwestern counties. Knowledge of the flora of Oklahoma improves daily with new state records of native and, more commonly, non-native species occurring several times a year. a Studies of Oklahoma's Flora—As might be expected, this diverse flora has long been of interest to taxonomists and our knowledge of it is the result of contributions by numerous explorers, surveyors, botanists, taxonomists, and academicians for more than 450 years (see Tyrl and Shryock 2013). Some individuals merely recorded their observations of the flora as they passed through, whereas, others collected plants, made systematic surveys of distinctive ecogeographic areas, or prepared treatments of different plant groups such as grasses and ferns. We traditionally cite the doctoral dissertation submitted to Harvard University by G.W. Stevens in 1916 as the first formal floristic study of the state's plants. Never formally published, his treatment included keys and generic descriptions, but lacked descriptions for the more than 1,600 listed species. In 1929, T.R. Stemen and W.S. Myers published a Spring Flora of Oklahoma with Keys which was followed by their more comprehensive Oklahoma Flora in 1937. In the latter book, they treated 147 families, 640 genera, and 1,626 species; however, they did not include the graminoid families Poaceae, Cyperaceae, and Juncaceae. Illustrations accompanied abbreviated descriptions of families, genera, and species. Taxonomic treatments of portions of Oklahoma's flora were prepared by G.J. Goodman a and D.A. McCoy. In 1958, Goodman published The Spring Flora Of Central Oklahoma. As the title indicates, the treatment was restricted both for area and season. A Study of Flowering Plants was published by McCoy in 1968. It is a treatment providing descriptions of the major families. In 1952, U.T. Waterfall undertook the writing of a w comprehensive treatment for the flora of Oklahoma. Recognizing that there was an immediate need for a means of identification, he first prepared keys to all of the state's plant species and personally published four editions of Keys to the Flora of Oklahoma (later editions are only reprintings of the fourth edition). Simultaneously with publication 11 S v r of the keys, he began to write diagnostic descriptions of each taxon. Sadly, this manuscript was not completed because of his death in 1971. S v Flora of Oklahoma Project—The last edition of Waterfall's Keys (1969) was the primary taxonomic reference for studies of the state's flora for more than two decades. During this time, numerous taxonomic revisions of families and genera were published, and many nomenclatural changes were made. In addition, additional native taxa were y discovered, and numerous introduced taxa have become naturalized. In the fall of 1983, botanists from throughout the state initiated an effort to write and publish a modem floristic treatment for Oklahoma's vascular plants. Initial steps v included the establishment of an editorial board, formation of a non-profit corporation, development of an editorial r format, production of a computerized data base for families, solicitation of contributors, and initiation of research and writing. In order to provide students and other individuals keys and descriptions as quickly as possible, the editorial y board decided to sequentially produce a key to families, descriptions of families, keys to the genera of each family, keys to their species, and finally genus and species descriptions. The key to families, titled Keys to the Vascular Plants of Oklahoma, appeared in 1994. Completion of the family descriptions resulted in 1997's Keys and 5 Descriptions for the Vascular Plant Families of Oklahoma. Concurrent with the publication of this second book, work was begun on keys to the genera and species of all families. As updates were completed and added to the book, V iterations were published, the last appearing in 2010. S y Flora of Oklahoma Keys & Descriptions—In the 21 years since the publication of our first iteration of the Keys, additional changes in the classification of families, genera, and species. These changes reflect the accumulation of S additional taxonomic data and new interpretations of characters in terms of phylogenetics. We have adopted, the r classifications of the Angiosperm Phylogeny Group (APGIII; www.mobot.org/MOBOT/research/APweb) (Stevens 2001 onwards) and Integrated Taxonomic Information System (ITIS; http://www.itis.gov). Our specific taxonomic 5 decisions as to family, genus, and species concepts are based on our field work, examination of Oklahoma herbarium v specimens, and review of the literature. Author citations appearing in the index are taken from ITIS, Brummitt and Powell (1992), and the International Plant Names Index (IPIN). S v `J USE OF KEYS AND DESCRIPTIONS Use—As noted above, the editorial board envisions that an individual will use this book in conjunction with other v taxonomic works to identify unknown plant species in the state. The following steps are recommended. 5 1. Examine the vegetative and floral characters of the plant at-hand. V 2. Key the unknown plant using the "Key to Groups." V V 3. Compare the characters of the plant with the brief diagnosis of the identified group. 4. Key the plant to family using the key to families of that group. 5. Compare the characters of the plant with the family descriptions that follow the "Key to Groups." we 6. Use the generic key that follows the family description to identify the genus of the unknown plant. y W 7. Compare the characters of the plant with the genus descriptions appearing in available taxonomic treatments such as Shinners & Mahler's Illustrated Flora of North Central Texas, Manual of the Vascular Plants of Texas, Flora of the Great Plains, or Gray's Manual of Botany. S y 8. Use the key appearing under the genus name to identify the species of the unknown plant. S 9. Compare the characters of the plant with the species descriptions appearing in the available taxonomic y treatments. 5 10. Use the glossary that appears at the end of this book to determine the meanings of unfamiliar terms. S iii r S a a a Punctuation—To facilitate use of the keys and descriptions, characters are separated from one another by periods and their first letter capitalized. Their different states are separated by semicolons. Commas are used as appropriate w for clarity. In addition, major characters are bold-faced in the descriptions for emphasis and ease of comparison. a The authors welcome any comments regarding the utility and accuracy of these keys, glossary, and index. Please inform us if you encounter problems in keying taxa, troublesome or poorly worded couplets and leads, omitted taxa, and characters that are inappropriate or inaccurate. Please send all comments and suggestions to Adam K. Rybum, Coordinating Editor, Department of Biology, Oklahoma City University, Oklahoma City, Oklahoma 73106 a ([email protected]). Information about this project is available at www.biosurvey.ou.edu/floraok. References Brummitt, R.K. and C.E. Powell (editors). 1992. Authors of Plant Names. Royal Botanic Gardens, Kew, England. Commission for Environmental Cooperation. 1997. Ecological Regions of North America. Toward a Common Perspective. Commission for Environmental Cooperation, Montreal, Quebec, Canada. Goodman, G.J. 1958. Spring Flora of Central Oklahoma. University of Oklahoma Press, Norman, OK. a Hoagland, B. 2000. The Vegetation of Oklahoma: A Classification for Landscape Mapping and Conservation a Planning. Southwestern Naturalist 45:385-420. Hoagland, B. W., A. K. Buthod, and T. D. Fagin. 2004-Present. Oklahoma Vascular Plants Database. (http://www.oklahomaplantdatabase.org/). Oklahoma Biological Survey, University of Oklahoma, Norman, Oklahoma, USA. Johnson, K. S., and K. V. Luza. 2008. Earth Sciences and Mineral Resources of Oklahoma. Educational Publication a 9. Oklahoma Geological Survey, University of Oklahoma, Norman, Oklahoma, USA. McCoy, D.A. 1968. A Study of Flowering Plants. Cameron University. Lawton, OK. Oklahoma Climatological Survey. 2018. Climate of Oklahoma. Website accessed 23 June 2018 (http://climate.ok.gov/index.php/site/page/climate_of_okahoma). Stemen, T.R. and W.S. Myers. 1929. Spring Flora of Oklahoma with Key. Harlow Publishing, Oklahoma City,OK. Stemen, T.R. and W.S. Myers. 1937. Oklahoma Flora. Harlow Publishing, Oklahoma City, OK. w Stevens, P.F. 2001 onwards. Angiosperm Phylogeny Website. Version 12, July 2012 [and more or less continuously updated since]. http://www.mobot.org/MOBOT/research/APweb/. fa The International Plant Names Index (2014). Published on the Internet http://www.ipni.org [accessed 2014]. Tyrl, RJ. and P. Shryock. 2013. A cavalcade offield botanists in Oklahoma--Contributors to our knowledge of the a flora of Oklahoma. Oklahoma Native Plant Record 13:55-100. Tyrl, R.J., S.C. Barber, P. Buck, W. Elisens, J.R. Estes, P. Folley, L.K. Magrath, C.E.S. Taylor, and R.A. Thompson. 2004. Keys and Descriptions for the Vascular Plants of Oklahoma. Flora Oklahoma Inc, Noble, OK Waterfall, U. T. 1969. Keys to the Flora of Oklahoma. Published by the author. Stillwater, OK. Woods, A. J., J. M. Omernik, D. R. Butler, J. G. Ford, J. E. Henley, B. W. Hoagland, D. S. Arndt, and, B. C. Moran. 2005, Ecoregions of Oklahoma (color poster with map, descriptive text, summary tables, and a photographs). U.S. Geological Survey, Reston, Virginia, USA (map scale 1:1,250,000). a S S fS a a A iv KEY TO GROUPS (page) I. Plants trees or shrubs or woody vines or arboreal subshrubs. 2. Plants woody vines or arboreal subshrubs. GROUP A (2) 2. Plants trees or shrubs. 3. Stems succulent; bearing spines in clusters. CACTACEAE (114) 3. Stems not succulent; not bearing spines in clusters. 4. Flowers or cones appearing before leaves. GROUP B (3) 4. Flowers or cones appearing with or after leaves. 5. Leaves opposite or whorled or fascicled or in rosettes. GROUP C (4) 5. Leaves alternate. 6. Leaves compound. GROUP D (6) 6. Leaves simple. 7. Leaf margins entire. GROUP E (7) 7. Leaf margins lobed or toothed. GROUP F (10) 1. Plants herbs, some woody at the base. 8. Plants aquatic floating on or submerged in or emergent from water. GROUP G (12) 8. Plants terrestrial or epiphytic. 9. Plants herbaceous epiphytes or herbaceous vines. GROUP H (18) 9. Plants not herbaceous epiphytes nor herbaceous vines. 10. Chlorophyll absent. Plants parasitic or saprophytic or mycotrophic. GROUP I (19) 10. Chlorophyll present. Plants autophytic. 11. Plants not producing flowers and seeds. Spores produced in strobili or son i or sporocarps or in aggregations of sporangia at ends of elongated stalks. GROUP J (19) 11. Plants producing flowers and seeds. Spores produced in anthers or ovaries. 12. Stems bearing spines and/or glochids in areoles; succulent. Foliage leaves absent. Ovaries inferior. Perianth parts 25 or more. Stamens 25 or more. CACTACEAE (114) 12. Stems not bearing spines or glochids in areoles; succulent or not succulent. Foliage leaves present or absent. Ovaries superior or inferior. Perianth parts of various numbers. Stamens of various numbers. 13. Plants acaulescent, aerial stems not apparent and leaves not cauline. GROUP K (22) 13. Plants caulescent, aerial stems apparent and leaves cauline. 14. Perianth parts absent. GROUP L (25) 14. Perianth parts present. 15. Perianth parts in I-series or parts all similar. 16. Perianth parts 3 or multiples of 3. GROUP M (26) 16. Perianth parts 1 or 2 or 4 or 5 or multiples of 4 or 5 or many. GROUP N (28) 15. Perianth parts in 2-series. couplet 17 17. Petals 3 or multiples of 3. GROUP O (31) 17. Petals I or 2 or 4 or 5 or multiples of 4 or 5 or many. 18. Corollas bilaterally symmetrical. 19. Petals free. GROUP P (32) 19. Petals fused at least at base or apex. 20. Ovaries inferior, wholly or partially. GROUP Q (33) 20. Ovaries superior. GROUP R (33) 18. Corollas radially symmetrical or asymmetrical. 21. Petals free. 22. Ovaries inferior, wholly or partially. GROUP S (35) 22. Ovaries superior. 23. Pistils or fruits 1 per flower. GROUP T (35) 23. Pistils or fruits 2 or more per flower. GROUP U (38) 21. Petals fused at least at base or apex. 24. Ovaries inferior, wholly or partially. GROUP V (38) 24. Ovaries superior. GROUP W (39) a a GROUP A: Plants woody vines or arboreal evergreen subshrubs. 1. Plants arboreal evergreen subshrubs. SANTALACEAE a L Plants woody vines. 2. Leaves opposite or whorled. 3. Plants climbing by tendrils or aerial rootlets, or prehensile petioles or petiolules. 4. Plants climbing by prehensile petioles or petiolules. Perianth in 1-series. Fruits achenes. RANUNCULACEAE is 4. Plants climbing by aerial rootlets or tendrils from leaf rachises (may be absent on scrambling-bushy forms). Perianth in 2-series. Fruits capsules. 5. Leaves compound. Petals fused; red-orange; 40-90 mm long. Capsules elongate. BIGNONIACEAE 5. Leaves simple. Petals free; green to cream; 2-5 mm long. Capsules globose. CELASTRACEAE A 3. Plants climbing by twining or scrambling stems. A 6. Leaf margins crenate or serrate. 7. Leaves compound. Fruits capsules. BIGNONIACEAE A 7. Leaves simple. Fruits berries. CAPRIFOLIACEAE A 6. Leaf margins entire. 8. Flowers radially symmetrical. Fruits follicles. Seeds comose. a Shoot exudate viscous; white. Axillary buds solitary. APOCYNACEAE 8. Flowers strongly to weakly bilaterally symmetrical. Fruits berries. a Seeds without hairs. Shoot exudate watery; clear. Axillary buds multiple. CAPRIFOLIACEAE 2. Leaves altemate. 9. Plants climbing by tendrils or aerial rootlets. 10. Venation parallel-convergent. Tendrils paved. Inflorescences umbels. Pith absent. SMILACACEAE a 10. Venation palmate or pinnate or pinnipalmate. Tendrils solitary or absent. Inflorescences panicles or cymes. Pith present. r`►„ 11. Leaves compound. 12. Leaves I-pinnately compound. (poison ivy) ANACARDIACEAE 12. Leaves palmately compound or 2- or 3-pinnately compound. 13. Leaves 2-or 3-pinnately compound; leaflets numerous. VITACEAE 13. Leaves palmately compound. 14. Leaflets always 3 per leaf. [Leaves 1-pinnately compound, but rachises occasionally short or almost absent; hence leaves falsely appearing palmately compound]. Inflorescences a panicles. Pistillates 3-carpellate. Fruits drupes; white. (poison ivy) ANACARDIACEAE 14. Leaflets typically 5 or rarely 3 due to abscission of 2 leaflets. Inflorescences cymes. Pistils 2-carpellate. a Fruits berries; dark blue to black. YITACEAE a I I. Leaves simple. 15. Leaves ovate or oblong-ovate; margins entire. Inflorescences racemose panicles. Flowers perfect. Calyces deeply 5-parted. Corollas absent. Fruits achenes; subtended by persistent sepals. POLYGONACEAE 15. Leaves cordate or rotund to broadly ovate; margins toothed. Inflorescences cymes. Flowers functionally imperfect. Calyces slightly to shallowly 4-lobed. Corollas present, may be caducous. Fruits berries. VITACEAE 9. Plants climbing by twining stems. a 16. Leaves compound. 17. Stems bearing prickles. Pistils numerous. Fruits aggregates of drupelets or achenes. ROSACEAE a 17. Stems not bearing prickles. Pistils I. Fruits berries or legumes. 18. Leaflets 3-7 cm long. Flowers radially symmetrical. Corollas ~ greenish; 1-3 mm long. Berries bluish purple or green. VITACEAE 18. Leaflets 10-30 cm long. Flowers bilaterally symmetrical. A Corollas purplish or lilac or blue; 15-20 mm long. Legumes reddish brown. FABACEAE 16. Leaves simple. 2