microsoft word onpsrecord8.doc oklahoma native plant record volume 8, number 1, december 2008 3 foreword this has been a very busy year for our authors, reviewers, and editors. thank you for waiting patiently for volume 8. i think you will agree that it was worth the wait. susan barber has provided our historic article for 2008. her thesis, “a floristic study of the vascular plants of the gypsum hills and redbed plains area of southwestern oklahoma”, is long overdue to be published . she researched the relationships between soil and vegetation types, just one of the underlying causes for the great biodiversity in oklahoma. her thorough work provides much more to the reader than the title reveals. it is a data-rich source for future botany research, and we know you’ll enjoy it. “updated flora of the wichita mountains wildlife refuge” by keith carter, pablo rodriguez, and michael dunn marks a new step in botanical research in oklahoma. the herbarium at cameron university [camu] is now housing the refuge’s plant specimens, thanks to a grant and a lot of work by students, faculty, and staff at cameron university. this is the first effort to update information regarding species at the refuge since we published the late paul buck’s 1977 checklist of the flora in 2002. hopefully, it will spur interest in keeping the refuge list up-to-date and bring recognition to a very deserving state institution’s herbarium. we also hope that this will mark the beginning of a cooperative relationship between the society and our state institutions’ herbaria. one of the main goals of the record is the initiation of new sources of data for biodiversity research in oklahoma, and this paper is evidence that we are reaching that goal. it’s been several years since we’ve published clark ovrebo’s popular paper about lawn mushrooms. “spring mushrooms of oklahoma” by ovrebo and nancy weber is a new, enlightening and enjoyable article with colorful photos from which we can learn a great deal more about the intriguing kingdom of fungi. we’ve also been waiting several years for “ferns and rare ferns in oklahoma” by bruce smith. it’s finally here with photos to help identify them. hopefully, a checklist of oklahoma ferns will be forthcoming. finally, we have a memorial to paul buck, long-time board member and promoter of the society. constance murray has provided us with a look at what it was like to have a professional, as well as a personal relationship, with someone so many of us have known and respected, someone who had a tremendous impact on the study of botany and ecology in oklahoma. sheila strawn, editor 2017 oklahoma native plant record oklahoma native plant record 3 volume 17, december 2017 foreword the search for historic articles that would be important to botanical research often leads us down surprising pathways to sources that are sometimes hidden in plain sight. lynn nabb mentioned her grandmother’s 1959 master’s thesis from the university of tulsa in a facebook post that honored her work and her life. until then, maxine clark’s “a study of the flowering plants of tulsa county, oklahoma” had been quietly sitting in the university library for almost 60 years. maxine was a student of dr. ralph kelting and a friend of dr. harriet barclay. we are grateful to the university’s library staff who helped us obtain the thesis. this year the record offers a number of important works from a wide variety of sources. it brings together several articles that have to do with species interactions and a couple of articles that offer readers the opportunity to know something “first.” we chose paul buck’s “allelopathy” from a previous issue of the gaillardia for our “critic’s choice essay,” because it explores deeply the “war in the garden” and the “vicious world in nature.” dr. buck’s 2004 article may help us better understand several issues discussed in “laboratory studies of allelopathic effects of juniperus virginiana on five species of native plants” by erica corbett and andrea lashley. this work involved undergraduate students in an important research opportunity and explores possible negative interactions between plant species. as for the firsts, urban species have historically been overlooked by botanists because their habitats had been altered by human activity. researchers have changed that perspective. urban studies are now valued because they address the effects that humans have had on species. urban parks can be surprisingly biodiverse. “vascular flora of e. c. hafer park, edmond, oklahoma” is from gloria caddell and students katie christoffel, carmen esqueda, and alonna smith at the university of central oklahoma. it is the first species list for edmond’s hafer park, which was established in 1979. for another first, clark ovrebo reports on an interesting earth star fungus that, until now, was known only in texas and japan! be on the lookout for it. evidently, it has a much wider distribution than previously thought. speaking of wide distribution… the oklahoma native plant record is getting more widely distributed every year. because it is listed in the directory of open access journals and abstracted by the centre for agricultural bioscience international, it can be accessed by researchers around the world. and here’s another “first.” the record has a new editor this year. gloria caddell is joining the editorial board as co-editor. it’s time for a change of leadership, and she has graciously agreed to begin taking over the helm and her new responsibilities. sheila strawn and gloria caddell co-editors oklahoma native plant society the purpose of the onps is to encourag e the study, protection, propag ation, appreciation and use of the native plants of oklahoma. m embership in onps shall be open to any person who supports the aims of the society. onps offers individual, student, family, and life membership. officers and board president: pat folley vice-president: chad cox secretary: maurita nations treasurer: mary korthase board members: berlin heck iris mcpherson sue amstutz jim elder paul reimer larry magrath northeast chapter chair: jim elder central chapter chair: judy jordan cross-timbers chapter chair: ron tyrl historian: lynn allen conservation chair: berlin heck publicity co-chairs: ruth boyd & betty culpepper marketing chair: larry magrath photo contest: paul reimer ann long award chair: paul reimer harriet barclay award chair: connie taylor onps service award chair: sue amstutz newsletter editor: chad cox librarian: bonnie winchester website manager: chad cox managing editor: sheila strawn technical editor: pat folley technical advisor: bruce hoagland cover: cercis canadensis (redbud) photo courtesy of charles lewallen. “that man is truly ethical who shatters no ice crystal as it sparkles in the sun, tears no leaf from a tree…” albert schweitzer articles (c) the authors journal compilation (c) oklahoma native plant society except where otherwise noted, this work is licensed under a creative commons attributionnoncommercial-sharealike4.0 international license, https://creativecommons.org/licenses/ by-nc-sa/4.0/, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly attributed, not used for commercial purposes, and, if transformed, the resulting work is redistributed under the same or similar license to this one. https://doi.org/10.22488/okstate.17.100001 oklahoma native plant record volume 1, number 1, december 2001 oklahoma native plant record volume 1, number 1 table of contents forward ...................................................................................................................... 2 ms. pat folley, onps president the spermatophyta of oklahoma county, oklahoma ....................................... 3 masters thesis of dr. u. t. waterfall floristic list for oklahoma county ..................................................................... 25 dr. bruce w. hoagland native orchids of oklahoma ............................................................................... 39 dr. lawrence k. magrath galium parisiense var. leiocarpum tausch, new for oklahoma ............................ 67 dr. lawrence k. magrath checklist of the ferns, natural falls state park ................................................. 68 dr. bruce a. smith onps critic’s choice essay : the limestone glade ............................................. 72 mr. jim norman journal of the oklahoma native plant society volume 12, december 2012 oklahoma native plant record 3 volume 12, december 2012 foreword this year’s oklahoma native plant record is all about learning from history. publishing dr. marilyn semtner’s 1972 master’s thesis this year offers us an opportunity to gain another perspective on why some introduced species become invasive in natural habitats where others do not. as the oklahoma invasive plant council, formed in 2008, seeks ways to guide our state agencies to determine best practices for preserving our native plant species, research like this, both old and new, can inform policy decisions. mr. randall ledford has collected extensive information regarding use of oklahoma’s native plant species by the pawnee native americans. he gives us a preliminary plant list that is sure to become part of an important resource that can be used in pawnee cultural education and by ethnobotanists. with full respect for the pawnee culture, his list includes scientific names as well as pawnee names and descriptions of uses that have been carefully researched and whose content has been approved by the pawnee elders. we are hoping to build interest and anticipation in this area of social and botanical research overlap. his goal is to collect and organize a larger body of this little known ethnobotany for wider dissemination dr. gloria caddell and ms. kristi rice have provided us with the long anticipated flora of alabaster caverns state park. it also compares flora in those gypsum outcrops with two other previous studies done ten years ago at selman living lab in woodward county and on a ranch in major county. teaching and inspiring botany students at mcloud high school and at the university of oklahoma biological station, dr. bruce smith has contributed several articles to the record in the past. this year he offers us a comparison study of two oak forests based on data collected by his students. engaging his students in plant distribution and ecological studies, he strives to fulfill our need to collect and preserve data for the future — a future history, to be used by future scientists. this year our “critics’ choice essay” is from dr. wayne elisens. he tells us how new software and digitization methods are bringing new light to historical collections, virtually. herbaria are making specimen data and images globally accessible. we will be able to see and learn from historical and current collections from all over the world. the oklahoma native plant record will keep passing on the science and building on what we know. we do not want to lose, or fail to learn, what future generations will need to know to keep oklahoma’s native plant species thriving in oklahoma. as our practice of publishing historical, unpublished work shows, we believe in the importance of historical studies and how they can inform current science policies and future research. moving into the future, all previous volumes of the record are now available on the internet at http://ojs.library.okstate.edu/osu/index.php/index, and it is listed on the directory of open access journals through http://www.doaj.org. sheila strawn managing editor http://ojs.library.okstate.edu/osu/index.php/index http://www.doaj.org/ journal of the oklahoma native plantsociety, volume 3, number 1, december 2003 oklahoma native plant record journal of the oklahoma native plant society 2435 south peoria tulsa, oklahoma 74114 volume 3, number 1, december 2003 issn 1536-7738 managing editor: sheila strawn technical editor: patricia folley technical advisor: bruce hoagland cd-rom producer: chadwick cox website: http://www.usao.edu/~onps/ the purpose of onps is to encourage the study, protection, propagation, appreciation and use of the native plants of oklahoma. membership in onps shall be open to any person who supports the aims of the society. onps offers individual, student, family, and life memberships. officers and board members president: james elder vice-president: constance murray secretaries: kimberly a. shannon tina julich treasurer: mary korthase past president: patricia folley board members: paul buck kay gafford melynda hickman lawrence magrath elfriede miller paul reimer northeast chapter chair: constance murray central chapter chair: susan chambers cross-timbers chapter chair: suzanne mcallister mycology chapter: clark ovrebo historian: carla and dale chlouber ann long award chair: paul buck harriet barclay award chair: constance taylor onps service award chair: sue amstutz conservation chair: chadwick cox publicity co-chairs: ruth boyd betty culpepper publications co-chairs: sheila strawn constance taylor marketing co-chairs: lawrence magrath susan chambers photo contest co-chairs: patricia and chadwick cox newsletter editor: chadwick cox librarian: bonnie winchester website manager: chadwick cox mailing committee chair: karen haworth color oklahoma committee chair: constance murray wildflower workshop chair: larry magrath cover: phoradendron serotinum, mistletoe in flower, by charles lewallen. articles (c) the authors journal compilation (c) oklahoma native plant society except where otherwise noted, this work is licensed under a creative commons attributionnoncommercial-sharealike4.0 international license, https://creativecommons.org/licenses/by-ncsa/ 4.0/, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly attributed, not used for commercial purposes, and, if transformed, the resulting work is redistributed under the same or similar license to this one. https://doi.org/10.22488/okstate.17.100016 oklahoma native plant record volume 3, number 1, december 2003 . 2 oklahoma native plant record volume 3 number 1 table of contents forward.........................................................................................3 mr. james f. elder, onps president black mesa flora study…………………………………………4 dr. james k. mcpherson black mesa state park flora update……………………………19 ms. patricia a. folley vascular flora of the keystone wildlife management area……23 dr. bruce w. hoagland and ms. amy k. buthod floristic survey of the nature conservancy’s preserve in johnston county, oklahoma …………….…..……….38 ms. kimberly a. shannon historical accounts of the transformation of a prairie town….51 mr. todd d. fagin and ms. melissa scott brown three bird orchid and crane-fly orchid in oklahoma………….68 dr. lawrence k. magrath take time to watch, not just smell the wildflowers…………..73 dr. gloria m. caddell journal of the oklahoma native plant society, volume 4, number 1, december 2004 oklahoma native plant record journal of the oklahoma native plant society 2435 south peoria tulsa, oklahoma 74114 volume 4 number 1, december 2004 issn 1536-7738 managing editor, sheila a. strawn technical editor, patricia folley technical advisor, bruce hoagland cd-rom producer, chadwick cox website: http://www.usao.edu/~onps/ the purpose of the onps is to encourage the study, protection, propagation, appreciation and use of the native plants of oklahoma. membership in onps shall be open to any person who supports the aims of the society. onps offers individual, student, family, and life membership. 2004 officers and board members president: james elder onps service award chair: sue amstutz vice-president: constance murray conservation chair: chadwick cox secretaries: publicity chair: kimberly a. shannon publications co-chairs: tina julich sheila strawn treasurer: mary korthase constance taylor past president: patricia folley marketing co-chairs: board members: lawrence magrath paul buck susan chambers kay gafford photo contest co-chairs: melynda hickman patricia and chadwick cox monica macklin newsletter editor: chadwick cox elfriede miller librarian: bonnie winchester stanley rice website manager: chadwick cox northeast chapter chair: constance murray mailing committee chair: karen haworth central chapter chair: sharon mccain color oklahoma committee chair: cross-timbers chapter chair: constance murray suzanne mcallister wildflower workshop chair: larry magrath mycology chapter chair: clark ovrebo historians: carla and dale chlouber ann long award chair: patricia folley cover photo: sorghastrum nutans, indian harriet barclay award chair: grass, our state grass by charles lewallen constance taylor articles (c) the authors journal compilation (c) oklahoma native plant society except where otherwise noted, this work is licensed under a creative commons attribution-noncommercialsharealike4.0 international license, https://creativecommons.org/licenses/by-nc-sa/4.0/, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly attributed, not used for commercial purposes, and, if transformed, the resulting work is redistributed under the same or similar license to this one. https://doi.org/10.22488/okstate.17.100025 2 oklahoma native plant record volume 4, number 1, december 2004 oklahoma native plant record volume 4, number 1 table of contents forward ...................................................................................................... 3 mr. james f. elder, onps president ecological factors affecting the distribution of woody vegetation near the arkansas river, tulsa county with special reference to the smoke-tree .............................................................................. 4 masters thesis of dr. anne wanamaker long cotinus obovatus raf. (smoke-tree ) in oklahoma ................................ 24 dr. bruce w. hoagland giant cane and southeastern indian baskets ..................................... 26 wichita mountains wildlife refuge survey 2000 ms. julia a. jordan vascular flora of the chouteau wildlife management area wagoner county, oklahoma ......................................................... 30 dr. bruce w. hoagland and dr. forrest l. johnson (dec.) status and habitat characteristics of cypripedium kentuckiense (kentucky lady’s slipper) in southeastern oklahoma ................ 40 ms. amy k. buthod and dr. bruce w. hoagland common lawn and garden fungi of central oklahoma ................ 48 dr. clark l. ovrebo critic’s choice essay: support your local herbarium ..................... 55 dr. wayne j. elisens editorial: why do species’ names change? ...................................... 57 ms. patricia a. folley oklahoma native plant record volume 7, number 1, december 2007 3 foreword and forward while i always look “forward” to preparing each volume for you, i haven’t always gotten the “foreword” right. in fact, the wrong word is used in the table of contents of the first four volumes. it is misspelled in both the table of contents and in the section headings of the last two volumes. ruth boyd, who has proof-read the journal with me every year since its inception in 2001 is most likely doing summersaults in her grave because i missed the error in the title for this section for 6 years in a row. i will do better. but had it not been for ruth’s keen eye and sharp editing pencil our journal would not have become the respected source of botanical research that it has. we remain indebted to her for correcting my many other errors. in this foreword to volume seven, the oklahoma native plant record mourns the passing of ruth boyd and of larry magrath, two of the society’s long-time members. larry magrath was one of our major contributers of scientific papers. had it not been for larry’s willingness to submit significant articles and encourage others to do so, we would have had a very thin journal for the first three years. both larry and ruth experienced poor health for several years, but continued to work with the record, giving me time to learn how to manage without being overwhelmed by editorial responsibilities. with the passing of ruth and larry, we will all have to step up and accept more responsibility for passing on the legacy of botanical research in oklahoma and our new staff of proof-readers will do their best to get it right. yes, it takes more than one to replace ruth. the record will always need new authors, reviewers, proof-readers and editors. if we don’t step up and do it, no one else will. it’s time for each of us, perfect or not, to move forward, doing the most and the best that we can. to build a larger legacy for oklahoma botany, one that is built on the best practices of research, we need to be open to allowing others to see our work and give us advice. with that comes responsibility. we must respect ownership of ideas. that’s why oklahoma native plant journal does not seek to own the work of our authors. we publish the articles while authors retain ownership and decide who else can use it. we believe in open sources and encourage open research. we look forward to receiving articles submitted to us in the future. in this volume bruce hoagland presents more articles based on data from the oklahoma natural heritage inventory. one gives us an updated perspective on charles s. wallis’ work vascular plants of the oklahoma ozarks, which represents our historical article this year. wallis was born in 1911 and compiled this flora for his phd thesis at oklahoma state university in 1959. hoagland’s other contribution this year was done with amy buthod, as an inventory of vascular plants at the new oklahoma centennial botanical garden in osage county. as part of our goal to encourage new authors, we enthusiastically present caleb stott’s the need for savanna restoration in the cross timbers. it is a review of relevant literature regarding one of oklahoma’s most endangered ecosystems. it is co-authored with mike palmer and kelly kindischer. in another article, mike palmer has also given us a great new research tool. it is a checklist for oklahoma floras. he has gathered all the known published floras of oklahoma and catalogued them in tabular form, referencing geographic, topographic, and taxonomic data to a bibliography of 85 references for oklahoma flora. with this volume, the oklahoma native plant record continues to bring you interesting and valuable scientific works which will enhance the purpose of the society, to promote the study, protection, propagation, appreciation, and use of native plants of oklahoma. thank you for your support. sheila strawn, editor oklahoma native plant record 67 volume 1, number 1, december 2001 magrath, l.k. https://doi.org/10.22488/okstate.17.1000066 galium parisiense var. leiocarpum tausch, new for oklahoma lawrence k. magrath curator, usao (ocla) herbarium chickasha, oklahoma73018-5358 while doing some routine plant collecting in chickasha in june of 1999 i found a galium with which i was not familiar. however, when i tried to key it out in smith (1994) it immediately keyed to galium parisiense l. var. leiocarpum tausch. it also keyed out in fernald (1950), britton (1907), and munz and keck (1963) and hickman (1993). this species is originally from europe. oklahoma: grady county: chickasha, walmart plaza on ponderosa drive; t6n, r7w, sw 1/4 sec 3; open disturbed grassy area on west side of street, red clay and some sand; elevation ca 1100 ft; scattered to locally abundant in ca 5 acre area; flowers greenish-white, some with red-brown corolla lobes; 9 june 1999; l.k. magrath 20590 (ocla). additional collections at same site: 25 june 1999; l.k. magrath, pete taylor, et al 20693 (ocla). 27 may 2001; l.k. magrath 21449 (ocla). 24 june 2001; l.k. magrath 21452 (ocla). plants dead but with some seeds persisting. oklahoma: oklahoma county: oklahoma city, will rogers park and garden center at nw 36 th street and i-240; grassy area near edge of wooded area southeast of rose garden; scattered; most plants already dead, but with some seeds persisting; 16 june 2001; l.k. magrath 21451 (ocla) also found at the same site: galium pedemontanum (bell.) all. oklahoma: grady county: chickasha, walmart plaza on ponderosa drive; t6n, r7w, sw 1/4 sec 3; open disturbed grassy area on west side of street, red clay and some sand; elevation ca 1100 ft; scattered to locally abundant in ca 5 acre area; flowers yellow to greenish-yellow; 25 april 2001; l.k. magrath, stephen garvin, val maseykin et al. 21344 (ocla); another collection at same site: 27 may 2001; l. k. magrath 21448 (ocla). significance: galium parisiense l. var. leiocarpum tausch represents an extension west from north central arkansas (baxter, fulton, newton & pope counties). this is a rather significant jump to the west from ozarkian woodlands habit to a relatively open mid-grass prairie habitat. the galium pedemontanum (bell.) all. is an extension northward and westward from southern and eastern oklahoma (taylor & taylor 1994). i would guess that both species arrived in chickasha via freight being delivered to the walmart shopping center. the oklahoma city location is near an area with numerous visitors, and next to an interstate highway, so there are several possibilities that might account for its presence at this location. it may be worth checking similar locations in other cities in oklahoma for these species and other possible introductions. references: britton, n. l. 1907. manual of the flora of the northern states and canada. 2nd ed. henry holt & co. ny. fernald, m. l. 1950. gray's manual of botany. 8 th ed. american book co. ny. gleason, h. a. and a. cronquist. 1963. manual of vascular plants of northeastern united states and adjacent canada. d. van nostrand co., inc. princeton, n.j. hickman, j. c. ed. 1996. the jepson manual: higher plants of california. univ. of california press. berkeley, ca munz, p. a. & d. d. keck. 1963. a california flora. univ. of california press. berkeley, ca. smith, e. b. 1978. an atlas and annotated list of the vascular plants of arkansas. dept. of botany & bacteriology, univ. of arkansas at fayetteville; fayetteville, ar. smith, e. b. 1994. keys to the flora of arkansas. univ. arkansas press; fayetteville, ar. taylor, r. j. & c. e. s. taylor. 1994. an annotated list of the ferns, fern allies, gymnosperms and flowering plants of oklahoma. biology dept. herbarium, southeastern oklahoma state univ.; durant, ok journal of the oklahoma native plant society, volume 11, december 2011 five year index to oklahoma native plant record volume 6 4 the lichens of north central oklahoma, darvin w. keck 51 annotated nomenclatural update to keck (1961), douglas m. ladd 53 vascular flora of a red sandstone hills site, canadian county, oklahoma, bruce w. hoagland and amy k. buthod 69 vascular flora of a riparian site on the canadian river, cleveland county, oklahoma, lacy burgess and bruce w. hoagland. 80 critic’s choice essay: cedar-apple rust, clark l. ovrebo volume 7 4 vascular plants of the oklahoma ozarks, charles s. wallis 21 updated oklahoma ozark flora, bruce w. hoagland 54 the vascular flora of the oklahoma centennial botanical garden site osage county, oklahoma, bruce w. hoagland and amy buthod 67 vascular plant checklists from oklahoma, michael w. palmer 78 the need for savanna restoration in the cross timbers, caleb stotts, michael w. palmer, and kelly kindscher 91 botanizing with larry magrath, patricia a. folley volume 8 4 a floristic study of the vascular plants of the gypsum hills and redbed plains area of southwestern oklahoma, 1975 m. s. thesis, susan c. barber 37 updated list of taxa for vascular plants of the gypsum hills and redbed plains area of southwestern oklahoma, susan c. barber 45 updated flora of the wichita mountains wildlife refuge keith a. carter, pablo rodriguez, and michael t. dunn 57 common spring mushrooms of oklahoma, clark l. ovrebo and nancy s. weber 61 fern habitats and rare ferns in oklahoma, bruce a. smith 67 tribute to paul buck, constance murray volume 9 4 vascular plants of southeastern oklahoma from san bois to the kiamichi mountains, 1969 ph.d. dissertation, f. hobart means 38 composition and structure of bottomland forest vegetation at the tiak research natural area, mccurtain county, oklahoma bruce w. hoagland and newell a. mccarty 59 is seedling establishment very rare in the oklahoma seaside alder, alnus maritima ssp. oklahomensis?, stanley a. rice and j. phil gibson 64 whatever happened to cheilanthes horridula and cheilanthes lindheimeri in oklahoma? bruce a. smith 70 critic’s choice essay: invasive plants versus oklahoma’s biodiversity, chadwick a. cox volume 10 4 the identification of some of the more common native oklahoma grasses by vegetative characters, 1950 m. s thesis, william franklin harris 34 the vascular flora of hale scout reservation, leflore county, oklahoma bruce w. hoagland and amy k. buthod 54 the toxicity of extracts of tephrosia virginiana (fabaceae) in oklahoma mary gard 65 four western cheilanthoid ferns in oklahoma, bruce a. smith 77 critic’s choice essay: being a method proposed for the ready finding ... to what sort any plant belongeth, ronald j. tyrl oklahoma native plant society c/o tulsa garden center 2435 south peoria tulsa, oklahoma 74114 _________________________________________________________________________ in this issue of oklahoma native plant record volume 11, december 2011: _________________________________________________________________________ 4 survey of the vascular flora of the boehler seeps and sandhills preserve, ph.d. dissertation linda gatti clark 22 schoenoplectus hallii, s. saximontanus, and the putative s. hallii x s. saximontanus hybrid: observations from the wichita mountains wildlife refuge and the fort sill military reservation from 2002 – 2010 marian smith and paul m. mckenzie 33 spatial genetic structure of the tallgrass prairie grass dichanthelium oligosanthes (scribner’s panicum) molly j. parkhurst, andrew doust, margarita mauro-herrera, janette a. steets, and jeffrey m. byrnes 43 the effects of removal of juniperus virginiana l. trees and litter from a central oklahoma grassland jerad s. linneman, matthew s. allen, and michael w. palmer 61 the changing forests of central oklahoma: a look at the composition of the cross timbers prior to euro-american settlement, in the 1950s, and today richard e. thomas and bruce w. hoagland 75 critic’s choice essay: some thoughts on oklahoma plants and summer 2011’s exceptional drought leslie e. cole five year index to oklahoma native plant record – inside back cover journal of the oklahoma native plant society volume 12, december 2012 five year index to oklahoma native plant record volume 7 4 vascular plants of the oklahoma ozarks, charles s. wallis 21 updated oklahoma ozark flora, bruce w. hoagland 54 the vascular flora of the oklahoma centennial botanical garden site osage county, oklahoma bruce w. hoagland and amy buthod 67 vascular plant checklists from oklahoma, michael w. palmer 78 the need for savanna restoration in the cross timbers caleb stotts, michael w. palmer, and kelly kindscher 91 botanizing with larry magrath, patricia a. folley volume 8 4 a floristic study of the vascular plants of the gypsum hills and redbed plains area of southwestern oklahoma, 1975 m. s. thesis, susan c. barber 37 updated list of taxa for vascular plants of the gypsum hills and redbed plains area of southwestern oklahoma, susan c. barber 45 updated flora of the wichita mountains wildlife refuge keith a. carter, pablo rodriguez, and michael t. dunn 57 common spring mushrooms of oklahoma, clark l. ovrebo and nancy s. weber 61 fern habitats and rare ferns in oklahoma, bruce a. smith 67 tribute to paul buck, constance murray volume 9 4 vascular plants of southeastern oklahoma from san bois to the kiamichi mountains, 1969 ph. d. dissertation, f. hobart means 38 composition and structure of bottomland forest vegetation at the tiak research natural area, mccurtain county, oklahoma, bruce w. hoagland and newell a. mccarty 59 is seedling establishment very rare in the oklahoma seaside alder, alnus maritime ssp. oklahomensis? stanley a. rice and j. phil gibson 64 whatever happened to cheilanthes horridula and cheilanthes lindheimeri in oklahoma? bruce a. smith 70 critic’s choice essay: invasive plants versus oklahoma’s biodiversity, chadwick a. cox volume 10 4 the identification of some of the more common native oklahoma grasses by vegetative characters, 1950 m. s. thesis, william franklin harris 34 the vascular flora of hale scout reservation, leflore county, oklahoma bruce w. hoagland and amy k. buthod 54 the toxicity of extracts of tephrosia virginiana (fabaceae) in oklahoma, mary gard 65 four western cheilanthoid ferns in oklahoma, bruce a. smith 77 critic’s choice essay: being a method proposed for the ready finding ... to what sort any plant belongeth, ronald j. tyrl volume 11 4 survey of the vascular flora of the boehler seeps and sandhills preserve, ph. d. dissertation, linda gatti clark 22 schoenoplectus hallii, s. saximontanus, and the putative s. hallii x s. saximontanus hybrid: observations from the wichita mountains wildlife refuge and the fort sill military reservation from 20022010, marian smith and paul m. mckenzie 33 spatial genetic structure of the tallgrass prairie grass dichanthelium oligosanthes (scribner’s panicum), molly j. parkhurst, andrew doust, margarita mauro-herrera, janette a. steets, and jeffrey m. byrnes 43 the effects of removal of juniperus virginiana l. trees and litter from a central oklahoma grassland, jerad s. linneman, matthew s. allen, and michael w. palmer 61 the changing forests of central oklahoma: a look at the composition of the cross timbers prior to euro-american settlement, in the 1950s and today, richard e. thomas and bruce w. hoagland 75 critic’s choice essay: some thoughts on oklahoma plants and summer 2011’s exceptional drought, leslie e. cole oklahoma native plant society c/o tulsa garden center 2435 south peoria tulsa, oklahoma 74114 _________________________________________________________________________ in this issue of oklahoma native plant record volume 12, december 2012: _________________________________________________________________________ 4 possible mechanisms of the exclusion of johnson grass by tall grass prairies, m. s. thesis marilyn a. semtner 33 a preliminary pawnee ethnobotany checklist c. randy ledford 43 vascular flora of alabaster caverns state park, cimarron gypsum hills, woodward county, oklahoma gloria m. caddell and kristi d. rice 63 a comparison of the composition and structure of two oak forests in marshall and pottawatomie counties bruce smith 69 critic’s choice essay: virtual herbaria come of age wayne elisens five year index to oklahoma native plant record – inside back cover journal of the oklahoma native plant society, volume 15, december 2015 oklahoma native plant record 3 volume 15, december 2015 foreword after 15 years, we are more than pleased with the variety of excellent articles submitted and accepted for publication in the oklahoma native plant record. this year, as most years, together, they meet all onps goals. “encouraging the study of native plants.” we never know how the record of a single study will encourage future research, but we are sure our historic article will be of special value to today’s botanists and ecologists studying historic species distributions and environmental changes. in 1934, ben osborn may not have been aware of how valuable his list of flowering dates would be to the issue of global warming, but his article, “first flowering dates for central oklahoma” fills that role. in his preface to that article, dr. wayne elisens contributes the history that puts that data into perspective. floristic surveys, like that of black mesa by amy buthod and bruce hoagland from the oklahoma biological survey, and descriptions like those in forest structure and fire history at lake arcadia by chad king, from the university of central oklahoma, make future comparative studies not only possible, but likely. “encouraging the protection of native plants.” kudzu (pueraria montana) has long been described as an invasive species, but like many exotic species that have been introduced without thought of how they would interact with native species, it didn’t start out that way. marli claytor and karen hickman from oklahoma state university summarize the current extent of kudzu and what might be done to protect our native species. “encouraging the propagation of native plants.” the risks of monoculture plantings and the benefits of planting multiple species within gardens is the topic of the article by oklahoma state university’s bonner, rebek, cole, kahn, and steets. this research is important for landscapers and gardeners because of plant species’ effects on arthropod abundance, a main point of douglas tallamy’s recent presentations at the society’s events in tulsa and oklahoma city. their article provides the data and reasons to heed his advice. “encouraging the appreciation of native plants.” for enthusiasts and plant lovers, this year we have started a new tradition, by chosing our critic’s choice essay from previous “botanist’s corner” articles published in the gaillardia, the society’s newsletter. this year’s essay, by the late paul buck, about an often maligned native species, is entitled “mistletoe, phoradendron serotinum”. “encouraging the use of native plants.” in the past, we have published articles about how native americans used native plant species. “antifungal activity in extracts of plants from southwestern oklahoma against aspergillus flavus” shows us how plants can be used for more current medicinal purposes. it is also a great example of research projects that can inspire students who are involved to continue in botany. this year’s student research project is from tahzeeba frisby and her students at cameron university in lawton. as you can see, articles for all interest groups of our membership (gardeners, academic faculty, landscapers, and enthusiasts) are represented. it is the wide variety of authors who contribute to our journal that helps us bring those many interests together in ways that best promote our goals. why not consider submitting your manuscript next year? remember that our editorial board includes a manuscript editor, dr. mark fishbein, who can find help for first time and citizen-scientist authors. tell us about your ideas and submit your articles early, so we can see that your work gets the most helpful reviews and comments. don’t forget that the oklahoma native plant record is a professionally reviewed publication, listed globally in the “directory of open access journals”, and our abstracts are indexed in the “centre for agricultural bioscience international”, which is based in the u.k. sheila strawn, managing editor journal of the oklahoma native plant society, volume 7, number 1, december 2007 91 oklahoma native plant record volume 7, number 1, december 2007 editorial botanizing with larry magrath sunday, october 4, 1998. a field trip for two doesn’t take much planning – a phone call will do: “one of my students has brought in a collection of scirpus hallii. want to go with me on sunday to verify the site?” well, of course! larry was one of the state’s most ardent collectors, and s. hallii (name since changed to schoenoplectus hallii) is a sedge. that makes it a plant i need to know. just after 8 a.m. that sunday i picked up larry and his gear in chickasha, and we headed southwest. but first – he’d thought of another lake that was “almost on the way”, and there were exposed mud flats just covered with sedges. so we went due west for maybe 10 miles, to lake burtschi. there were thousands of inch-tall sedges of several different species; cyperus surinamensis, c. aristatus, fimbristylis autumnalis, and fuirena simplex, mostly. they lay on the damp sand like a city lawn; tiny annual species doing their best to set seed before frost. there we also collected samples of arundo donax, a grass that grows in shallow water, and can reach more than two meters tall. then, “since we are in the neighborhood” we stopped at a private property called williams’ wilderness, whose owner had given permission. there we found an orchid, hexalectris spicata and some other goodies. traveling sw on sh19, we stopped along the south edge of apache near lakeside village to see how lake ellsworth had fared. that lake was down ten feet, and had exposed acres of sandy bottom, much of it covered with the tiny annual sedges. all were in furious bloom. there we collected a sedge-like grass, eragrostis reptans; as well as fimbristylis vahlii, cyperus odoratus, amannia coccinea, and a strange liverwort called riccocarpus natans. the upper edges were banked with a vigorous morning glory with small white flowers, ipomoea lacunosa. finally we reached jed johnson lake in the wichita mountain wildlife reservation. there, an expanse of mud flats some 4 meters wide and ten meters long had been exposed by the low water. the shoreline was composed of broken red-granite gravels and sand, much disturbed by fishermen. scirpus hallii was there in abundance and in bloom or fruit. larry counted 114 plants, and each of us collected a specimen for our herbarium. our trip had been both entertaining and successful. think it was over? you’ve never been on a field trip with larry! we were free to wander as far and wide as our strength and the day lasted. we checked rush lake, also on the reserve, and found it embedded in a huge stand of eleocharis quadrangulata. while i took pictures, a curious armadillo came right up and sniffed near-sightedly at my shoe. lunch with larry was always a challenge: it had to be fast, and it had to be vegetarian. veggie fast food isn’t readily available along country roads. we settled for sub sandwiches at the love’s station on sh49. dodging traffic through lawton, we took sh 7 east to sh 65, then went south through temple, turned east there on sh 5, and soon arrived at moneke park near lake waurika. hiking through an open forest community, we found the other relative of poke-weed, rivina humilis, and in bloom. first time i’d ever seen that. it was a real treat. our day was coming to an end. the cloud bank that had hovered to the west all day long grew higher and darker. we reluctantly headed north on us 81, but soon https://doi.org/10.22488/okstate.17.100056 92 oklahomanative plantrecord volume 6,number 1,december 2006 had to make a stop just north of addington. there, beside the highway, is a large prairiedog community, and we enjoyed their company until it grew too dark for photography. by the time we reached chickasha, it was pitch-black except for the lightning that was almost intense enough to drive by. larry unloaded his prizes in a heavy downpour, and i headed for home by sh9. again, lightning and heavy, heavy rain accompanied the trip. by way of the evening news, i learned that ninnekah, just south of chickasha, had been struck by a tornado right after we drove through, and that a swarm of them had produced the lightning that made the passage so interesting. the tornadoes had covered a large swath of central oklahoma that night while larry and i were busy pressing the plants and writing up our notes. over the years there were many such field trips with larry, most of them with the oklahoma native plant society or the nature conservancy. each of them was “floriferous” and interesting. the photo below is from one of our trips to round mountain in leflore county with jim norman and charles lewallen, who set up the remote photo. patricia folley, 1 june 2007 onps larry magrath, botanizing with patricia folley, charles lewallen, and jim norman. oklahoma native plant record, volume 11, number 1, december 2011 1 oklahoma n ative plant record journal of the oklahoma native plant society 2435 south peoria tulsa, okla homa 74114 volume 11, december 2011 issn 1536-7738 http://ojs.library.okstate.edu/osu/ managing editor: sheila strawn production editor: paula shryock electronic production editor: sandy graue technical advisor: bruce hoagland editorial assistants: patricia folley, kristi rice the purpose of onps is to encourage the study, protection, propagation, appreciation and use of the native plants of oklahoma. membership in onps is open to any person who supports the aims of the society. onps offers individual, student, family, and life memberships. 2011 officers and board members president: lynn michael vice-president: marilyn stewart secretary: sandy graue treasurer: mary korthase membership coordinator: tina julich historian: sharon mccain past president: kim shannon board members: clare miller buddy miller marilyn stewart ron tyrl brooke bonner janette steets central chapter chair: joe roberts cross-timbers chapter chair: ron tyrl mycology chapter chair: steve marek northeast chapter chair: alicia nelson color oklahoma chair: tina julich conservation chair: chadwick cox gaillardia editor: chadwick cox website manager: chadwick cox http://projects.usao.edu/~onps/index.html harriet barclay award chair: rahmona thompson anne long award chair: gloria caddell onps service award chair: sue amstutz photo poster curators: sue amstutz & marilyn stewart photography contest chair: kim shannon nominations chair: sheila strawn librarian: karen haworth mailings chair: karen haworth publicity chairs: kim shannon & marilyn stewart wildflower workshop chair: lynn michael cover photo by carolyn lilly “fritillary and thistle”, 1st place photo contest winner, special category: flora and fauna, 2011. articles (c) the authors journal compilation (c) oklahoma native plant society except where otherwise noted, this work is licensed under a creative commons attributionnoncommercial-sharealike4.0 international license, https://creativecommons.org/licenses/by-ncsa/4.0/, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly attributed, not used for commercial purposes, and, if transformed, the resulting work is redistributed under the same or similar license to this one. https://doi.org/10.22488/okstate.17.100079 http://ojs.library.okstate.edu/osu/� http://projects.usao.edu/~onps/index.html� oklahoma native plant record volume 11, december 2011 2 oklahoma native plant record volume 11 table of contents foreword ................................................................................................................................................. 3 survey of the vascular flora of the boehler seeps and sandhills preserve, ph.d. dissertation ................................................................................... 4 dr. linda gatti clark schoenoplectus hallii, s. saximontanus, and the putative s. hallii ×s. saximontanus hybrid: observations from the wichita mountains wildlife refuge and the fort sill military reservation 2002 – 2010 .......................................................... 22 dr. marian smith and dr. paul m. mckenzie spatial genetic structure of the tallgrass prairie grass dichanthelium oligosanthes (scribner’s panicum) ........................................................................................... 33 ms. molly j. parkhurst, dr. andrew doust, dr. margarita mauro-herrera, dr. janette a. steets, and dr. jeffrey m. byrnes the effects of removal of juniperus virginiana l. trees and litter from a central oklahoma grassland .............................................................................................. 43 mr. jerad s. linneman, dr. matthew s. allen, and dr. michael w. palmer the changing forests of central oklahoma: a look at the composition of the cross timbers prior to euro-american settlement, in the 1950s, and today ....................................................................................................... 61 dr. richard e. thomas and dr. bruce w. hoagland critic’s choice essay: some thoughts on oklahoma plants and summer 2011’s exceptional drought ............................................................................................................ 75 leslie e. cole, d.v.m. editorial policies and procedures ..................................................................................................... 77 five year index to oklahoma native plant record ........................................... inside back cover oklahoma native plant record, journal of the oklahoma native plant society, volume 11, december 2011 title page table of contents foreword oklahoma native plant record, volume 15, number 1, december 2015 4 oklahoma native plant record volume 15, december 2015 wayne elisens https://doi.org/10.22488/okstate.17.100110 preface to first flowering dates for central oklahoma wayne elisens professor of plant biology curator of the bebb herbarium (okl) department of microbiology and plant biology university of oklahoma norman, ok elisens@ou.edu global climate change is predicted to have deleterious effects on human health and welfare including frequency of extreme weather events, sea level rise and coastal flooding, decreased agricultural productivity, fluctuating biotic interactions and range shifts, and altered seasonality and phenology (ipcc 2014). phenology, the study of cyclic and seasonal natural phenomena such as flowering and animal migrations, is especially important as an indicator of changing climates and ecosystem changes (e.g., diez et al. 2012). for plants, tracking of firstor peak-flowering events has been a common approach to investigate species’ responses to climatic factors. individuals, organizations, and botanical gardens have recorded flowering times for a wide range of species over many years (tooke & battey 2010). currently, academic as well as citizen scientists are actively engaged in gathering plant phenological data. schools, online communities, and native plant societies are often involved in phenological tracking activities (e.g., haggerty and mazer 2008) by partnering with agencies such as the usa national phenology network (www.usanpn.org). below is a privately printed but unpublished report of first flowering dates for a variety of species in central oklahoma from 1927–1929 and 1933. much of the baseline data was gathered by lois gould in 1927–1929 (gould 1928, 1929a, 1929b) in central oklahoma as part of a comparative study among 14 midwestern colleges and universities. in 1933, ben osborn added observations from norman and oklahoma city and organized the compiled data chronologically to provide a 3-year record of flowering phenology by earliest and average flowering date, species name, common name, location, year of observation, and observer (gould or osborn). mr. osborn typed this report and deposited a copy in the library of the robert bebb herbarium of the university of oklahoma as “separate no. 27” where it has remained until this printing. the present report has not been published formally to the best of our knowledge. we hope this summary of first flowering in central oklahoma in the early twentieth century will assist present-day investigations of the biological effects of climate change by providing a valuable plant phenological benchmark. lois h. gould, daughter of dr. charles gould who was the former director of the oklahoma geological survey, received a b.a. in botany from the university of oklahoma in 1930. her passions included art, plants, and birds. ms. gould married the canadian entomologist dr. ralph d. bird, who taught at the university of oklahoma from 1929 to 1933. mrs. (gould) bird moved to canada when her husband accepted a position at a canadian federal entomology laboratory (anonymous 1972). oklahoma native plant record 5 volume 15, december 2015 wayne elisens ben o. osborn received the first bachelor’s degree in agricultural journalism from oklahoma a&m college (oklahoma state university) in 1931. he began his career as a copy editor with the oklahoma livestock news and was a news and radio script writer for the oklahoma agricultural extension service. he then embarked on a 36-year career with the usda as a soil conservationist, information specialist, speechwriter, and editor for the journal of soil and water conservation (anonymous 1999 ). with co-author elizabeth barkley, mr. osborn published a list of the vascular plants of pottawatomie county, oklahoma (barkley and osborn 1933). literature cited anonymous. 1972, march 4. ralph durham bird obituary. manitoba free press. anonymous. 1999, november 9. ben o. osborn obituary. the washington post. barkley, e.a. and b. osborn. 1933. a preliminary list of the vascular plants of pottawatomie county, oklahoma. norman (ok): university of oklahoma. diez, j.m., i. ibañez, a.j. miller-rushing, s.j. mazer, t.m. crimmins, m.a. crimmins, c.d. bertelsen, and d.w. inouye. 2012. forecasting phenology: from species variability to community patterns. ecology letters 15:545−553. gould, l. 1928. notes on the phenology of middle western spring flowers. proceedings of the oklahoma academy of science 8:59−62. gould, l. 1929a. phenological notes for 1928 on middle western spring flowers. proceedings of the oklahoma academy of science 9:39−42. gould, l. 1929b. lateness of the spring of 1928 as determined by comparison of first blooming dates. proceedings of the oklahoma academy of science 9:43−46. haggerty, b.p. and s.j. mazer. 2008. the phenology handbook. santa barbara (ca): university of california. ipcc, intergovernmental panel on climate change. 2014. climate change 2014: impacts, adaptation, and vulnerability. part a: global and sectoral aspects. field, c.b, v.r. barrow, d.j. dokken, k.j. mach, m.d. mastrandrea, t.e. bilir, m. chatterjee, k.l. ebi, y.o. estrada, r.c. genova, b. girma, e.s. kissel, a.n. levy, s. maccracken, p.r. mastrandrea, and l.l. white (eds.). new york: cambridge university press. tooke, f. and n. h. battey. 2010. temperate flowering phenology. journal of experimental botany 61:2853−2862. preface to first flowering dates for central oklahoma by dr. wayne elisens oklahoma native plant record, volume 16, number 1, december 2016 oklahoma native plant record 79 volume 16, december 2016 paul buck https://doi.org/10.22488/okstate.17.100124 critic’s choice essay a conversation with a small beetle reprinted from gaillardia, fall 2000 paul buck, deceased professor emeritus department of biological science university of tulsa tulsa, ok 74104 today we find our lives filled with technological innovations such as personal computers, the internet and email, supersonic aircraft, space probes, interspecies gene transfers, and on and on. yet, you think we lead an unusual life! let me tell you what happened recently to an insect acquaintance. she related her tale of woe as i sat out back watching the tall phlox pat folley gave me grow taller. first, a number of gardeners in the neighborhood grow arum italicum mill., an arum lily, for its large, attractive, light veined leaves and clusters of beautiful, bright red berries which appear late in the growing season. introduced to north america, the species is native to southern europe and in some areas of italy is considered a common weed. in oklahoma, flowering takes place in may, and in arum the reproductive structure is actually not the typical flower but an inflorescence surrounded by a large leaf. on our oklahoma native plant society field trips we have seen numerous jack-in-the-pulpit plants, and the floral system is quite similar. the erect flowering stalk (spadix) is enclosed in an enveloping bract (spathe). the flowers are unisexual with the pistillate (female) at the base of the spadix and the staminate (male) above. over those two sets of fertile flowers is a whorl of sterile flowers which, when inflated, form a barrier between the floral chamber within the spathe and the open area above. my friend, a small, dark beetle, said her recent experience started one warm afternoon while foraging when she sensed what she felt was the aroma of food (you and i would probably say it smelled like a combination of carrion and urine). she followed her “nose” to a large plant (we later identified it as arum italicum) and landed on the open throat of the spathe. a large number of beetles, gnats, and blowflies had already gathered. she sensed the aroma was welling up from the tubular spathe and, again following her “nose,” walked to the opening. she reported slipping at the edge on tiny oil droplets and falling through some bristles into the depths of the chamber. there she found the stigmas of the pistillate flowers covered with a sweet, slimy fluid. she noted the inflated bristles that so readily permitted her fall were keeping large insects out. they were forced to fly off, seeking food elsewhere. once at the bottom of the pit, her first thought was of escape. however, she discovered the walls of the lower chamber were just as slippery as the upper spathe surface, and climbing out was impossible until she realized she could climb over the lower female flowers. as she did, she noticed others with pollen on their backs losing those grains to the sticky surfaces of the female flowers as 80 oklahoma native plant record volume 16, december 2016 paul buck they labored upward. unfortunately when the group reached the base of the bristles, they encountered downward pointing hairs which prevented further progress. she lamented, “what to do?”, but only briefly. the chamber was warm and out of the rain, abundant food was being produced by the flowers, and about half the crowd was male. there was but one thing to do — party! with the setting of the sun, my friend and the others settled down. i do not know if it was the darkness, full bellies and party fatigue, or simply bedtime for little beetles. however, during the night the staminate flowers matured and rained pollen from above. with dawn and the rising sun, everyone awoke to find themselves coated with pollen adhering to the sticky exudate from the stigmas. once again, how to escape? lo and behold, the downward pointing hairs had wilted along with the bristles, and it was possible to climb up and over their wrinkled surfaces to the throat of the spathe and freedom. interestingly, the upper portion of the spathe (appendix) had lost the carrion aroma, and my friend with her pollen-laden companions, previously prisoners of the night, flew away. however, the escaping insects picked up the aroma of another arum inflorescence and agreed to drop by for a visit, only to be trapped in a new prison chamber. this time, one where the pollen on their backs would be transferred to the flowers, and pollination would take place. when i last saw my friend, she was joining a group headed toward yet another arum plant. there is an additional feature of arum i would like to mention before closing. the terminal portion of the spadix, the appendix, is the source of the aroma, unpleasant to you and me but attractive to my beetle friend. while the chemical producing the aroma is being released, the appendix tissue generates heat to the point that it may be as much as 36 degrees warmer than the surrounding air. research suggests the temperature elevation serves to volatilize the smelly compound, increasing the speed with which it is spread into the atmosphere. for most of us, that is an interesting aspect of the overall process, and we quickly see the reason behind it. for you chemists, it raises another question. what metabolic pathways are utilized by the plant to produce such significant energy release? how do the plants do it? what an interesting story and introduction to pollination ecology right in the back yard. you see there is a benefit to taking a few minutes to chat with a small beetle. onps critic’s choice essay: a conversation with a small beetle. gaillardia, fall 2000 by dr. paul buck oklahoma native plant record, volume 11, number 1, december 2011 oklahoma native plant record volume 11, december 2011 cole, l.e. https://doi.org/10.22488/okstate.17.100086 75 some thoughts on oklahoma plants and summer 2011’s exceptional drought leslie e. cole, d.v.m. oklahoma has just had a summer of incredible heat and exceptional drought (d4), the worst such designation possible from the national drought monitor and marked by a menacing dark red on the drought maps they make. the impacts of this tough climate event can be seen everywhere one looks from the dormant or dead blackjacks and eastern redcedars to the thin young crows, and felt in the economy and our communities. we can set the stage for these musings by quoting mr. gary mcmanus, associate state climatologist, oklahoma climatological survey. oklahoma is “… just coming off one of our driest you-name-the-period on record. the current drought originated in september 2010 with the arrival of la nina in the equatorial pacific waters. this water year, which ended september 2011, finished as the second driest on record for oklahoma with a statewide average precipitation total of 20.26 inches, 16.43 inches below normal. the driest such period on record was 18.69 inches from the 1955-1956 water year. for the panhandle, west central, central and southwestern parts of the state, it was easily the driest water year on record. southwest oklahoma’s water year average of 12.68 inches was more than 18 inches below normal and nearly 5 inches drier than the previous record low total of 17.45 inches, again from the 1955-56 water year.” and to gain perspective on the record heat of summer 2011, also from mr. mcmanus… “according to data from the oklahoma mesonet, the state’s climatological summer, june 1 through august 31, ended with a statewide average of 86.8 degrees, obliterating the previous state record of 85.2 degrees from the summer of 1934.” oklahoma has won the prize for hottest summer for any state since records began. oklahoma agriculture has experienced nearly $2 billion dollars in losses due to the current drought, according to estimates by oklahoma state university’s division of agricultural sciences and natural resources. the oklahoma department of agriculture, food, and forestry estimates crop losses of more than $953 million and cattle losses of about $1 billion. much of oklahoma is still prairie grassland and well suited to produce protein from grazing herbivores like cattle. oklahoma is home to the second largest beef cattle herd in the united states and the drought, with its withered grasses and forage and dried ponds, has hit the state’s beef producers hard, particularly those west of interstate 35 and in southwest oklahoma, according to the oklahoma cattlemen’s association. these agricultural losses will be the highest ever recorded in a single year for our state. as a veterinarian, i suspect that a lot of less than ideal hay will be fed and less than ideal areas will be grazed in oklahoma this fall and winter. we could see increased oral injury in livestock due to the presence of higher numbers of mature grass seed heads in graze and late cut hay (like setaria or foxtail grass) with stiff barbed bristles that easily penetrate flesh and are kept there by the barbs. relatively small amounts of these types of oklahoma native plant record volume 11, december 2011 cole, l. e. 76 bristled seed heads in hay can produce these lesions (1.8% for setaria). a significant volume of hay is being shipped into oklahoma from out of state. there could be toxic plants and seeds associated with this out of state hay that we don’t normally see in oklahoma. those concerned about livestock health and oklahoma native plants should be on the lookout for these aliens. drought stressed plants can accumulate or produce toxins not usually present under more “normal” conditions. cyanogenetic glycosides that yield hydrocyanic acid (hcncyanide) upon hydrolysis are a concern in stressed sorghum spp. (sudan grass, johnsongrass, etc.), prunus spp. and others. nitrate levels in stocks, other forage or some hay are a concern as well. plants that accumulate or produce toxins are usually avoided by browsers and grazers; however, when said browsers and grazers are faced with eating dirt or that less than palatable toxic plant, the toxic plant gets eaten. oklahoma has many toxic plants that can cause problems if ingested by livestock; for example, the loco weeds (astragalus spp. and oxytropis spp.), selenium accumulators (astragalus spp. again and stanleya spp.), oxylate accumulators (rumex spp. and chenopodium spp.) and saponin producers (phytolacca and sesbania spp.). even some of oklahoma’s smallest plants have been important in this past summer’s heat and drought. “blooms” or explosive reproduction of bluegreen algae in warm state waters this summer have made the headlines. individual cells of these organisms are microscopic but they are collected into colonies, filaments or masses of filaments. cattle, sheep, horses, swine, dogs, cats, fowl, geese, wild and domestic ducks, game and song birds, fish, rodents, and small game have been killed by ingesting these smallest of plants. lack of water can eventually lead to catastrophic biological failures and death in plants. one can think about the wilting of leaves and decreased turgor pressure, hydraulic failure with stomata closure and reduced photosynthesis and the myriads of other adaptations, mechanisms and strategies that plants employ to survive the stresses of oklahoma’s wild climate and unique ecology. what i remember most about this year’s drought was the amazing green-up of the grasses and the abundant fall flowers that followed the relatively meager fall rains. the native plants of oklahoma are scientifically fascinating, true survivors, and incredibly beautiful. kingsbury, john m. 1964. poisonous plants of the united states and canada. prentice hall: englewood cliffs, nj. national agricultural statistics service (nass). oklahoma field office and the oklahoma department of agriculture, food, and forestry (odaff). oklahoma agricultural statistics 2011. onps critic’s choice essay: some thoughts on oklahoma plants and summer 2011’s exceptional drought by leslie e. cole, d.v.m. journal of the oklahoma native plant society, volume 8, number 1, december 2008 57 oklahoma native plant record volume 8, number 1, december 2008 ovrebo & weber https://doi.org/10.22488/okstate.17.100062 common spring mushrooms of oklahoma clark l. ovrebo nancy s. weber department of biology department of forest ecosystems university of central oklahoma and society edmond, ok 73034 oregon state university e-mail: covrebo@uco.edu corvallis, oregon 97331 introduction springtime brings a resurgence of greenery and wildflowers to the landscape. for those interested in fungi it is time to look for mushrooms as well. in oklahoma spring mushrooms appear for approximately two to three-weeks from late march to mid-april. the exact time depends on temperature, moisture, and in which corner of the state you hunt mushrooms. collectors might encounter a few of the “gilled mushrooms”, basidiomycota, during the spring, but it is the members of the phylum ascomycota, often referred to as “ascomycetes”, that are the most prominent and popular of the spring fungi. here, we present a selection of common ascomycetes in the order pezizales, the morels and related cup-fungi that you may encounter in the spring woods. the ascomycota includes a diverse group of fungi ranging from yeasts to devastating plant pathogens, to edible wild mushrooms. members of this phylum bear their spores in microscopic sac-like structures called asci. some ascomycetes form fruiting bodies called ascocarps or ascomata. in the order pezizales, the basic form of the ascocarp is called an apothecium. apothecia resemble a cup or saucer with the asci lining the cup or covering the upper surface of the saucer; these fungi are often referred to as “cup-fungi.” in some species the apothecium has a stalk or stem. in others the apothecium may be recurved or contorted into any of a number of shapes including thimbles and pitted or wrinkled caps. below we provide photographs and brief descriptions of the more commonly encountered spring-fruiting members of the pezizales. more details can be found in most mushroom field guides. common names, where known, are given in parentheses. we are hesitant to provide information on edibility. great care must be taken to be absolutely sure of a mushroom’s identity. only after becoming proficient at identifying mushrooms, and only then, can one determine edibility. vouchers for specimens described here are housed in the mycological collection of the university of central oklahoma’s herbarium (csu). order pezizales morchella esculenta (common, yellow or tan morel) this is by far the most popular spring mushroom collected for consumption. it is recognized by its pitted cap with light tan or gray pits separated by creamy-white ribs when young. the ribs do not blacken at maturity. the entire length of the cap is attached to the stalk and both the cap and stalk are hollow. two slightly different forms of this species are illustrated. one has a more rounded cap (fig. 1) and the other is more tapered (fig. 2). the morphological variation within this species needs further investigation, so we cannot be certain whether these variations represent distinct species. websites such as www.mushroomexpert.com/morchella_yello w.html can offer more information. the common morel is usually found in wooded areas. river bottom forests seem to be good places in oklahoma for finding morels. we have frequently found morels near eastern red cedar trees as well. don’t count out metropolitan areas. the first author has found them in his own yard and on the university of central oklahoma campus. 58 oklahoma native plant record volume 8, number 1, december 2008 ovrebo, c.l. figure 1 morchlla esculenta with rounded cap. figure 2 morchella esculenta with tapered cap. morchella semilibera (half-free morel) it fruits at about the same time as the common morel and differs from the common morel by the way that the cap is attached to the stalk (fig. 3). the lower half of the cap is free from the stalk and resembles a skirt. the ribs of the cap turn dark brown to black with age and the caps are often darker and smaller than those of the common morel. figure 3 morchella semilibera gyromitra caroliniana gyromitra c. is by far the largest spring mushroom found in oklahoma (fig. 4). the cap is brownish red and convoluted or brainlike. the stalk is robust with the exterior formed into irregular, rounded ridges separated by irregular grooves. the inside tissue of the cap and stalk appears to be stuffed with folded or convoluted tissue (fig. 5). figure 4 gyromitra caroliniana external view figure 5 gyromitra caroliniana showing internal structure 59 oklahoma native plant record volume 8, number 1, december 2008 ovrebo & weber verpa conica (bell morel) it is recognized by the brown, smooth to slightly wavy, bell-shaped apothecium that is attached only at the stalk apex (fig. 6), resembling a thimble sitting on a finger. for that reason it is also called the thimble morel. figure 6 verpa conica helvella acetabulum this differs in outward appearance from the previous because the apothecium is cup-shaped (fig. 7). the inside of the apothecium is brown to grayish brown. a very short stalk may be present or absent. its surface has sharp-edged ribs that extend onto the lower surface of the apothecium, sometimes nearly to the margin of the cup. figure 7 helvella acetabulum helvella stevensii this is a relatively small fungus. the spore-bearing surface of its apothecium is ivory to pale tan at maturity. in some views mature apothecia often resemble pies with a missing wedge (fig. 8) or have three lobes, but in young apothecia the margins are rolled over the spore-bearing surface. the undersurface of the apothecium is covered with short hairs that can be seen with a hand lens. the stalk is round in outline or slightly flattened. figure 8 helvella stevensii urnula craterium (devil’s urn) the apothecium of this fungus is shaped more like a water or wine goblet than a drinking cup because of its long stalk (fig. 9). apothecia are dark brownish black overall and typically arise in clusters from, or adjacent to, downed logs. urnula craterium is generally the first fungus to appear in the spring, often well in advance of the morels. figure 9 urnula craterium 60 oklahoma native plant record volume 8, number 1, december 2008 ovrebo, c.l. sarcoscypha occidentalis (stalked scarlet cup) this cup fungus makes its first appearance in late spring and fruits throughout the summer and into the early fall. it is a small fungus with the apothecium seldom being larger than one cm across (fig. 10). the apothecium is bright red and the stipe is white. this fungus appears to fruit on the soil but is actually attached to buried wood. figure 10 sarcoscypha occidentalis oklahoma native plant record, volume 12, number 1, december 2012 oklahoma native plant record volume 12, december 2012 bruce smith https://doi.org/10.22488/okstate.17.100092 63 a comparison of the composition and structure of two oak forests in marshall and pottawatomie counties bruce a. smith mcloud high school mcloud, ok 74851 key words: forest, composition, crosstimbers, science education, veg etation structure abstract in october 2011, high school students from mcloud high school sampled an oak forest in earlsboro, pottawatomie county. in july, 2012, students in the pre-collegiate field studies camp at the university of oklahoma biological station sampled the marshall county forest at the buncombe creek camp ground, located approximately 100 miles south of the earlsboro forest and 1 mile north of the university of oklahoma biological station. one component of each botany course was to study the composition and structure of an oak forest. these 2 forests were chosen to compare because of their similarity in composition and physical distance apart. they found 10 hardwood species in the marshall county forest and 9 in the pottawatomie county forest, with 6 species common to both. quercus stellata was most important in both forests and most frequent in the pottawatomie forest where the total density was 0.141/m2. quercus stellata and ulmus alata were most frequent in the marshall county forest where the total density was 0.107/m2. introduction the best way to learn how to identify the trees, shrubs, woody vines, and herbaceous plants of a forest, is to make frequent visits and practice field identification. high school students from mcloud high school and the pre-collegiate field studies camp at the university of oklahoma biological station (uobs) did just that; they made frequent visits, but to different forests. the mcloud high school students sampled a local forest, as well as a forest near earlsboro, oklahoma. after spending time in the forests, students learned to recognize the different shades of green, shapes and colors of tree bark, growth habits, blade complexity, leaf phyllotaxy, leaf margins, leaf shapes, leaf textures, leaf odors, and even the taste of leaves of different species. by walking through the woods, i have learned the taste and effects of prickly ash– strong and bitter; numbing. i have learned the texture of hackberry leaves–scabrous and rough one way, smooth another. i have felt the barks of trees. all this i have learned by walking through the woods. cindy do mcguiness high school oklahoma city, oklahoma in october, 2011, mcloud high school students studied an oak forest near earlsboro in central pottawatomie county (35.425˚, -97.0875˚). in july, 2012, precollegiate uobs students studied an oak forest at the buncombe creek camp ground (33.52˚, -96.48˚), 100 miles south and 20 miles east of the earlsboro forest, near the biological station in marshall county. the two forests provide an oklahoma native plant record volume 12, december 2012 bruce smith 64 interesting comparison and contrast due to their similarity in composition and 100 mile north to south difference in location. students determined the composition of the forest by first learning to identify species within each of the quadrats. students then collected data that can be used in long-term ecological studies. the structure of the forest was determined by calculating density, relative density, frequency, relative frequency, basal area, relative basal area, and importance values of those trees and shrubs in the forest. by measuring relative importance and frequencies of hardwood species, rather than calculating leaf area indices or other seasonal changes, their comparison of data taken in july in marshall county to data taken in october in pottawatomie county is still valid. methods students set up eighteen 10 x 10 meter quadrats in each forest at a maximum distance from each other. this increased the likelihood of encountering a greater variety of habitats. in each quadrat, trees and shrubs were identified to species or genus, and then diameters of living woody stems 4 cm or greater at breast height (dbh) were measured. the more traditional method for measuring dbh has been to include stems 7.62 cm (3 in.) or greater (greller et al. 1979, phillippi et al.1988, rudnicky and mcdonnell 1989, stalter 1981). including stems of 4 cm or greater will include more individual woody plants and yield a more complete data set than most traditional studies. a more recent study in new york (glaeser 2006) measured dbh of woody plants that were 2 cm or greater. measuring dbh at 4 cm or greater in this study may make direct comparisons with other studies using traditional measurements problematic, but a more accurate comparison of these 2 sets of forest data is possible. with the number of student data collectors in a field class and the use of computers which can handle greater sets of data, this can be a cost-effective way to improve data collection for long-term studies. students were taught to determine density, relative density, frequency, relative frequency, basal area, and relative basal area for individual species using a simple calculator. to save time and improve accuracy, data from the forests were entered in an excel 2010 program for 18 quadrats from each forest. importance values were calculated by adding the three relative values for each species results in the marshall county forest, 10 species were identified in the 1800 m2 sampling area. in the pottawatomie forest, 9 species were found in the 1800 m2 sampling area. the 2 forests had 6 species in common: quercus stellata (post oak), q. marilandica (black jack oak), carya texana black hickory, fraxinus americana (white ash), ulmus alata (winged elm), and juniperus virginiana (eastern redcedar). u. alata had the highest density in the marshall county forest. q. stellata had the highest density in the pottawatomie forest. q. stellata and u. alata had the highest frequency in the marshall county forest. q. stellata had a frequency of 1.00, the highest frequency in the pottawatomie forest. q. stellata had the highest basal area in both forests. the 2 trees with the highest importance values respectively in both forests were q. stellata and u. alata. the total density for the pottawatomie forest was 0.141 trees/m2 and the marshall county forest was 0.107 trees/m2. the total basal area for the pottawatomie county was 21.2 cm2/m2. the total basal area for the marshall county forest was 23.3 cm2/m2. the 6 common species in both forests had a relative importance of 0.944 for the marshall county forest and 0.954 for the pottawatomie county. oklahoma native plant record volume 12, december 2012 bruce smith 65 figure 1 buncombe creek forest, marshall county, oklahoma table 1 density, frequency, basal area, and importance values for the buncombe creek forest, marshall county. species density, trees/m2 frequency basal area cm2/m2 importance value quercus stellate 0.0233 0.944 14.3 1.06 ulmus alata 0.0422 0.944 2.14 0.710 jumiperus virginiana 0.0161 0.722 1.31 0.378 quercus marilandica 0.0106 0.500 2.82 0.338 carya texana 0.00222 0.222 0.751 0.106 fraxinus americana 0.00778 0.444 1.42 0.239 morus rubra 0.00222 0.222 0.0850 0.0768 vaccinium spp. 0.000556 0.0556 0.00698 0.0186 prunus mexicana 0.000556 0.0556 0.00698 0.0187 quercus velutina 0.00111 0.111 0.461 0.0565 oklahoma native plant record volume 12, december 2012 bruce smith 66 figure 2 earlsboro forest, central pottawatomie county, oklahoma table 2 density, frequency, basal area, and importance values for the earlsboro forest, pottawatomie county. species density, trees/m2 frequency basal area cm2/m2 importance value quercus stellata 0.111 1.00 18.9 2.05 ulmus alata 0.0183 0.667 1.56 0.454 jumiperus virginiana 0.00278 0.222 0.0938 0.108 quercus marilandica 0.00278 0.222 0.233 0.114 carya texana 0.000556 0.0556 0.0109 0.0253 fraxinus americana 0.00222 0.222 0.241 0.110 amelanchier spp. 0.00111 0.111 0.0650 0.0526 celtis spp. 0.00167 0.111 0.0785 0.0572 quercus shumardii 0.000556 0.0556 0.0279 0.0261 oklahoma native plant record volume 12, december 2012 bruce smith 67 discussion the relative importance values for the 6 common species show 2 very similar forests even though they are separated by at least 100 miles. at the same time, they are very different in terms of their composition of shrubs, understory trees, vines, and herbaceous plants of the forest floor. the marshall county forest has a much denser forest floor, understory layer, and shrub layer than does the pottawatomie county forest (figures 1 and 2). another major difference in the 2 forests is the dominance of post oak in the pottawatomie forest, where quercus stellata had the highest density, frequency, basal area, and importance value. the density of post oaks in the pottawatomie forest is almost five times greater and the importance value is nearly two times greater than the post oaks in the marshall county forest even though the post oak basal area did not differ much. future studies might reveal the cause for these differences. as a part of a field learning experience, students are able to collect large data sets over a long period of time, which might otherwise be prohibitively expensive to obtain. furthermore, getting students into the field provides them with a depth of knowledge they could not possibly learn from reading a text or looking at dried specimens. while these studies provided an opportunity to begin a long-term ecological research project that involved students in field research, student identification of species in the field could be inaccurate to the point that it renders data useless. however, we found that allowing students time in the field to learn species identification (using more than a key and dried specimens) before beginning the field study, appeared to increase their accuracy. students received immediate feedback regarding the accuracy of their species identification from instructors and teaching assistants, who were in the field with them. the ecological value of this student research is that it creates baseline data for further research, to track changes in the 2 forests with possible links to changes in species due to global climate change. the greater value of this research is the invaluable experience for high school students, increasing their knowledge of nature and science aptitude by actually being in the natural environment (louv 2011). they learn more than facts. they learn how to learn from the forest. as i was walking through the forest; sun shining, elm leaves fluttering, birds flying, critters bustling, it occurred to me; mother nature teaches the purest kind of wisdom: you don’t need to be in a classroom to learn. knowledge is everywhere. magen clark and caitlyn carr mcloud high school mcloud, oklahoma beginning this long term study will also provide a beginning set of data to test hypotheses regarding how students learn in the field, versus how they learn in the lab or classroom. while i am confident that students have learned to identify trees during this project, future field studies should be accompanied by assessment of student identification skills comparing both field and laboratory experiences. this outdoor experience meets c3 pass standards 1 and 2 (oklahoma pass 2006) for general biology. acknowledgments i would like to give special thanks to the 2011-2012 mcloud high school botany class and students in the 2012 pre-collegiate field studies camp. i would also like to thank alonna price, evan smith, and jimmie manyanga for their assistance in the buncombe creek forest. thank you to austin carroll and colby brackett for oklahoma native plant record volume 12, december 2012 bruce smith 68 providing the statistical program. thank you, debbie and jay mize, for allowing us to sample your wonderful forest in earlsboro. thank you, us army corps of engineers at buncombe creek, for allowing us to study and collect samples in the forest for many years. thank you, crissy smith and the corps of engineers, for help with the gps coordinates. thank you, richard butler, for looking over my rough draft. thanks to the reviewers of this article. and finally thank you, ron tyrl. it was in the summer of 1990 that dr. tyrl shared with me how to study the composition and structure of forests using simple statistics. you cannot imagine how much mileage i got from that simple lesson. literature cited glaeser, c. w. 2006. the floristic composition and community structure of the forest park woodland, queens county, new york. urban habitats 4(1):102-126. greller, a. m. 1979. a vascular flora of the forested portion of cunningham park, queens county. new york. corrections and additions i. bulletin of the torrey botanical club 106:45. louv, richard. 2011. the nature principle. chapel hill (nc): algonquin books of chapel hill. phillippi, m. a., e. i. collins, j. l. bruner, and r. j. tyrl. 1988. succession changes in a salix nigra (salicaceae) forest in south-central oklahoma, usa. transactions of the illinois academy of science 81:61-70. oklahoma priority academic student skills, 2006 oklahoma. [cited 2012 nov]. available from: http://ok.gov/sde/sites/ok.gov.sde/file s/c3%20pass%20sci.pdf rudnicky, j. l. and mcdonnell, m. j. 1989. forty-eight years of canopy change in a hardwood-hemlock forest in new york city. bulletin of the torrey botanical club 116:52-64. stalter, r. 1981. a thirty-nine year history of the arborescent vegetation of alley park, queens county, new york. bulletin of the torrey botanical club 108:485-487. http://ok.gov/sde/sites/ok.gov.sde/files/c3%20pass%20sci.pdf http://ok.gov/sde/sites/ok.gov.sde/files/c3%20pass%20sci.pdf a comparison of the composition and structure of two oak forests in marshall and pottawatomie counties by dr. bruce smith oklahoma native plant record, volume 14, number 1, december 2014 oklahoma native plant record 43 volume 14, december 2014 stanley a. rice and sonya l. ross https://doi.org/10.22488/okstate.17.100104 oklahoma deciduous trees differ in chilling enhancement of budburst stanley a. rice sonya l. ross department of biological science physical plant department southeastern oklahoma state university southeastern oklahoma state university durant, ok 74701 durant, ok 74701 srice@se.edu sross76@se.edu keywords: climate change, phenology abstract in many tree species, winter chilling accelerates budburst in response to spring warmth. global climate change has already accelerated budburst in deciduous tree species around the world. but as global climate change leads to milder winters, trees species also experience less chilling, which may actually delay spring budburst in some species. we hypothesized that reduced duration of winter chilling would delay spring budburst in sycamore (platanus occidentalis) and pecan (carya illinoinensis), but would not delay it in sweetgum (liquidambar styraciflua). we tested this hypothesis experimentally by manipulating the number of weeks of chilling from 0 to 6 weeks. lack of winter chilling did not delay budburst in sweetgum but did delay it in sycamore and pecan, in agreement with the hypothesis. mild winters in oklahoma may eventually favor the growth of sweetgums at the expense of sycamores and pecans. introduction earlier spring budburst in deciduous trees is widely recognized as one of the consequences of global climate change. it has been occurring for the last century and a half and has continued in recent decades (schwartz et al. 2006; ibañez et al. 2010; polgar and primack 2011). this conclusion is based upon several sources of information: comparison of recent with historical budburst dates, including the records of henry david thoreau at walden pond, and comparisons of recent with historical herbarium specimens and photographs (primack et al. 2004; millerrushing et al. 2006; primack 2014); satellite imagery during recent decades (liang et al. 2011); yearly records of individual woody plants during recent decades (schwartz 1994; rice and schwartz, in prep.); and functional models (e.g. morin et al. 2009). the first author of this paper has maintained an ongoing record of budburst times for about 400 individuals of 22 deciduous tree species in durant, oklahoma, starting in 2006. by observing each tree at least weekly, and usually more often, the first author determined budburst date for each individual using a protocol similar to that of the globe program (globe.gov 2014). the data clearly indicate earlier budburst during the nine-year period, particularly from 2008-2012, during which time several tree species advanced their budburst time about two days per year. this did not occur in all species, however. in particular, budburst did not change in american elm (ulmus americana l.) and became later each year in silver maple (acer saccharinum l.), probably in association with summer drought and heat damage that either directly, or indirectly through pathogens, killed many of these trees (rice and schwartz, in prep). 44 oklahoma native plant record volume 14, december 2014 stanley a. rice and sonya l. ross it is, however, invalid to extrapolate the trend toward earlier budburst for most tree species, because many woody species benefit from chilling for timely budburst (schwartz and hanes 2010). chilling induces the development of structures within buds and/or alters the concentration of plant growth substances such as cytokinins (hewett and wareing 1973), a process sometimes called vernalization. if winters in some areas (such as southern oklahoma) become brief and warm, the buds of some species may experience insufficient chilling and therefore reduced fitness (luedeling et al. 2011). some tree species also require a minimum daylength for budburst (e.g., heide 1993a). in general, we would expect tree species that open their buds early in the spring to have floral and vegetative structures already well-formed within the buds prior to winter, while these structures may have to develop during the winter in tree species that open their buds later in the spring. the tree species in the latter group may require chilling to initiate and complete the process of bud development. we therefore hypothesized that tree species that open their buds early in the spring do not have as much chilling enhancement of budburst as tree species that open their buds later in the spring. specifically, we expected a negative association between time of budburst and chilling enhancement. we used three species to test this hypothesis in oklahoma: sweetgum (liquidambar styraciflua l., altiginaceae), which opens its buds earliest of these three species, often in february; sycamore (platanus occidentalis l., platanaceae), which opens its buds later, often in march; and pecan (carya illinoinensis (wangenh.) k. koch, juglandaceae), which opens its buds last of these three species, often in april. numerous studies have examined the effect of chilling on budburst, but most of these studies have been conducted at higher latitudes (e.g., hunter and lechowicz 1992; heide 1993b; chuine 2000). we wanted to test the hypothesis using oklahoma trees, which may differ genetically from trees of the same species that live in other locations. for example, research in other parts of the world show that trees such as pecans (kuden et al. 2013) have a chilling enhancement of budburst, but we cannot conclude from this that oklahoma trees of these species have a similar chilling enhancement. methods we selected five individual trees at least 10 meters in height that are in the long-term data set from each of the three species. all were in parks or along streets in durant, oklahoma (fig. 1). we originally also included post oak (quercus stellata wangenh.), but mortality of twigs during the experiment reduced the sample size to only one twig in two of the treatments. from each tree, we obtained six twigs with intact terminal and axillary buds, two for each of the three chilling treatments described below, resulting in 30 twigs for each species (total of 90 twigs). we gathered twigs on 18 november 2013, after leaf senescence was well advanced but before the first frost. we labeled all twigs with masking tape. for each tree, we placed two twigs in a plastic food storage container with wet paper towels and stored them in a refrigerator at about 10º c for three weeks, and we stored two other twigs for six weeks. six weeks is considerably less than the average of approximately 18 weeks between first (about november 9) and last (about march 28) frost during the previous four decades in durant, oklahoma (fig. 2). at the end of chilling, we removed the twigs and placed them in warm conditions to allow budburst to begin. we also placed two twigs from each tree immediately into warm conditions (0 weeks). these unchilled twigs were the control. the warm conditions were in a temperature-controlled laboratory oklahoma native plant record 45 volume 14, december 2014 stanley a. rice and sonya l. ross (about 25º c) under continuous fluorescent illumination. once we exposed the buds to warm conditions, we kept the cut ends of the twigs continuously submerged in water. we checked each twig at least twice a week for signs of budburst, defined as green tissue showing through separated bud scales (fig. 3). we also changed the water and cleansed the cut ends of the twigs with a brush to prevent decomposers, living off of sap, from blocking the xylem. eighteen of the original 90 twigs failed to burst their buds during this experiment and were presumed dead. figure 1 the general habitat of the trees used in this study in november 2013 46 oklahoma native plant record volume 14, december 2014 stanley a. rice and sonya l. ross figure 2 average first and last freeze dates in oklahoma, 1961-2010 average. maps modified and used with permission from oklahoma climatological survey (http://climate.ok.gov/index.php/climate). figure 3 buds of sweetgum (liquidambar styraciflua) at various stages of budburst. the top bud has not yet opened, and the bud at the lower right is just beginning to open. durant oklahoma native plant record 47 volume 14, december 2014 stanley a. rice and sonya l. ross results and discussion results shown in the following table indicate that chilling greatly reduced the budburst time in sycamore (p < 0.001) and pecan (p = 0.012) but not in sweetgum (p = 0.089). because the data distribution was skewed toward early budburst dates, we used separate kruskal-wallis analyses for each species to obtain these values (ibm spss 2011). all three species burst their buds quickly following six weeks of chilling. budburst of unchilled sycamore and pecan buds were significantly delayed while unchilled sweetgum buds burst quickly after exposure to warm temperatures. table mean number of days of exposure to warmth that induced budburst in three species of trees in southeast oklahoma. values in parentheses indicate the number of twigs that were exposed to different levels of the treatment (= chilling). asterisk (*) indicates significant difference using kruskal-wallis test at p < 0.05. weeks of chilling species zero three six p-value liquidambar styraciflua 26.3 (7) 29.8 (9) 24.7 (6) 0.089 platanus occidentalis 63.0 (6) 38.0 (10) 21.3 (9) 0.001* carya illinoinensis 53.3 (6) 32.6 (10) 23.4 (9) 0.012* discussion the data confirmed the association between time of budburst and chilling enhancement, based on oklahoma specimens of three tree species. we would expect sweetgum to respond the most to warmer winters. gunderson et al. (2012) reported that experimentally-imposed warmer temperatures caused earlier budburst in sweetgum than in three other tree species, consistent with our results. global climate change has been associated not only with earlier spring budburst in deciduous trees but also earlier flowering in spring wildflowers and earlier spring activity in many kinds of animals (miller-rushing et al. 2008; willis et al. 2008; primack 2014). the extent to which different species respond to global climate change may alter the species makeup of an ecological community (miller-rushing and primack 2008). if future global climate change should cause forests in the southern united states, such as those in southern oklahoma, to have very mild winters, not all deciduous tree species will continue their trend toward earlier budburst. instead, some tree species—such as the sycamores and pecans in this study—may reverse their trend toward earlier budburst and instead have later budburst. flexibility of phenological response appears to be an important contributor to survival in a world of global climate change. the eventual loss of chilling temperatures may alter the relative growth patterns of deciduous tree species in oklahoma. 48 oklahoma native plant record volume 14, december 2014 stanley a. rice and sonya l. ross acknowledgement the authors would like to thank an anonymous reviewer for helpful comments and suggestions for this manuscript. references chuine, i. 2000. a unified model for budburst of trees. journal of theoretical biology 207(3):337-347. globe.gov 2014. budburst protocol. http://www.globe.gov/documents/356 823/2538681/earth_prot_budburst.pdf. accessed 19 june 2014. gunderson, c.a., n.t. edwards, a. v. walker, k.h. o’hara, c.m. campion, and p.j. hanson. 2012. forest phenology and a warmer climate— growing season extension in relation to climatic provenance. global change biology 18(6):2008-2025. heide, o.m. 1993a. dormancy release in beech buds (fagus sylvatica) requires both chilling and long days. physiologica plantarum 89(1):187-191. heide, o.m. 1993b. daylength and thermal time responses of budburst during dormancy release in some northern deciduous trees. physiologia plantarum 88(4):531-540. hewett, p.f. and e.w. wareing. 1973. cytokinins in populus×robusta: changes during chilling and bud burst. physiologia plantarum 28(3):393-399. hunter, a.f. and m.j. lechowicz. 1992. predicting the timing of budburst in temperate trees. journal of applied ecology 29(3):597-604. ibáñez, i., r.b. primack, a.j. millerrushing, e. ellwood, h. higuchi, s.d. lee, h. kobori, and j.a. silander. 2010. forecasting phenology under global warming. philosophical transactions of the royal society b365:3247-3260. ibm corp. released 2011. ibm spss statistics for windows, version 20.0. armonk, ny: ibm corp. kuden, a.b., o. tuzcu, s. bayazit, b. yildirim, and b. imrak. 2013. studies on the chilling requirement of pecan nut (carya illinoensis koch) cultivars. african journal of agricultural research 8(24):31593165. liang, l., m.d. schwartz, and s. fei. 2011. validating satellite phenology through intensive ground observation and landscape scaling in a mixed seasonal forest. remote sensing of environment 115:143-157. luedeling, e., e.h. girvetz, m.a. semenov, and p.h. brown. 2011. climate change affects winter chill for temperate fruit and nut trees. plos one 6(5):e20155. miller-rushing, a.j., r.b. primack, d. primack, and s. mukunda. 2006. photographs and herbarium specimens as tools to document phenological changes in response to global warming. american journal of botany 93(11):16671674. miller-rushing, a.j. and r. b. primack. 2008. global warming and flowering times in thoreau’s concord: a community perspective. ecology 89(2):332-341. miller-rushing, a.j., t.l. lloyd-evans, r.b. primack, and p. satzinger. 2008. bird migration times, climate change, and changing population sizes. global change biology 14(9):1959-1972. morin, x., m.j. lechowicz, c. augspurger, j. o’keefe, d. viner, and i. chiune. 2009. leaf phenology in 22 north american tree species during the 21st century. global change biology 15(4):961975. polgar, c.a. and r.b. primack. 2011. leafout phenology of temperate woody plants: from trees to ecosystems. new phytologist :doi: 10.1111/j.14698137.2011.03803.x. primack, r.b. 2014. walden warming: climate change comes to thoreau’s woods. [chicago]: university of chicago press. oklahoma native plant record 49 volume 14, december 2014 stanley a. rice and sonya l. ross primack, d., c. imbres, r.b. primack, a.j. miller-rushing, and p. del tredici. 2004. herbarium specimens demonstrate earlier flowering times in response to warming in boston. american journal of botany 91(8):12601264. schwartz, m.d. 1994. monitoring global change with phenology: the case of the spring green wave. international journal of biometeorology 38(1):18-22. schwartz, m.d., r. ahas, and a. aasa. 2006. onset of spring starting earlier across the northern hemisphere. global change biology 12:343-351. schwartz, m.d. and j.m. hanes. 2010. continental scale phenology: warming and chilling. international journal of climatology 30:1595-1598. willis, c.g., b. ruhfel, r.b. primack, a.j. miller-rushing, and c.c. davis. 2008. phylogenetic patterns of species loss in thoreau's woods are driven by climate change. proceedings of the national academy of sciences usa 105(44):17,029-17,033. . oklahoma deciduous trees differ in chilling enhancement of budburst by dr. stanley rice and ms. sonya ross oklahoma native plant record, volume 15, number 1, december 2015 96 oklahoma native plant record volume 15, december 2015 marli claytor and karen r. hickman https://doi.org/10.22488/okstate.17.100116 kudzu, pueraria montana (lour.) merr. abundance and distribution in oklahoma marli claytor karen r. hickman natural resource ecology and management oklahoma state university 008c ag hall stillwater, ok 74078 karen.hickman@okstate.edu key words: invasion, invasive species, mapping, federal noxious weed abstract invasive species are a growing problem in the united states, and kudzu (pueraria montana) (lour.) merr. is one of the most well documented invaders of southeastern states. documenting the invasion of kudzu in oklahoma, however, has not been a targeted focus in previous studies; thus, maps of its occurrence differ among sources. our primary objective was to locate and confirm the presence of kudzu throughout oklahoma. specifically, we attempted to confirm previously recorded populations of kudzu and estimate the extent of the invasion at those sites. in addition, we wanted to locate stands of kudzu within oklahoma that had not been recorded and to assess the extent of invasion. a survey was sent to state and county officials to acquire information on locations and general knowledge of kudzu. points of occurrence and estimated extent of invasion in hectares were then placed in arcmap programming to create a consolidated map of kudzu. samples were collected, pressed, and placed in the university of oklahoma’s bebb herbarium (okl). we determined the majority of kudzu locations are in the southeastern portion of the state and total a minimum of 32.4 hectares. results of the survey indicated half of the respondents polled were unaware of kudzu’s presence in the state. introduction invasive species are a growing concern in the united states, as well as across the globe. there are approximately 17,000 native species of vascular plants in the u. s., compared to a continually increasing estimate of 6,000 nonnative species (forseth and innis 2004). invasive species can be detrimental to the environments they occupy and cause major ecosystem changes (mitich 2000). kudzu, pueraria montana (lour.) merr. (fabaceae) is an introduced, leguminous vine which causes major changes in areas in which it invades. kudzu is listed as one of the world’s 100 worst invasive species of all time (sage et al. 2009). first introduced at the 1876 centennial exposition in pennsylvania, kudzu has since made a lasting impact on the southeastern u. s. (brown 2010). upon introduction, the vine was sold to the public to aid with soil erosion control and as forage for livestock; additionally, the soil conservation service (currently natural resources conservation service) and other national agencies encouraged the planting of kudzu (forseth and innis 2004). eventually, evidence indicated that the vine overtopped mature trees, took over native plant dominated areas, buildings, and disturbed areas, and became a financial burden to those who mailto:karen.hickman@okstate.edu oklahoma native plant record 97 volume 15, december 2015 marli claytor and karen r. hickman tried to control and eradicate the invader. kudzu has been found to alter a landscape abruptly as it can grow up to 30 cm a day and between 10 to 30 m in one growing season (mitich 2000). additionally, kudzu fixes nitrogen and releases isoprene into the environment, which can create pollution in the atmosphere, further reducing environmental value (hickman et al. 2010). kudzu is one of the worst invasive species in the u. s. and is continuing its spread across the country (fig. 1). it has been estimated that the vine covers 2.83 million hectares in the southeast, in 1955 was declared a weed by the u. s. department of agriculture (alderman 1998) and declared a federal noxious weed in 1999 (mitich 2000). kudzu has a wide climatic range which facilitates its ability to continue spreading northward (mitich 2000). it has been suggested that kudzu is limited in its range by annual rainfall, which needs to be a minimum of 100 cm a year (mitich 2000). the vine is also considered to be limited in its distribution by lack of hardiness; however, it has exceeded many expert predictions in range expansion (mitich 2000). this area includes oklahoma, which was once believed to be unsuitable habitat for kudzu (mitich 2000). figure 1 distribution map of kudzu across the united states, in the usda plants database http://plants.usda.gov/core/profile?symbol=pumo kudzu is present in oklahoma but has not been the focus of a targeted survey in the state. because of the variability in data, maps illustrating the distribution of kudzu are inconsistent among sources (e.g., state and national agencies). thus, a need for an updated map has arisen for future management of the species. for instance, the oklahoma vascular plant database map, whose data are based on herbarium records, indicates 22 counties with kudzu (fig. 2), while a map from early detection and distribution mapping systems (eddmaps) includes 12 counties http://plants.usda.gov/core/profile?symbol=pumo http://plants.usda.gov/core/profile?symbol=pumo 98 oklahoma native plant record volume 15, december 2015 marli claytor and karen r. hickman (oklahoma vascular plant database 2014; eddmaps 2014). while some of the occurrences overlap, there are some inconsistencies. importantly, none of these maps are based on a compilation of reliable field observations and specimens that have been critically examined by experts. thus, we attempted to confirm previously known locations, obtain information about new sightings, and collect specimens for confirmation. a survey was utilized to cover oklahoma as a whole and to gather as much information as possible about the plant from knowledgeable persons primarily within the oklahoma state university extension service. surveys have been found to be a useful tool when other forms of data sources or collection methods are not adequate, and in this case it was not practical to reach as many people through other methods (innovation insights 2006). survey reports were then confirmed by groundtruthing and utilized to create a detailed map of kudzu locations and the extent of invasion at each site. figure 2 oklahoma vascular plant database map of kudzu occurrence by county http://www.oklahomaplantdatabase.org methods kudzu location, extent of invasion, and date of record were obtained from available records, which included the ovpd records of herbarium specimens, information collected by the oklahoma invasive plant council (okipc; k. hickman, unpublished), directed contact with botanists in the state, and through a survey sent to osu extension personnel, land managers known to have experience with kudzu, oklahoma department of wildlife conservation employees, and the okipc. the survey provided the majority of data collected. a link to the kudzu survey, which was created through survey monkey (www.surveymonkey.com), was sent out through email. five questions were asked regarding the respondent’s knowledge of kudzu and its presence in oklahoma. http://www.oklahomaplantdatabase.org/ http://www.oklahomaplantdatabase.org/ http://www.surveymonkey.com/ oklahoma native plant record 99 volume 15, december 2015 marli claytor and karen r. hickman questions asked in the survey included: 1) what county of oklahoma are you currently working or residing in? 2) have you seen or heard of kudzu inhabiting land in oklahoma? 3) if so, please provide the locations of the kudzu sightings. 4) in acres, how large of an area would you estimate that the infestation is at each site? 5) please provide contact information for verification and/or additional inquiries. approximately two hundred invitations were emailed to osu county extension offices, oklahoma department of wildlife conservation, oklahoma department of transportation, and the oklahoma invasive plant council members. these agencies and organizations were chosen based on previous experience we have had with them concerning invasive species and the ability to send mass emails to the group. also, individuals were included who had knowledge of oklahoma vegetation and ecosystems, or who dealt with invasive species frequently. we used arcgis arcmap v. 10.1 (esri, redlands, ca) software to create distribution maps. a state overview illustrating counties with kudzu present was created, along with more detailed maps of the counties displaying extent of the invasion of kudzu. estimates of the extent invaded were made on sites (19), approximated from googleearth imagery (4), or reported in the surveys (5). points were added to the map for individual stands of kudzu across oklahoma, illustrating area invaded for each location within the county. for our map, we included sites that were confirmed to have kudzu; we did not include locations of kudzu that we visited and confirmed kudzu was not present. mapped points (table 1) only include confirmed locations of kudzu, but not sites in ovpd that were not confirmed via a visit or sites visited where no plants were found. samples of kudzu were collected from all confirmed sites visited (16) to create herbarium voucher specimens. we traveled to some, but not all of the locations, due to time constraints of the project (see table 1). sites chosen to visit were those with larger infestations reported or those reported in the survey. samples of individual plants were cut in sections including leaves, flowers, and pods (if available, as samples were taken throughout the project year). specimens were deposited at the university of oklahoma’s bebb herbarium (okl). results the survey received 52 responses from the approximately 200 emails sent, which indicates a return rate of close to 25%. of those, over 50% (28) respondents had knowledge of kudzu in oklahoma, while 46% (24) reported having not seen or heard of the vine’s encroachment within the state. of those surveyed, 17 provided locations, and 10 estimated dimensions of the area invaded of kudzu. of those reported, 9 locations were new, previously unrecorded sites of kudzu. maps (figs. 3, 4) were created using data from the survey and previously known locations (confirmed by groundtruthing) of kudzu (see table 1). if kudzu was confirmed as absent from a site, then it was removed from the map. a gray scale was utilized to illustrate the extent of invasion of kudzu in each county. figure 3 presents specific locations of kudzu in the state along with their corresponding extent of invasion, while figure 4 illustrates presence and extent by county. based on our results, at least 32.4 hectares of land are invaded by kudzu in oklahoma across 28 sites. 100 oklahoma native plant record volume 15, december 2015 marli claytor and karen r. hickman table 1 locations of kudzu identified from previous documentation (oklahoma vascular plants database), survey results, or on-site discoveries. kudzu was confirmed present or absent via site visits or previous documentation. estimates of the extent of kudzu invasion were obtained during on-site visits using gps or googleearth imagery. site name longitude latitude source of location data source of extent of invasion status of kudzu on site idabel -94.709 33.896 discovered by marli claytor google earth to estimate coverage confirmed present claremore -95.599 36.299 from survey site visit confirmed present antlers -95.637 34.233 discovered by marli claytor google earth to estimate coverage confirmed present p st. & springdale rd. , ardmore -97.108 34.159 previous documentation site visit confirmed present marsden rd. love co. -97.195 34.070 site visit site visit confirmed present tater hill rd. ardmore -97.008 34.144 previous documentation site visit confirmed present shawnee -96.962 35.333 previous documentation /survey google earth to estimate coverage confirmed present haskell -95.611 35.754 from survey google earth to estimate coverage confirmed present eufaula -95.339 35.281 previous documentation site visit confirmed present cleveland county -97.164 35.233 previous documentation /survey site visit confirmed present dickson -96.928 34.188 from survey from survey inconclusive red river -95.500 33.877 previous documentation unavailable inconclusive oklahoma native plant record 101 volume 15, december 2015 marli claytor and karen r. hickman untitled placemarkhulbert -95.226 35.869 previous documentation site visit confirmed present shoals -95.398 33.968 previous documentation site visit inconclusive north eufuala -95.387 35.391 previous documentation site visit confirmed present norman -97.156 35.232 previous documentation / from survey site visit confirmed present okemah -97.399 35.430 previous documentation site visit confirmed present washita river tributary -97.510 34.779 previous documentation site visit confirmed present fittstown -96.635 34.618 previous documentation site visit confirmed present durant -96.410 34.056 previous documentation site visit confirmed present duncan -97.986 34.594 previous documentation site visit confirmed present stillwater -97.063 36.113 previous documentation site visit confirmed present osage -96.304 36.242 from survey from survey confirmed present osage -96.282 36.246 from survey from survey confirmed present adair from survey unavailable inconclusive caddo -98.324 35.464 previous documentation confirmed absent marshall -96.685 34.148 previous documentation confirmed absent pontotoc -96.634 34.579 previous documentation confirmed absent 102 oklahoma native plant record volume 15, december 2015 marli claytor and karen r. hickman figure 3 distribution map of counties with confirmed kudzu invasion, showing acres invaded. acres represent total acres for all sites within each county. map created using arcmap. figure 4 locations of kudzu across the state of oklahoma, featuring area invaded for each site. map was created using arcmap programming with data from the survey. oklahoma native plant record 103 volume 15, december 2015 marli claytor and karen r. hickman discussion the survey was successful in acquiring important information on kudzu throughout the state. nearly half of the respondents had no knowledge of kudzu being present in the state, which indicates very little familiarity with the vine even from knowledgeable professionals. close to 30% of all sightings reported were new locations in the state. this prompts the question: if we had sent out more surveys, how many more new locations would have been documented? the new distribution map aids in assessing current and future invasion of kudzu. in comparison to the ovpd, our map includes 22 counties reported while the other has only 20; additionally, not all ovpd counties are included in the new map as some reports could not be confirmed or old populations were found to no longer exist as determined through our site visits (see figs. 2, 3). it can be observed that kudzu currently exists primarily in the eastern portion of the state. climatic restrictions are most likely limiting the range of kudzu (jarnevich and stohlgren 2009. once kudzu has invaded an ecosystem it is very difficult to eradicate, further facilitating its spread across oklahoma. it is likely that kudzu will continue not only its coverage north, but also invade more hectares where stands currently persist (jarnevich and stohlgren 2009). currently there are at least 32.4 hectares invaded with kudzu in oklahoma, which is extremely small in comparison to the total seven million hectares invaded in the united states (eskridge and alderman 2010). this does not mean we can ignore the problem, but presents our state with an opportunity to stop a problem while we can. if our state began an early detection and rapid response (edrr) program for kudzu, it would be possible to limit the future spread of the vine and keep our state and economy safe from the detriment of invasion. edrr programs work to develop a system of effectively addressing issues of invasive species through the steps of: early detection and reporting of new plants, identification and collection of specimens, verification of new plant records, archival of new records where appropriate, rapid assessment of new records, and rapid response to new records determined to be invasive (westbrooks 2004). to stop this problem now would save the state financially in the long run. more studies need to be conducted on kudzu, and there is a current study on viability of kudzu seeds in oklahoma (zoeller and hickman, unpublished). this study will be crucial in estimating to what extremes kudzu can further invade oklahoma. education for the state needs to occur to stop the further expansion of kudzu. the creation of our updated map will aid in educating citizens on where the vine resides and if they should be on alert for presence in their area. to inform the public, the first step will be to train county and state officials to properly identify kudzu and instruct citizens on how to handle the issue. kudzu has caused major damage in the southeastern united states, but this destruction can be reduced through proper education and effectively implementing an edrr program. acknowledgements we would like to thank those who participated in the survey or contributed a location. thanks to oklahoma state university natural resource ecology and management department and also the oklahoma state university honor’s college. finally, we would like to thank the others who helped complete this project: 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to make a major ecosystem threat. critical reviews in plant sciences 23:401–413. hickman, j.e., s. wu, l.j. mickley, and m.t. lerdau. 2010. kudzu (pueraria montana) invasion doubles emissions of nitric oxide and increases ozone pollution. proceedings of the national academy of sciences, usa 107:10115– 10119. innovation insights. 2006. using surveys for data collection in continuous improvement. http://www.opia.psu.edu/sites/default/ files/insights014.pdf (7 august 2014). jarnevich, c. and t. stohlgreen. 2009. near term climate projections for invasive species distributions. biological invasions 11:1373–1379. mitich, l. 2000. kudzu [pueraria lobata (willd.) ohwi]. weed technology 14:231– 235. oklahoma vascular plant database. 2014. distribution map of pueraria montana (kudzu). http://www.oklahomaplantdatabase.org (29 november 2014). sage, r.f., h.a. coiner, d.a. way, g.b. runion, s.a. prior, h.a. torbert, r. sicher, and l. ziska. 2009. kudzu [pueraria montana (lour.) merr. variety lobata]: a new source of carbohydrate for bioethanol production. biomass and bioenergy 33:57–61. usda. nrcs. 2014. the plants database. greensboro (nc): national plant data team. http://plants.usda.gov/core/profile?sy mbol=pumo (17 oct 2014). waldron, g.e. and b.m.h. larson. 2012. kudzu vine, pueraria montana, adventive in southern ontario. the canadian fieldnaturalist 126:31–33. westbrooks, r.g. 2004. new approaches for early detection and rapid response to invasive plants in the united states. weed technology 18:1468–1471. http://www.bloomberg.com/news/articles/2014-07-25/kudzu-that-ate-u-s-south-heads-north-as-climate-changes http://www.bloomberg.com/news/articles/2014-07-25/kudzu-that-ate-u-s-south-heads-north-as-climate-changes http://www.bloomberg.com/news/articles/2014-07-25/kudzu-that-ate-u-s-south-heads-north-as-climate-changes http://www.eddmaps.org/distribution/usstate.cfm?sub=2425%20 http://www.eddmaps.org/distribution/usstate.cfm?sub=2425%20 http://www.opia.psu.edu/sites/default/files/insights014.pdf http://www.opia.psu.edu/sites/default/files/insights014.pdf http://www.oklahomaplantdatabase.org/ http://plants.usda.gov/core/profile?symbol=pumo%20 http://plants.usda.gov/core/profile?symbol=pumo%20 kudzu, pueraria montana (lour.) merr. abundance and distribution in oklahoma by ms. marli claytor and dr. karen r. hickman 2 oklahoma native plant record volume 1, number 1, december 2001 foreword the oklahoma biological survey is in the process of making an inventory of the plant specimens that have made their way into the herbaria housed at our universities. they will eventually make that information available on the world wide web. all kinds of information will be available, electronically, from those dried specimens. they are a priceless treasure, recording our past and the efforts made to understand it. putting that data on the web will be a way of making it accessible to people who have no physical access to the herbaria, and little time to extract it. other kinds of plant information are stored in the minds of our members and scientists. possibly, the files stored in computers will outlast them, maybe not. who knows? one thing we do know: people have been interested in the flora of oklahoma for more than a hundred years. some of their observations have been recorded but, for the most part, not in published form. believing that those records are important for the understanding of our current flora, the oklahoma native plant society has determined to bring some of those records to your eye in a more durable form. for many years, dr. u. t. waterfall’s keys to the flora of oklahoma has been the only statewide source of identification keys. few know that his first attempt to catalog the plants of oklahoma was a master’s thesis that includes a list of the plants he found in oklahoma county in the 1930’s. what a difference there is in oklahoma county between that time and this! to put that survey into perspective, we are including a working copy of the biological survey’s list for oklahoma county, made in the year 2000. this journal intends to publish in each issue, a previously unpublished historic study, which may serve as a baseline for your own investigations into your local flora. we will also include recent studies, student papers, current plant lists, and essays of permanent value. in the future, we hope to be able to publish either little’s or bebb’s catalog of plants of muskogee county, and other such lists as may be discovered. for that purpose, we challenge the members of oklahoma native plant society to offer records and observations which they may have available. not that there will ever be a complete record of the life in any one place, because life refuses to be reduced to a score-card, but because we believe that a knowledge of the plants of a specific location is a fundamental part of understanding other life that may be found there, including our own. you, our readers, will know of current or historic information that should be included in future issues. we hope you will share those with us. patricia folley, president oklahoma native plant society april, 2001 oklahoma native plant record, volume 3, number 1, december 2003 oklahoma native plant record 3 volume 3, number 1, december 2003 foreword over the years our state’s ecology has been affected by many changes. new dams, turnpikes, urban sprawl, forestry practices, highway right of away mowing, and excessive use of herbicides are just a few of the activities that have had a profound impact upon our state’s native vegetation, many within my lifetime. concerned oklahomans founded the oklahoma native plant society (onps) in 1987 with the goal of encouraging the study, protection, use, and appreciation of our native plants. many of our citizens are unaware of the unique geographic and biological characteristics of oklahoma. botanists and ecologists have debated where oklahoma should be placed on the biodiversity scale. states such as california, texas, north carolina, and florida have more species. california and texas have more bioregions. when you consider our geographic location, numbers of species, and bioregions, oklahoma is considered by many to be number three in terms of biodiversity. many western and eastern, as well as northern and southern species intermingle here. our extensive river system, moving primarily from the northwest to the southeast, further divides the state into unique regions. the sand dunes and the great salt plains are classic examples. early explorers and botanists made extensive use of these waterways, washington irving and thomas nuttall are two of the most prominent. the state’s elevation and annual rainfall amounts change as you move from west to east, resulting in different bioregions across our state. as a youngster growing up in oklahoma city, i would often spend time with a map of oklahoma and wonder about the regions in our state. black mesa, the high plains prairies, the forests of eastern oklahoma, and the ozark, quachita, wichita, quartz, and arbuckle mountain regions were some of the areas i would dream of someday visiting. eventually, through family vacations, ecology field trips in college, and my association with onps, i was able to visit many of these places. our state is truly diverse. much of my appreciation of this is due to my associations with onps and as a student in harriet barclay’s ecology class. the new field guide to oklahoma plants, by ron tyrl, terrence bidwell, and ronald masters has an excellent introductory section covering the ecogeography and vegetation of oklahoma. this guide has fine maps and explanations on the geography, soils, and vegetation types. rolling hills, plains, and unique mountain regions characterize the state. six forest types are recognized: oak-hickory, oak-pine, post oak-blackjack oak, loblolly pine, cypress bottoms, and bottom land (flood plain). dr. david stahle, from the tree-ring laboratory at university of arkansas, states that the cross-timbers region (post oak-blackjack oak ) in oklahoma is one of the largest remaining old growth forests in north america. our state also consists of tallgrass, shortgrass, and mixed grass prairie regions. the field guide recognizes five different shrubgrassland types; sandsage grassland, mesquite grassland, shinnery oak–grassland, stabilized dunes, and pinon-juniper mesa. the oklahoma native plant record has covered some of these areas in past issues. it is becoming an excellent forum for discussing our state’s unique diversity. much of the journal’s success goes to those original onps founding members, whose foresight and concerns about our state have been an inspiration to us all. this journal is a monument to their efforts. i encourage everyone to support and contribute to its success. james elder onps president june 2003 oklahoma native plant record, volume 17, number 1, december 2017 1 oklahoma native plant r ecord journal of the okla hom a native plant society p. o. box 14274 tulsa, oklahoma 74159-1274 volume 17, december 2017 issn 1536-7738 http://ojs.library.okstate.edu/osu/ co-editors: sheila strawn and gloria caddell production editor: paula shryock electronic production editor: sandy graue manuscript editor: mark fishbein technical advisor: erica corbett the purpose of onps is to encourage the study, protection, propagation, appreciation, and use of the native plants of oklahoma. membership in onps is open to any person who supports the aims of the society. onps offers individual, student, family, and life memberships. officers and board members president: bruce smith vice-president: bill farris secretary: connie murray treasurer: mary korthase board members: barbara klein mary gard kathy supernaw mary-helen hagge derek mccall alyssa whiteman chapter chairs: central: patrick bell cross timbers: elaine lynch northeast: lynn michael historian: fran stallings publicity/merchandise chair: alicia nelson conservation chair: chadwick cox tulsa garden club liaison: sue amstutz awards chair: sue amstutz membership database: tina julich photo contest chair: lynn michael mailings/printings chair: sandy graue color oklahoma chair: pearl garrison gaillardia editor: marilyn stewart website manager: adam ryburn http://www.oknativeplants.org cover photo: nelumbo lutea willd. (american lotus) by sally webb for the 2016 onps photo contest articles (c) the authors journal compilation (c) oklahoma native plant society except where otherwise noted, this work is licensed under a creative commons attributionnoncommercial-sharealike4.0 international license, https://creativecommons.org/licenses/ by-nc-sa/4.0/, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly attributed, not used for commercial purposes, and, if transformed, the resulting work is redistributed under the same or similar license to this one. https://doi.org/10.22488/okstate.18.100001 http://ojs.library.okstate.edu/osu/ http://www.oknativeplants.org/ 2 oklahoma native plant record volume 17, december 2017 oklahoma native plant record volum e 17 table of contents foreword .................................................................................................................................................... 3 a study of the flowering plants of tulsa county, oklahoma, exclusive of the grasses, sedges, and rushes. m. s. thesis ...................................................................................... 4 maxine b. clark† laboratory studies of allelopathic effects of juniperus virginiana l. on five species of native plants ................................................................................................................ 37 erica a. corbett and andrea lashley vascular flora of e. c. hafer park, edmond, oklahoma ................................................................ 53 gloria m. caddell, katie christoffel, carmen esqueda, and alonna smith first record of chorioactis geaster from oklahoma .............................................................................. 69 clark l. ovrebo and sheila brandon critic’s choice essay: allelopathy .............................................................................................................. 72 paul buck† editorial policies and procedures ......................................................................................................... 74 five year index to oklahoma native plant record ............................................... inside back cover † indicates an author who is deceased oklahoma native plant record, journal of the oklahoma native plant society, volume 17, december 2017 title page table of contents foreword journal of the oklahoma native plant society, volume 7, number 1, december 2007 1 oklahoma native plant record journal of the oklahoma native plant society 2435 south peoria tulsa, oklahoma 74114 volume 7, number 1, december 2007 issn 1536-7738 managing editor: sheila strawn technical editor: patricia folley technical advisor: bruce hoagland cd-rom producer: chadwick cox website: http://www.usao.edu/~onps/ the purpose of onps is to encourage the study, protection, propagation, appreciation and use of the native plants of oklahoma. membership in onps shall be open to any person who supports the aims of the society. onps offers individual, student, family, and life memberships. 2007 officers and board members president: kim shannon vice-president: gloria caddell secretary: paula shryock treasurer: mary korthase membership database: tina julich past president: constance murray board members: paul buck ron tyrl lynn michael monica macklin elfriede miller stanley rice central chapter chair: lou duke/ marilyn stewart cross-timbers chapter chair: paul richardson mycology chapter chair: clark ovrebo northeast chapter chair: sue amstutz gaillardia editor: chadwick cox harriet barclay award chair: constance taylor ann long award chair: patricia folley onps service award chair: sue amstutz historian: sharon mccain librarian: bonnie winchester website manager: chadwick cox photo poster curators: sue amstutz & marilyn stewart color oklahoma chair: tina julich conservation chair: chadwick cox field trip chair: patricia folley mailings chair: karen haworth merchandise chair: susan chambers nominating chair: paula shryock photography contest chair: tina julich publications chair: sheila strawn publicity chairs: kim shannon & marilyn stewart wildflower workshop chair: constance murray cover photo: courtesy of patricia folley. “this opuntia polyacantha was blooming away on a rocky shore on jed johnson lake in the wichita mountains wildlife refuge. the photo was taken with a nikon coolpix camera about the size of a deck of cards, and no tripod. cactus flowers are wonderful for holding still!” articles (c) the authors journal compilation (c) oklahoma native plant society except where otherwise noted, this work is licensed under a creative commons attribution-noncommercialsharealike4.0 international license, https://creativecommons.org/licenses/by-nc-sa/4.0/, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly attributed, not used for commercial purposes, and, if transformed, the resulting work is redistributed under the same or similar license to this one. https://doi.org/10.22488/okstate.17.100049 2 oklahoma native plant record volume 7, number 1 table of contents foreword. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .3 vascular plants of the oklahoma ozarks . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 ph.d. dissertation dr. charles s. wallis updated oklahoma ozark flora . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21 dr. bruce w. hoagland the vascular flora of the oklahoma centennial botanical garden site . . . . 54 osage county, oklahoma dr. bruce w. hoagland and ms. amy buthod vascular plant checklists from oklahoma . . . . . . . . . . . . . . . . . . . . . . . . . . . . 67 dr. michael w. palmer the need for savanna restoration in the cross timbers . . . . . . . . . . . . . . . . 78 mr. caleb stotts, dr. michael w. palmer, and dr. kelly kindscher botanizing with larry magrath . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .91 editorial ms. patricia folley five-year index to oklahoma native plant record . . . . . . . . . inside back cover journal of the oklahoma native plant society, volume 11, december 2011 oklahoma native plant record 3 volume 11, december 2011 foreword looking back over the last 10 years of publishing the oklahoma native plant record gives us an honest sense of accomplishment. it has been an uphill struggle to establish our journal, but with the turning of the decade, the record has also turned the corner. as of this year all volumes are available online through oklahoma state university’s edmon low library as an e-journal publication. it can be accessed globally at http://ojs.library.okstate.edu/osu/. this year the historic paper is one of the chapters of linda gatti clark’s 1997 ph.d. dissertation for oklahoma state university (osu). we will have to wait for an update to this flora of boehler seeps from another source, but it should provide an important comparison of changes in species over time in this unique habitat. marian smith is from southern illinois university and paul mckenzie is endangered species biologist and coordinator for the us fish and wildlife service. their paper on hybridization of two local species of sedges is the first to be submitted entirely online, a good sign that we are finally getting established. incorporating online submission and publishing seems to be one of the most effective ways to connect with out-of-state scientists. yet, this article is also a link to the past, being inspired by dr. larry magrath, late member of oklahoma native plant society. molly parkhurst, andrew doust, margarita mauro-herrera, jeffrey byrnes, and janette steets from osu have introduced a brand new topic for the record; a population genetics study of scribner’s panicum, one of our native grasses. this up-to-date molecular research paper is likely to be cited in larger journals and is yet another sign of our progress. jerad linneman, one of michael palmer’s former students, addressed some of the redcedar controversies in his m.s. thesis from osu, but was hired by the u. s. government before he could publish it. matthew allen, also from osu, was recruited to update and co-author it for our journal. we appreciate michael palmer’s initiative and assistance in acquiring the manuscript. it is very timely, considering the redcedar controversies and their role in recent wildfires. this paper discusses the effects of removing redcedar from old field grasslands. richard thomas’s paper is also a “hot” topic. it is an interdisciplinary study based on climate change and biogeographic interaction. this article can be used by local botanists and teachers to relate environmental science and climate change to local consequences. it is a comparison of the composition of the cross timbers before euro-american settlement. remember and tell everyone you know that the record is now available online. if you want a printed copy of any of our future volumes, get your order in early. only 50 copies will be printed each year. sheila strawn managing editor oklahoma native plant record, volume 12, number 1, december 2012 1 oklahoma native plant record journal of the oklahoma native plant society 2435 south peoria tulsa, oklahoma 74114 volume 12, december 2012 issn 1536-7738 http://ojs.library.okstate.edu/osu/ managing editor: sheila strawn production editor: paula shryock electronic production editor: sandy graue technical advisor: bruce hoagland the purpose of onps is to encourage the study, protection, propagation, appreciation and use of the native plants of oklahoma. membership in onps is open to any person who supports the aims of the society. onps offers individual, student, family, and life memberships. 2012 officers and board members president: adam ryburn vice-president: lynn michael secretary: sandy graue treasurer: mary korthase membership coordinator: tina julich historian: jeanie coley past president: lynn michael board members: brooke bonner elaine lynch buddy miller clare miller janette steets jay walker central chapter chair: joe roberts cross-timbers chapter chair: ron tyrl mycology chapter chair: steve marek northeast chapter chair: alicia nelson color oklahoma chair: pearl garrison conservation chair: chadwick cox gaillardia editor: chadwick cox website manager: adam ryburn http://www.oknativeplants.org harriet barclay award chair: rahmona thompson anne long award chair: gloria caddell onps service award chair: sue amstutz photography contest chair: kim shannon librarian: karen haworth mailings chair: karen haworth publicity chair: alicia nelson cover photo: erythronium sp. (dogtooth lily) by lynn michael, past president of onps . articles (c) the authors journal compilation (c) oklahoma native plant society except where otherwise noted, this work is licensed under a creative commons attributionnoncommercial-sharealike4.0 international license, https://creativecommons.org/licenses/ by-nc-sa/4.0/, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly attributed, not used for commercial purposes, and, if transformed, the resulting work is redistributed under the same or similar license to this one. https://doi.org/10.22488/okstate.17.100087 http://ojs.library.okstate.edu/osu/ http://www.oknativeplants.org/ oklahoma native plant record volume 12, december 2012 2 oklahoma native plant record volume 12 table of contents foreword ................................................................................................................................................. 3 possible mechanisms of the exclusion of johnson grass by tall grass prairies m. s. thesis ................................................................................................................................ 4 dr. marilyn a. semtner a preliminary pawnee ethnobotany checklist .............................................................................. 33 mr. c. randy ledford vascular flora of alabaster caverns state park, cimarron gypsum hills, woodward county, oklahoma .......................................................................................... 43 dr. gloria m. caddell and ms. kristi d. rice a comparison of the composition and structure of two oak forests in marshall and pottawatomie counties ................................................................................. 63 dr. bruce smith critic’s choice essay: virtual herbaria come of age ....................................................................... 69 dr. wayne elisens editorial policies and procedures ..................................................................................................... 72 five year index to oklahoma native plant record ........................................... inside back cover oklahoma native plant record, journal of the oklahoma native plant society, volume 12, december 2012 title page table of contents foreword oklahoma native plant record, volume 15, number 1, december 2015 oklahoma native plant record journal of the oklahoma native plant society p. o. box 14274 tulsa, oklahoma 74159-1274 volume 15, december 2015 issn 1536-7738 http://ojs.library.okstate.edu/osu/ managing editor: sheila strawn production editor: paula shryock electronic production editor: sandy graue manuscript editor: mark fishbein technical advisor: kristi rice the purpose of onps is to encourage the study, protection, propagation, appreciation, and use of the native plants of oklahoma. membership in onps is open to any person who supports the aims of the society. onps offers individual, student, family, and life memberships. 2015 officers and board members president: joe roberts vice-president: sheila strawn secretary: sandy graue treasurer: mary korthase historian: adam ryburn past president: adam ryburn board members: mike dunn pearl garrison elaine lynch jay pruett bruce smith lara souza chapter chairs: central: adam ryburn cross timbers: mark fishbein northeast: connie murray southwest: doug kemper publicity/merchandise chair: alicia nelson conservation chair: chadwick cox tulsa garden club liaison: sue amstutz betty kemm service award: sue amstutz awards chair: gloria caddell membership database: tina julich photo contest chair: lynn michael mailings/printings chair: karen haworth gaillardia editors: adam ryburn, marilyn stewart website manager: adam ryburn http://www.oknativeplants.org cover photo: euonymous americana (hearts-a-bustin; strawberry bush) by connie arnold articles (c) the authors journal compilation (c) oklahoma native plant society except where otherwise noted, this work is licensed under a creative commons attributionnoncommercial-sharealike4.0 international license, https://creativecommons.org/licenses/ by-nc-sa/4.0/, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly attributed, not used for commercial purposes, and, if transformed, the resulting work is redistributed under the same or similar license to this one. https://doi.org/10.22488/okstate.17.100108 http://ojs.library.okstate.edu/osu/ http://www.oknativeplants.org/ 2 oklahoma native plant record volume 15, december 2015 oklahoma native plant record volume 15 table of contents foreword .................................................................................................................................................... 3 preface to first flowering dates for central oklahoma .................................................................... 4 dr. wayne elisens first flowering dates for central oklahoma ....................................................................................... 6 mr. ben osborn forest structure and fire history at lake arcadia, oklahoma county, oklahoma (1820–2014) ......................................................................................................................... 19 dr. chad king interplanting floral resource plants with vegetable plants enhances beneficial arthropod abundance in a home garden. ..................................................................... 31 ms. chrisdon b. bonner, dr. eric j. rebek, dr. janet c. cole, dr. brian a. kahn, and dr. janette a. steets contributions to the flora of cimarron county and the black mesa area ................................... 49 ms. amy k. buthod and dr. bruce w. hoagland antifungal activity in extracts of plants from southwestern oklahoma against aspergillus flavus. .......................................................................................................................... 78 dr. tahzeeba frisby and cameron university students kudzu, pueraria montana (lour.) merr. abundance and distribution in oklahoma. ................... 96 ms. marli claytor and dr. karen r. hickman critic’s choice essay: mistletoe, phoradendron serotinum (raf.) johnston ........................................... 105 dr. paul buck editorial policies and procedures ....................................................................................................... 107 five year index to oklahoma native plant record ............................................... inside back cover oklahoma native plant record, journal of the oklahoma native plant society, volume 15, december 2015 title page table of contents foreword five year index to oklahoma native plant r ecord volume 9 4 vascular plants of southeastern oklahoma from san bois to the kiamichi mountains, 1969 ph. d. dissertation, f. hobart means 38 composition and structure of bottomland forest vegetation at the tiak research natural area, mccurtain county, oklahoma, bruce w. hoagland and newell a. mccarty 59 is seedling establishment very rare in the oklahoma seaside alder, alnus maritime ssp. oklahomensis? stanley a. rice and j. phil gibson 64 whatever happened to cheilanthes horridula and cheilanthes lindheimeri in oklahoma? bruce a. smith 70 critic’s choice essay: invasive plants versus oklahoma’s biodiversity, chadwick a. cox volume 10 4 the identification of some of the more common native oklahoma grasses by vegetative characters, 1950 m. s. thesis, william franklin harris 34 the vascular flora of hale scout reservation, leflore county, oklahoma bruce w. hoagland and amy k. buthod 54 the toxicity of extracts of tephrosia virginiana (fabaceae) in oklahoma, mary gard 65 four western cheilanthoid ferns in oklahoma, bruce a. smith 77 critic’s choice essay: being a method proposed for the ready finding ... to what sort any plant belongeth, ronald j. tyrl volume 11 4 survey of the vascular flora of the boehler seeps and sandhills preserve, ph. d. dissertation, linda gatti clark 22 schoenoplectus hallii, s. saximontanus, and the putative s. hallii x s. saximontanus hybrid: observations from the wichita mountains wildlife refuge and the fort sill military reservation from 20022010, marian smith and paul m. mckenzie 33 spatial genetic structure of the tallgrass prairie grass dichanthelium oligosanthes (scribner’s panicum), molly j. parkhurst, andrew doust, margarita mauro-herrera, janette a. steets, and jeffrey m. byrnes 43 the effects of removal of juniperus virginiana l. trees and litter from a central oklahoma grassland, jerad s. linneman, matthew s. allen, and michael w. palmer 61 the changing forests of central oklahoma: a look at the composition of the cross timbers prior to euro-american settlement, in the 1950s and today, richard e. thomas and bruce w. hoagland 75 critic’s choice essay: some thoughts on oklahoma plants and summer 2011’s exceptional drought, leslie e. cole volume 12 4 possible mechanisms of the exclusion of johnson grass by tall grass prairies, m. s. thesis, marilyn a. semtner 33 a preliminary pawnee ethnobotany checklist, c. randy ledford 43 vascular flora of alabaster caverns state park, cimarron gypsum hills, woodward county, oklahoma, gloria m. caddell and kristi d. rice 63 a comparison of the composition and structure of two oak forests in marshall and pottawatomie counties, bruce smith 69 critic’s choice essay: virtual herbaria come of age, wayne elisens volume 13 4 ecology and taxonomy of water canyon, canadian county, oklahoma, m. s. thesis, constance a. taylor 29 a checklist of the vascular flora of the mary k. oxley nature center, tulsa county, oklahoma, amy k. buthod 48 smoke-induced germination in phacelia strictaflora, stanley a. rice and sonya l. ross 55 critic’s choice essay: a calvacade of oklahoma botanists in oklahoma – contributors to our knowledge of the flora of oklahoma, ronald j. tyrl and paula a. shryock oklahoma native plant society p.o. box 14274 tulsa, oklahoma 74159-1274 _________________________________________________________________________ in this issue of oklahoma native plant record volume 14, december 2014: _________________________________________________________________________ 4 flora of kiowa county, oklahoma, m. s. thesis lottie opal baldock 38 gardens of yesteryear sadie cole gordon 43 oklahoma deciduous trees differ in chilling enhancement of budburst stanley a. rice and sonya l. ross 50 mapping distribution in oklahoma and raising awareness: purple loosestrife (lythrum salicaria), multiflora rose (rosa multiflora), and japanese honeysuckle (lonicera japonica) katherine e. keil and karen r. hickman 67 non-twining milkweed vines of oklahoma: an overview of matelea biflora and matelea cynanchoides (apocynaceae) angela mcdonnell 80 critic’s choice essay: pollination ecology of our native prairie plants gloria m. caddell five year index to okla homa native plant r ecord – inside back cover oklahoma native plant record, volume 12, number 1, december 2012 oklahoma native plant record 33 volume 12, december 2012 c. randy ledford https://doi.org/10.22488/okstate.17.100090 a preliminary pawnee ethnobotany checklist c. randy ledford oklahoma state parks, biologist, retired crloutandabout@yahoo.com pawnee, oklahoma 74058 keywords: pawnee, ethnobotany, plant use, plants distribution abstract this document contains excerpts from a work in progress focusing on the ethnobotany of the pawnee native americans. the effort being made is to consolidate research findings to provide a written record specifically addressing plant use by the pawnee. the majority of the information gained was through literature reviews which provided a historic perspective. however, living among the pawnee for twenty-two years has provided some insight into modern uses of some plants. a priority at the onset was to identify and describe the broad-ranging application of plants within their culture. all the ethnobotanical examples here are based on plants that have been documented in oklahoma. each plant is related to its currently known biogeography in kansas and nebraska which was regionally part of their historic homeland until their removal to oklahoma beginning in 1875. introduction loss of land to encroaching euroamericans, inept government policies, disease, and warfare all contributed to the cultural degradation of the four bands or divisions of the pawnee. the bands are known as the chawi, kitkahahki, pitahawirata, and skiri. the pawnee belong to the caddoan language family which also includes the arikara, caddo, and wichita tribes. historically, the bands had linguistic differences and it is especially noted today when comparing the skiri dialect to the often called, “south bands”. well before the tribe was relocated to what is now known as pawnee, oklahoma, cultural fragmentation had begun. for an account of the chronological history of the pawnee see the pawnee indians by george e. hyde (1951). the floristic influence that enveloped and sustained the pawnee culture in their homeland arose from prairie plant associations and riparian environments linked to major rivers including the loup, platte, republican, and their tributaries. in close proximity to their villages with earth lodges were gardens where they cultivated crops such as beans, corn, squash, and tobacco. many plants were gathered from the surrounding areas to meet a variety of needs. in addition to farming, a summer and winter bison hunt was undertaken. their survival and religious practices were critically dependent on a deep connection to the natural world. their new land allocation in oklahoma consisted of different soils, a different climate and astronomical position, and plant life that further changed their life ways as pressure of acclimation and assimilation mounted. as a result, the botanical knowledge necessary to carry out ceremonies and other life ways continued to wane. as elders passed away, some loss of plant use knowledge accompanied them with each succeeding generation. although information sources dating in the 1800s were found that contributed to oklahoma native plant record volume 12, december 2012 c. randy ledford 34 the enthnobotany research, significant material was extracted from the works of james r. murie (half pawnee), melvin r. gilmore (married into the tribe), george a. dorsey, and others around the turn of the 20th century. the intent of the research was not to limit the focus to plants used for food and medicine. other relationships such as the use of plants for ceremonies, games, and materials were investigated to provide a broader representation as well as enhance an understanding of the complexity of pawnee culture. with each plant species listed, an attempt to document its location in kansas, nebraska, and oklahoma was made primarily using the united states department of agriculture (usda) and oklahoma vascular plants database (ovpd) websites. the results revealed that certain plants were documented in all three states, other plants had limited or restricted ranges in one or more states, and some were not found in oklahoma. the presentation here consists of only a sample of plant species that are found in oklahoma, reported to have been used by the pawnee. providing a written record of consolidation specifically addressing the use of plants by the pawnee may contribute to educational and cultural interests of the pawnee nation, individual tribal members, and others. oklahoma native plant record 35 volume 12, december 2012 c. randy ledford pawnee ethnic botany plant listing each plant is listed by family, genus, and species; former scientific name in parenthesis; one or more common names, including the pawnee language name and meaning (if known); distribution of the plant; and information regarding usage of the plant. cupressaceae juniperus virg inana l. eastern redcedar or “mother cedar” (as referred to by the pawnee) tawatsaako eastern redcedar grows in a variety of soil conditions and thrives in kansas, oklahoma, and nebraska (usda). true cedar, such as western redcedar, of the genus thuja, is not native to kansas, oklahoma, and nebraska. this evergreen tree, commonly called “cedar”, was used by many native americans for a variety of purposes [moerman 1998 (pp.290-291)]. in regard to the pawnee, smoke inhaled from burned twigs is used as a remedy for colds; a decoction of fruits and leaves is used as a cough medicine and given to horses for same purpose; and boughs are put on tepee poles to ward off lightning (ibid). historically, juniper trees were used in ceremonies including the skiri doctors and bear dances [murie 1981 (pp.170, 336)]. skiri bear society participants used the leaves for ceremonial smudges, and if a thunderstorm should threaten, a smudge was made to protect the lodge (earth lodge) [murie 1914 (p.604)]. murie did not define “smudge” in the text cited. “cedar poles” were fashioned for use as lances associated with the two lance society [ibid (p.561)]. gilmore [1919 (p.12)] noted that a smoke offering of cedar twigs was used as a remedy for nervousness and bad dreams. based on my observations, i report that the pawnee currently burn juniper leaves/needles as a ceremonial incense and/or prayer smoke offering, including use in the native american church. as i have witnessed, the smoke caused by placing juniper leaves on coals has been used to suffuse a person or object and in some situations only to allow smoke to permeate the surroundings. prior to the practice, it is often said “going to burn some cedar” and the word “smudge” is not used. nothing further will be added here since the focus of this paper is not to reveal ceremonial details. according to weltfish [1965 (p.387)], juniper was used for tepee poles by the pawnee, but not necessarily as first choice if cottonwood was available. juniper also had a place in ghost dance hand game paraphernalia. examples include hand game leaders mark rudder using a “cedar wood cross topped with a soft eagle feather and three cedar sticks topped with four red-painted crow feathers radially affixed to the top”, and barclay white having used tally or counting sticks that were made of “cedar” in a ceremony [lesser 1933 (pp.267-268, 288)]. also, emmett pierson’s hand game set included a “sack of crumpled cedar leaves” that was part of the contents of a bundle. at the onset of the ceremony, a handful of the leaves were placed oklahoma native plant record volume 12, december 2012 c. randy ledford 36 on coals [lesser 1933 (pp.274, 278)]. at one time, i was overseer of the pierson collection, on behalf of skiri band member ms. maude chisholm, which included a cloth bag containing juniper leaves. the collection on loan is housed in the pawnee bill ranch museum, pawnee, oklahoma, and also includes hand game sticks as well as other items. agavaceae yucca g lauca nutt. yucca or soapweed yucca chakida-kahtsu or chakila-kahtsu this perennial can be found growing in prairies and on hillsides in kansas and nebraska (usda). it has been documented across oklahoma (ovpd). the yucca is known for the roots being used as soap, especially for washing hair, and the fibers from the leaves used by the pawnee to make twine or cordage, with the leaf ends used as needles [gilmore 1919 (p.19)]. i have made cordage from the leaves and found it to be quite strong, especially if primary use is for binding material. liliaceae allium cana densis l. (=a. mutabile) wild onion or meadow garlic osidawa this perennial herb can be found growing in prairies, open woods, roadsides, and lawns. allium can be found in kansas and nebraska (usda). a. canadensis and other species of onion can be found throughout oklahoma (ovpd). my personal experience of using “wild onions” as food is that a little can go a long way. moerman [1998 (p.57)] cited at least thirty species of allium used by different native american tribes, with reference to the above named species associated with the pawnee for use as a spice, sauce, and relish. it is a species cited by gilmore [1919 (p.19)], but under the older name, a. mutabile. reportedly, the plant was eaten raw, cooked to flavor meat and soup, and also fried (ibid). poaceae arundinaria g ig antea (walter) muhl. river cane or giant cane usda database lists the plant in kansas, but not in nebraska. more than 20 counties in eastern oklahoma host the plant (ovpd). river cane is the “bamboo” of north america. river cane served many purposes for tribes, especially the southeastern woodland cultures, in the making of arrows, blow guns, whistles, construction materials, fishing items, and in basketry [moerman 1998 (p.104)]. the woody grass inhabits moist bottomlands and forest understories and can reach 20 or more feet in height. oklahoma native plant record 37 volume 12, december 2012 c. randy ledford stewart culin [1975 (p.99)], described pawnee gaming sets containing four dice each that were made from split cane ranging from 8 to 16.5 inches (20.32 cm to 41.91 cm) in length, with some sets painted, and a set with a small feather tied to the end of each piece of cane. according to tuttle [1838 (p.41)] a type of flute was made out of cane which she noted as “sugar cane”. in my opinion, the flute was more likely made out of river cane. moerman [1998 (p.499)] only listed the seminole as using sugar cane as a food. a small cane whistle was included in the warrior’s bundle belonging to eagle flying under the heavens [murie 1981 (p.190)]. h esperostipa spartea (trin.) barkworth (=stipa spartea) porcupine or needle grass pitsuts (hair brush) or paari pitsuts (pawnee hairbrush) porcupine grass prefers dry prairies and open woods with a geographic range that includes kansas and nebraska. according to the usda database and ovpd, the grass has been documented in three northern oklahoma counties: kay, osage, and washington. pawnee county joins osage county to the north. the grass was prepared as a brush after the stiff awns and stalks were tightly bound into a small rounded-bundle, followed by burning off the pointed grains [gilmore 1919 (p.14)]. gilmore, citing alice fletcher, noted that the grass brush was used in the pawnee hako pipe ceremony. included in the publication by fletcher [1904 (p.220)], regarding the hako ceremony, a section titled “explanation by the ku’rahus” (old man or priest) is as follows: “the grass of which the brush is made is gathered during a ceremony belonging to the rain shrine. it represents toharu, the living covering of mother earth. the power which is in toharu gives food to man and the animals so that they can live and become strong and able to perform the duties of life. this power represented by the brush of grass is now standing before the little child”. the grass brush was also described by weltfish [1965 (p.363)]. typhaceae t ypha latifolia l. cat-tail or broadleaf cattail hawahawa and kirit-tacharush (meaning “eye itch” with reference to down getting into eyes) the plant grows in a wide-spread range and conditions of moist sites and wetland environments. it can be found in kansas, oklahoma, and nebraska (usda and ovpd). the pawnee used the down to make dressings for burns and scalds [moerman 1998 (p.574)]. gilmore [1919 (p.12)] related that the down was used on infants to prevent chafing, as we use talcum; as a filling for pillows; and as padding for cradle boards, as well as in quilting baby wrappings. a great quantity of down was gathered in advance to have readied for the placement oklahoma native plant record volume 12, december 2012 c. randy ledford 38 of a newborn infant on the down. with the lack of cotton diapers in the olden times, pads of down were used (ibid). cat-tail leaves were used in the making of woven mats as an alternative to bulrush [weltfish 1965(p.404)]. anacardiaceae r hus g labra l. smooth sumac nuppikt, sour top smooth sumac is a shrub that often grows in prairies, fields, and edges of woodlands. it is common across oklahoma (ovpd) and much of kansas and nebraska (usda). the pawnee name is in reference to the sour-tasting red fruits that develop in summer. in the fall when the leaves turn red, they were gathered and dried for smoking [moerman 1998 (p.472)]. gilmore [1919 (p.47)] also noted the use of the red leaves for smoking. in relation to a chawi doctor ceremony, sumac leaves were mixed with tobacco for smoking [murie 1981 (p.203)]. the fruits were boiled to make a remedy for dysmenorrhea and also for bloody flux [gilmore 1919 (p.47)]. the use of its leaves, bark, and roots to make a black dye included the application to bison hides [moerman 1998 (p.472)]. araceae arisaema triphyllum (l.) schott jack-in-the pulpit or indian turnip nikso kororik kahtsu nitawau; medicine or herb, that bears, what resembles, an ear of corn (the ripe fruits) it grows in moist woodlands and is known to exist in kansas and nebraska (usda). it has been documented in more than 17 counties in oklahoma (ovpd). if you eat the un-cooked corm (root), you will certainly reap the unpleasant sensation due to the calcium oxalate crystals inherent in this poisonous plant. the corm was not reportedly used as a food source of the pawnee, but was pulverized and used as medicine. it was used to treat headaches by dusting the top of the head and temples and applied as a counterirritant for rheumatism and similar pains. the seeds were placed in gourd shell rattles [gilmore 1919 (p.17)]. asteraceae grindelia squarrosa (push) dun. curly cup or curly top gum weed bakskititis, stick-head (bak, head; skitits, sticky) gumweed is a perennial plant that grows in fields, along roadsides, and in waste places. the plant exudes a sticky substance which is true to the pawnee name. it is spread across the oklahoma native plant record 39 volume 12, december 2012 c. randy ledford northern half of kansas and found across the state of nebraska (usda). in oklahoma, the plant is mostly situated in western counties (ovpd). according to a pawnee informant, the tops and leaves were boiled to make a wash for saddle galls and sores on horses [gilmore 1919 (p.81)]. this species has the longest listing of uses by native americans of the grindelia genus [moerman 1998 (pp.252-253)]. h elianthus annuus l. common sunflower kirik-tara-kata, yellow eyes (kirik, eye; tara, having; kata, yellow) the annual plant is often found in fields and along roadsides. the common sunflower is listed for every county in kansas, and in nebraska, it is distributed across the entire state (usda). it has been documented in the majority of counties in oklahoma (ovpd). gilmore [1919 (p.78)] noted that he could not find that the plant was ever cultivated by any of the nebraska tribes, but there was evidence of such by some of the eastern tribes and the arikara (linguistic neighbors to the north). a pawnee informant of gilmore’s reported that the seeds were pounded up with certain roots (not identified or disclosed) and were taken in the dry form without further preparation, by women who became pregnant while still suckling a child for the reason that the suckling child should not become sick (ibid). h elianthus tuberosa l. jerusalem artichoke kisu-sit (kisu, tapering; sit, long) the perennial plant has been reported for mainly the eastern three-quarters of kansas and nebraska (usda). in oklahoma, it is erratically distributed mostly in the eastern half of the state (ovpd). it grows in wet soils of prairies, open woods, disturbed areas, and roadsides. the people of the nebraska tribes say they never cultivated the plant, but used its tubers for food [gilmore 1919 (p.79)]. the pawnee reportedly ate them only raw, but the others, according to their own statement, ate them raw, boiled, or roasted (ibid). in a pawnee tale, “coyote and the artichoke”, artichoke (presumably jerusalem artichoke), was mentioned; whereas, coyote ate too many artichokes which caused intestinal distress [dorsey 1906 (p.464)]. wonder what the moral of that story part is? fabaceae apios americana medik. (=glycine apios) groundnut or indian potato its the habitat of the perennial twining herb includes pond and stream banks, moist thickets, and wet meadows. it can be found in kansas and nebraska (usda) and mostly in central, eastern, and some southwestern counties in oklahoma (ovpd). oklahoma native plant record volume 12, december 2012 c. randy ledford 40 two parts of the plant were a food source for the pawnee. the tubers were eaten raw or cooked, preferably gathered in the fall, and the seeds of summer were consumed like peas [kindscher 1987 (pp.48-49)]. groundnut is a common native food plant of temperate and eastern north america. it is possible that the plant was propagated by the cheyenne and other tribes and its range extended westward (ibid). weltfish [1965 (p.415)] noted that the tubers were an important food provision for the winter bison hunt. gilmore [1919 (p.42)] reported that the tubers of the plant were used as a food source by all the tribes within its range and prepared by boiling or roasting. pediomelum esculenta (pursh) rydb. (=psoralea esculenta) indian breadroot or pomme blanche patsuroka the perennial herbaceous plant prefers prairies and has a scattered occurrence in about 16 oklahoma counties (ovpd). it is scattered through much of kansas and across nebraska (usda). the french name, “pomme blanche”, means white fruit. the plant’s root was an important substance of the vegetal diet of the plains tribes and after being peeled was eaten fresh, cooked, or stored to dry for use during the winter. the roots were braided in long strings by the tapering ends. when the women and children went to the prairie to gather the roots, on finding a plant the mother tells the children to note the directions which the several branches point and a child is sent in the general direction of each branch to look for another plant, for they say the plants “point to each other” [gilmore 1919 (p.40)]. nelumbonaceae nelumbo lutea willd. water lily or water chinquapin tukawiu, skiri band word and tut, chawi band the aquatic plant currently has a range in nebraska limited to 3 counties and is scattered in more than 20 counties in kansas. it has been documented across oklahoma (ovpd), but only in 12 counties by the usda. the plant was considered to be one invested with mystic powers. it was an important food source with use of the seeds and tubers (shaped somewhat like a banana). the hard, nutlike seeds were cracked and used with meat for making soup. the peeled tubers were cut up and cooked with meat or with hominy [gilmore 1919 (p.27)]. ranunculacea aquileg ia canadensis l. wild columbine or red columbine skalikatit or skarikatit, black seed the plant prefers growing in moist, well-drained, shady or partly shaded sites. it has been documented in some eastern kansas counties and some northern and eastern counties in oklahoma native plant record 41 volume 12, december 2012 c. randy ledford nebraska (usda). in oklahoma, the columbine is mainly in 17 northeastern and eastern counties (ovpd). according to an account of the seeds being used as a perfume and a love charm, seeds are pulverized and rubbed in the palms of the suitor, who then contrives to shake hands with the desired one, whose fancy it is expected will thus be captivated [gilmore 1919 (pp.30-31)]. also, historically, seeds were crushed in an elm mortar by a pestle made of the same wood, with the resulting powder being added to hot water and the infusion being drunk for fever and headache (ibid). conclusion i have made an attempt to provide a checklist of plant usage as it relates to the pawnee. before publication, a “preliminary pawnee ethnobotany checklist” was reviewed by mr. stephen bird (b.s., m.s.) and three other pawnees. my goal is to complete the larger paper which may include more than sixty species of plants, not including species associated with agriculture. when that is done, a pawnee review committee will be offered the opportunity to respond to the findings of the research. ethnobotany has many applications. along with the existing pawnee endeavors involving agriculture, linked to historic corn varieties and other cultivated plants of olden times, herbaceous native plants could also be grown in an ethnobotany garden to contribute to horticultural skills development, cultural education, and the actual use of the plants. also, information gained through the research could be applied to the arts and in the sanctioned reproduction of certain artifacts. it is like filling ones tow sack with pieces of lost earthly connections to possess in order to bring elements of the past to the present. lastly, i share an excerpt from “origin of the chaui”, also written “chawi”, as told by roaming chief-hereditary chief of the chaui (band) in about 1906 and recorded by dorsey [1997(p.13)]: the earth i give you, and you are to call her ‘mother’, for she gives birth to all things. the timber that shall grow upon the earth you shall make use of in many ways. some of the trees will have fruit upon them. shrubs will grow from the ground and they will have berries upon them. all these things i give you and you shall eat of them. never forget to call the earth ‘mother’, for you are to live upon her. you must love her, you must walk upon. literature cited culin, stewart. 1975. games of the north american indians. new york: dover publications, inc. dorsey, george. 1997 (first published 1906). the pawnee mythology. lincoln: university of nebraska press. fletcher, alice. 1996. the hako: song, pipe, and unity in a pawnee calumet ceremony. reprint of 1904 report. bison books edition. university of nebraska press. gilmore, melvin. 1977 (first published 1919). uses of plants of the indians of the missouri river region. lincoln: university of nebraska press. hyde, george. 1974 (first published 1951). the pawnee indians. norman: university of oklahoma press. kindscher, kelly. 1987. edible wild plants of the prairie. [lawrence]: university press of kansas. lesser, alexander. 1978 (originally published 1933). the pawnee ghost dance oklahoma native plant record volume 12, december 2012 c. randy ledford 42 hand game. [madison]: university of wisconsin press. moerman, daniel. 1998. native american ethnobotany. portland: timber press. murie, james. 1914. pawnee indian societies. anthropological papers of the american museum of natural history, volume 11, part 7. murie, james, edited by douglas r. parks. 1981. ceremonies of the pawnee. lincoln: university of nebraska press. oklahoma vascular plants database (ovpd). 2011. oklahoma biological survey. university of oklahoma, norman. tuttle, sarah. 1838. history of the american mission to the pawnee indians. boston: massachusetts sabbath school society. usda-nrcs plants database. 2011. plant profiles. united states department of agriculture. weltfish, gene. 1965. the lost universepawnee life and culture. lincoln: university of nebraska press. . a preliminary pawnee ethnobotany checklist by mr. c. randy ledford journal of the oklahoma native plant society, volume 15, december 2015 five year index to oklahoma native plant record volume 10 4 the identification of some of the more common native oklahoma grasses by vegetative characters, 1950 m. s. thesis, william franklin harris 34 the vascular flora of hale scout reservation, leflore county, oklahoma bruce w. hoagland and amy k. buthod 54 the toxicity of extracts of tephrosia virginiana (fabaceae) in oklahoma, mary gard 65 four western cheilanthoid ferns in oklahoma, bruce a. smith 77 critic’s choice essay: being a method proposed for the ready finding ... to what sort any plant belongeth, ronald j. tyrl volume 11 4 survey of the vascular flora of the boehler seeps and sandhills preserve, ph. d. dissertation linda gatti clark 22 schoenoplectus hallii, s. saximontanus, and the putative s. hallii x s. saximontanus hybrid: observations from the wichita mountains wildlife refuge and the fort sill military reservation from 20022010, marian smith and paul m. mckenzie 33 spatial genetic structure of the tallgrass prairie grass dichanthelium oligosanthes (scribner’s panicum), molly j. parkhurst, andrew doust, margarita mauro-herrera, janette a. steets, and jeffrey m. byrnes 43 the effects of removal of juniperus virginiana l. trees and litter from a central oklahoma grassland, jerad s. linneman, matthew s. allen, and michael w. palmer 61 the changing forests of central oklahoma: a look at the composition of the cross timbers prior to euro-american settlement, in the 1950s and today, richard e. thomas and bruce w. hoagland 75 critic’s choice essay: some thoughts on oklahoma plants and summer 2011’s exceptional drought, leslie e. cole volume 12 4 possible mechanisms of the exclusion of johnson grass by tall grass prairies, m. s. thesis marilyn a. semtner 33 a preliminary pawnee ethnobotany checklist, c. randy ledford 43 vascular flora of alabaster caverns state park, cimarron gypsum hills, woodward county, oklahoma, gloria m. caddell and kristi d. rice 63 a comparison of the composition and structure of two oak forests in marshall and pottawatomie counties, bruce smith 69 critic’s choice essay: virtual herbaria come of age, wayne elisens volume 13 4 ecology and taxonomy of water canyon, canadian county, oklahoma, m. s. thesis, constance a. taylor 29 a checklist of the vascular flora of the mary k. oxley nature center, tulsa county, oklahoma amy k. buthod 48 smoke-induced germination in phacelia strictaflora, stanley a. rice and sonya l. ross 55 critic’s choice essay: a calvacade of oklahoma botanists in oklahoma – contributors to our knowledge of the flora of oklahoma, ronald j. tyrl and paula a. shryock volume 14 4 flora of kiowa county, oklahoma, m. s. thesis, lottie opal baldock 38 gardens of yesteryear, sadie cole gordon 43 oklahoma deciduous trees differ in chilling enhancement of budburst stanley a. rice and sonya l. ross 50 mapping distribution in oklahoma and raising awareness: purple loosestrife (lythrum salicaria), multiflora rose (rosa multiflora), and japanese honeysuckle (lonicera japonica) katherine e. keil and karen r. hickman 67 non-twining milkweed vines of oklahoma: an overview of matelea biflora and matelea cynanchoides (apocynaceae), angela mcdonnell 80 critic’s choice essay: pollination ecology of our native prairie plants, gloria m. caddell oklahoma native plant society p.o. box 14274 tulsa, oklahoma 74159-1274 _________________________________________________________________________ in this issue of oklahoma native plant record volume 15, december 2015: _________________________________________________________________________ 4 preface to first flowering dates for central oklahoma, wayne elisens 6 first flowering dates for central oklahoma, ben osborn 19 forest structure and fire history at lake arcadia, oklahoma county, oklahoma (1820–2014), chad king 31 interplanting floral resource plants with vegetable plants enhances beneficial arthropod abundance in a home garden. chrisdon b. bonner, eric j. rebek, janet c. cole, brian a. kahn, and janette a. steets 49 contributions to the flora of cimarron county and the black mesa area amy k. buthod and bruce w. hoagland 78 antifungal activity in extracts of plants from southwestern oklahoma against aspergillus flavus, tahzeeba frisby and cameron university students 96 kudzu, pueraria montana (lour.) merr. abundance and distribution in oklahoma marli claytor and karen r. hickman 105 critic’s choice essay: mistletoe, phoradendron serotinum (raf.) johnston, paul buck five year index to oklahoma native plant record – inside back cover 68 oklahoma native plant record volume 1, number 1, december 2001 checklist of the ferns, natural falls state park bruce a. smith biology instructor, mcloud high school mcloud, ok 74851 natural falls state park, formerly known as dripping springs is located in northeast oklahoma. the park’s natural beauty and flora have attracted visitors since 1907. in a 1988 visit to the oklahoma state university herbarium, i noticed that several herbarium sheets of ferns were collected from dripping springs. this was intriguing and made me want to visit the area. due to my interest in floristics and taxonomy, natural falls state park seemed the perfect place to create a checklist of ferns. thus, the objective of this study was to create an inventory of the ferns of dripping springs using my collection and the collections and observations from earlier botanists. a systematic collection of the ferns of dripping springs was conducted on august 7, 1998, october 15, 1998, and october 20, 2001. using standard taxonomic methods, each plant was identified to species and subsequently inventoried. in three days of collecting, 17 species from 6 families and 12 genera were encountered. since 1925 a total of 19 species from 6 families and 12 genera have been reported to occur. introduction natural falls state park, formerly known as dripping springs is located in northeast oklahoma. the park’s natural beauty and flora have attracted visitors since 1907. botanists such as harriet barclay, charles wallis, edgar wherry and several others have taken a special interest in the park because of its interesting flora, especially the ferns. in the 1980’s the privately owned park was closed due to the poor condition of the buildings on the property (1). in 1990 dripping springs was purchased by the oklahoma tourism and recreation department with plans to make it once again accessible to the public (2). in a 1988 visit to the oklahoma state university herbarium i noticed that several herbarium sheets of ferns were collected from dripping springs. this was intriguing and made me want to visit the area. in july 1989, while on vacation in northeast oklahoma my family and i visited the park only to find that it had closed. in 1998 i learned through a fellow botanist that the area was once again open to the public. due to my interest in floristics and taxonomy, natural falls state park seemed to be the perfect place to create a checklist of ferns. thus, the objective of this study was to create an inventory of the ferns of dripping springs by using my collection and the collections and observations from earlier botanists. the study area dripping springs is located west of siloam springs in the southeast corner of delaware county r25e, t20n, sec. 32 (3). the most eye-catching feature in the park is the 25-meter waterfall (2). the surrounding area above the waterfall and ravine below was the site of this study and past studies by other botanists. as is common in the ozarks, the surface rock contains chert that gives rise to acid soils. underlying the spring is a stratum of limestone of both fernvale and fite formations. thus the soils at the lower levels (ravine) are somewhat alkaline (4). soils in the sampling areas include clarkesville very cherty silt loam, clarkesville stony silt loam, and staser gravelly loam. climate of the area is moist and warm temperature (5). smith, b.a. https://doi.org/10.22488/okstate.17.100007 oklahoma native plant record 69 volume 1, number 1, december 2001 smith, b.a. inventory of ferns a systematic collection of the ferns of dripping springs was conducted on august 7, 1998 and october 20, 2001. collecting focused on the ferns that grew in the ravine floor and rock surfaces. using standard taxonomic methods, each plant was identified to species and subsequently inventoried. specimens typically were collected in fertile condition with the exception of botrychium virginianum. nomenclature for the taxa follows flora of north america (6). vouchers will be deposited in the oklahoma state university herbarium (okla). flora of natural falls state park (dripping springs) in three days of collecting, 17 species from 6 families and 12 genera were encountered (table 1). fern collections and recordings from 1925-1977 are listed in tables 2 and 3. from 1925-2001 a total of 19 species from 6 families and 12 genera have been reported to occur. ferns designated as rare by the oklahoma natural heritage inventory (7) were asplenium bradleyi (s1). acknowledgements a special thanks to the staff at natural falls state park and tom crider for allowing me to have this opportunity. i hope the checklist will be useful. i also thank dr. ron tyrl for his advice on this study. finally i want to thank the mcloud high school botany class for accompanying me on the october 20, 2001 field trip references 1. the delaware county historical society. heritage of the hills: a delaware county history. jay, ok (1979). 2. news release oklahoma tourism and recreation department. oklahoma city, ok (1997). 3. the roads of oklahoma. map. fredericksburg, tx (1997). 4. wherry, e. “ferns of dripping springs, oklahoma.” american fern journal 18:61 (1928). 5. cole, e., soil survey of cherokee and delaware counties, oklahoma, u.s. government printing office, washington, d.c. (1970). 6. flora of north america editorial committee, flora of north america, vol. 2, pteridophytes and gymnosperms. oxford university press, new york (1933). 7. oklahoma natural heritage inventory. short working list of rare plants 22 may 2001. oklahoma biological survey, norman (2001). 70 oklahoma native plant record volume 1, number 1, december 2001 smith, b.a. table 1. checklist of ferns, natural falls state park (dripping springs) collected by bruce smith august 7, 1998 and october 15, 1998, and october 20, 2001 aspleniaceae asplenium bradleyi d.c. eaton. bradley’s spleenwort, august 7, 1998 3590 (okla) asplenium platyneuron (l.) britton, sterns, & poggenb. ebony spleenwort, 3578 (okla) asplenium rhizophyllum l. walking fern 3562 (okla) asplenium trichomanes l. maidenhair spleenwort 3619 (okla) dryopteridaceae cystopteris bulbifera (l.) bernh. bulblet bladder fern 3589 (okla) cystopteris tennesseensis shaver tenessee bldder fern 3573 (okla) dryopteris marginalis (l.) a. gray marginal shield fern 3568 (okla) onoclea sensibilis l. sensitive fern 3774 (okla) polystichum acrostichoides (michx.) schott. christmas fern 3580 (okla) woodsia obtusa (spreng.) torr. blunt lobed woodsia 3622 (okla) ophioglossaceae botrychium virginianum (l.) sw. rattlesnake fern 3773 (okla) polypodiaceae pleopeltis polypodioides (l.) e.g. andrews & windham resurrection fern 3772 (okla) pteridaceae adiantum capillus-veneris l. southern maidenhair fern 3565 (okla) adiantum pedatum l. northern maidenhair fern3576 (okla) cheilanthes lanosa (michx) d.c. eaton. hairy lip fern 3584 (okla) pellaea atropurpurea (l.) link purple cliff brake fern 3582 (okla) thelypteridaceae phegopteris hexagonoptera (michx.) fee broad beech fern 3571 (okla) table 2. checklist of ferns natural falls state park (dripping springs) reported by john k. small and edgar t. wherry may 3,1925 (4) aspleniaceae asplenium bradleyi d.c. eat. bradley’s spleenwort asplenium platyneuron (l.) britton, sterns, & poggenb. ebony spleenwort asplenium resiliens kunze little ebony spleenwort asplenium rhizophyllum l. walking fern asplenium trichomanes l. maidenhair spleenwort dryopteridaceae cystopteris bulbifera (l.) bernh. bulblet bladder fern cystopteris fragilis (l.) bernh. brittle fern dryopteris marginalis (l.) a. gray marginal shield fern polystichum acrostichoides (michx.) schott. christmas fern pteridaceae adiantum capillus-veneris l. southern maidenhair fern cheilanthes lanosa (michx.) d.c.eaton hairy lip fern oklahoma native plant record 71 volume 1, number 1, december 2001 smith, b.a. table 3. checklist of ferns natural falls state park (dripping springs) collected from 1928-1977 aspleniaceae asplenium platyneuron (l.) britton, sterns & poggenb. june 30, 1957 charles wallis 4454 (okla) asplenium rhizophyllum l. may 4, 1928 robert stratton 798 (okla) july 17, 1929 robert stratton (okla) july 7, 1950 u.t. waterfall 9571 (okla) june 30, 1957 charles wallis 4456 (okla) june 16, 1972 john and connie taylor 10792 (okla) asplenium trichomanes l. may 7, 1938 milton hopkns 3250 (okla) july 7, 1950 u.t. waterfall 9561 (okla) dryopteridaceae cystopteris bulbifera (l.) bernh. june 30, 1957 charles wallis 4449 (okla) cystopteris tennesseensis shaver july 7, 1950 u.t. waterfall 9569 (okla) june 16, 1972 john and connie taylor 10791 (okla) dryopteris marginalis (l.) a. gray may 4, 1928 t.a. tripp 137 (okla) may 7, 1938 milton hopkins 3235 (okla) june 16, 1972 john & connie taylor 10794 (okla) september 4, 1977 t.a. zanoni 3349 (okla) polystichum acrostichoides (michx.) schott june 30, 1957 charles wallis 4451 (okla) woodsia obtuse (spreng.) torr. july 16, 1929 robert stratton 1722 (okla) july 8, 1957 charles wallis 4724 (okla) polypodiaceae pleopeltis polypodioides (l.) e.g. andrews & windham june 30, 1957 charles wallis 4455 (okla) ophioglossaceae botrychium virginianum (l.) sw. april 28, 1957 charles wallis 3660 (okla) pteridaceae adiantum capillus-verneris l. june 2, 1963 charles wallis 8760 (okla) july 7, 1950 u.t. waterfall 9570 (okla) june 16, 1972 john and connie taylor 10788 (okla) adintum pedatum l. may 4, 1928 robert stratton 804 (okla) may 7, 1938 milton hopkins 3254 (okla) cheilanthes lanosa (michx.) d.c. eaton august 11, 1932 featherly and still (okla) may 7, 1938 milton hopkins 3251 (okla) pellaea atropurpurea (l.) link july 7, 1950 u.t. waterfall 9560 (okla) thelypteridaceae phegopteris hexagonoptera (michx.) fee july 18, 1929 robert stratton (okla) oklahoma native plant record, volume 15, number 1, december 2015 oklahoma native plant record 19 volume 15, december 2015 chad b. king https://doi.org/10.22488/okstate.17.100112 forest structure and fire history at lake arcadia, oklahoma county, oklahoma (1820–2014) chad b. king department of biology university of central oklahoma edmond, ok 73034 cking24@uco.edu keywords: fire history, cross timbers, dendroecology abstract evidence indicates that the structure of oklahoma cross timbers forests are in transition due to changing climate, land-use patterns, and fire suppression efforts. however, only a handful of studies have addressed the history of fire across the oklahoma cross timbers landscape. this research adds to the body of literature by studying the contemporary forest structure and fire history at lake arcadia in oklahoma county, oklahoma. results demonstrate that post oak (quercus stellata wangenh.) and blackjack oak (q. marilandica münchh.), two common species in oklahoma cross timbers, dominate the forest. however, several mesophytic tree species are found in the overstory as well as the sapling layer of the forest. a total of 25 fire events (mean fire interval = 4.14 years) were documented during the 20th century using fire-scar analysis of q. stellata trees and remnant wood (stumps, snags, recently dead trees). high fire frequencies in the early to mid-20th century corresponded to the recruitment of q. stellata and q. marilandica. wet conditions (pdsi > 0) during the late 20th century and no fires after 1985 corresponded to the recruitment of non-oak, mesophytic species at the study site. the results of this study suggest that changes in fire frequency and moisture availability are contributing to changes in tree density and species composition at the study site. introduction fire has long been recognized as an important driver of forest dynamics (pyne 1982). in eastern north america, fire was a likely contributor to the development and sustainment of oak (quercus spp.) forests (abrams 1992). anthropogenic fire likely played a role in promoting upland oak forests, as well as changes in these forests (guyette et al. 2002). several upland quercus species benefit from and are adapted to frequent surface fires for their regeneration and recruitment in forests (abrams 1992). however, fire suppression during the 20th century has led to increasing densities of fire-sensitive, mesophytic tree species and a decline in quercus density (nowacki and abrams 2008). understanding the frequency of historic fires has an important role in explaining changes to contemporary forests. the result is a rich history of studies of fire history across the eastern united states (shumway et al. 2001; guyette et al. 2006; mcewan et al. 2007; king and muzika 2014; muzika et al. 2015; among others). one of the common patterns found in these studies is that surface fires were often frequent events prior to euro-american settlement of the area and that fire remained frequent during early euro-american settlement prior to fire suppression efforts in the early and mid-20th century. several interacting factors likely contributed to changing fire frequencies in eastern north american forests, including human density, topography, drought, and mailto:cking24@uco.edu 20 oklahoma native plant record volume 15, december 2015 chad b. king climate change (guyette et al. 2002; mcewan et al. 2011). recently, research on forest structure and dynamics in oklahoma cross timbers forests and savannas has highlighted the increase in fire-sensitive tree density and decrease in quercus density since the 1950s attributed to drought and fire suppression efforts (desantis et al. 2011). fire history studies in the oklahoma cross timbers have demonstrated frequent fires prior to euro-american settlement and a continued presence of fire on the landscape into the mid-20th century (shirakura 2003; clark et al. 2007; stambaugh et al. 2009; desantis et al. 2010a; allen and palmer 2011). this research adds to the growing body of literature of forest dynamics and fire history in the oklahoma cross timbers. preliminary investigation of the arcadia conservation education area in northeast oklahoma county revealed the presence of fire scarred trees and remnant wood indicative of historic fires at the site. this research had two objectives: 1) describe the contemporary forest structure by analyzing species composition, density, basal area, and age structure in the overstory and sapling layers of the forest and 2) relate the forest structure to the frequency of historic fires using dendrochronology. methods study site lake arcadia is an approximately 736 ha recreational and water supply lake located in northeastern oklahoma county. the army corps of engineers constructed the lake beginning in 1980 with the lake pool filling by 1987. the study site was located on the south side of the lake at the arcadia conservation education area (acea) (35o37’29”n, 97o23’16”w). the acea is an approximately 226 ha area administered by the oklahoma department of wildlife conservation since 1996; prescribed fire is not utilized at the site (d. griffith, area manager, pers. comm.). mean annual temperature is 15.63oc, and mean annual precipitation is 91.4 cm. annual precipitation is bimodal with the greatest amounts of precipitation during may-june and september-october (oklahoma climatological survey, www.mesonet.org). soils in this area are classified as stephenville-darnell-niotaze, characterized by shallow sandy to loamy soils (dominick 2003; carter and gregory 2008). elevations at the study site range from 308.5 m at the lake edge to 323.4 m at the southern boundary of the acea. preliminary investigation revealed fire scarred trees and remnant wood within a 43 ha area of the acea. the focus of this research was within the 43 ha area to study the fire history and forest composition and structure. forest composition, age structure, and radial growth stand structure data were collected on twenty 0.04 ha fixed-area plots located randomly within the 43 ha study area. within each plot, the diameter at breast height (dbh) of all overstory trees (dbh >10 cm) was measured, and trees were identified to species. for each species in the overstory, estimates of relative density (trees/ha), relative dominance (basal area/ha), and relative importance were calculated to describe the contemporary composition of the forest overstory. increment cores were collected at 30 cm above the ground from two to four of the largest overstory trees per plot for estimates of age structure and radial growth at the study site. tree selection was based on the development of the longest tree-ring chronology for the site which can limit age structure interpretation. a total of 71 increment cores were collected from acea. two 0.01 ha fixed-area subplots were established in each 0.04 ha overstory plot to analyze the species composition and density of saplings (dbh <10 cm, >1.37 m height). http://www.mesonet.org/ oklahoma native plant record 21 volume 15, december 2015 chad b. king saplings were identified to species and counted within each subplot. cross-sections of one to two saplings were collected from paired subplots to study the age structure and radial growth of saplings. increment cores were returned to the university of central oklahoma where they were mounted and sanded with progressively finer sandpaper (80-grit to 1200-grit) in order to see individual tree-ring boundaries and cellular structure (stokes and smiley 1996). cross-dating procedures were used to confidently assign calendar years to each tree-ring on an increment core. individual ring-width measurements, to the nearest 0.01mm, were collected on each sample using a velmex ta unislide system (velmex, inc., bloomfield, ny), binocular microscope, and j2x measurement software (voortech consulting, holderness, nh). tree-ring series measurements were compared graphically, using the list method (yamaguchi 1991), and statistically using the program cofecha (holmes 1983; grissino-mayer 2001a). following crossdating and assignment of calendar years to each tree-ring, pith dates were recorded for age estimation at coring height and tree cohort establishment at the study site. in the event that the pith was missed in an increment core, the methods of duncan (1989) were used to estimate the number of tree-rings missed to the pith of the tree. fire history cross-sections were collected selectively from q. stellata remnant wood to study the fire history of the site. quercus stellata has been used successfully for fire history studies in oklahoma cross timbers (clark et al. 2007; stambaugh et al. 2009; desantis et al. 2010b; allen and palmer 2011). the analysis approach of guyette and stambaugh (2004) was used to identify fire scars in q. stellata. in their study, fire scars were identified based on bark fissure patterns, common in oak species (smith and sutherland 2001), and scarring that occurs across multiple samples during the same year. a total of 21 samples exhibited scarring associated with surface fires, including 13 recently dead q. stellata, two saplings that demonstrated fire scars, and six snags. three samples could not be successfully cross-dated. all samples were sanded with progressively finer sandpaper (80-grit to 1200-grit). ring-widths for each remnant sample were measured using the velmex ta unislide system (velmex, inc., bloomfield, ny), binocular microscope, and j2x measurement software (voortech consulting, holderness, nh). based on cross-dating, calendar years for each treering on fire-scarred samples were assigned using correlation analysis with a master treering chronology created from 39 cross-dated q. stellata tree-ring series from the study site. calendar years were assigned to each identified fire scar on a sample. a fire chronology was created based on all fire scars for analysis of fire frequency (mean fire interval) and fire severity (fire years in which >25% samples were scarred) using the program fhx2 (grissino-mayer 2001b). superposed epoch analysis (grissino-mayer 2001b) was used to test the association of fire year and drought. instrumental palmer drought severity index (palmer 1965) data for the time period 1895 to 2013 from oklahoma climate region 5 were used to associate fire year and drought. an average was calculated for reconstructed palmer drought severity index (cook et al. 2004) for gridpoint 178 and 179 for purposes of comparing drought and growth of trees prior to 1895. reconstructed palmer drought severity indices are reconstruction models based on the association of instrumental palmer drought severity indices and regional tree-ring chronologies (cook et al. 1999). 22 oklahoma native plant record volume 15, december 2015 chad b. king results a total of nine species were identified in the overstory at lake arcadia. quercus stellata was the most dominant species, but q. marilandica had the highest density. overall, these two species accounted for 88% of the basal area and 68% of the overstory tree density at the study site. two celtis species (c. laevigata willd.; c. occidentalis l.) combined had the third highest relative tree density (15.1%) and relative dominance (5.51%) (table 1). table 1 overstory (dbh >10 cm) statistics and sapling (dbh <10 cm, >1.37 m height) density at lake arcadia, oklahoma county, oklahoma. relative importance value = (relative density + relative dominance)/2. species trees/ha relative density basal area (m2/ha) relative dominance relative importance value sapling density (stems/ha) quercus stellata 95 29.2 14.5 70.1 49.7 576 quercus marilandica 126 38.8 3.66 17.7 28.3 480 celtis laevigata 28 8.62 0.71 3.47 6.04 192 juniperus virginiana 25 7.69 0.78 3.77 5.73 384 celtis occidentalis 21 6.46 0.42 2.05 4.25 1056 ulmus rubra 15 4.62 0.50 2.41 3.51 192 sideroxylon lanuginosum 7 2.15 0.06 0.27 1.21 --ulmus americana 4 1.23 0.03 0.12 0.68 96 sapindus drummondii 4 1.23 0.01 0.05 0.64 --cornus drummondii ----------384 cercis canadensis ----------192 quercus muehlenbergii ----------96 celtis reticulata ----------96 total 325 100 20.6 100 100 3744 a total of 11 species was found in the sapling layer (see table 1). the sapling layer was rather dense (3,744 stems/ha). celtis occidentalis had the highest sapling density (1,056 stems/ha), and the three celtis species accounted for 35% of the sapling density at lake arcadia. approximately 78% of the overstory tree species was also found in the sapling layer; the exceptions were sapindus drummondii hook & arn. and sideroxylon oklahoma native plant record 23 volume 15, december 2015 chad b. king lanuginosum michx. two species, that have the potential of growing up to the existing overstory, were identified in the sapling layer but were not found in the overstory (c. reticulata torr.; q. muehlenbergii engelm.). a total of 137 samples from nine species was collected for analysis of forest age structure, radial growth, and fire history at lake arcadia. quercus stellata and q. marilandica accounted for 71% (n = 97) of the samples. increment cores were collected from q. stellata (n = 39), q. marilandica (n = 16), s. drummondii (n = 1), ulmus americana (l.) (n = 1), u. rubra (muhl.) (n = 3), s. lanuginosum (n = 2), c. laevigata (n = 3), and c. occidentalis (n = 6). sapling cross-sections were collected from q. stellata (n = 5), q. marilandica (n = 16), s. lanuginosum (n = 2), c. laevigata (n = 8), c. occidentalis (n = 10), and juniperus virginiana (l.) (n = 4) for estimates of sapling age and radial growth. a total of 21 q. stellata samples exhibited scarring associated with surface fires. two q. stellata saplings exhibited fire scars. the largest diameter tree in our study plots was a q. stellata that measured 67.5 cm dbh. the oldest tree was a q. stellata that was 193 years old (1821–2014). however, only 23.3% of q. stellata trees dated prior to the 20th century (fig. 1). the oldest q. marilandica in our study plots was 108 years old (1906–2014). q. stellata demonstrated continuous recruitment beginning in the 1880s, with the 1910s having the recruitment of a large cohort (see fig. 1). q. marilandica also demonstrated continuous recruitment during the early and mid-20th century. the oldest non-quercus individual in the overstory was a c. occidentalis that was 62 years old (1952–2014). the age structure of the non-quercus species in the overstory (s. drummondii, u. americana, u. rubra, s. lanuginosum, c. laevigata, c. occidentalis) indicated continuous recruitment beginning in the 1950s and peaking during the 1980s (see fig. 1). figure 1 age structure of q. stellata, q. marilandica, and non-oak species. non-oak species include s. lanuginosum, c. laevigata, c. occidentalis, s. drummondii, u. americana, u. rubra. arrows indicate the year of a fire. bottom graph represents reconstructed (dashed line) and instrumental (full line) palmer drought severity index (pdsi) for central oklahoma. 24 oklahoma native plant record volume 15, december 2015 chad b. king the oldest sapling in the understory of the study plots was a q. marilandica that was 62 years old (1952–2014). approximately 49% (n = 19) of non-oak saplings recruited during the 1980s (fig. 2). establishment of non-oak species appeared to correspond to periods of above-average pdsi following the 1960s (see figs. 1, 2). figure 2 age structure of q. stellata, q. marilandica, and non-oak saplings at lake arcadia. non-oak species includes s. lanuginosum, c. laevigata, c. occidentalis, and j. virginiana. arrows represent years of fire. bottom graph is the instrumental pdsi for central oklahoma (1952-2014). fifty-one fire scars were identified and dated, that occurred from 25 different fire events (fig. 3). the earliest fire occurred in 1844 with a range of fire years from 1844 to 1985. however, the 1844 fire scar was not used in any of the fire analyses due to a low sample depth during that time period it and being represented on only one sample. approximately 29.7% of years 1898 to 1985 had a fire. the most severe fire years (based on percentage of trees scarred) included 1898 (33.3%), 1912 (55.6%), 1922 (41.7%), and 1955 (41.7%). the mean fire interval (mfi) for all fires from 1898 to 1985 was 4.14 years (sd ± 2.22, range 2–9 years). superposed epoch analysis was conducted to test the association between drought and any fire year. results indicated no significant association between any fire year (1898 to 1985) and drought (fig. 4). severe fires oklahoma native plant record 25 volume 15, december 2015 chad b. king during 1912 and 1955 were associated with extreme drought conditions (pdsi < -2.99). for the period 1898 to 1985, 13 fires occurred during dry conditions (pdsi < 0), and 11 fires occurred during wet conditions (pdsi > 0). discussion changes in historic fire regimes and land-use patterns often lead to changes in forest structure and composition. in the cross timbers region of oklahoma, changes in forest structure and composition are apparent in terms of increasing tree density, particularly increases in fire sensitive tree species (desantis et al. 2010a, 2011). the result of changing historic forest dynamics is the “mesophication” (nowacki and abrams 2008) of cross timbers forests. this study demonstrates the continued dominance of q. stellata and q. marilandica in the overstory of this cross timbers forest. however, this study also highlights the effect of a changing fire regime on forest structure at the study site. total basal area for this study is similar to other studies across multiple sites in the oklahoma cross timbers (desantis et al. 2010a, 2011) and arkansas cross timbers (bragg et al. 2012). desantis et al. (2010a) demonstrate increases in non-oak basal area and tree density across multiple sites in oklahoma between the 1950s and 2000s. this study shows that celtis species collectively make up the third most important group at the study site (see table 1). the two celtis species, juniperus virginiana and ulmus species, in this study along with the other non-oak species are sensitive to fire as seedlings and saplings (coladonato 1992, 1993; sullivan 1993; anderson 2003; gucker 2011). generally, only the most severe fires will kill overstory trees of these species. figure 3 fire history graph for lake arcadia in northeastern oklahoma county, oklahoma. horizontal lines represent the length of the tree-ring record for each sample (n = 18). vertical dashes represent the year of a fire scar in each tree-ring record. the composite fire chronology is represented by the fire years. 26 oklahoma native plant record volume 15, december 2015 chad b. king figure 4 superposed epoch analysis for all fires from 1898 to 1985 compared to pdsi (drought). this program analyzes the relationship between any fire year and drought (grissino-mayer 2001b). year “0” is the year of any fire year; year “-1” indicates the departure from the mean pdsi one year prior to any fire year. horizontal lines represent 95% confidence interval based on 1000 monte carlo simulations. this study highlights the recruitment of a large number of non-oak trees during the 1980s (see figs. 1, 2). there are two factors which likely contributed to this recruitment. the last fire that was documented at lake arcadia occurred in 1985 (see fig. 3). additionally, following the drought during the late 1970s and early 1980s in the central oklahoma region was an 18 year period (1982–2000) of above-average pdsi (see figs. 1, 2). this 18 year period along with no fires after 1985 likely contributed to recruitment of these non-oak species. the data also show recruitment of non-oak trees following the 1950s drought (see figs. 1, 2). desantis et al. (2011) found increases in species recruitment following drought during the 1950s and decreases in quercus recruitment associated with fire suppression. clark et al. (2007) indicated increased recruitment of j. virginiana during fire free periods and increased recruitment of quercus species following frequent fires. the results of this study also suggest that fire-free periods (between 1955 and 1964; post-1985) (see fig. 3) contributed to non-oak recruitment at lake arcadia. the 1964 and 1967 fires are represented on only one sample, which may suggest that these fires were of low severity and had little effect on non-oak recruitment during this time oklahoma native plant record 27 volume 15, december 2015 chad b. king period. recruitment of q. stellata during the early 20th century occurred during high fire frequency (1905–1926, mfi = 2.62 years). following 1926, the number of fires declined to seven in a 38 year period (1926– 1964). the current q. marilandica overstory recruited during the mid-20th century period, which coincided with surface fires. the fire frequency at lake arcadia (mfi = 4.14 years) is within the range of other studies in the cross timbers and other mixed-species forests of oklahoma. desantis et al. (2010b) in okmulgee county reported an mfi equal to 2.7 years for the time period 1750 to 2005. when considering a similar time period to this study, they report an mfi of 2 years. clark et al. (2007) indicated a range of fire frequency between 2.5 and 6 years (1770– 2002) based on the aspect of the forest stand at sites in osage county. allen and palmer (2011) report an mfi for all fires of 2.3 years (1729–2005) at a different site in osage county. stambaugh et al. (2009) at the wichita mountains national wildlife refuge found an mfi of 4.7 years for all fires between 1722 and 2001. at the nickel family nature and wildlife preserve in northeastern oklahoma, stambaugh et al. (2013) found a fire frequency of 2.6 years in a mixed oak-pine (quercus-pinus) forest. masters et al. (1995) in a study of fire history in mccurtain county reported a mean fire interval of 3.8 years. comparing the association between drought and fire year revealed no significant association at lake arcadia (see fig. 4). this result is similar to other studies in the oklahoma cross timbers (allen and palmer 2011; desantis et al. 2010b; stambaugh et al. 2009) and contrary to that reported by clark et al. (2007). three of four severe fire years (1898, 1912, 1955) coincided with below average pdsi (drought) conditions. the exception was the 1922 severe fire year which coincided with above average pdsi. in all previous studies of fire history in the oklahoma cross timbers, fires were found to be frequent events prior to euroamerican settlement (<1890) and after euroamerican settlement (>1890). there is a noticeable lack of fires between 1844 and 1898 (see fig. 3). there are several possible explanations for the absence of fire scars. not every fire which occurs at a site will result in the formation of a fire scar. most remnant samples had only heartwood present that often resulted in too few tree-rings to accurately cross-date. decomposition of the heartwood was also a common feature of the trees at lake arcadia that possibly resulted in the loss of fire scars that were present during the mid and late 19th century. however, even with the limited temporal scope of the fire history, this study demonstrates frequent fires at the lake arcadia area during the 20th century. conclusions fire was likely an important factor that sustained the dominance of quercus species in upland forests (abrams 1992). while this study has some limitations, it does highlight quercus recruitment coincided with frequent fires during the 20th century. changes in fire frequency after 1985 and fire-free periods promoted non-oak recruitment in the understory, similar to other studies in the oklahoma cross timbers (clark et al. 2007) and across other upland quercus forests in the eastern united states (abrams 1992). studies of fire history are important for understanding forest development, the historical role of humans on the landscape, and the development of management guidelines for sites which utilize prescribed fire. this study adds to the growing knowledge of historic fire frequency in the oklahoma cross timbers. fires were frequent events that shaped the historic cross timbers, and often the high frequencies continued into the mid and late 20th century. comparatively, the number of 28 oklahoma native plant record volume 15, december 2015 chad b. king studies that have specifically addressed fire history in the oklahoma cross timbers is limited. other sites should be selected and studied to further expand the knowledge of historic fire on the cross timbers landscape. acknowledgements i would like to thank justin cheek and shey ramsey at the university of central oklahoma for their assistance in the field and laboratory. i would like to graciously thank daniel griffith at the arcadia conservation education area for permission to collect samples at the study site. thanks to two anonymous reviewers for constructive comments and suggestions on an earlier version of this manuscript. this research was supported by the office of research and grants at the university of central oklahoma. literature cited abrams, m.d. 1992. fire and the development of oak forests. bioscience 42:346–353. allen, m.s. and m.w. palmer. 2011. fire history of a prairie/forest boundary: more than 250 years of frequent fire in a north american tallgrass prairie. journal of vegetation science 22:436–444. anderson, m.d. 2003. juniperus virginiana. in: fire effects information system. u.s. department of agriculture, forest service, rocky mountain research station, fire sciences laboratory (producer). http://www.fs.fed.us/database/feis/ (16 july 2015). bragg, d.c., d.w. stahle, and k.c. cerny. 2012. structural attributes of two oldgrowth cross timbers stands in western arkansas. american midland naturalist 167:40–55. carter, b.j. and m.s. gregory. 2008. soil map of oklahoma. norman (ok): oklahoma geological survey. clark, s.l., s.w. hallgren, d.m. engle, and d.w. stahle. 2007. the historic fire regime on the edge of the prairie: a case study from the cross timbers of oklahoma. pp. 40–49. in: masters, r.e., k.e.m. galley (eds.). proceedings of the 23rd tall timbers fire ecology conference: fire in grassland and shrubland ecosystems. tallahassee (fl): tall timbers research station. coladonato, m. 1992. ulmus americana. in: fire effects information system. u.s. department of agriculture, forest service, rocky mountain research station, fire sciences laboratory (producer). http://www.fs.fed.us/database/feis/ (16 july 2015). coladonato, m. 1993. ulmus rubra. in: fire effects information system. u.s. department of agriculture, forest service, rocky mountain research station, firesciences laboratory (producer). http://www.fs.fed.us/database/feis/ (16 july 2015). cook, e.r., d.m. meko, d.w. stahle, and m.k. cleaveland. 1999. drought reconstructions for the continental united states. journal of climate 12:1145– 1162. cook, e.r., d.m. meko, d.w. stahle, and m.k. cleaveland. 2004. north american summer pdsi reconstructions. igbp pages/world data center for paleoclimatology data contribution series. boulder (co): noaa ngdc paleoclimatology program. desantis, r.d., s.w. hallgren, t.b. lynch, j.a. burton, and m.w. palmer. 2010a. long-term directional changes in upland quercus forests throughout oklahoma, usa. journal of vegetation science 21:606– 615. http://www.fs.fed.us/database/feis/ http://www.fs.fed.us/database/feis/ http://www.fs.fed.us/database/feis/ oklahoma native plant record 29 volume 15, december 2015 chad b. king desantis, r.d., s.w. hallgren, and d.w. stahle. 2010b. historic fire regime of an upland oak forest in south-central north america. fire ecology 6:45–61. desantis, r.d., s.w. hallgren, and d.w. stahle. 2011. drought and fire suppression lead to rapid forest composition change in a forest-prairie ecotone. forest ecology and management 261:1833–1840. dominick, m.d. 2003. soil survey of oklahoma county. united states department of agriculture, natural resources conservation service. duncan, r.p. 1989. an evaluation of error in tree age estimates based on increment cores in kahikatea (dacrycarpus dacrydioides). new zealand natural sciences 16:31–37. grissino-mayer, h.d. 2001a. evaluating crossdating accuracy: a manual and tutorial for the computer program cofecha. tree-ring research 57:205– 221. grissino-mayer, h.d. 2001b. fhx2 software for analyzing temporal and spatial patterns in fire regimes from tree rings. tree-ring research 57:115–124. gucker, c.l. 2011. celtis occidentalis. in: fire effects information system. u.s. department of agriculture, forest service, rocky mountain research station, fire sciences laboratory (producer). http://www.fs.fed.us/database/feis/ (16 july 2015). guyette, r.p, r.m. muzika, and d.c. dey. 2002. dynamics of an anthropogenic fire regime. ecosystems 5:472–486. guyette, r.p. and m.c. stambaugh. 2004. post oak fire scars as a function of diameter, growth, and tree age. forest ecology and management 198:183–192. guyette, r.p., d.c. dey, m.c. stambaugh, and r.m. muzika. 2006. fire scars reveal variability and dynamics of eastern fire regimes. pp. 20–39. in: dickinson, m.b. (ed.). proceedings of the conference: fire in eastern oak forests: delivering science to land managers. vol. gtrnrs-p-1. columbus (oh): ohio state university. u.s. department of agriculture, u.s. forest service, northern research station. holmes, r.l. 1983. computer-assisted quality control in tree-ring dating and measurement. tree-ring bulletin 43:69– 78. king, c.b. and r.m. muzika. 2014. historic fire and canopy disturbance dynamics in an oak-pine (quercus-pinus) forest of the missouri ozarks. castanea 79:78–87. masters, r.e., j.e. skeen, and j. whitehead. 1995. preliminary fire history of mccurtain county wilderness area and implications for red-cockaded woodpecker management. pp. 290–302. in: kulhavy, d.l., r.g. hooper, and r. cost (eds.). red-cockaded woodpecker symposium iii: recovery, ecology and management, 24–28 january 1993, north charleston, south carolina. nacogdoches (tx): stephen f. austin state university, college of forestry, center for applied studies in forestry. mcewan, r.w., t.f. hutchinson, r.p. long, d.r. ford, and b.c. mccarthy. 2007. temporal and spatial patterns of fire occurrence during the establishment of mixed-oak forests in eastern north america. journal of vegetation science 18:655–664. mcewan, r.w., j.m. dyer, and n. pederson. 2011. multiple interacting ecosystem drivers: toward an encompassing hypothesis of oak forest dynamics across eastern north america. ecography 34:244–256. muzika, r.m., r.p. guyette, m.c. stambaugh, and j.m. marschall. 2015. fire, drought, and humans in a heterogeneous lake superior landscape. journal of sustainable forestry 34:49–70. nowacki, g.j. and m.d. abrams. 2008. the demise of fire and “mesophication” of http://www.fs.fed.us/database/feis/ 30 oklahoma native plant record volume 15, december 2015 chad b. king forests in the eastern united states. bioscience 58:123–138. palmer, w.c. 1965. meteorological drought. research paper 45. washington (dc): u.s. department of commerce, weather bureau. pyne, s.j. 1982. fire in america: a cultural history of wildland and rural fire. princeton (nj): princeton university press. shirakura, f. 2003. tornado damage and fire history in the cross timbers of the tallgrass prairie preserve, oklahoma [master’s thesis]. stillwater (ok): oklahoma state university. shumway, d.l., m.d. abrams, and c.m. ruffner. 2001. a 400-year history of fire and oak recruitment in an old-growth forest in western maryland, u.s.a. canadian journal of forest research 31:1437–1443. stambaugh, m.c., r.p. guyete, r. godfrey, e.r. mcmurry, and j.m. marschall. 2009. fire, drought, and human history near the western terminus of the cross timbers, wichita mountains, oklahoma, usa. fire ecology 5:51–64. smith, k.t. and e.k. sutherland. 2001. terminology and biology of fire scars in selected central hardwoods. tree-ring research 57:141–147. stokes, m.a. and t.l. smiley. 1996. an introduction to tree-ring dating. chicago (il): university of chicago press. sullivan, j. 1993. celtis laevigata. in: fire effects information system. u.s. department of agriculture, forest service, rocky mountain research station, fire sciences laboratory (producer). http://www.fs.fed.us/database/feis/ (16 july 2015). yamaguchi, d.k. 1991. a simple method for cross-dating increment cores from living trees. canadian journal of forest research 21:414–416. http://www.fs.fed.us/database/feis/ forest structure and fire history at lake arcadia, oklahoma county, oklahoma (1820–2014) by dr. chad king journal of the oklahoma native plant society, volume 8, number 1, december 2008 45 oklahoma native plant record volume 8, number 1, december 2008 carter, et al. https://doi.org/10.22488/okstate.17.100061 an updated flora of the wichita mountains wildlife refuge keith a. carter, pablo rodriguez, and michael t. dunn 1. department of biological sciences, cameron university, lawton, oklahoma 73505 1 author for correspondence: phone 580-581-2287; e-mail: michaeld@cameron.edu the herbarium collections of the wichita mountains wildlife refuge have been transferred to the cameron university herbarium (camu) so that they could be safely curated, and electronically databased and still remain accessible to refuge personnel while for the first time becoming readily available to other interested researchers. this paper is a report on the initial inventory of the specimens. the 1784 specimen collection includes 101 families, 339 genera, and 634 species that have been physically repaired and taxonomically updated as needed, accessioned into the camu collections, and entered into the specify database. introduction the wichita mountains are some of the oldest exposed mountains in the world and because the area was too rocky to plow, they formed a natural refugium that preserved what is arguably the largest remaining intact tract of southern mixed-grass prairie in existence. the mountains were part of the kiowa-comanche-apache reservation in the late 19th century. when the reservation was opened to settlement in 1901, the land was set aside by the federal government. originally administered by the department of the interior, jurisdiction was transferred to the forest service in 1905, and in 1935 management of the wichita mountains wildlife refuge (wmwr) was transferred to what is now the fish and wildlife service (morgan, 1973). in 1907 bison were reintroduced to the refuge and in 1927 congress issued a mandate to preserve the bloodline of texas longhorn cattle. elk, which had been extirpated by 1875 were transplanted from jackson hole, wyoming, and today in addition to the buffalo, longhorn cattle, and elk that get most of the public’s attention, the refuge is home to over 50 mammal species including prairie dogs, coyotes, bobcats, and mountain lions (tyler, 2005). in addition, over 240 bird, 64 reptile and amphibian, and 36 fish species have been identified. eight hundred and six vascular plants have been identified at the wmwr (eskew, 1938; osborn and allan, 1949; buck, 1977). much of the natural history of the refuge is recorded in herbarium collections that were housed in the basement of the headquarters building. in 2005, refuge management recognized the need to protect the specimens, and make the data available to the scientific community as well as the general public, but still keep the data accessible to refuge biologists and technicians. the only facility that met all of the criteria was the cameron university herbarium (camu), and in 2006 the specimens were transferred to camu as a permanent loan. in 2008 the national science foundation (nsf) provided funding to procure additional cabinets and equipment and to hire student workers to enter the specimens into the specify database. this paper is the first report of the inventory of these specimens and will serve as a benchmark for future studies that will update the complete flora of the refuge. the 23,885 hectare wichita mountains wildlife refuge is located in northwestern comanche county, oklahoma (fig.), ranging from 34°41’n to 34°50’n and 98°48’30”w to 98°30’30”w. elevation ranges from 404 m (1330 ft) where cache creek crosses the wmwr southern boundary to 756 m (2479 ft) at the summit of mt. pinchot. the mountains themselves are predominantly cambrian igneous rock and the surrounding 46 oklahoma native plant record volume 8, number 1, december 2008 carter, et al. plains are predominantly permian sedimentary rock (price and gilbert, 1996). the ecoregion is categorized as great plains steppe shrub province (bailey, 1995) or central great plains (woods et al., 2005) and receives on average 86.84 cm (34.19 in) of precipitation annually, with may the wettest month (mean 13.03 cm [5.13 in]) and january the driest (mean 3.50 cm [1.38 in]). mean annual temperature is 22.220c (72.00f) (oklahoma climatological survey, 2007). materials and methods a total of 1784 specimens were accepted as a permanent loan from wmwr to camu in june 2006. nsf-biological research collections funds were awarded in 2008 and were used to purchase new herbarium cabinets and begin curation during that summer. specimens were first triaged for damage and physically repaired as necessary. preliminary identification and taxonomic updating were the responsibility of kac. specimens were then entered into the specify database by kac and pr. all identifications and taxonomic updates were then verified by mtd before annotations were added and specimens were fumigated and curated into separate color coded genus folders. because taxonomy for many of the specimens is ambiguous and no completed treatment of the flora of oklahoma was broadly accepted, a combination of mcgregor et al., (1986), diggs et al (1999) and judd et al., (2008) was used to update the taxonomy of the specimens (details available upon request). results and discussion three of the 1784 specimens were collected outside the wmwr proper but all three have duplicates collected on the refuge. the collection includes 101 families, 339 genera, and 634 species (appendix), including: 1 family of charophyceans, 6 families of seedless vascular plants, 2 gymnosperm families, 1 basal angiosperm, 16 monocot families and 75 eudicot families. the largest families are asteraceae with 88 species and poaceae with 99 species. the largest monocot genera are carex and eragrostis with 12 species each. the largest herbaceous and woody eudicots are respectively polygonum with nine and quercus with eight species each. now that these preliminary data have been compiled, we hope to expand the project by updating the taxonomy of the classic buck (1977) report on the flora of wmwr to enable direct comparison with this assemblage, quantify the percentage of exotics in the collections, and with the permission of wmwr biologists and administrators, begin surveying the refuge for some of the rarer plants in the collection to identify those taxa in danger of extirpation. acknowledgements we would like to acknowledge nsf-dbibrc grant 0749657 for funding this research, and sam waldstein (former refuge manager), ralph bryant acting refuge manager (2006), jeff rupert (refuge manager) and walter munsterman (wildlife biologist) for their support of this project. in addition, donna lohr, amber roy, and tom sodhi assisted as student researchers. references bailey rg 1995. description of the ecoregions of the united states. www.fs.fed.us/land/ecosysmgmt/index .html buck p 1977. vascular plants of the wichita mountains wildlife refuge 1977. unpublished informational handout of the wichita mountains wildlife refuge. reprinted in: oklahoma native plant record, 2 (1): 4-21, 2002. diggs gm jr., lipscomb bl, and o’kennon rj. 1999. shinners & mahler’s illustrated flora of north central texas. botanical research institute of texas, fort worth, tx: 1626p. eskew ct 1939. the flowering plants of the wichita mountains wildlife refuge, oklahoma. american midland naturalist, 20: 695-703. judd ws, campbell cs, kellogg ea, stevens pf, and donoghue mj. 2008. plant systematics, a phylogenetic approach, 3rd ed., sinauer associates, sunderland, massachusetts: 611p. 47 oklahoma native plant record volume 8, number 1, december 2008 carter, et al. mcgregor rl, barkley tm, brooks re, and schofield ek. 1986. flora of the great plains. university of kansas press, lawrence, kansas: 1402p. morgan eb, 1973. the wichita mountains, ancient oasis of the prairies. texan press, waco, texas: 253p. oklahoma climatological survey. 2007. oklahoma climatological data. university of oklahoma, norman. (www.ocs.ou.edu). osborn b and allan pf. 1949. vegetation of an abandoned prairie dog town in tallgrass prairie. ecology, 30:322-332. price jd and gilbert mc. 1996. geologic map of the mount scott area, eastern wichita mountains, oklahoma. u.s.g.s. national cooperative mapping program. specify software project, biodiversity research center, university of kansas, lawrence, ks 66045. www.specifysoftware.org tyler jd 2005. birds of southwestern oklahoma and north central texas. transcript press norman, oklahoma: 119p. woods aj, omernik jm, butler dr, ford jg, henley je, hoagland bw, arndt ds, and moran bc. 2005. ecoregions of oklahoma (color photo with map, descriptive text, summary tables, and photographs). u.s. geological survey, reston, virginia. figure location of wichita mountains wildlife refuge, comanche county, oklahoma 48 oklahoma native plant record volume 8, number 1, december 2008 carter, et al. appendix green algae characeae chara vulgaris l. seedless vascular plants isoetaceae isoetes melanopoda gay & dur. equisetaceae equisetum laevigatum a. braun aspleneaceae asplenium trichomanes l. dryopteridaceae dryopteris marginalis (l.) a. gray woodsia obtusa (spreng.) torr. marsileaceae pilularia americana a. braun marsilea vestita hood. and grev. pteridaceae cheilanthes eatoni baker cheilanthes lanosa (michx.) d.c. eaton cheilanthes tomentosa link. pellaea atropurpurea (l.) link. pellaea wrightiana hook. gymnosperms cupressaceae juniperus virginiana l. pinaceae pinus elliottii englm. angiosperms monocots agavaceae manfreda virginica (l.) salisb. ex rose alismataceae echinodorus berteroi (spreng.) fassett sagittaria latifolia willd. sagittaria montevidensis cham. & schlecht. araceae arisaema dracontium (l.) schott commelinaceae commelina erecta l. tradescantia occidentalis (britt.) smyth tradescantia ohiensis raf. cyperaceae bulboschoenus maritimus (l.) palla bulbostylis capillaris (l.) kunth ex c.b. clarke carex amphibola steud. carex annectens (bickn.) bickn. carex austrina mack. carex blanda dewey carex emoryi dewey carex festucacea schkuhr. ex willd. carex frankii kunth. carex gravida bailey carex grisea wahl. carex microrhyncha mack. carex muehlenbergii schkuhr ex willd. carex vulpinoidea michx. cyperus acuminatus torr. & hook. ex torr. cyperus echinatus (l.) wood cyperus erythrorhizos muhl. cyperus esculentus l. cyperus lupulinus (spreng.) marcks. cyperus odoratus l. cyperus pseudovegetus steud. cyperus schweinitzii torr. cyperus setigerus torr. & hook cyperus squarrosus l. cyperus strigosus l. eleocharis acutisquamata buckley eleocharis compressa sulliv. eleocharis engelmanni steud. eleocharis montevidensis kunth. eleocharis obtusa (willd.) schult. eleocharis palustris (l.) roem. & schult. eleocharis parvula (roemer & j.a. schutles) link ex bluff, nees & schaeur eleocharis quadrangulata fern. eleocharis tenuis (willd.) schult. eleocharis wolfii (a.gray) a.gray ex britton fimbristylis puberula (michx.) vahl. fimbristylis vahlii (lam.) link. 49 oklahoma native plant record volume 8, number 1, december 2008 carter, et al. fuirena simplex vahl. lipocarpha micrantha (vahl.) tucker schoenoplectus acutus (muhl ex bigelow) a. love & d. love schoenoplectus pungens (vahl.) palla schoenoplectus tabernaemontani (k.c. gmel.) palla scleria pauciflora muhl. scirpus atrovirens muhl. scirpus pendulus muhl. hydrocharitaceae najas guadalupensis (spreng.) morong. iridaceae sisyrinchium angustifolium mill. juncaceae juncus acuminatus michx. juncus dudleyi wiegand. juncus interior wieg. juncus marginatus rostk. juncus tenuis woot. & standl. juncus torreyi coville lemnaceae lemna minor l. liliaceae allium canadense l. allium drummondii regel. allium stellatum ker. androstephium coeruleum (scheele) greene camassia scilloides (raf.) cory cooperia drummondii herbert erythronium americanum ker. nothoscordum bivalve greene ex rydb. orchidaceae spiranthes magnicamporum sheviak. poaceae agrostis hyemalis (walter) b.s.p. agrostis elliottiana schult. alopecurus carolinianus walt. andropogon gerardii vitman. aristida dichotoma michx aristida longespica poir. aristida oligantha michx. aristida purpurea nutt. bothriochloa barbinodes (lag.) herter. bothriochloa laguroides (d.c.) herter. bouteloua curtipendula (michx) torr. bouteloua gracilis (willd ex kunth) lag. ex griffiths bouteloua hirsuta lag. bouteloua rigidiseta (steud.) hitchc. bromus arvensis l. bromus catharticus vahl bromus commutatus schrad. bromus japonicus thunb. ex murray bromus pubescens muhl. ex willd. buchloe dactyloides (nutt.) engelm. cenchrus spinifiex cav. chasmanthium latifolium (mickx.) h.o. yates chloris verticillata nutt. chloris virgata sw. coelorachis cylindrica (michx.) nash cynodon dactylon (l.) pers. dactylis glomerata l. dichanthelium acuminatum (sw.) gould & c.a. clark dichanthelium depauperatum (muhl.) gould dichanthelium linearifolium (scribn. ex nash) gould dichanthelium malacophyllum (nash) gould dichanthelium oligosanthes (j.a. schultes) gould digitaria californica (benth.) henr. digitaria cognata (schultes) pilger digitaria sanguinalis (l.) scop. echinochloa crus-galli (l.) beauv. eleusine indica (l.) gaertn. elymus canadensis l. elymus virginicus l. eragrostis capillaris (l.) nees eragrostis cilianensis (all.) vignalo ex janch eragrostis curtipedicellata buckley eragrostis hypnoides (lam.) b.s.p. eragrostis intermedia hitchc. eragrostis pectinacea (mickx.) nees ex steud. eragrostis pilosa (l.) beauv. eragrostis reptans (michx.) nees eragrostis secundiflora j. presl. eragrostis sessilispica buckley eragrostis spectabilis (pursh.) steud. eragrostis trichodes (nutt.) a.w. wood eriochloa contracta hitchc. eriochloa sericea (scheele) munro ex vasey erioneuron pilosum (buckley) nash festuca versuta beal hordeum pusillum nutt. 50 oklahoma native plant record volume 8, number 1, december 2008 carter, et al. leersia oryzoides (l.) sw. leptochloa fascicularis (lam.) a. gray limnodea arkansana (nutt.) l.h. dewey lolium pratense (huds.) s.j.darbyshire melica nitens (scribn) nutt. ex piper muhlenbergia capillaris (lam.) trin. muhlenbergia mexicana (l.) trin. muhlenbergia racemosa (michx) b.s.p. panicum anceps michx. panicum capillare l. panicum dichotomiflorum michx. panicum obtusum kunth. panicum philadelphicum bernh. ex trin. panicum virgatum l. pascopyrum smithii (rydb.) a. love paspalum dilatatum poir. paspalum pubiflorum rupr. paspalum setaceum michx. phalaris caroliniana walter poa annua l. poa arachnifera torr. poa compressa l. schedonnardus paniculatus (nutt.) trel. setaria parviflora (poir) kerguélen setaria viridis (l.) p. beauv. schizachyrium scoparium (michx.) nash sorghastrum nutans (l.) nash sorghum halepense (l.) pers. spartina pectinata link sphenopholis obtusata (michx) scribn. sporobolus airoides torr. sporobolus clandestinus (biehler) hitchc. sporobolus compositus (poir.) merr. sporobolus cryptandrus (torr) a. gray sporobolus neglectus nash sporobolus pyramidatus (lam.) hitchc. sporobolus vaginiflorus (torr ex a. gray) a.w. wood tridens albescens (vasey) wooton & standl. tridens flavus (l.) hitchc. tridens muticus (torr) nash tridens strictus (nutt) nash tripsacum dactyloides (l.) l. vulpia octoflora (walter) rydb. pontederiaceae heteranthera limosa (sw.) willd potamogetonaceae potamogeton ampifolius tuckerm. potamogeton diversifolius raf. potamogeton nodosus poir potamogeton pusillus l. smilacaceae smilax bona-nox l. smilax pseudochina l. smilax rotundifolia l. smilax tamnoides l. typhaceae typha domengensis l. typha latifolia l. eudicots acanthaceae justicia americana (l.) vahl ruellia caroliniensis (j.f. gmel) steud. ruellia humilis nutt. amaranthaceae alternanthera caracasana kunth amaranthus albus l. amaranthus hybridus l. amaranthus retroflexus l. amaranthus rudis sauer froelichia floridana (nutt.) moq. froelichia gracilis (hook.) moq. gossypianthus lanuginosus (poir.) moq guilleminea densa (humb. & bonpl. ex willd.) moq. anacardiaceae rhus glabra l. rhus trilobata nutt. toxicodendron radicans (l.) kuntze apiaceae ammoselinum popei torr. & gray chaerophyllum tainturieri hook. cicuta maculata l. daucus pusillus michx. eryngium leavenworthii torr. & gray lomatium foeniculaceum (nutt.) coult. & rose polytaenia nuttallii dc. ptilimnium nuttallii (dc.) britt. sanicula canadensis l. spermolepis divaricata (walt.) raf. ex ser. spermolepis echinata (nutt. ex dc.) heller 51 oklahoma native plant record volume 8, number 1, december 2008 carter, et al. apocynaceae apocynum cannabinum l. amsonia ciliata walt. asclepiadaceae asclepias asperula (dcne.) woods. asclepias latifolia engelm. & gray asclepias pumila (gray) vail asclepias stenophylla gray asclepias tuberosa l. asclepias verticillata l. asclepias viridis walt. asteraceae achillea millefolium l. ambrosia artemisiifolia l. ambrosia psilostachya dc. ambrosia trifida l. antennaria parlinii fern. aphanostephus pilosus buckl. aphanostephus ramosissimus dc. aphanostephus skirrhobasis (dc.) trel. artemisia dracunculus l. artemisia filifolia torr. artemisia longifolia nutt. artemisia ludoviciana nutt. aster ericoides l. aster oblongifolius nutt. aster patens ait. aster subulatus michx. baccharis salicina torr. & gray bidens cernua l. bidens frondosa l. brickellia eupatorioides (l.) shinners centaurea americana nutt. chaetopappa asteroides nutt. ex dc. cirsium ochrocentrum gray cirsium undulatum (nutt.) spreng. conyza canadensis (l.) cronq. conyza ramosissima cronq. coreopsis grandiflora hogg ex sweet coreopsis tinctoria nutt. dysodiopsis tagetoides (torr. & gray) rydb. echinacea angustifolia dc. eclipta prostrata (l.) l. engelmannia peristenia (raf.) goodman & c.a. watson erigeron strigosus muhl. ex willd. eupatorium serotinum michx. evax prolifera nutt. ex dc. evax verna raf. gaillardia aestivalis (walt.) h. rock gaillardia pulchella foug. gaillardia suavis (gray & engelm.) britt. & rusby gamochaeta purpurea (l.) cabrera grindelia papposa mckelvey grindelia squarrosa (pursh) dunal gutierrezia dracunculoides (dc.) s.f. blake helenium amarum (raf.) h. rock helenium microcephalum d.c. helianthus annuus l. helianthus hirsutus raf. helianthus maximiliani schrad. helianthus pauciflorus nutt. helianthus petiolaris nutt. heterotheca canescens (dc.) shinners heterotheca stenophylla (gray) shinners hieracium longipilum small hymenopappus scabiosaeus l'hér. hymenopappus tenuifolius pursh iva annua l. krigia caespitosa (raf.) k.l. chambers krigia dandelion (l.) nutt. lactuca canadensis l. lactuca serriola l. lactuca tatarica (l.) c.a. mey liatris aspera michx. liatris punctata hook. liatris scariosa (l.) willd. machaeranthera pinnatifida (hook.) shinners nothocalais cuspidata (pursh) greene packera plattensis (nutt) w.a. weber & a. love palafoxia sphacelata (nutt. ex torr.) cory pluchea camphorata (l.) dc. polanisia dodecandra (l.) d.c. pseudognaphalium canescens dc. pseudognaphalium stramineum (kunth) w.a. weber pyrrhopappus grandiflorus (nutt.) nutt. ratibida columnifera (nutt.) wooton & standl. rudbeckia hirta l. silphium asteriscus l. silphium laciniatum l. silphium radula nutt. solidago arguta ait. solidago gigantea ait. solidago missouriensis nutt. solidago petiolaris ait. tetraneuris linearifolia (hook.) greene thelesperma filifolium (hook.) gray townsendia exscapa (richards.) potter vernonia baldwinii torr. 52 oklahoma native plant record volume 8, number 1, december 2008 carter, et al. xanthium strumarium l. xanthisma texanum dc. boraginaceae buglossoides arvensis (l.) i.m. johnston lithospermum incisum lehm. mysotis verna nutt. brassicaceae draba brachycarpa nutt. ex torr. & a. gray draba cuneifolia nutt. ex torr. & a. gray draba reptans (lam.) fern. descurainia pinnata (walt.) britt. erysimum capitatum (dougl. ex hook.) greene lepidium oblongum small lepidium virginicum l. lesquerella auriculata (engelm & a. gray) s. watson lesquerella engelmannii (a.gray) s. watson lesquerella gracilis (hook.) s. watson lesquerella ovalifolia rydb. ex britt. rorippa nasturtium-aquaticum (l.) hayek rorippa palustris (l.) bess. rorippa sessiliflora (nutt.) hitchc. sibara virginica (l.) rollins callitrichaceae callitriche heterophylla pursh. campanulaceae lobelia appendiculata a. dc. lobelia cardinalis l. triodanis leptocarpa (nutt.) nieuwl. triodanis perfoliata (l.) nieuwl. capparaceae cleomella angustifolia torr. caprifoliaceae viburnum rufidulum raf. symphoricarpos orbiculatus moench caryophyllaceae cerastium brachypodum (engelm. ex gray) b.l. robins. minuartia michauxii (fenzl.) farw. minuartia patula (michx.) mattf. paronychia jamesii torr. & a. gray paronychia virginica spreng. sagina decumbens (ell.) torr & a. gray silene antirrhina l. ceratophyllace ceratophyllum demersum l. chenopodiaceae chenopodium album l chenopodium leptophyllum (moq.) nutt. chenopodium simplex (torr.) raf. chenopodium standleyanum aell. salsola tragus l monolepis nuttalliana (schult.) greene cistaceae lechea tenuifolia michx. clusiaceae hypericum drummondii (grev. & hook) torr. & a. gray hypericum mutilum l. convolvulaceae convolvulus arvensis l. evolvulus nuttallianus schult. cornaceae cornus drummondii c.a. mey cuscutaceae cuscuta gronovii willd. cuscuta coryli engelm. crassulaceae sedum nuttallianum raf. cucurbitaceae cyclanthera dissecta (torr. & gray) arn. cucurbita foetidissima kunth ibervillea lindheimeri (gray) greene melothria pendula l. ebenaceae diospyros virginiana l. euphorbiaceae acalypha gracilens a. gray acalypha ostryifolia riddell chamaesyce glyptosperma (engelm) small chamaesyce maculata (l.) small chamaesyce missurica (raf.) shinners chamaesyce nutans (lag.) small chamaesyce prostrata (aiton) small croton capitatus michx. croton glandulosus l. 53 oklahoma native plant record volume 8, number 1, december 2008 carter, et al. croton lindheimerianus scheele. croton monanthogynus michx. croton texensis (klotzch.) muell. arg. euphorbia commutata engelm. euphorbia corollata l. euphorbia dentata michx. euphorbia marginata pursh. euphorbia spathulata lam. phyllanthus caroliniensis walt. phyllanthus polygonoides nutt. ex spreng. stillingia sylvatica garden ex l. tragia ramosa torr. fabaceae acacia angustissima (mill.) kuntze amorpha canescens pursh. amorpha fruticosa l. apios americana medik. astragalus crassicarpus nutt. astragalus plattensis nutt. baptisia australis (l.) r. br. baptisia bracteata muhl. ex elliot baptisia sphaerocarpa nutt. cercis canadensis l. chamaecrista fasciculata (michx.) greene clitoria mariana l. crotalaria sagittalis l. dalea aurea nutt. ex pursh. dalea candida willd. dalea enneandra nutt. dalea multiflora (nutt.) shinners dalea purpurea vent. dalea tenuis (j.m. coult) shinners desmanthus illinoensis (michx.) macm desmanthus leptolobus torr & a. gray desmodium ciliare dc. desmodium nudiflorum (l.) dc. desmodium paniculatum (l.) dc. desmodium sessilifolium (torr.) torr. & a.gray galactia volubilis (l.) britt. glycyrrhiza lepidota (nutt.) pursh. gymnocladus dioicus (l.) koch. hoffmanseggia glauca (ortega) eifert indigofera miniata ortega lespedeza capitata michx. lespedeza procumbens michx. lespedeza virginica (l.) britt. lotus unifoliolatus (hook.) benth. melilotus albus medik. neptunia lutea (leavenw.) benth. pediomelum esculentum (pursh.) rydb. pediomelum linearifolium (torr. & a. gray) j.w. grimes psoralidium tenuiflorum (pursh.) rydb. senna marilandica (l.) link. strophostyles helvula (l.) elliott strophostyles leiosperma (torr. & a.gray) piper strophostyles umbellatum (muhl ex willd.) britt. stylosanthes biflora (l.) britton, sterns. & pogg. vicia ludoviciana nutt. fagaceae quercus buckleyi nixon & dorr. quercus macrocarpa michx. quercus marilandica muench. quercus muehlenbergii engelm. quercus shumardii buckl. quercus stellata wang. quercus velutina lam. quercus virginiana mill. fumariaceae corydalis aurea willd. gentianaceae sabatia campestris nutt. geraniaceae geranium carolinianum l. grossulariaceae ribes aureum pursh haloragaceae myriophyllum pinnatum (walter) britton, stens & poggenb. myriophyllum spicatum l. hydrophyllaceae nama hispidum a. gray juglandaceae carya illinoinensis (wang.) k. koch juglans major (torr.) heller juglans microcarpa berland juglans nigra l. krameriaceae krameria lanceolata torr. 54 oklahoma native plant record volume 8, number 1, december 2008 carter, et al. lamiaceae hedeoma hispida pursh. lycopus americanus muhl. monarda citriodora cerv. ex lag. monarda clinopodioides a. gray monarda fistulosa l. monarda pectinata nutt. nepeta cataria l. salvia azurea michx. ex lam. salvia reflexa hornem. scutellaria drummondii benth. scutellaria wrightii a. gray scutellaria resinosa torr. teucrium canadense l. teucrium laciniatum torr. trichostema brachiatum l. lentibulariaceae utricularia gibba l. linaceae linum berlandieri hook. linum hudsonioides planch. linum imbricatum (raf.) shinners linum rigidum pursh linum sulcatum riddell loasaceae mentzelia oligosperma nutt. mentzelia nuda (pursh.) torr & a. gray lythraceae ammannia auriculata willd. ammannia coccinea rottb. lythrum alatum pursh. malvaceae callirhoe involucrata (torr. & a. gray) a. gray sphaeralcea coccinea nutt. menispermaceae cocculus carolinus (l.) dc. molluginaceae mollugo verticillata l. moraceae morus rubra l. nyctaginaceae mirabilis nyctaginea (michx.) macm. mirabilis linearis (gray) greene mirabilis glabra (s. wats.) standl. mirabilis hirsuta a. nels. mirabilis albida (walter) heimerl. oleaceae fraxinus pennsylvanica marsh. onagraceae calylophus serrulatus (nutt.) p.h. raven gaura coccinea nutt. ex pursh. gaura parviflora douglas ex lehm. gaura sinuate nutt. gaura suffulta engelm. ex a. gray gaura triangulata buckley ludwigia alternifolia l. ludwigia decurrens (walt.) dc. ludwigia peploides (kunth) raven ludwigia repens j.r. forst oenothera biennis l. oenothera brachycarpa a. gray oenothera grandis (britton) smith oenothera jamesii a. gray oenothera laciniata hill oenothera linifolia nutt. oenothera macrocarpa (a. gray) w.l. wagner oenothera triloba nutt. stenosiphon linifolius (nutt. ex e. james) heynh. oxalidaceae oxalis violacea rydb. oxalis stricta l. papaveraceae argemone polyanthemos (fedde) g.b. ownbey pedaliaceae proboscidea louisianica (mill.) thell. phytolaccaceae phytolacca americana l. plantaginaceae plantago aristata michx. plantago elongata pursh. plantago patagonica jacq. plantago virginica l. plantago wrightiana dcne. 55 oklahoma native plant record volume 8, number 1, december 2008 carter, et al. polemoniaceae ipomopsis rubra fern. polygalaceae polygala alba nutt. polygala verticillata l. polygonaceae eriogonum annum nutt. eriogonum longifolium nutt. polygonum amphibium l. polygonum convolvulus l. polygonum hydropiperoides michx. polygonum lapathifolium l. polygonum pensylvanicum l. polygonum punctatum ell. polygonum ramosissimum michx. polygonum scandens l. polygonum tenue michx. rumex altissimus wood rumex crispus l portulacaceae claytonia virginica l. portulaca pilosa l. portulaca umbraticola kunth. portulaca oleracea l. talinum calycinum engelm. talinum parviflorum nutt. primulaceae adrosace occidentalis l. dodecatheon meadia l. samolus valerandi l. ranunculaceae anemone berlandieri pritz delphinium carolinianum walt. myosurus minimus l. rhamnaceae ceanothus herbaceus raf. rosaceae agrimonia parviflora aiton crataegus reverchonii sarg. crataegus viridis l. geum aleppicum jacq. geum canadense jacq. potentilla arguta pursh prunus angustifolia marsh. prunus gracilis engelm. & a. gray prunus mexicana s. watson prunus virginiana l. rubus aboriginum rydb. rubus occidentalis l. rubiaceae cephalanthus occidentalis l. diodia teres walt. galium aparine l. galium pilosum aiton galium texense a. gray galium virgatum nutt. hedyotis nigricans (lam.) fosberg houstonia caerulea l. houstonia pusilla schoepf rutaceae ptelea trifoliata l. salicaceae populus alba l. populus deltoides marsh salix caroliniana michx. salix nigra marsh. santalacaceae comandra umbellata (l.) nutt. sapindaceae aesculus glabra willd. acer grandidentatum nutt. acer saccharinum l. acer saccharum marsh. sapindus saponaria l. . sapotaceae sideroxylon lanuginosum michx. scrophulariaceae agalinis fasciculata pennell bacopa rotundifolia (michx.) wettst. castilleja purpurea (nutt.) g. don. castilleja sessiliflora pursh. collinsia violacea nutt. gratiola virginiana l. leucospora multifida (michx) nutt. linaria canadensis (l.) dumont. lindernia dubia (l.) pennell 56 oklahoma native plant record volume 8, number 1, december 2008 carter, et al. penstemon cobaea nutt penstemon oklahomensis pennell scrophularia lanceolata pursh. veronica arvensis l. veronica peregrine l. simaroubaceae ailanthus altissima (p. mill.) swingle solanaceae physalis cinerascens (dunal) hitchc. physalis longifolia nutt. quincula lobata (torr.) raf. solanum dimidiatum raf. solanum elaeagnifolium cav. solanum ptycanthum dunal solanum rostratum dunal ulmaceae celtis laevigata willd. ulmus americana l. ulmus rubra muhl urticaceae boehmeria cylindrica (l.) sw. parietaria pensylvanica muhl. ex willd. valerianaceae valerianella radiata (l.) dufr. verbenaceae glandularia canadensis (l.) nutt. glandularia pumila (rydb.) umber lippia cuneifolia (torr.) steud lippia nodiflora (l.) michx. verbena bracteata lag. & rodr. verbena stricta vent. verbena urticifolia l. violaceae hybanthus verticillatus (ortega) baill viola bicolor pursh viola missouriensis greene viola sororia willd. vitaceae ampelopsis cordata michx. cissus incisa des moul. parthenocissus quinquefolia (buckley) britton ex small vitis cinerea (englm.) millardet. vitis riparia michx. vitis rupestris scheele zygophyllaceae kallstroemia californica (s. wats.) vail tribulus terrestris l. oklahoma native plant record, volume 11, number 1, december 2011 oklahoma native plant record volume 11, december 2011 smith, m. & mckenzie, p. m. 22 sch oenoplectus h allii, s. saxim on t anus, and the putative s. h allii × s. saxim ontanus hybrid: observations from the wichita mountains wildlife refuge and the fort sill military reservation 2002 – 2010 marian smith paul m. mckenzie southern illinois state university u.s. fish and wildlife service edwardsville, il 62025 101 park deville drive, suite a msmith@siue.edu columbia, mo 65203 keywords: schoenoplectus, hybridization, manag ement abstract schoenoplectus hallii, s. saximontanus, and the putative s. hallii × s. saximontanus hybrid are obligate wetland sedges that occur in the sparsely vegetated margins of ponds, ditches or swales with fluctuating water levels. the species are amphicarpic and have easily identified differences between spikelet and basal achenes. we surveyed selected sites at the refuge in 2001, 2002, and 2007 – 2010, surveyed 4 sites on the fort sill military reservation in 2009 and 2010, and collected voucher specimens from all populations. scanning electron microsope (sem) photographs of spikelet and basal achenes indicate distinct morphological differences between species and the presence of “winged” ridges on s. saximontanus. field observations indicated that populations at all sites vary in size and species distribution annually, and that both parental species appeared to be declining in number. we concluded that in populations where s. hallii and s. saximontanus co-occur, hybridization may be a threat to one or both parental species. the distribution of achenes by waterfowl and ungulates indicates that management to prevent establishment of mixed populations, and therefore hybridization, is not practical. we recommend that s. hallii be evaluated for federal listing under the endangered species act, a range-wide assessment be completed for s. saximontanus, and that all sites with mixed populations should be examined for the presence of hybrids. introduction schoenoplectus hallii (a. gray) s .g. sm. and s. saximontanus (fernald) raynal are sedge species that were once thought to be separated geographically, with s. hallii present in the midwest and eastern u. s. and s. saximontanus largely confined to the west (gleason and cronquist 1991, beatty et al. 2004). schoenoplectus hallii has a global ranking of g2 (imperiled). it is listed as “critically imperiled” in eight of the 12 states in which it occurs (natureserve 2010) and as “imperiled” or “vulnerable” in three other states. herbarium records indicate that it had been reported from georgia and massachusetts prior to 1981 (mckenzie et al. 2007), but those populations are thought to have been extirpated (natureserve 2010, mckenzie et al. 2007). schoenoplectus saximontanus has a global ranking of g5 (secure) (natureserve 2010), but it is listed as “critically imperiled” in british columbia as well as in seven of the 12 states where it occurs. it has been reported from two states in mexico (flora of north america 2002). throughout its range, s. saximontanus is considered to be an uncommon species https://doi.org/10.22488/okstate.17.100082 oklahoma native plant record volume 11, december 2011 smith, m. & mckenzie, p. m. 23 whose distribution is scattered (flora of north america 2002). schoenoplectus hallii and s. saximontanus are obligate wetland species that have similar habitat requirements: most often sandy, rocky, or gravelly soil, occasionally clay, around the margins of ponds, ditches and swales with fluctuating water levels, and a scarcity of other plants as competitors (flora of north america 2002, mckenzie et al. 2007). they most commonly complete their life cycle as annuals, but short-lived perennials have been reported from texas (o’kennon and mclemore 2004). both species have 2-3 small basal leaves and tufted stems ~4-40 cm long with small, inconspicuous rhizomes. the species are amphicarpic (having two distinct types of achenes), with numerous inflorescences on aerial stems containing perfect flowers and occasional, pistillate flowers enclosed in a leaf sheath at the plant base. it is difficult to distinguish between the species from vegetative characteristics alone. achenes of s. hallii are 2-sided, and flowers have 2lobed styles; whereas, achenes of s. saximontanus are distinctly 3-sided, and flowers have 3-lobed styles (flora of north america 2002). achenes of both species have transverse ridging, and magrath (2002) reported that the ridges on s. saximontanus were “winged;” whereas, those on s. hallii were smooth. this character had not been reported prior to his 2002 publication. both species have been reported from five states (ks, mo, ne, ok, and tx) (o’kennon and mclemore 2004, mckenzie et al. 2007, natureserve 2010); however, only oklahoma (magrath 2002), kansas (c. freeman, pers. comm. 2006), and texas (bob o’kennon, pers. comm. 2007) have sites with mixed populations (magrath 2002, smith et al. 2004). although s. saximontanus occurs in eight counties in ok, it only co-occurs with s. hallii in comanche county (oklahoma vascular plant database 2006). in 2000, 134 sites at the wichita mountains wildlife refuge (wmwr) in comanche county were surveyed for s. hallii and s. saximontanus by dr. larry magrath and personnel from the refuge (magrath 2002). in august 2001, m. smith and p. mettler-cherry re-examined the population sites surveyed by magrath in 2000, and in subsequent years, 2002 and 2007 – 2010, the authors conducted surveys of selected sites at the refuge and fort sill military reservation (fsmr). during the 2001 survey, m. smith noted what appeared to be plants containing achenes that were intermediate between the two species, i.e., some appeared to be 2sided like those of s. hallii, except the usually flat or convex side contained a conspicuous bulge, and the achenes often had the “winged” appearance reported by magrath (2002). some plants had both 2 and 3-sided achenes, some with, and some without “winged ridges.” other individuals produced only a few viable-looking achenes, with the majority of inflorescences bearing a preponderance of aborted achenes. smith interpreted these anomalies as suggestive of hybridization between s. hallii and s. saximontanus. the objectives of this report are to discuss the results of a seed bank study for three sites conducted in 2001; to provide photographic documentation of the wingedridge appearance of s. saximontanus achenes reported by magrath (2002); to summarize field observations made during visits to wmwr in 2002 and 2007 – 2010 and fsmr in 2009 and 2010; and to discuss the presence of putative hybrids of s. hallii and s. saximontanus and potential conservation concerns associated with hybridization among rare species. botanical nomenclature listed in this report follows yatskievych and turner (1990) except for marsilea vestita which follows diggs et al. (1999), eleocharis coloradoensis which follows smith (flora of north america 2002), and eleocharis ovata which follows yatskievych (1999). oklahoma native plant record volume 11, december 2011 smith, m. & mckenzie, p. m. 24 characteristics of spikelet and basal achenes in july 2001, spikelet and basal achenes were collected from individuals of s. hallii and s. saximontanus for scanning electron microscopy (sem) examination of achene surfaces and cross-sectional shape. seeds were mounted onto aluminum stubs using double-stick transfer tabs (electron microscopy sciences) and examined with a hitachi s2460n variable pressure sem at 30 kv and 20 pa pressure. images were digitally recorded using a noran voyager iii interface. for achene cross-sectional views, plastic blocks containing embedded specimens were sectioned until a transverse median section was obtained. both species exhibit transverse ridges (figure 1 a – h) as described in flora of north america (flora of north america 2002) and flora of the great plains (great plains flora association 1986), except that the distinct difference between ridge morphology in s. hallii compared to s. saximontanus (see figure 1 a – h) is not discussed in either account. in flora of the great plains (1986), the achenes of both species were described simply as having “transverse ridges,” and no description of the amphicarpic (basal) achenes was provided. in flora of north america (2002), spikelet achenes of both species were described as having “mostly sharp ridges.” the basal achenes of s. hallii were described as “rugose with rounded edges,” and this is confirmed by the sems in this article, but basal achenes of s. saximontanus were characterized in flora of north america (2002) as “with obscure to evident horizontal ridges.” the sems portrayed in figure 1 (a – h) illustrate a distinctive species-difference in ridge shape on spikelet achenes and depict the basal achenes in s. saximontanus as having distinct, sharp ridges (see figure 1 f, h). ridges on spikelet achenes in s. saximontanus, compared to s. hallii, are sharper and more elaborate in design, resulting in the “winged” appearance described by magrath (2002). ridges on basal achenes are subtle and incomplete in s. hallii (see figure 1 e, g), but prominent, more complete, and “winged” in s. saximontanus (see figure 1 f, h). figure 1 surface views of front (a) and opposite (c) sides of spikelet achene of s. hallii and front (b) and opposite (d) sides of spikelet achene of s. saximontanus. front (e) and opposite (g) sides of basal achene of s. hallii and front (f) and opposite (h) sides of basal achene of s. saximontanus. bars = 130 μm figure 2 (a – b) illustrates the often described cross-sectional shape of spikelet achenes: “plano-convex” (flora of north america 2002) for s. hallii (see figure 2 a) oklahoma native plant record volume 11, december 2011 smith, m. & mckenzie, p. m. 25 and “equilaterally, sharply trigonous,” (flora of north america 2002) for s. saximontanus (see figure 2 b). magnified views of surface features (figure 2 e – h) in both types of achenes reinforce the differences in surface ridges between the two species shown in figure 1. ridges in s. hallii spikelet achenes (see figure 2 e) are not sharp compared to the elaborate, sharp-tipped wings of s. saximontanus (see figure 2 f). surfaces of the basal achene of s. hallii (see figure 2 g) are mostly absent of ridges, but those of s. saximontanus (see figure 2 h) are prominent and elaborate, although not quite as sharp as in the spikelet achene. figure 2 cross-sectional views of spikelet (a) and basal (c) achenes of schoenoplectus hallii and spikelet (b) and basal achenes (d) of s. saximontanus. bars = 100 μm. surface views of ridges of spikelet (e) and basal (g) achenes of s. hallii. surface views of ridges of spikelet (f) and basal (h) achenes of s. saximontanus. bars = 20 μm soil seed bank study of 2001 in 2001, 15 plants were selected at each of three sites that had been included in magrath’s 2002 report (boggy flat, quanah parker lake, and hollis lake at wmwr), and soil cores (1.75 cm × 8 cm) were collected and separated into 2 cm sections. achenes were recovered, counted, and separated by species and soil depth. achenes were tested for viability as in malone (1967); all achenes were viable. eighty-six percent of the achenes were contained within the first 2 cm of the soil. all three sites had achenes of both species present in the soil; however, at the time of the site visit, extant populations at boggy flats appeared to have only s. saximontanus and hollis lake had only s. hallii. both species were present in the extant population at quanah parker. the presence of viable achenes of s. hallii and s. saximontanus in the soil at all three sites indicates either the undetected presence of both species during the site visit or the existence of both species at each site in previous years. as achenes of these species may remain dormant and viable for extended periods (mcclain et al. 1997), their presence in the above-ground population might have occurred many years in the past and the current extant population may reflect an increase in one species and a decline or elimination of the other. field observations 2002, 2007 – 2010 in 2002, the authors visited selected sites at wmwr (table) and specimens of schoenoplectus were collected from five sites where s. hallii or s. saximontanus had been previously collected or reported (magrath 2002). field and laboratory observations indicated that the achenes of most spikelets of s. hallii and s. saximontanus were mature and exhibited characteristics typical for the species as described in the introduction; oklahoma native plant record volume 11, december 2011 smith, m. & mckenzie, p. m. 26 however, some achenes of the presumed s. hallii and s. saximontanus hybrids appeared to be abortive or malformed, while other putative hybrids produced both fully developed 2and 3-sided achenes. individuals of the putative hybrids were noticeably taller with longer inflorescences than either parent, and the spikelet scales were conspicuously brownish-orange (smith et al. 2004). specimens were sent to dr. alfred schuyler, dr. s. galen smith, and dr. anton rezniceik for verification. the presence of a putative hybrid (mckenzie #2028) was independently confirmed by each scientist based on morphological characters (smith et al. 2004). subsequent visits to wmwr and fsmr were made to collect material for a future genetic analysis. the only site to be visited every year from 2007 – 2010 was medicine tank at wmwr; therefore, we will discuss some apparent trends in population size and species distribution at that site. in 2002, s. hallii was abundant and concentrated on the west shore of the pond while s. saximontanus was abundant on the south and east shore (see table). the putative hybrid was scattered but present in various locations. in contrast, populations of all three species were abundant and widely distributed along the shore in 2007. in 2008, populations of s. hallii and s. saximontanus, although present at the site (see table), appeared to be smaller and more restricted than in 2002 and 2007. as in 2007, s. hallii and s. saximontanus were scattered along the entire pond margin without any noticeable concentration at different shore edges; however, the putative s. hallii × s. saximontanus hybrids were more abundant and concentrated along the northwest shore (see table). in 2009, the water level at medicine tank was lower than had been observed on previous visits, and there was a noticeable reduction in population size of s. hallii and s. saximontanus, especially the latter which was scattered and extremely rare. the putative hybrid was common and the population had expanded beyond the northwest shore (see table). schoenoplectus hallii was rare at medicine tank in 2010, and s. saximontanus, which had been present in previous years, was notably absent. conversely, the putative hybrid was abundant (see table) and had apparently overtaken habitat around the pond that had been previously occupied by s. hallii or s. saximontanus. in addition to medicine tank, we visited three other sites in 2009 at wmwr, and water levels were lower at all of them than in previous years. ponds that were composed mostly of hardened clay were lacking, or had very few, schoenoplectus individuals (ingram pond and rock dam), but ponds that had rocks, cobble, and/or sand had healthy flowering and fruiting plants (medicine tank, quanah parker lake), although in reduced numbers compared to 2007 and 2008 (see table). no bulrushes were observed at two sites where s. saximontanus was found in 2007 (boggy flat and unnamed pond ~1.5 mi east, see table). in 2010, we collected s. saximontanus from grama lake, which was the only time we documented any bulrush at this site during our visits between 2001 and 2010. other sites were visited and vouchers were collected at wmwr from 2007 – 2010 as noted in the table. we visited four sites at fsmr in 2009 and two sites in 2010 and documented the presence of s. hallii, s. saximontanus, and putative s. hallii × s. saximontanus hybrids at the reservation in both years. in 2009, s. hallii was found at all four sites, and s. saximontanus and the putative hybrid were present at two (see table). in 2010 s. hallii and s. saximontanus had disappeared from pottawatomie pond and the putative hybrid had increased in number (see table). schoenoplectus spp. observed at fsmr were all at ponds that had a sandy or gravelly shoreline. our collections constitute the first documented records of s. hallii, s. oklahoma native plant record volume 11, december 2011 smith, m. & mckenzie, p. m. 27 saximontanus, and the putative hybrids from fsmr and confirm the predictions of magrath (2002) that the two parent species would likely be discovered at fort sill. habitat and plant associates schoenoplectus hallii, s. saximontanus, and putative s. hallii × s. saximontanus hybrids occurred along the edges of ponds and lakes at wmwr and at fsmr that had receding shorelines. very little vegetation competed with the schoenoplectus spp. in the narrow marginal areas along the water’s edge. although the majority of populations of schoenoplectus occurred in sandy or gravelly/rocky soil, a few were found on clay substrates. many sites were heavily grazed by elk, bison, or longhorn cattle, and the soil was significantly trampled by ungulates or disturbed by foraging feral hogs. plant associates varied widely from site to site and included the following species: ammannia coccinea rottb., bacopa rotundifolia (michx.) wettst., bergia texana (hook.) seub. ex walp., cyperus acuminatus torr. & hook. ex torr., c. difformis l., c. setigerus torr. & hook., c. squarrosus l., eclipta prostrata (l.) l., eleocharis acicularis (l.) roem. & schult., e. atropurpurea (retz.) j. presl. & c. presl., e. coloradoensis (britt.) gilly, e. ovata (roth) roem. & schult., e. parvula (roem. & schult.) link ex bluff, nees & schauer, juncus spp., justicia americana (l.) vahl, lindernia dubia (l.) pennell, marsilea vestita hook. & grev., panicum scoparium lam., paspalidium geminatum (forssk.) stapf., var. geminatum, phyla nodiflora (l.) greene, pilularia americana a. braun, tribulus terrestris l., and xanthium strumarium l. discussion we observed culms of s. saximontanus, s. hallii, and the putative hybrid flowering and producing viable fruit from july through mid-october, indicating that they are able to do so anytime during the growing season when conditions for germination and growth are favorable (baskin et al. 2003). at all four sites where s. hallii and s. saximontanus occur in mixed populations, the putative hybrid was present (medicine tank, quanah parker lake, zania pond, and pottawatomie pond) (see table), suggesting that hybridization is a definite possibility in any mixed population. soil cores collected in 2001 from boggy flat and hollis lake appeared to contain achenes of both s. hallii and s. saximontanus, but as hybrids had not been verified at the time the cores were processed, it is possible that hybrid seed might have been present. in any case, it is likely that in some years individuals of both species (and possibly those of the putative hybrid) may emerge at those sites in the future. schoenoplectus hallii and s. saximontanus cooccur at rhodes lake, tx where they are on opposite ends of the reservoir (robert o’kennon, pers. comm. oct. 2007); however, the ease with which waterfowl may transport achenes for long distances (devlaming and proctor 1968; powers et al. 1978) suggests that the species may form a mixed population in the near future. historically, s. hallii and s. saximontanus were apparently allopatric and likely came in contact with one another via the muddy feet of migrating waterfowl, as suggested by mcclain et al. (1997) and beatty et al. (2004). it was postulated by magrath (2002) that large herbivores such as bison and other animals were dispersal agents for achenes of s. hallii and s. saximontanus at wmwr. our observations support the possibility of elk and bison as dispersal agents, as hoofprints of both were evident at every site. as neither elk nor bison are common at fsmr, we propose that achenes of the two species at the reservation are more likely to have been transported among ponds via waterfowl, white-tail deer, and feral hogs. toni hodgkins, naturalist at fsmr, reported that feral hogs equipped oklahoma native plant record volume 11, december 2011 smith, m. & mckenzie, p. m. 28 with radio transmitters were documented moving to and from the reservation and noted that 1200 pigs had been captured in less than a year (toni hodgkins, pers. comm. 2010). there is an historical site in kansas where s. hallii and s. saximontanus were known to be sympatric in 1997, but craig freeman at the university of kansas examined the voucher specimens from the site in 2006 and did not note any evidence of hybridization (craig freeman, pers. comm. 2006). nevertheless, it would be advisable to continue to monitor these areas in ks and tx to further assess the incidence of hybridization in mixed populations. we suggest that when viewing live material in the field, or when examining dried voucher specimens, a thorough evaluation of multiple spikelets from different plants is necessary for reliable identification of plants collected from populations where s. hallii and s. saximontanus co-occur. spikelets should be carefully examined for the presence of mixed style numbers, abortive achenes, or abnormally shaped, 2or 3-sided achenes. some spikelets that have achenes characteristic of one species may have an achene that would be better identified as the other species. thus, the failure to examine multiple spikelets from different plants may result in a premature determination of specimens collected from mixed populations. the presence of s. hallii × s. saximontanus hybrids at wmwr and fsmr may threaten the long term persistence of s. hallii and s. saximontanus in ok and constitute a threat to the conservation of the species in north america. according to conservation geneticists, the possible dangers of hybridization are numerous and pose a serious threat to the survival of rare species that hybridize with a closely related congener (levin et al. 1996). although the extinction of rare species typically is attributed to environmental change that renders the habitat unsuitable (harrison 1991; national research council 1995), hybridization may have a profound effect on the persistence of a species (rieseberg 1991; ellstrand 1992; rieseberg and linder 1999). hybrids compete for space and resources with parental species and reduce the potential for plants to replace themselves, thereby inhibiting the growth of their populations – the lower the rate of population growth, the greater the potential for extinction in a variable environment (menges 1992). the numerical disadvantage of a rare species is compounded by the proliferation of fertile hybrids (rhymer and symberloff 1996). if one of the species is rarer than the other, the addition of hybrids to a population containing congeners decreases the proportional representation of the less abundant parent. in time, backcrossing can result in the assimilation of the rare species whose genetic identity will become extinct, and, over evolutionary time, the dna of the former rare species may be lost from the gene pool altogether (rieseberg et al. 1996). in the case of s. hallii and s. saximontanus in ok, both of which appear to be rare, it is possible that although much of the dna of both species may survive, the two species may be subsumed into a new species with a new genetic identity. future actions because of the impossibility of controlling achene dispersal-agents at wmwr and fsmr, it is unlikely that management to prevent hybridization is possible. we recommend, however, that monitoring of the populations of schoenoplectus at wmwr and fsmr be continued to confirm or dismiss the importance of hybridization as a threat in the area. given the documented hybridization between s. hallii and s. saximontanus in oklahoma native plant record volume 11, december 2011 smith, m. & mckenzie, p. m. 29 oklahoma and the identification of new threats to s. hallii in illinois (mckenzie et al. 2010), the species should be re-evaluated as a possible candidate for federal listing under the endangered species act. no range-wide status assessment exists for s. saximontanus, and as noted in the introduction, the species is critically imperiled in seven of the 12 states where it has been documented. we recommend that a thorough analysis of the distribution and size of populations of s. saximontanus be made, and that potential threats to the species be assessed. further studies may provide evidence that this species may also warrant protection under the endangered species act. acknowledgments the observations reported in this paper could not have been made without the help of a large number of friends and colleagues. we thank dr. paige mettler-cherry and dr. nancy parker for their company during our many trips to oklahoma, their help in the field, and their advice on schoenoplectus issues in general. sam waldstein, former refuge manager, was essential to our project and, without his help during our first visit in 2001 our work at wmwr would not have been possible. dr. larry magrath served as our inspiration to undertake this work, generously shared his observations from 2000, and allowed us to view the voucher specimens at the usao herbarium. robert o’kennon of brit spent a day with us in pouring rain, cheerfully taking us to the schoenoplectus sites he had recently discovered in north central texas. volunteer donna phillips took the time during our subsequent visits to wmwr to help us locate small ponds in the research area. at fort sill, fsmr biologist toni hodgkins enabled us to visit ponds and search for schoenoplectus during our 2009 and 2010 visits to the lawton area. literature cited baskin, c. c., j. m. baskin, e. w. chester, and m. smith. 2003. ethylene as a possible cue for seed germination of schoenoplectus hallii (cyperaceae), a rare summer annual of occasionally flooded sites. american journal of botany 90:620-627. beatty, b. l., w. f. jennings, and r. c. rawlinson. 2004. schoenoplectus hallii (gray) s.g. sm. (hall’s bulrush): a technical conservation assessment. prepared for the usda forest service, rocky mountain region, species conservation report. peer review administered by center for plant conservation, st. louis, missouri. devlaming, v. and v. w. proctor. 1968. dispersal of aquatic organisms: viability of seeds recovered from the droppings of captive killdeer and mallard ducks. american journal of botany 55:20-26. diggs, g. m. jr., b. l. lipscomb, m. d. reed, and r. j. o’kennon. 1999. illustrated flora of north central texas. sida, botanical miscellany 16:1-1626. elstrand, n. c. 1992. gene flow by pollen: implications for plant conservation genetics. oikos 63:77-93. flora of north america editorial committee, eds. 2002. vol. 2. magnoliophyta: commelinidae (in part): cyperaceae. oxford university press, new york. gleason, h. a. and a. cronquist. 1991. manual of vascular plants of northeastern united states and adjacent canada, ed. 2. new york botanical garden, bronx. great plains flora association. 1986. r. l. mcgregor, coordinator; t. m. barkley, r. e. brooks, and e. k. schofield (eds.) flora of the great plains. university press of kansas, lawrence. harrison, s. 1991. local extinction in a metapopulation context: an empirical evaluation. biological journal of linnean society 3-88. oklahoma native plant record volume 11, december 2011 smith, m. & mckenzie, p. m. 30 levin, d. a., j. francisco-ortega, and r. k. jansen. 1996. hybridization and the extinction of rare plant species. conservation biology 10:10-16. magrath, l. k. 2002. schoenoplectus hallii and s. saximontanus. 2000 wichita mountain wildlife refuge survey. oklahoma native plant record 2:54-62. malone, c. r. 1967. a rapid method for enumeration of viable seeds in soil. weeds 15:381-382. mcclain, w. e., r. d. mcclain, and j. e. ebinger. 1997. flora of temporary sand ponds in cass and mason counties, illinois. castanea 62:65-73. mckenzie, p. m., s. g. smith, and m. smith. 2007. status of schoenoplectus hallii (hall’s bulrush) (cyperaceae) in the united states. journal botanical research institute of texas 1:457-481. mckenzie, p. m., m. smith, and t. kelley. 2010. observations of hall’s bulrush (schoenoplectus hallii) (cyperaceae) in mason county illinois in 2009. transactions of illinois state academy of science 103:97-108. menges, e. s. 1992. stochastic modeling of extinction in plant populations. in: fiedler and jain, eds. conservation biology, pp. 253-275. chapman and hall, new york. national research council. 1995. science and the endangered species act. national academy press, washington, dc. natureserve. 2010. natureserve explorer: an online encyclopedia of life [web application]. version 5.0 natureserve, arlington, virginia. available at: http://www.natureserve.org/explorer. (accessed: feb 2010) o’kennon, r. j. and c. mclemore. 2004. schoenoplectus hallii (cyperaceae), a globally threatened species new for texas. sida 21:1201-1204. oklahoma vascular plant database. 2006. http://www.coordinatesolutions.com/o vpd/ovpd.aspx. (accessed: feb 2010) powers, k. d., r. e. noble, and r. h. chabreck. 1978. seed distribution by waterfowl in southwestern louisiana. journal of wildlife management 42:598-605. rhymer, j. m. and d. simberloff. 1996. extinction by hybridization and introgression. annual review of ecological systems 27:83-109. rieseberg, l. h. 1991. hybridization in rare plants: insights from case studies in cerocarpus and helianthus. d. a. falk and k. e. holsinger, eds. in: genetics and conservation of rare plants, pp. 171-181. oxford university press, new york. rieseberg, l. h. and c. r. linder. 1999. hybrid classification: insights from genetic map-based studies of experimental hybrids. ecology 80:361370. rieseberg, l. h., b. sinervo, c. r. linder, m. c. ungerer, and d. m. arias. 1996. role of gene interactions in hybrid speciation: evidence from ancient and experimental hybrids. science 272:741645. smith, m., p. mckenzie, p. mettler-cherry, and s. g. smith. 2004. a putative hybrid of schoenoplectus saximontanus and s. hallii (cyperaceae) from oklahoma. sida 21:475-479. yatskievych, g. 1999. steyermark’s flora of missouri – volume 1 – revised ed. missouri department of conservation, jefferson city. yatskievych, g. and j. turner. 1990. catalogue of the flora of missouri. monograph systematic botany, missouri botanical garden 37:1-345. oklahoma native plant record volume 11, december 2011 smith, m. & mckenzie, p. m. 31 table schoenoplectus hallii (sh), s. saximontanus (ss), and putative hybrid (hy) at wichita mountain wildlife refuge (wmwr) and fort sill military reservation (fsmr) at selected sites: 2002, 2007-2010; boggy flat (bf), elmer thomas (et), engineer pond (ep), grama lake (gl), hollis pond (hl), ingram pond (ip), medicine tank (mt), pottawatomie pond (pp), quanah parker lake (qpl), rock dam (rd), zania pond (zp) date area site species collected *collection # abundance july 2002 wmwr hp sh 2023 ~100 flowering and fruiting culms mt sh 2029 abundant on w shore ss 2027 abundant on s & e shore hy 2028 scattered ip sh 2031 ~ 200 plants scattered et sh 2035 ~ 200 plants scattered bf ss 2026 abundant100,000s of plants oct 2007 wmwr mt sh 2315 common, widely distributed ss 2316 common, widely distributed hy 2317 common, scattered bf ss 2313 ~ 100 plants scattered rd ss 2318 ~ 10,000s plants pond e of rd ss 2314 ~ 100 plants scattered sep 008 wmwr mt sh 2349 thinly scattered ss 2350 thinly scattered hy 2351 concentrated along nw corner of pond aug 2009 wmwr mt sh 2391 uncommon, scattered ss 2392 rare hy 2393 common ip sh 2395 uncommon, scattered qpl sh 2406 rare ss hy 2407 2408 rare, scattered uncommon, scattered rd ss 2390 rare oklahoma native plant record volume 11, december 2011 smith, m. & mckenzie, p. m. 32 table continued date area site species collected *collection # abundance aug 2009 fsmr et sh 2397 30 plants, scattered ep sh 2399 50 plants, scattered zp sh 2400 rare ss 2401 rare hy 2402 common pp sh 2403 abundant ss 2404 uncommon, scattered hy 2405 abundant sep 2010 wmwr mt sh 2459 rare hy 2406 common rd ss 2457 uncommon, scattered; heavily grazed gl ss 2464 uncommon, scattered sep 2010 fsmr et sh 2462 uncommon, scattered pp hy 2461 ~10,000s plants *collections were sent to one or more of the following herbaria: brit, mich, mo, okl, wisc, umo schoenoplectus hallii, s. saximontanus, and the putative s. hallii ×s. saximontanus hybrid: observations from the wichita mountains wildlife refugeand the fort sill military reservation 2002 – 2010 by dr. marian smith and dr. paul m. mckenzie journal of the oklahoma native plant society, volume 8, number 1, december 2008 61 oklahoma native plant record volume 8, number 1, december 2008 smith, b.a. https://doi.org/10.22488/okstate.17.100063 fern habitats and rare ferns in oklahoma dr. bruce a. smith mcloud high school mcloud, oklahoma e-mail: fronds02@yahoo.com this paper features some of the more common fern habitats in oklahoma and provides information on four rare oklahoma ferns from two fern families: aspleniaceae and pteridaceae. surprisingly, ferns can be found in a variety of habitats across oklahoma. introduction with over 2500 species of vascular plants (taylor and taylor 1991), oklahoma is rich in both plant and habitat diversity. the vast majority of oklahoma’s vascular plants are flowering plants. less than 100 species are ferns and fern allies. needless to say, ferns and fern allies do not get the same attention as do flowering plants. one obvious reason is that they are not as showy and do not catch our eye as wildflowers do. secondly, we tend to visit wildflower habitats more often than fern habitats. ferns live in some of the most interesting places, however. if you are in the quartz mountains you may be staring at a western diamondback snake and a star cloak fern on the same rock. if you are hunting the netted chain fern in southeastern oklahoma you may be up to your ankles in mud. one of the objectives of this article will be to introduce you to some of the typical habitats and places that you can find ferns. you will also be introduced to some of the rare ferns of oklahoma. habitat information for some of the species is from the flora of north america (1993). rare species are those listed in the oklahoma natural heritage inventory (2005). collection dates and distribution information are from the oklahoma biological survey database and from personal encounters with the species. authority and common names are from taylor and taylor (1991) and the flora of north america (1993). technical descriptions of each species can also be obtained from the flora of north america. to distinguish between the different taxa i would encourage readers to use field guides and a good dichotomous key such as keys and descriptions for the vascular plants of oklahoma (tyrl et al. 2007) or the illustrated flora of north central texas (diggs et al. 1999). fern habitats one of the best places to look for ferns is on rock outcrops with mosses. rocks are great places to find ferns, no matter what part of the state you are in. ferns can even be found embedded in mosses on trees. if you can’t find them on rocks and trees, look for them in marshes, bogs, mudflats, woodland forests, areas with rocky soils, near waterfalls, and even floating on the water surface. the places you will likely not find them are in lawns or prairies. often, when someone has brought or described to me the leaf of a “fern” they found in such a habitat, it has been achillea millefolium l., the common yarrow. common yarrow is a flowering plant in the composite family asteraceae. do not let the rocky outcrop habitats in quartz mountain resort, or other islands of the wichita mountains (fig. 1), discourage you from looking for ferns. southwestern oklahoma is a great place to see several families of ferns including the aspleniaceae, dryopteridaceae, and especially the maidenhair family, pteridaceae. figure 1 rock outcrop, quartz mountain resort. 62 oklahoma native plant record volume 8, number 1, december 2008 smith, b.a. the overhang of the cave at robbers cave state park and lodge supports a healthy population of asplenium bradleyi d.c. eaton, bradley’s spleenwort, one of oklahoma’s rarer ferns (fig. 2). i have visited this same population many times over the years. the population appears to have grown and is healthier than ever. figure 2 rock outcrop overhang of robbers cave. limestone crevices can hold lichens as well as argyrochosma dealbata (pursh) windham, the powdery cloak-fern (fig. 3). the arbuckle mountains are great places to see several species of ferns, especially the maidenhair ferns. figure 3 argyrochosma dealbata in limestone crevice with lichens, turner falls in the arbuckle mountains. cheilanthes lanosa (michx.) d.c. eaton, the hairy lipfern, grows on rocky soil with a spike moss on elk mountain in the wichita mountains (fig. 4). other granitic rocks on which to find this fern are in the great plains state park and quartz mountain resort. figure 4 cheilanthes lanosa with spike moss. cheilanthes lanosa also grows on other rock types such as the limestone at beavers bend resort park and robbers cave state park and lodge. this fern is one of the few in oklahoma that has the ability to take over large patches of hillsides in open areas (fig. 5). figure 5 cheilanthes lanosa in an open area, beavers bend resort park. i do not know of any oklahoma epiphytic ferns other than pleopeltis polypodioides (l.) andrews & windham (figs. 6 & 7), the resurrection fern. it is common in eastern forests on both mossy covered rocks and mossy covered trees. it occurs as far west as johnston county. 63 oklahoma native plant record volume 8, number 1, december 2008 smith, b.a. figure 6 pleopeltis polypodiodes , the resurrection fern, in idabel city park. figure 7 p. polypodioides growing as an epiphyte. osmunda cinnamomea l., cinnamon fern, grows under the canopy of a mesic forest in choctaw county (fig. 8). cinnamon ferns can live in acidic soils, vernal seeps, and moist areas. cinnamon ferns can be seen on public land at ferndale bog in mcgee creek state park. the best time to visit them is in may when you can see their cinnamon colored fertile fronds. figure 8 cinnamon fern in choctaw county woodwardia areolata (l.) t. moore (fig. 9) grows in wet forest soils in choctaw county, but they can also be found in seeps and acidic bogs. look carefully at the erect fertile frond in the foreground. the elongated sori of each leaflet fit end to end forming a chain, thus the common name, netted chain fern. growing laterally in the background you can see the sterile fronds that do not produce sori. both fronds are part of the same rhizome. this is an interesting species to see, especially when both types of fronds are present. it is a southeastern oklahoma species that can be seen in at least five counties. figure 9 woodwardia areolata with fertile frond in foreground. onoclea sensibilis l. (fig. 10) grows in marshy soils on mccurtain county roadsides. sensitive ferns can be found in open swamps, thickets, 64 oklahoma native plant record volume 8, number 1, december 2008 smith, b.a. marshes, or lowland woods. like woodwardia areolata, the sensitive fern has separate fertile (brown) fronds and sterile (green) fronds. this species has a much wider distribution than the netted chain fern. the sensitive fern is seen as far west as creek county. figure 10 onoclea sensibilis with fertile brown fronds. mudflats like the one at the university of oklahoma biological station in marshall county are not the greatest habitat to look for ferns (fig. 11). however, marsilea vestita hook. and grev., water clover, was collected there in 2006. figure 11 mudflat habitat at ou biological station. on the falls and rocks in the creek area at price falls in falls creek baptist assembly (fig. 12) you can see adiantum capillus-veneris l., the southern maidenhair fern. the tissue thin fronds require moist cool air to survive. climb fifteen feet above the waterfall on the rock and away from the creek and you will not find it. falls creek is a wonderful place to find several species of the pteridaceae: adiantum capillus-veneris, argyochosma dealbata, astrolepis integerrima (hook.) benham & windham, cheilanthes tomentosa l., and pellaea atropurpurea (l.) link. figure 12 price falls at falls creek baptist assembly. rare ferns i do not remember when i first became a “pteridomaniac”. the spore must have begun developing in 1977 after enrolling in my first field botany course, plant taxonomy, under dr. doyle mccoy. since 1977 i have taken my share of botany field trips all over the state. in fact, i consider every day a botany field trip, always looking for that fern or other plant that i have never seen as well as those “old friends”, as dr tyrl would call them, like asplenium platyneuron (l.) britton, sterns & poggenb. (fig. 13) and woodsia obtusa (spreng.) torr. (fig. 14), two ferns that are as common as dandilions. figure 13 asplenium platyneuron, ebony spleenwort, a very common fern. 65 oklahoma native plant record volume 8, number 1, december 2008 smith, b.a. figure 14 woodsia obtusa, blunt-lobed cliff fern, another common fern the following species are relatively rare in oklahoma and are listed as “species of concern” by the oklahoma natural heritage inventory (2008). the first three are in aspleniaceae family and the last is in pteridaceae. i have had the good fortune to see each of them more than once, some in multiple locations, others in only one location. asplenium bradleyi d.c. eaton (fig. 15) common name: bradley’s spleenwort distribution: latimer county, also seen in atoka county. note: this is a difficult species to describe, but it can easily be identified using a field guide or dichotomous key. figure 15 asplenium bradleyi growing on sandstone rock, robbers cave state park and lodge. asplenium pinnatifidum nutt. (fig. 16) common name: lobed spleenwort distribution: latimer county. note: this species has only been reported at robbers cave state park and lodge, but there are several populations throughout the park, including robbers cave. figure 16 asplenium pinnatifidum, robbers cave state park and lodge. asplenium septentrionale (l.) hoffm. (fig. 17) common name: forked spleenwort distribution: cimarron county. note: this fern does not have the typical fern appearance. the novice might even mistake it for a grass. the fronds have a grass-like appearance with narrow linear blades. the blade apex can be forked, thus its common name. figure 17 asplenium septentrionale, north of black mesa state park and nature preserve. 66 oklahoma native plant record volume 8, number 1, december 2008 smith, b.a. cheilanthes wootonii maxon (fig. 18) common name: beaded lipfern distribution: cimarron, greer, and kiowa counties, but also seen in canadian county. note: cheilanthes species are difficult to identify. c. wootonii hook. can easily be mistaken for c. eatonii baker, c. tomentosa link, or even c. lindheimeri. you’ll need a good dichotomous key such as keys and descriptions for the vascular plants of oklahoma (tyrl et al. 2007) to identify members of this genus. figure 18 cheilanthes wootonii, methodist canyon camp. conclusion i hope you will visit a fern habitat on a future field trip. you do not need to wait until spring to see ferns because there are several species in our state that are evergreens. you will find ferns to be both fascinating and beautiful. if you are fortunate enough to come across one of these rare ferns, please practice good conservation by not collecting it and by protecting its habitat. acknowledgements my thanks to the following individuals: richard butler for accompanying me on many field trips the last five years and helping to edit portions of the article: mickey cooper for giving me my start in botany; doyle mccoy for giving me my start in oklahoma native plants; ron tyrl for training me as a botanist and providing so many great botanical opportunities; richard butler, catherine eimen, bruce hoagland, and sheila strawn, for helping to edit portions of this article; and to my wife, sharon, for helping to edit the article and for allowing me to pursue my passion. i also give my thanks for access to these properties: hoby family property; university of oklahoma biological station; falls creek baptist assembly; methodist canyon camp; southwest baptist assembly; turner falls; idabel city park; and the following state facilities: beavers bend resort park, black mesa state park and nature preserve, great plains state park, mcgee creek state park, quartz mountain resort, red rock canyon state park, and robbers cave state park and lodge. literature cited diggs gm jr., lipscomb bl, and o’kennon r. 1999. shinner’s & mahler’s illustrated flora of north central texas. botanical research institute of texas, sida, botanical miscellany no. 6. flora of north america, editorial committee. flora of north america. 1993. vol 2. pteridophytes and gymnosperms, new york, oxford university press. oklahoma natural heritage inventory. 2008. oklahoma natural heritage inventory working list of rare oklahoma plants. oklahoma biological survey. <www.oknaturalheritage.ou.edu/plants> accessed december 2008. taylor rj and taylor ces. 1991. an annotated list of the ferns, fern allies, gymnosperms and flowering plants of oklahoma. biological department herbarium, durant, southeastern oklahoma state university. tyrl rj, barber sc, buck p, elisens wj, folley pa, magrath lk, murray cl, smith ba, taylor ces, and thompson ra. 2007. keys and descriptions for the vascular plants of oklahoma. noble, flora of oklahoma, inc. journal of the oklahoma native plant society, volume 7, number 1, december 2007 78 oklahoma native plant record volume 7, number 1, december 2007 the need for savanna restoration in the cross timbers caleb stotts michael w. palmer kelly kindscher restoration technician botany department kansas biological survey tallgrass restoration oklahoma state university university of kansas and management, lawrence, ks stillwater, ok lawrence, ks along the prairie/forest transition zone oak savannas have been severely degraded by logging, clearing for agriculture, fire suppression, invasion of exotic plants, and excessive livestock grazing. savanna shares equal billing with tallgrass prairie as the most threatened plant community in the midwest. as such, there is increasing interest in restoring these communities. conservation criteria have not been developed for the post oak (querces stellata) and blackjack oak (querces marilandica) savanna of the cross timbers. oak savanna was arguably an important component of the historical cross timbers region. following settlement, overgrazing in conjunction with a decrease in fire frequency and/or intensity has increased the density of oak stands to the point where they resemble closed-canopy forests rather than savanna. this is a threat to the biodiversity of the cross timbers. proactive land management practices are recommended for restoring savanna communities. such efforts may require thinning-out areas of degraded oak savanna to help re-establish the herbaceous understory. fire is recommended to restore ecological processes that limit woody plant encroachment and promote biodiversity. further research should investigate the ecological dynamics and functions of oak savannas, as well as provide further guidelines for its conservation. introduction along the prairie/forest transition zone, oak savanna communities have been severely degraded by logging, clearing for agriculture, fire suppression, invasion of exotic plants, and excessive livestock grazing (abrams 1992). oak savanna shares equal billing with tallgrass prairie as the most threatened plant community in the midwest and among the most threatened in the world (henderson 1995). as such, there is increasing interest in restoring these communities (whitney and decant 2005). in the cross timbers region, however, there has been little effort to evaluate the conservation status of savannas or woodlands. community classification in the prairie/forest transition zone, upland communities are not always discrete entities separated by sharp lines. instead, they often blend into each other imperceptibly. even so, named communities are useful abstractions that help us think stotts, et al. https://doi.org/10.22488/okstate.17.100055 and communicate about various parts of the landscape (palmer and white 1994, packard and mutel 1997). definitions adapted from faber-langendeon (2001) and lauver et al. (1999) provide us with an operational classification for common midwestern upland communities: 1) prairie – areas dominated by herbaceous vegetation (grass and forbs); trees generally not exceeding 10% cover; 2) savanna – areas dominated with herbaceous vegetation and scattered trees with 10-25% cover; 3) woodland – areas dominated by an open stand of trees with 25-60% canopy cover and a herbaceous understory; and 4) forest – areas dominated by trees with 60-100% cover and little herbaceous vegetation. these communities are illustrated in fig. 1. savanna is maintained by frequent fire. along the prairie-forest transition zone, certain species of oaks are the only trees that were historically savanna. this is in a large part due to their physiological adaptations to fire, which include thick bark, prolific resprouting and resistance to rotting after scarring (abrams 1992). 79 oklahoma native plantrecord volume 7,number 1,december 2007 stotts et al. just what the understory and ground layer vegetation of oak savanna was like historically is largely unknown (henderson 1995). while no plant species is known to be endemic to oak savanna (nuzzo 1985), there are species that are considered savanna specialists in the midwest (packard 1988). historically, the savanna community was probably a slowly shifting mosaic of plant species associations that had varying degrees of shade and sun tolerance (henderson 1995). cross timbers savannas the cross timbers region is located in portions of oklahoma, texas, kansas, and arkansas (fig. 2). it is characterized by a mosaic of upland communities including prairie, savanna, woodland and forest (fig. 3). post oak (quercus stellata) and blackjack oak (quercus marilandica) are the dominant tree species throughout in the wooded systems. kuchler (1964) defined the potential natural vegetation of the cross timbers as savanna-like, characterized by tallgrass prairie with low broadleaf deciduous trees scattered singly or in groves of varying size. these groves often occur with an open canopy cover and grassy understory (kuchler 1974). the herbaceous understory of cross timbers savanna is similar in composition to the surrounding prairie (dyksterhuis 1948; kuchler 1964, 1974, palmer unpublished data). savanna also occurs in the cross timbers region as a gradual transition between closed-canopy forests and grasslands, with a margin of isolated trees (dyksterhuis 1957, penfound 1962). this sort of edge can be tens of meters wide. classifying some cross timbers sites as savanna can be problematic due to the tendency of post oak and blackjack oak to root sprout and produce groupings of trees with interlocking crowns (hoagland et al. 1999). in the cross timbers region, woodlands have a similar species composition as savanna (palmer, unpublished data). as such, we recognize that many properties of savanna are likely to be shared with woodlands, and we treat the two as largely synonymous in this paper. restoration of midwest oak savannas nuzzo (1985) estimated that oak savannas in eight states in the midwest probably covered 11 to 13 million hectares at the time of settlement and have been reduced in extent by 99.98%. packard (1988) found that several plants that were historically associated with savanna communities are now uncommon. populations of these ‘savanna specialists’ have been successfully established through restoration efforts. largely because of these findings, the conservation value of savanna communities has been recognized and restoration efforts are increasing. the ultimate goal is to help replace the loss of habitat that is leading to the gradual disappearance of plant and animal species (packard 1988). midwest oak savanna vs. cross timbers savanna the cross timbers and certain areas of the midwest occupy a transition zone between the great plains and the eastern deciduous forest. despite this, the savannas of the cross timbers are considered distinct from midwest oak savannas to their north. (mcpherson 1997). the midwest is characterized by its former glaciation, relatively mesic soils and northern plant affinities, while the cross timbers region is characterized by its largely sandy soils, generally rough topography and southern plant affinities. furthermore, the cross timbers has not experienced the extent of sod-busting that the midwest has, and 80 oklahomanative plantrecord volume 7,number 1,december 2007 stotts, et al. includes substantial areas of native tallgrass prairie and old-growth forest. despite these distinctions, there is very little difference in ecosystem classification. küchler (1964) described regions of oak savanna in the midwest as being nearly identical to that of the cross timbers in vegetation type; characterized by tallgrass prairie with broadleaf deciduous trees scattered singly or in groves. historical and current extent of cross timbers savanna the extent to which we can understand the structure of pre-settlement vegetation is limited. despite this, analysis of historical accounts, early photographs, early land surveys, and existing vegetation have provided much insight into historical vegetation. numerous authors have described historical vegetation communities throughout the cross timbers region as savanna-like (bruner 1931, dyksterhuis 1957, 1948, lathrop 1958, rice and penfound 1959, penfound 1962, kuchler 1974, 1964, johnson and risser 1975, smiens and diamond 1986, hoagland et al. 1999, francaviglia 2000). this is not to conclude that savanna was the dominant vegetation type in the cross timbers. it does indicate, however, that savanna was a well-represented component within a mosaic of prairie and forest during the time of settlement. many authors conclude that, during post-settlement, overgrazing in conjunction with a decrease in fire frequency and / or intensity has increased the density of oak stands to the point where they resemble closed-canopy forests rather than savanna (dyksterhuis 1948, 1957, lathrop 1958, rice and penfound 1959, penfound 1962, bell and hulbert 1974, johnson and risser 1975, smiens and diamond 1986, abrams 1992, hoagland et al. 1999). this conversion has been at the expense of the herbaceous understory and the associated biodiversity. unlike the midwest oak savannas, there are no reliable estimates as to how much cross timbers savanna actually existed at the time of settlement or how much has been lost since settlement. despite this, these studies indicate that savannas were important aspects of the historical cross timbers region and now represent only a remnant of a vast vegetation type. biodiversity and natural heritage the mosaic of communities in the cross timbers provide for a wide variety of habitat for plants and animals (costello 1969, oklahoma biodiversity plan 1993), and savannas contribute to this habitat diversity (fig. 4). savannas may produce an edge effect, where interfaces between community types support species from both communities, resulting in elevated species composition. as in the midwest, there may be savanna specialists in the cross timbers, species that prefer the distinct habitat offered by an open stand of trees. cross timbers savanna should be valued in regards to their conservation status for their contribution to the natural heritage of the united states. this is especially true for post oak trees that have reached the age of 200+ years (fig. 5). according to the oklahoma biodiversity plan (1993), foremost among the threats to plant diversity in oklahoma is a dramatic change in the fire regime from what occurred historically. as the result of an altered fire regime, the encroachment of woody species into savannas is indeed a threat to the diversity of the cross timbers (rice and penfound 1959, johnson and risser 1975, johnson 1986, archer 1995, hoagland et al. 1999). 81 oklahoma native plantrecord volume 7,number 1,december 2007 stotts et al. figure 1 schematic diagram showing the changes in structure along a gradient from prairie to forest. this structural gradient is often reflective of a fire frequency gradient, with prairies maintained by more frequent fires (faber-langendeon 2001). figure 2 location of the cross timbers region. (adapted from küchler 1964). figure 3 a cross timbers mosaic of prairie, savanna and forest communities. 82 oklahomanative plantrecord volume 7,number 1,december 2007 stotts, et al. figure 4 a blackjack oak savanna. these scattered trees provide for habitat diversity. figure 5 this old-growth post oak tree has low, horizontal branches. this type of architecture may be indicative of its having grown in an open-canopy environment. 83 oklahoma native plantrecord volume 7,number 1,december 2007 stotts et al. restoration recommendations restoration is the work of enhancing ecological quality. high quality communities have most natural processes intact and are rich in conservative plant species; those that are restricted to intact, natural remnants. disrupted or degraded systems (those that have been plowed, overgrazed, protected from fire, etc.) lose those conservative species. the principal challenge in remnant restoration is to reinstate or speed up the processes that allow these remnantdependent species of plants and animals to regain their important roles in the system (packard and ross 1997). several authors have commented on the need for proactive land management to combat woody encroachment in the cross timbers (dyksterhuis 1948, smiens and diamond 1986, engle et al. 1996, 2006, francaviglia 2000). proactive land management practices are indeed recommended for restoring savanna. in degraded savannas, a combination of treatments is recommended for restoring an open-stand of trees with a grassy understory. mechanical removal of trees with tree-clipping devices and/or chainsaws may be used to thin dense stands. for areas thick with shrubs, mowing treatments may be used. fire should be used as a process to re-establish native grasses and forbs, with a long-term goal of promoting plant diversity and limiting woody encroachment. there are many acres of private land in the cross timbers with degraded oak savanna. a major obstacle to restoring natural diversity on private lands has been the lack of economic incentive. savanna restorations, however, may provide increased forage and combat further loss of forage due to woody encroachment. light to moderate grazing can be compatible with maintaining the plant structure needed by many savanna species (henderson 1995). in addition to providing optimum habitat for many plant and wildlife species, oak savanna was probably the optimum habitat for many game species (e.g., bobwhite quail, turkey, deer, and rabbits) (henderson 1995). thus, management for oak savanna is compatible with traditional wildlife management and hunter interests. the ultimate goal should be to help restore habitats, the loss of which, has lead to the gradual disappearance of plant and animal species (packard 1988). for example, the black-capped vireo is a native to the cross timbers region. this federally endangered species prefers to nest in open savanna vegetation, and the decrease in open savanna vegetation has had negative impacts on the population (hoagland et al. 1999). this is a prime example of how savanna restoration efforts could increase biodiversity by providing habitat for a target species. currently, savannas are not well represented throughout the cross timbers. much of the cross timbers vegetation is now characterized by a mosaic of prairie and closed canopy forest. by restoring savanna communities, the structural diversity of the landscape is increased. these efforts will likely lead to higher compositional and functional diversity. mendelson et al. (1992), however, criticize what they believe is a rush to create savannas on forested sites that never supported savannas. most crucially for the cross timbers, there are old-growth forests in the region that have never been savannalike. such forests are clearly not a target for savanna restoration. careful research should be used to plan and implement any particular savanna restoration project (see packard and mutel 1997). managers need to understand the characteristics of the site and the potential impacts of restoration techniques. analysis of the site’s existing plant communities and 84 oklahomanative plantrecord volume 7,number 1,december 2007 stotts, et al. rare plant or animal populations is crucial. inference of pre-settlement vegetation through analysis of government land office (glo) surveys, soils, and topography should help guide the process. environmental factors influencing cross timbers savannas savanna represents one component of a complex and dynamic ecosystem. within the cross timbers, there are several interacting environmental factors influencing vegetation for a given area. these include 1) climate; 2) soil; 3) topography; 4) grazing; and 5) fire. understanding how all of these factors influence the relative abundance of woody and herbaceous plants is fundamental to managing for and restoring native savanna communities (mcpherson 1997). climate the cross timbers is home to a dynamic climate that is capable of supporting grassland or forest. there have been long-term ‘dry’ and ‘moist’ events, punctuated with shorter-term cyclic variations in climatic conditions (dean et al.1984). the climate of the cross timbers has varied substantially even over the last few centuries, where changes in rainfall patterns have caused east-west shifts in the ecotone (shaw and lee 1995). interannual and decadal variability in precipitation and temperature have been naturally high at both local and regional scales (mcpherson 1997). as precipitation regimes shifted, so did community composition and structure (wright 1963). extreme climatic events may be more important than shifts in means (katz and brown 1992) for changes in cross timbers savannas. “pulses” of tree recruitment may occur during relatively brief periods of high soil moisture (mcpherson 1997). wet fuels decrease the likelihood of fire and allow for trees to take advantage of the higher soil moisture. subsequent growth of woody plants, may transform prairie into savanna or savanna into forest (jameson 1987). on the other hand, the fine fuels which accumulate during these periods of high precipitation may also dispose the system to intense fire and thereby limit tree recruitment (scholes and archer 1997). significant destruction of cross timbers trees during long periods of drought have been documented (rice and penfound 1959). while grasses are also damaged by drought, they may rapidly reestablish areas due to their propagation by rhizomes once there is sufficient soil moisture (weaver 1968). major droughts in the cross timbers region occur at unpredictable intervals. such droughts may increase the chance of fire due to dry fuels (axelrod 1985), however, it may decrease fire intensity due to decreased fuel production (skarpe 1992). due to the effects of a variable precipitation and fire regime, cross timbers savannas have possibly experienced a high degree of shifting on the landscape, as well as conversions to full prairie or forest. present vegetation may represent one phase of a continually changing assembly of communities (wethington 1994). this information is important for predicting how a natural savanna community might respond to changes in climate. soils the very existence of cross timbers trees is largely traceable to certain geologic units from which the sandy soils are derived (dyksterhuis 1948). these alternating materials have formed different soil associations that are characterized by coarse-textured sandy loam soils and by fine-textured clay loam soils. these are generally associated with savanna or forest, and grassland respectively (dyksterhuis 1948, smeins and diamond 1986). 85 oklahoma native plantrecord volume 7,number 1,december 2007 stotts et al. studies in the cross timbers have indicated that soil moisture availability is the primary factor controlling species composition (clark 2003, johnson and risser 1972, rice and penfound 1959). the higher moisture-retaining capacity of coarsetextured soils is largely responsible for supporting the higher water demands of trees where rainfall is marginal for tree survival (bell and hulbert 1974). finetextured soils may reduce water availability to woody plants below thresholds necessary for survival in the dry summers (mcauliffe 1994). the usda (2007) characterizes certain soil types in the cross timbers as ‘savanna’ range site. these are the most likely locations in which to restore a degraded savanna. topography topography influences the ‘fire probability pattern’ (grimm (1984) that results from frequent fires superimposed on landscape features that include fire-prone topographic regions as well as natural fire barriers. frequency of fires for a prairieforest ecotone in pre-settlement times was largely determined by topographic relief and the distribution of firebreaks, such as waterways (anderson 1990). because fire frequency was determined by the roughness of landscape features, the density of trees on a landscape can often be viewed as a function of surface roughness (anderson 1990). old-growth forest in the cross timbers is highly related to steep and rocky slopes (therrell and stahle 1998). much of the cross timbers forest prior to settlement was likely associated with a fire-protected landscape. as previously mentioned, old growth forests are not the place for savanna restoration. grazing native herbivores influenced the proportion of woody and herbaceous plants by disproportionately consuming or damaging more of one vegetation type than the other (mcpherson 1997). as such, herbivores may interact with competition patterns between woody and herbaceous vegetation as well as with fire regimes, and may thus be involved in large-scale physiognomic dynamics of savannas (skarpe 1991). ungulates like bison, elk, deer and pronghorn antelope, among other herbivores were all present on the historical prairie/forest transition. of these, bison may have had the greatest impact on woody plant establishment in terms of their huge numbers and their alteration of fire intensity (shaw and lee 1995). high grass biomass can affect tree biomass by fueling fires. bison grazing could have reduced the fuel load and reduced fire frequency, intensity, or continuity of spread (baisan and swetham 1990). however, bison herds are believed to have existed in low numbers in the cross timbers (shaw and lee 1995). the effects of overgrazing cattle likely differed drastically from historical bison grazing in the cross timbers. in the absence of heavy cattle grazing, a considerable quantity of litter was produced between established trees. when fires started with these heavy fuel loadings, small trees and saplings were knocked back. the result was an open stand of timber (penfound 1962). in managing for savanna communities, overgrazing should not be allowed to reduce the fuel loading to the point where fire cannot suppress woody plants. fire fire has influenced plant communities for millions of years. fires are thought to be important for the origin and maintenance of grassland, savanna, and woodland community physiognomies by limiting woody plant establishment (anderson 1990, sullivan 1995, dorney and dorney 1989). native americans have been in the southern plains for more than 10,000 years 86 oklahomanative plantrecord volume 7,number 1,december 2007 stotts, et al. (kay 1998), during which they set frequent fire to the tallgrass prairie landscape (shaw and lee 1995, moore 1995). fire may promote grasses or woody plants in cross timbers savannas, as both vegetation types are well-adapted to fire. fire frequency, fire intensity, and fire season interact to shape the response of vegetation to fire (wright and bailey 1982, engle et al. 1996). a given fire may favor either grasses or trees depending on the nature of these interactions. the frequency of fire plays a critical role. in savanna ecosystems, a decrease in fire frequency leads to woody encroachment, while more frequent fires may favor a relatively stable community (scholes and archer 1997). frequent fires, however, do little to suppress woody plant development if they are of low intensity (briggs et al. 2005). fire intensity varies as a function of weather, stage of plant development, fuel load, topography, soil type, and previous management (bidwell et al. 2004). generally, a well managed rangeland with plenty of fine fuels will produce a high intensity fire that may effectively control woody plant establishment. this underscores the importance of the current vegetation in not only shaping the fire environment, but also in the response of vegetation to a given fire (engle et al. 1996). the season of a fire is very important for the relative effect on grasses and woody plants. the way species respond to a fire depends heavily on the timing of the fire relative to their phenological development. in general, plants that are actively growing, flowering, or setting seed at the time of the fire, tend to decline over time (davidson and kindscher 1999). burning at different times of the year is recommended to inhibit certain species from dominating the community and to promote biodiversity. to control woody plants, burning following bud break and full leaf-out is the most effective time (bidwell et al. 2004). once a savanna is re-established, carefully prescribed burns can maintain open stands of cross timbers oaks for long periods of time (engle et al. 2006). used wisely, prescribed fire can enhance biodiversity, combat tree encroachment, reduce danger of catastrophic wildfires, and improve range conditions for livestock. research needs the current extent of high-quality savanna stands should be assessed throughout the cross timbers. judgments must be made as to the degree to which stands of vegetation appear to be functioning under natural ecological processes. plant identification in highquality stands of oak savanna should be used to provide information on flora composition, richness and physiognomy. lists of fauna that utilize and prefer these communities should be compiled. this information can be used to assess the integrity and functions of savanna communities, to analyze their contribution to the biodiversity of the cross timbers, and as reference information for restoration efforts. while numerous studies indicate that savannas were important components of the historic cross timbers, their actual extent is uncertain. assessing the actual past extent of savanna remains a top research priority. if savanna historically dominated the cross timbers region and are now very poorly represented, their conservation would be a very high priority. if savannas were originally rare and transient, they would deserve less attention than if they are the last remnant of a vast vegetation type. unfortunately, tools for assessing past extent of savanna vegetation are limited. glo surveys are perhaps the best available tool. early land survey records have contributed significantly to our understanding of the structure of north america’s pre-settlement ecosystems. by 87 oklahoma native plantrecord volume 7,number 1,december 2007 stotts et al. way of summary, land surveys have been used to determine: 1) species compositions of pre-settlement savannas and woodlands; 2) landscape-level disturbance processes; 3) site-specific determinants; 4) species associations and community classification, and; 5) vegetation types for mapping purposes (egan and howell 2005). this information has figured prominently in the restoration of a number of historic ecosystems (egan and howell 2005). schroeder (1981), for instance, created a statewide map of glo surveys from missouri that described a mosaic of forest, woodland, savanna, and prairie landscapes. the map serves as a foundation for the missouri department of natural resources efforts to restore savanna ecosystems in that state’s parks (mccarty 1998). this information is commonly used as a reference for restoration efforts, and numerous post oak savanna restorations have occurred with success in missouri. the plat maps used for mapping, however, were made up solely on the basis of data written in the early surveyor’s notes, which have certain biases and limitations (king 1978). furthermore, we should view this information as but one snapshot of past vegetation patterns that were constantly shifting with an ever-changing climate, native american activities (batek et al. 1999), and grazing patterns. also, early settlers may have cut down trees before the survey was completed. as such, we are forced to consider just how representative they are as a true picture of the “presettlement” vegetation (noss 1985). the dynamics of savannas are not well known because landscape-level processes have been radically, and sometimes irreversibly altered by recent human activities. (rebertus and burns 1997) further research should increase our understanding of the mechanisms of the cross timbers ecosystem. elucidation of the interactions, dynamics and determinants, and identification of robust generalizations that can be broadly applied to savanna ecosystems would benefit ecological theory, modeling and land management (house et al. 2003). fundamental questions include: what controls the relative abundance of woody and herbaceous plants for a given set of conditions at given site? how do the vegetation types interact with each other? is a given woody-herbaceous ratio dynamically stable and persistent under a particular set of conditions (house et al. 2003). finally, circumstances under which restoration techniques are effective or ineffective need to be identified. as such, restoration efforts should be monitored. conclusion oak savannas throughout the cross timbers region have been degraded by woody encroachment. savanna restoration efforts are recommended to combat this threat to biodiversity. the ultimate goal is to restore ecological processes and help replace lost habitat that is leading to the gradual disappearance of plant and animal species. there is, however, much that is unknown about the ecological dynamics and functions of savanna communities. it is hoped that with research and restoration of savanna communities, some answers will be provided. acknowledgments the authors recognize the invaluable contributions of the following: the stotts family, jim minnerath, daniel dyer, the usfws eastern kansas district fire crew, and the stotts. literature cited abrams, m.d. 1992. fire and the development of oak forests. bioscience 52: 346-353. anderson, r.c. 1990. the historic role of fire in the north american grassland. in: collins, s. l. and l.l. wallace. fire in north american tallgrass prairies. 88 oklahomanative plantrecord volume 7,number 1,december 2007 stotts, et al. norman and london: university of oklahoma press. p 8-18. archer, s. 1995. tree-grass dynamics in a prosopis-thornscrub savanna parkland: reconstructing the past and predicting the future. ecoscience 2: 83-99. axelrod, d.l. 1985. rise of the grassland biome, central north america. the botanical review 51: 163-201. batek, m.j. 1999. reconstruction of early nineteenth-century vegetation and fire regimes in the missouri ozarks. journal of biogeography 26: 397-412. bell, e. and hulbert, l. 1974. effect of soil on occurrence of cross timbers and prairie in southern kansas. transactions of the kansas academy of science. baisan, c.h. and t.w. swetnam. 1990. fire history on a desert mountain range: rincorn mountain wilderness, arizona, usa. canadian journal of forestry restoration 20: 1559-1569. briggs, j.m., a.k. knapp, j.m. blair, j.l. heisler, g.a. hoch, m.s. lett and j.k. mccarron. 2005. an ecosystem in transition: causes and consequences of the conversion of mesic grassland to shrubland. bioscience 55: 243-254. bruner, w.e. 1931. the vegetation of oklahoma. ecological monographs 1(2): 100-113. clark, s. l. 2003. stand dynamics of an oldgrowth oak forest in the cross timbers of oklahoma. 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https://doi.org/10.22488/okstate.17.100084 43 the effects of removal of juniper us virginiana l. trees and litter from a central oklahoma grassland jerad s. linneman1 1usda matthew s. allen2 animal & plant health inspection service michael w. palmer2 4700 river rd., unit 153, 5d-06g mike.palmer@okstate.edu riverdale, md 20737-1228 2oklahoma state university department of botany 301 physical sciences stillwater, ok 74078-3013 keywords: biodiversity, experiment, invasion, ordination abstract we studied species composition after juniperus virginiana tree and litter removal in a central oklahoma grassland. tree removal had the most significant effect on stems per quadrat and vegetation cover. litter removal effects were not as strong. however, stems per quadrat and vegetation cover in litter removal treatments were higher than in litter intact treatments. species richness increased for all treatments in the first year post-treatment, after which species richness declined at every sampling period and in every treatment for the duration of the study. absolute cover of typical prairie species increased in the cut with no litter treatment whereas cover of woody forest species increased in the no cut with no litter treatment. we suggest that even without prescribed fire, redcedar tree removal may result in a return of prairie vegetation. however, additional efforts besides tree removal may be required to restore some invaded grasslands. introduction for the last several decades, there has been a growing interest in management techniques required to maintain and/or restore vegetation. the two most common problems faced in grassland restoration are habitat destruction and the loss of native species diversity due to the encroachment of woody species. concerns about decreased diversity and the invasion of exotic woody species have spurred extensive study throughout the world including argentina (ghersa et al. 2002), australia (costello et al. 2000, whiteman and brown 1998), canada (peltzer and köchy 2001), french prealps (barbaro et al. 2001), south africa (holmes et al. 2000, holmes and marais 2000) and the united states (petranka and mcpherson 1979, callaway and aschehoug 2000, fitch et al. 2001, briggs et al. 2002b, van els et al. 2010). in the united states, two examples of fire adapted vegetation types that have received much attention regarding restoration are the longleaf pine sandhill vegetation of northwestern florida (kush et al. 1999, provencher et al. 2000, provencher et al. 2001) and the tallgrass prairie of the eastern great plains (axmann and knapp 1993, briggs et al. 2002a, briggs et al. 2002b). in both instances the elimination of fire has caused a decrease in species richness and facilitated their conversion into forests. tallgrass prairie researchers have suggested that reductions in abundance and altered community composition are related to a oklahoma native plant record volume 11, december 2011 linneman, j. s., et al. 44 multitude of environmental factors associated with woody invasion. examples of such altered environmental factors include soil moisture (engle et al. 1987, facelli and pickett 1991b), solar radiation (smith and stubbendieck 1990, facelli and pickett 1991a & b) and soil temperature (weaver and rowland 1952, hulbert 1969). in addition, leaf litter from woody species may alter grassland litter dynamics (facelli and pickett 1991b). within the tallgrass prairie region, eastern redcedar (juniperus virginiana l.) has increased dramatically, converting millions of hectares of grassland to woodland or closed canopy forest (schmidt and leatherberry 1995, briggs et al. 2002a). redcedar invasion is not restricted to impacted or degraded sites and exhibits high survivorship in diverse native grasslands (ganguli et al. 2008). typical control methods include mechanical felling via chainsaws, large cutting machinery, or cabling and prescribed fire. although felling and prescribed fire are effective in reducing redcedar abundance in prairies, the continuous application of this management technique has left a significant gap in our understanding about the role redcedar litter plays in tallgrass prairie restoration. in particular, we do not understand the role of the overstory tree versus the leaf litter in determining species composition. we conducted this study to disentangle the effects of redcedar overstory canopy and accumulated litter on prairie species richness and composition. elucidating these effects will allow for a more informed approach to redcedar removal and prairie restorations. methods study site we conducted this experiment at the james k. mcpherson botanical preserve located 16 km west of stillwater, oklahoma (36°06'00"n, 97°12'30"w). after a brief period of row crop agriculture, the site was converted into pastureland and grazed until the 1960's. oklahoma state university (osu) purchased the land and managerial control was turned over to the department of botany. in 1995, the department of botany introduced a burning regime, consisting of a three to five year return interval, to the northwestern half of the preserve with the goal of stimulating the return of a native tallgrass prairie community. t ree selection and classification we selected 47 potential study trees based on several criteria including tree isolation, minimization of surrounding tree effects, the existence of an intact litter layer underneath the tree, and tree size. we recorded canopy diameter in the northsouth and east-west direction, height, stem diameter at both 10 cm and diameter-atbreast-height (dbh), and gender. for those trees with multiple stems, we recorded separate diameter measurements for each primary stem, which we later converted into basal area (ba) at 10 cm and dbh, respectively. we randomly assigned all trees into two groups (cut and no cut); ten study trees were then randomly selected from each group. sampling desig n sampling design was based on a two by two factorial design of tree removal and litter removal. underneath each study tree, we positioned two 50 cm × 50 cm quadrats so that each quadrat was completely under the canopy of the overstory redcedar. in addition, we positioned the two quadrats in such a way to maintain homogenous litter cover between quadrats and to minimize inter-quadrat variation in vegetation. after permanently marking each quadrat, we randomly assigned a litter removal treatment to one of the two quadrats under each tree. we conducted an initial vegetation sampling in may 2001, prior to treatment application. oklahoma native plant record volume 11, december 2011 linneman, j. s., et al. 45 all subsequent sampling occurred biennially in may and september of 2002 – 2003. sampling of species composition consisted of identifying each plant species rooted inside the quadrat and estimating its percent cover to the nearest percent for any cover less than 5% and to the nearest 5% for any cover over 5%. we marked unknown species for later identification. species nomenclature and code symbols follow that of the usda plants database (usda 2004). in addition, at several locations within this paper we refer to the response of j. virginiana redcedar seedlings and not the study tree or any of its structures. experimental treatments the tree removal treatment was applied using a chainsaw and pruning shears between 17 and 19 may, 2001. we removed crowns and branches from the top down, with the aid of rigging equipment, to minimize the amount of disturbance to the litter layer and vegetation in the quadrats. we removed litter from litter removal quadrats by hand, taking care to minimize disturbance to vegetation. however, plants that had germinated in the litter layer and had not reached the soil surface were removed along with the litter during the initial treatment. the litter removal treatment was applied between 21 and 24 may, 2001. treatment acronyms for tree and litter removal are: cut with no litter (cn), cut with litter (cl), no cut with no litter (nn), and no cut with litter (nl); i.e. the control. at each post-treatment sampling, we removed newly accumulated litter from the litter removal quadrats after observing vegetation. on a few occasions we removed branches from surrounding trees that started to grow over the tree removal quadrats. statistical analyses statistical analysis included the use of both anova and ordination techniques. we performed repeated measures anova using proc mixed for each environmental variable recorded using sas (version 8). for each environmental variable, initial (pre-treatment) observations were used as a baseline for all subsequent samplings (post-treatment). preliminary analyses included tree gender as an explanatory variable. however, because gender showed no significant main or interaction effects, we removed gender and re-ran all anovas. we analyzed compositional data using direct gradient analysis. direct gradient analysis uses species data and directly relates it to measured environmental variables, in this case dummy variables representing the treatments. we selected partial redundancy analysis (prda) because it is generally considered more appropriate in short-term experimental studies where species responses are believed to be linear and over relatively short gradients. all ordinations were conducted using canoco for windows 4.5 (ter braak and šmilauer 2002) on absolute cover of each species within a sample. we developed a priori hypotheses about the potential affect of treatment application on species cover. we hypothesized that tree removal and litter removal would have a positive effect on stems per quadrat, vegetation cover and species richness. in addition, the combination of tree removal with litter removal, conditions most similar to open prairie (cn), would have the largest effect; whereas, the combination of no tree removal and no litter removal, the control condition (nl), would have no effect or the least positive effect on species. we have not included any correction factors for statistical problems associated with multiple comparisons (legendre and legendre 1998, hallgren et al. 1999). oklahoma native plant record volume 11, december 2011 linneman, j. s., et al. 46 results density and richness there were significant differences in stem density (p<0.001) between all quadrats prior to treatment application. however, the difference between the means of the densest and sparsest treatments was only 2.5 stems per quadrat. both the cut with litter (cl) and cut with no litter (cn) treatments had the lowest stems per quadrat prior to treatment application. stem density increased for all treatments except no cut with litter (nl) treatment by the second sampling. this increase was roughly 2-2.5 fold thus resulting in an increase of 10-13 stems per treatment (figure 1). significant differences (p=0.0052) in density between nl & nn (no cut-no litter) only occurred in may 2002. on the other hand, there were significant differences in stems density between litter treatments within the cut treatment, cl and cn, in september 2002 (p=0.0366) and 2003 (p=0.0483). the cut treatment had a much more pronounced effect on density regardless of litter treatment. in september and may 2002 2003, there were significant differences between both cn and nn (p=0.006, 0.004, 0.001 respectively) and cl and nl (p=0.0052, 0.003, 0.0159 respectively). as with density, there were significant differences in initial species richness (p<0.001) between all quadrats prior to treatment application. again, the magnitude of the mean difference was quite small, fewer than 1.0 species per treatment. additionally, the cl and cn treatments again had the lowest richness. the increase in species richness by the second sampling was not as dramatic as that observed in stems per quadrat by the same sampling. generally increases in mean species richness were in the order of 0.4-1.25 species per quadrat (figure 2). significant differences in species richness between nl and nn only occurred in september 2002 (p=0.0244); however may 2002 was marginally insignificant (p=0.0533). conversely, significant differences in species richness between cl and cn occurred in both may 2002 (p=0.0381) and september 2002 (p=0.0026). the cut treatment had a slightly weaker influence on species richness as compared to stems per quadrat. significant differences in species richness were observed between cn and nn in september 2002 (p=0.0055) and 2004 (p=0.0007). significant differences in species richness were also observed between cl and nl in september 2002 (p=0.0457) and september 2003 (p=0.0358). veg etation cover there was no significant difference in total vegetation cover prior to treatment application. there was a substantial increase in total cover through samplings two and three in both the cl and cn treatments (figure 3). this increase in total cover was in the order of 8.75-11.25%. on the other hand, total cover in both the nn and nl treatments only increased by ~2%. no significant differences in total cover were observed between the nl and nn treatments at any sampling. on the other hand, there was a significant difference between the cl and cn treatments in september 2003 (p=0.0024). although litter removal did not have a major effect on total cover, tree removal did. significant differences between cn and nn were observed in september and may 2002-2003 (p=0.001, 0.0023, <0.001 respectively). in addition, significant differences between cl and nl were also observed in september and may 2002-2003 (p=0.0071, 0.0075, 0.0318 respectively). unlike total vegetation cover, there were significant differences (p<0.001) in initial mean forb cover between treatments; however these differences were only 0.125%. forb cover in both of the cut treatments, cl and cn, increased over the duration of the study although both no cut treatments, nl and nn, were relatively oklahoma native plant record volume 11, december 2011 linneman, j. s., et al. 47 static throughout the study (figure 4). there were no significant differences in forb cover for nl or nn treatments at any time, whereas a significance difference between cl and cn only occurred in september 2002 (p=0.0056). the tree removal treatment yielded a significant difference between cn and nn in may 2002 (p<0.0001) and september 2002 (p=0.0486), whereas a significant difference between cl and nl occurred only in may 2003 (p=0.0131). graminoid cover responded similarly to forb cover with significant differences in initial mean graminoid cover between treatments (p=0.0164). once again, the differences between treatments were small (0.15%). graminoid cover increased over the first post-treatment sampling for all treatments (figure 5). graminoid cover was not significantly affected by litter in nl or nn treatments. however, litter had a significant effect in september 2003 (p=0.0012) in the cl and cn treatment. the tree removal treatment had a stronger affect with significant differences in graminoid cover between cn and nn in september and may 2002-2003 (p=0.0253, 0.0092, <0.0001 respectively) and between cl and nl in september 2002 (p=0.0133). marginal insignificance was also observed between cl and nl in may, 2003 (p=0.052). significant differences in woody cover (p=0.0197) were also present at the onset of this study. however, differences in mean woody cover between treatments were once again small (0.15%). woody cover increased in all treatments over the duration of this study although these increases were only in the 0.5-2.0% range (figure 6). in fact, no significant differences were found between any combination of litter removal and/or tree removal treatments at any sampling. direct gradient analysis partial redundancy analysis (prda) was conducted to test a priori hypotheses regarding the effects of tree removal, litter removal and their interaction at each sampling. results of prda only showed significant differences in absolute species cover between litter removal treatments in may, 2002 and september, 2002 (p<0.001). conversely, prda showed significant differences (p<0.001) in absolute species cover between tree removal treatments at every post-treatment sampling period. the litter removal with tree removal interaction effect was only significant in september 2002 (p=0.029). therefore, it appears that tree removal does have a stronger effect on species composition over time than litter removal. when treatment centroids by sampling period are plotted in ordination space three items become apparent: first, tree removal results in an increased magnitude of movement of treatment centroids over time (figure 7 a, b). second, litter removal also results in an increased magnitude of movement of treatment centroids over time (see figure 7 a, b). finally, the overall amount of compositional change of cut treatments was greater than litter removal treatments. a prda scatter plot of absolute species cover, treatment centroids and passive environmental variables based on all posttreatment samplings is displayed in figure 8. the four dummy treatment variables accounted for 5.4% of the total explained species variance. although woody cover was not significantly affected by tree removal or litter removal treatments at any sampling, woody forest species such as cercis canadensis, celtis occidentalis, parthenocissus quinquefolia, quercus stellata, juniperus virginiana seedlings, and ulmus rubra all dominated the no cut treatments with a slightly higher cover in the litter treatment (nl). alternatively, grasses typical of the open prairie such as tridens flavus, eragrostis oklahoma native plant record volume 11, december 2011 linneman, j. s., et al. 48 spectabilis, dicanthelium oligosanthes, sorghastrum nutans, bothriochloa saccharoides, and sporobolus compositus dominated the tree removal treatments. in addition, each one of these graminoids (with the exception of t. flavus) also had higher absolute cover in the litter removal treatment (cl). sedges such as carex festucacea and c. bushii both dominated the nn treatment. on the other hand, forb species typically associated with pastures such as ambrosia sp., a. psilostachya, amphiachyris dracunculoides, acalypha gracilens, and croton monanthogynus dominated the cn treatment. discussion increases in stem density and species richness were expected as a result of litter removal and tree removal treatments. our results are similar to those of monk and gabrielson (1985) who observed a stronger influence of overstory cover compared to litter cover on old field vegetation. for all manipulated quadrats (cl, cn, and nn) increased stems per quadrat is more likely to be due to increased perennial graminoid stems than woody or forb stems. reductions in stems per quadrat in nl and nn treatments after september 2002 are likely the result of continued overstory tree presence and its associated reductions in solar radiation. studies by monk and gabrielson (1985), yager and smeins (1999), and joy and young (2002) have all suggested that reductions in light similar to those observed in this study resulted in significant decreases in plant density and cover. on the other hand, we believe that reductions in stems per quadrat in september 2003 for cl and only the slight increase for cn were caused by relatively little precipitation received in 2002 – 2003. total precipitation recorded at the marena mesonet station, located approximately 4 km from the study site, was 63.0 cm from october 2002 to september 2003. this precipitation total is only 64-69% of annual precipitation for the site of 91.4-99.1 cm (oklahoma mesonet, oklahoma climatological survey). by comparison, the decreases in species richness over the course of this study suggest relatively little recruitment of new species occurred regardless of treatment. provencher et al. (2000) found that species richness also decreased after the application of felling and slash burning in florida’s sandhill vegetation. however, provencher et al. (2000) observed an increase in species richness two years after treatment application. results from prda (see figure 8) suggest a transition from pretreatment species composition dominated by mesic or forest species to post-treatment tallgrass prairie species. it is possible that during this transition, forest species were lost faster than prairie species were added; therefore, we observe a decrease in species richness. however, the majority of species present in each treatment’s cumulative species pool were, on average, not present in each quadrat. generally only 10-20% of each treatment’s cumulative species pool was observed in each quadrat (see figure 2). it should be noted that species richness may be strongly linked to density (i.e. rarefaction effect) and thus the richness-per-quadrat should not be interpreted independently of stem counts (palmer et al. 2000). this suggests that given more time species richness may increase as these rare species become more universally distributed into cut quadrats. linneman and palmer (2006) suggested that species composition underneath redcedar trees may be a random subset of the species from the surrounding matrix. the results from this study suggest that this subset of species is nonrandom and comprised of two main types. the first group appears to be remnant prairie grasses, and the second is disturbance-tolerant forbs. the absolute cover of almost all graminoid species increased as a result of tree removal. of particular interest is that the most oklahoma native plant record volume 11, december 2011 linneman, j. s., et al. 49 abundant graminoid species were native tallgrass prairie species such as sorghastrum nutans and sporobolus compositus. conversely, the positive response of disturbance favoring forb species like amphiachyris dracunculoides, ambrosia psilostachya and croton monanthogynus may lead to further reductions in species richness if they become dominant. several researchers, including clary (1971), clary and jameson (1981), brockway et al. (1998), and provencher (2000), have all observed increases in graminoid and forb cover following overstory tree removal. in this study, annual species increased in cut treatments; however, few annuals dominated cover in any treatment. although the increase of disturbance-tolerant forbs may be inhibitive in the short term, the observed increases in absolute cover of native tallgrass prairie species suggest that even without subsequent prescribed fire treatments, community composition may return to its pre-invasion condition with time. the long-term effects of eastern redcedar in grasslands are unclear. the results from this study suggest the continued presence of eastern redcedar in grasslands may (1) facilitate the forestation of grasslands or at least (2) continue to reduce the tallgrass prairie species pool in invaded grasslands. briggs et al. (2002a) determined that species present in the prairie were not consistently different from those found in a closed canopy redcedar forest. however, both this study and linneman and palmer (2006) show an apparent shift in community composition away from tallgrass prairie species toward forest tree species such as cercis canadensis, celtis occidentalis, juniperus virginiana, quercus stellata and ulmus rubra. these same tree species frequently occur under redcedar canopies in nearby cross timbers forest environments as well (van els et al. 2010) and, it should be noted, the cedars studied here were in relatively close proximity to cross timbers stands. additionally, it is possible that the dynamics of cedar invasion may differ in old fields (studied here) from those in previously undisturbed prairie. although complete extirpation of native tallgrass prairie species is not likely in the short term, areas with extensive invasion and subsequent tree removal may require seeding of prairie species to encourage the return of characteristic prairie vegetation. this will inevitably increase the cost of restoration beyond the already high cost of tree removal (bidwell et al. 2002). areas with less than 75% cover of redcedar, however, have greater potential for recovery, as most tallgrass prairie species persist in inter-tree spaces until this point (limb et al. 2010). continued invasion by eastern redcedar in the great plains has serious implications not only for the existence of native grasslands but also for biodiversity and potential future restorations. as shown here, removal of redcedar, even in the absence of subsequent prescribed fire, has the potential to increase the number of stems per quadrat and increase species richness for several years post-treatment. litter removal, either by mechanical means or prescribed fire, should further benefit and accelerate the return of tallgrass prairie vegetation. without tree removal, these grasslands will continue to lose native prairie species in favor of mesic and/or forest species. in the absence of broad-scale control efforts, redcedar will continue to fragment and replace native grasslands, perhaps to the extent that future prairie restoration efforts may require seed inputs beyond what is available from surrounding sources via natural dispersal. acknowledgments we thank sam fuhlendorf for his editorial comments. in addition, jared laufenberg aided with tree removal. mark payton’s help with sas analyses and programming was greatly appreciated. the oklahoma native plant record volume 11, december 2011 linneman, j. s., et al. 50 payne county audubon society and the james k. mcpherson fund, administered by the department of botany at oklahoma state university, provided funding for this study. literature cited axmann, b. d. and a. k. knapp. 1993. water relations of juniperus virginiana and andropogon gerardii in an unburned tallgrass prairie watershed. southwestern naturalist 38:325-330. barbaro, l., t. 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m. and c. marais. 2000. impacts of alien plant clearance on vegetation in the mountain catchments of the western cape. south african forestry journal 189:113-117. holmes, p. m., d. m. richardson, b. w. van wilgen, and c. gelderblom. 2000. recovery of south african fynbos vegetation following alien woody plant clearing and fire: implications for restoration. australian ecology 25:631639. hulbert, l. c. 1969. fire and litter effects in undisturbed bluestem prairie in kansas. ecology 50:874-877. joy, d. a.and d. r. young. 2002. promotion of mid-successional seedling recruitment and establishment by juniperus virginiana in a costal environment. plant ecology 160:125-135. kush, j. s., r. s. meldahl, and w. d. boyer. 1999. understory plant community response after 23 years of hardwood control treatments in natural longleaf pine (pinus palustris) forests. canadian journal of forest resources 29:1047-1054. legendre, p. and l. legendre. 1998. numerical ecology. elsevier, amsterdam. linneman j. s. and m. w. palmer. 2006. the effect of juniperus virginiana l. on plant species composition in an oklahoma grassland. community ecology 7: 235-244. limb, r. f., d. m. engle, a. l. alford, and e. c. hellgren. 2010. tallgrass prairie plant community dynamics along a canopy cover gradient of eastern redcedar (juniperus virginiana l.). rangeland ecology managment 63: 638-644. monk, c. d. and f. c. gabrielson. 1985. effects of shade, litter and root competition on old-field vegetation in south carolina. bulletin of the torrey botanical club 112:383-392. palmer, m. w., d. b. clark, and d. a. clark. 2000. is the number of tree species in small tropical forest plots nonrandom? community ecology 1:95-101. peltzer, d. a. and m. köchy. 2001. competitive effects of grasses and woody plants in mixed-grass prairie. journal of ecology 89:519-527. petranka, j. w. and j. k. mcpherson. 1979. the role of rhus copallina in the dynamics of the forest-prairie ecotone in north-central oklahoma. ecology 60:956965. provencher, l., b. j. herring, d. r. gordon, h. l., rodgers, g. w. tanner, l. a. brennan and j. l. hardesty. 2000. restoration of northwest florida sandhills through harvest of invasive pinus clausa. restoration ecology 8:175-185. provencher, l., b. j. herring, d. r. gordon, h. l., rodgers, g. w. tanner, l. a. brennan and j. l. hardesty. 2001. effects of hardwood reduction techniques on longleaf pine sandhill vegetation in northwest florida. restoration ecology 9:13-27. schmidt, t. l. and e. c. leatherberry. 1995. expansion of eastern redcedar in the lower midwest. northern journal of applied forestry 12: 180-183. smith, s. d. and j. stubbendieck. 1990. production of tall grass prairie herbs below eastern redcedar. prairie naturalist 22:13-18. ter braak, c. j. f. and p. šmilauer. 2002. canoco for windows 4.5. biometris-plant research international. wageningen, netherlands. usda, nrcs. 2004. the plants database, version 3.5 (http://plants.usda.gov). national plant data center, baton rouge, la. van els, p., r. e. will, m. w. palmer, and k. r. hickman. 2010. changes in forest understory associated with juniperus encroachment in oklahoma, usa. applied vegetation science 13:356-368. weaver, j. e. and n. w. rowland. 1952. effect of excessive natural mulch on the development, yield, and structure of a native grassland. botanical gazette 114:119. oklahoma native plant record volume 11, december 2011 linneman, j. s., et al. 52 whiteman, g. and j. r. brown. 1998. assessment of a method for mapping woody plant density in a grassland. journal of arid environments 38:269-282. yager, l. y.and f. e. smeins. 1999. ashe juniper (juniperus ashei: cupressaceae) canopy and litter effects on understory vegetation in a juniper-oak savanna. southwestern naturalist 44:6-16. figure 1 mean stems per 0.25 m2quadrat in tree removal and litter removal treatments for 2.5 years. the data points have been staggered to increase visibility of 95% confidence intervals (determined for each treatment at each sampling). cl=cut with-litter, cn=cut with no litter, nl=no cut with litter, nn=no cut with no litter. 0 5 10 15 20 25 30 35 40 may-01 jul-01 oct-01 jan-02 apr-02 jul-02 oct-02 jan-03 apr-03 jul-03 oct-03 sampling date s te m s p er q u ad ra t cl cn nl nn oklahoma native plant record volume 11, december 2011 linneman, j. s., et al. 53 figure 2 mean species richness per 0.25 m2 and cumulative species richness of tree removal and litter removal treatments for 2.5 years. the data points have been staggered to increase visibility of 95% confidence intervals (determined for each treatment at each sampling). cl=cut with litter, clc=cumulative cut with litter, cn=cut with no litter, cnc=cumulative cut with no litter, nl=no cut with litter, cnl=cumulative no cut with litter, nn=no cut with no litter, cnn=cumulative no cut with no litter of tree removal and litter removal treatments for 2.5 years. 0 5 10 15 20 25 30 35 may-01 jul-01 oct-01 jan-02 apr-02 jul-02 oct-02 jan-03 apr-03 jul-03 oct-03 sampling date s pe ce is r ic hn es s cl cn nl nn clc cnc nlc nnc oklahoma native plant record volume 11, december 2011 linneman, j. s., et al. 54 figure 3 mean percent total cover of tree removal and litter removal treatments for 2.5 years. the data points have been staggered to increase visibility of 95% confidence intervals (determined for each treatment at each sampling). cl=cut with litter, cn=cut with no litter, nl=no cut with litter, nn=no cut with no litter. 0 2 4 6 8 10 12 14 16 18 20 may01 jul-01 oct-01 jan-02 apr-02 jul-02 oct-02 jan-03 apr-03 jul-03 oct-03 sampling date p er ce nt t ot al c ov er cl cn nl nn oklahoma native plant record volume 11, december 2011 linneman, j. s., et al. 55 figure 4 mean percent forb cover of tree removal and litter removal treatments for 2.5 years. the data points have been staggered to increase visibility of 95% confidence intervals (determined for each treatment at each sampling). cl=cut with litter, cn=cut with no litter, nl=no cut with litter, nn=no cut with no litter. 0 1 2 3 4 5 6 7 8 9 10 may-01 jul-01 oct-01 jan-02 apr-02 jul-02 oct-02 jan-03 apr-03 jul-03 oct-03 sampling date p er ce n t f o rb c o ve r cl cn nl nn oklahoma native plant record volume 11, december 2011 linneman, j. s., et al. 56 figure 5 mean percent graminoid cover of tree removal and litter removal treatments for 2.5 years. the data points have been staggered to increase visibility of 95% confidence intervals (determined for each treatment at each sampling). cl=cut with litter, cn=cut with no litter, nl=no cut with litter, nn=no cut with no litter. 0 2 4 6 8 10 12 14 may-01 jul-01 oct-01 jan-02 apr-02 jul-02 oct-02 jan-03 apr-03 jul-03 oct-03 sampling date p er ce n t g ra m in o id c o ve r cl cn nl nn oklahoma native plant record volume 11, december 2011 linneman, j. s., et al. 57 figure 6 mean percent woody cover of tree removal and litter removal treatments for 2.5 years. the data points have been staggered to increase visibility of 95% confidence intervals (determined for each treatment at each sampling). cl=cut with litter, cn=cut with no litter, nl=no cut with litter, nn=no cut with no litter. 0 1 2 3 4 5 6 7 8 may-01 jul-01 oct-01 jan-02 apr-02 jul-02 oct-02 jan-03 apr-03 jul-03 oct-03 sampling date p er ce n t w o o d y c o ve r cl cn nl nn oklahoma native plant record volume 11, december 2011 linneman, j. s., et al. 58 figure 7 prda trajectory of tree and litter removal treatment centroids for each sampling period. the two figures are from the same analysis but were separated to increase legibility. cl=cut with litter, cn=cut with no litter, nl=no cut with litter, nn=no cut with no litter. axes 1 and 2 are displayed in both figures. -0.3 0.4 -0 .2 0. 3 cl1 cn1 cl2 cn2 cl3 cn3 cl4 cn4 cl5 cn5 axis 1 ax is 2 -0.3 0.4 -0 .2 0. 3 nl1 nn1 nl2 nn2 nl3 nn3 nl4 nn4 nl5 nn5 axis 1 ax is 2 oklahoma native plant record volume 11, december 2011 linneman, j. s., et al. 59 figure 8 prda triplot of species codes, treatment centroids and supplemental environmental variables. all post-treatment samplings are included and axes 1 and 2 are displayed. species codes represent the relative multi-dimensional position of each species in ordination space based on absolute cover of each species. species codes are indexed in appendix 1. arrow length indicates the relative strength of supplemental variables. cl=cut with litter, cn=cut with no litter, nl=no cut with litter, nn=no cut with no litter. -1.0 1.5 -1 .0 1. 0 acgr2 ambro amdr amps bosa cabu5 cafe3 cani3 ceca4 ceoc crmo6 diac2 ersp gapu3 juvi lecu opma2 pape5 paqu2 poar7 qust rhco sonu2 spco16 teca3 trfl2 ulru nn nl cn cl ba.10cm canopy cedar litter soil rock axis 1 ax is 2 oklahoma native plant record volume 11, december 2011 linneman, j. s., et al. 60 appendix species names and usda plant codes species usda code acalypha gracilens acgr2 ambrosia psilostachya amps ambrosia sp. ambro amphiachyris dracunculoides amdr bothriochloa saccharoides bosa carex bushii cabu5 carex festucacea cafe3 carex nigromarginata cani3 celtis occidentalis ceoc cercis canadensis ceca4 croton monanthogynus crmo6 dichanthelium acuminatum diac2 eragrostis spectabilis ersp gamochaeta purpurea gapu3 juniperus virginiana juvi lespedeza cuneata lecu opuntia macrorhiza opma2 oxalis stricta oxst parietaria pensylvanica pase5 parthenocissus quinquefolia paqu2 quercus stellata qust rhus copallinum rhco sorghastrum nutans sonu2 sporobolus compositus spco16 teucrium canadense teca3 tridens flavus trfl2 ulmus rubra ulru the effects of removal of juniperus virginiana l. trees and litter from a central oklahoma grassland by mr. jerad s. linneman, dr. matthew s. allen, and dr. michael w. palmer oklahoma native plant record, volume 12, number 1, december 2012 oklahoma native plant record volume 12, december 2012 wayne elisens https://doi.org/10.22488/okstate.17.100093 69 critic’s choice essay virtual herbaria come of age wayne elisens department of microbiology and plant biology oklahoma biological survey university of oklahoma these are exciting times for natural history collections. an international effort is underway to make images and data of biological specimens available in electronic format via digitization. these initiatives are an effort to bring natural history collections out of the dark of museum and herbarium cabinets and into the light of public access for use by stakeholders in government, academia, biodiversity organizations, business, and k-12 education. the democratization of information contained in natural history collections through images and online databases is an important new development to better investigate our natural world and solve important social and environmental problems (scoble 2010). for herbarium collections, digitized images and data from specimens are generally referred to as a virtual herbarium. what exactly do we mean by digitization of natural history collections? for plants, digitizing collections transforms herbarium specimens into digital images and label data sorted (parsed) into its component units such as names, locations, collectors, dates, habitats, and reproductive state. all data and images are fully searchable and distributed in electronic format, such as virtual herbaria. there are several outstanding examples of virtual herbaria already online, such as australia’s virtual herbarium (http://avh.ala.org.au/) and the new york botanical garden’s virtual herbarium (http://sciweb.nybg.org/science2/virtualher barium.asp.html). according to the us interagency working group on scientific collections (iwgsc 2007), plant specimen data distributed via virtual herbaria would have a profound impact on science education and investigations of environmental change and quality, invasive species, public health, national security, bioscience research, and many other issues (niba 2010). why digitize natural history collections? think for a moment about the incomparable treasure trove of biodiversity information contained in the world’s natural history collections. if we focus just on plants, herbarium specimens document most of what is known about the world’s plant species diversity and represent a 200+ year record of what species were present at a given location and at a given time. herbaria collections not only document the different kinds of plants constituting a flora, but they record valuable information about where they occurred and when they were flowering or fruiting. plant specimens provide a spatial and temporal window into the dynamic processes of plant diversity, introduction and spread of exotics, expansion and contraction of species ranges, and changes in time of flowering and fruiting. is a digitized herbarium specimen as valuable as the specimen itself? certainly not, and this is one of the main reasons for maintaining natural history collections in museums and herbaria. the primary rationale for digitizing specimens is access. scientists throughout the world will have greatly enhanced access to digitized specimens, which greatly adds to their value for research and education. in the us, a key component is in place to assist efforts to digitize biological research collections (e.g., herbarium specimens) – the integrated digitized biocollections resource http://avh.ala.org.au/ http://sciweb.nybg.org/science2/virtualherbarium.asp.html http://sciweb.nybg.org/science2/virtualherbarium.asp.html oklahoma native plant record volume 12, december 2012 wayne elisens 70 (idigbio; www.idigbio.org). tools and training provided by idigbio are funded by the national science foundation, who also established a 10-year funding program entitled advancing digitization of biological collections (adbc) to aid conversion of biodiversity collections into electronic formats. these advances in funding and infrastructure were established using recommendations of the national science and technology council, who recognized the importance of biocollections for national science infrastructure. with an estimated 90 million herbarium specimens in u.s. herbaria (tulig et al. 2012), is it feasible to construct a us virtual herbarium comparable to australia’s virtual herbarium based on “only” 6 million specimens? at a minimum, digitization of biocollections involves specimen imaging, image processing, electronic data capture, and georeferencing of locality descriptions (nelson et al. 2012). mass digitization methods continue to be refined and automated (beaman and cellinese 2012), but it is unlikely that all us herbarium specimens can be digitized in a 10-year timeframe. however, a recent survey conducted by the us virtual herbarium project (barkworth and murrell 2012) indicated that ca. 30% of herbarium specimen labels were already databased. while it appears that much digitization has occurred at the individual herbarium or regional level, there has been no coordinated national effort to expedite digitization of biocollections. however, the idigbio mission aims to fill that void and has a major objective to facilitate access to us biocollection data. this goal is certainly feasible, especially since a global portal for digitized images and data from natural history collections already exists – gbif, the global biodiversity information facility (www.gbif.org). gbif currently serves up more than 300 million specimen records. herbaria throughout the country are actively engaged in efforts to image and database information and to present them in searchable online formats. international standards and best practices for data capture have been established and are being implemented by the collections community nationwide. luckily, botanists in oklahoma demonstrated great foresight by establishing a data portal for digitized herbarium label data for specimens collected in oklahoma – the oklahoma vascular plants database (ovpd; hoagland et al. 2004). with the ovpd as a firm foundation and a collaborative network in place among curators in the state and region, oklahoma herbaria are poised to expand their digitization efforts. this endeavor will help develop the concept of a virtual herbarium to maturity and will undoubtedly enhance the value and access of real herbaria. literature cited barkworth, m. e. and z. e. murrell. 2012. the us virtual herbarium: working with individual herbaria to build a national resource. zookeys 209:55-73. beaman, r. s. and n. cellinese. 2012. mass digitization of scientific collections: new opportunities to transform the use of biological specimens and underwrite biodiversity science. zookeys 209:7-17. hoagland b. w., a. k. buthod, i. h butler, p. h. c. crawford, a. h. udasi, w. j. elisens, and r. j. tyrl. 2004. oklahoma vascular plants database. oklahoma biological survey, university of oklahoma, norman. available from: http://www.oklahomaplantdatabase.org ). iwgsc, interagency working group on scientific collections. 2007. scientific collections: mission-critical infrastructure for federal science agencies. available from: http://www.whitehouse.gov/files/docum ents/ostp/nstc%20reports/revision_1 -22_09_cl.pdf). nelson, g., d. paul, g. riccardi, and a. r. mast. 2012. five task clusters that enable http://www.idigbio.org/ http://www.gbif.org/ http://www.oklahomaplantdatabase.org/ http://www.oklahomaplantdatabase.org/ http://www.whitehouse.gov/files/documents/ostp/nstc%20reports/revision_1-22_09_cl.pdf http://www.whitehouse.gov/files/documents/ostp/nstc%20reports/revision_1-22_09_cl.pdf http://www.whitehouse.gov/files/documents/ostp/nstc%20reports/revision_1-22_09_cl.pdf oklahoma native plant record volume 12, december 2012 wayne elisens 71 efficient and effective digitization of biological collections. zookeys 209:19-45. niba, network integrated biocollections alliance. 2010. a strategic plan for establishing a network integrated biocollections alliance. available from: http://digbiocol.wordpress.com/brochur e/ scoble, m. j. 2010. natural history collections digitization: rationale and value. biodiversity informatics 7:77-80. tulig, m., n. tarnowsky, m. bevans, a. kirchgessner, and b. m. thiers. 2012. increasing the efficiency of digitization workflows for herbarium specimens. zookeys 209:103-113. onps http://digbiocol.wordpress.com/brochure/ http://digbiocol.wordpress.com/brochure/ critic’s choice essay: virtual herbaria come of age by dr. wayne elisens oklahoma native plant record, volume 14, number 1, december 2014 oklahoma native plant record 67 volume 14, december 2014 angela mcdonnell https://doi.org/10.22488/okstate.17.100106 non-twining milkweed vines of oklahoma: an overview of matelea biflora and matelea cynanchoides (apocynaceae) angela mcdonnell botany department oklahoma state university 301 physical sciences stillwater, ok 74078 (608) 698-1217 angela.mcdonnell@okstate.edu key words: distribution, ecology, morphology, taxonomy abstract matelea (apocynaceae, asclepiadoideae) is a genus of approximately 225 species in milkweed subtribe gonolobinae. this new world genus is predominantly found in tropical to subtropical regions and is represented in oklahoma by four species. two of these, m. biflora and m. cynanchoides, are closely related, non-twining perennial herbs that have long confused amateur and professional botanists alike due to similar morphological features. this paper includes a brief review of their taxonomic history and describes the morphology, ecology, and distribution of these species in oklahoma and neighboring states. photographs, a distribution map, and a key to the species of matelea in oklahoma are included. introduction milkweeds in oklahoma from apocynaceae subfamily asclepiadoideae (the former asclepiadaceae) display an array of diversity. species include erect and prostrate herbs and herbaceous vines in five genera (asclepias l., cynanchum l., funastrum e. fourn., gonolobus, and matelea aubl.). in addition to variation in growth form, they exhibit a variety of corolla forms and variation in the distinctive features of the milkweed subfamily: fusion of male and female floral whorls forming a gynostegium, and an additional floral whorl, the corona. matelea is a large genus of approximately 225 species in the milkweed subtribe gonolobinae. this new world genus consists mostly of vines and is found in tropical and subtropical regions. matelea is known to be broadly polyphyletic (i.e., consisting of multiple lineages that are not necessarily closely related) and is a good candidate for taxonomic dissolution (krings, thomas, and xiang 2008; parks 2008; mcdonnell and fishbein, in prep). in oklahoma, matelea is represented by four species that form two morphologically distinct pairs; matelea baldwyniana (sweet) woodson and matelea decipiens (alexander) woodson are vines most common in the eastern part of the state, and matelea biflora (nutt. ex raf.) woodson and matelea cynanchoides (engelm. and a. gray) woodson are prostrate to decumbent species, present throughout much of the eastern two-thirds of the state. pending additional phylogenetic and morphological study, the four species will likely be placed in two genera, distinct from matelea in the strict sense, which will be restricted to species native to central and south america. one of these oklahoman matelea species pairs, the non-twining herbaceous species m. biflora (figs. 1a, 1c) and 68 oklahoma native plant record volume 14, december 2014 angela mcdonnell m. cynanchoides (figs. 1b, 1d), are closely related and possess similar morphological features. both species currently reside in matelea subgenus chthamalia, a group of approximately 30 milkweeds that are apparently adapted to arid habitats, have a center of diversity in northern mexico, and are the focus of my dissertation research. this paper will clarify the taxonomic history and morphological differences between the species and will also provide a key to identify the species native to oklahoma. figure 1 matelea biflora and matelea cynanchoides. (a) m. biflora habit, note prostrate stem. photo by mark fishbein. (b) m. cynanchoides habit, note decumbent-ascending stem. (c) m. biflora flowers, note pubescent corolla and reflexed corolla margins. (d) m. cynanchoides flowers and buds, note glabrous corolla and planar corolla margins. photo by mark fishbein. oklahoma native plant record 69 volume 14, december 2014 angela mcdonnell methods specimen records (336 total, 205 of which were viewed, see appendix for list of viewed specimens) for matelea biflora and m. cynanchoides were downloaded from online data repositories including: the global biodiversity information facility (gbif http://www.gbif.org); tropicos (http://tropicos.org); seinet (http://swbiodiversity.org/portal/index.ph p); and herbarium websites, such as the oklahoma vascular plants database (ovpd http://www.oklahomaplantdatabase.org). specimen loans (abbreviations follow thiers [2014]) were obtained from the us national herbarium (us), the new york botanical garden (ny), the missouri botanical garden (mo), harvard university herbaria (a, econ, gh), arizona state university (asu), university of texas at austin (tex, ll), kansas state university (ksc), university of arizona (ariz), university of new mexico (unm), and louisiana state university (lsu). specimens at the oklahoma state university herbarium (okla), botanical research institute of texas (brit), sul ross state university (srsc), and the university of oklahoma (okl) were examined on visits to those herbaria. additional data were obtained from my field collections and the unaccessioned collections and database of mark fishbein (oklahoma state university). occurrence data were curated manually to confirm or change species identifications and for georeferencing. the resulting specimen database was used to plan fieldwork across the range of each species. fieldwork in oklahoma and texas was carried out in the summers of 2011, 2012, and 2013. for each population located in the field, specimens were collected and the following data recorded: latitude and longitude coordinates obtained with a handheld gps device (usually a garmin® gpsmap 76), elevation obtained by gps and checked in google earth®, substrate, relative local abundance, vegetation type, co-occurring species, occurrence of interacting arthropods (flower visitors and herbivores), and morphological notes. specimens obtained from loans and field collections were used for morphological study. measurements of floral and vegetative characters were carried out using olympus® cellsens entry 1.6 imaging software and an olympus® szx10 dissecting microscope outfitted with an olympus® sc30 cmos color camera. a distribution map (fig. 2) for both species was produced using a combination of google earth®, adobe® illustrator, and adobe® photoshop software. the points on the map include specimens examined and records downloaded from databases for which specimens were not examined. due to imprecise locality data, not all records could be accurately mapped. records with ambiguous or incomplete locality data were excluded. 70 oklahoma native plant record volume 14, december 2014 angela mcdonnell figure 2 distribution map showing ranges of matelea biflora (black) and m. cynanchoides (white) results and discussion matelea biflora (nutt. ex raf.) woodson gonolobus biflorus nutt. ex raf. gonolobus biflorus nutt. ex torr., nom. illeg. chthamalia biflora (nutt. ex raf.) decne. gonolobus biflorus nutt. ex raf. var. wrightii a. gray purple milkweed vine, star milkvine, twoflowered milkvine taxonomic history the type specimens of what would eventually be named matelea biflora (see figs. 1a, 1c) were collected by intrepid english botanist thomas nuttall near the red river in the arkansas territory during his travels between october 2, 1818, and february 18, 1820. the collection date was not recorded by the collector or by subsequent taxonomists working with the material. the specimens were probably collected in the summer of 1819, the only time during his trip when flowering specimens were likely abundant. at the time, the arkansas territory included all of present day arkansas and most of present day oklahoma (the northernmost counties and the panhandle of oklahoma were excluded). according to his journal and the interpretations of later scholars, nuttall doesn’t appear to have crossed the border into texas, which was then owned by spain. the specimens were likely collected on the oklahoma side of the red river, in either choctaw or mccurtain county (lottinville 1980; tyrl and shryock 2014). the specimens were labeled in nuttall’s handwriting “gonolobus *biflorus nutt”. the asterisk denotes his convention of marking a species name as new (mclean 1980; stuckey 1966). many of the gonolobus biflorus specimens nuttall collected received additional labels and were distributed to several herbaria. currently, there are at least eight duplicate oklahoma native plant record 71 volume 14, december 2014 angela mcdonnell sheets held at herbaria of the academy of natural sciences, philadelphia (ph); royal botanic gardens, kew (k); smithsonian institution (us); and the new york botanical garden (ny). significantly, g. biflorus was never mentioned in nuttall’s collections towards a flora of the territory of arkansas (1837), the publication in which he describes many new taxa from the region, nor in any of his other publications. thus, the name indicated as new on nuttall’s labels was never published by him. like other species discovered and named but not published by nuttall, g. biflorus was apparently validated by john torrey (1859) in his report on the united states and mexican boundary survey. therefore, some sources cite the authority for this species as g. biflorus nutt. ex torr. however, even before nuttall’s (1837) report on the flora of the arkansas territory was published, constantine samuel rafinesque, a self-educated professor of botany and natural history who elicited considerable controversy from his contemporaries (boewe 2003; warren 2004), published a new flora of north america (1836). in this work, rafinesque was the first to describe and validly publish gonolobus biflorus from a specimen he saw at the herbarium of zaccheus collins, a philadelphia merchant and avid collector of herbarium specimens. according to correspondence held by the american philosophical society, the two men were friendly and discussed botanical findings, travels, reading habits, and finance (collins 1805–1827, redfield 1876). in 1833, two years after collins’ death, most of his herbarium was sold to rev. lewis david von schweinitz, and a small portion of the collection was sold to rafinesque shortly thereafter (stuckey 1971). rafinesque does not describe the morphology of the specimen in his publication. he also fails to cite the collector of the specimen he studied. he does state that the plant is from “the red river in arkanzas and texas”, nearly the precise locality from which nuttall collected, except for the inclusion of texas. however, there are no records showing that nuttall traveled in texas. notably, rafinesque used the exact epiphet, “biflora” indicated by nuttall on the slips accompanying his specimens. collins seems to be the link between rafinesque and nuttall. nuttall named the plantaginaceae genus collinsia for him in 1817 and called collins “a gentleman whose talents as a botanist and a mineralologist are deservedly acknowledged”. during nuttall’s trip to the arkansas territory, he and collins exchanged letters (lawson 2004), and after the trip, collins received a complete set of duplicates (stuckey 1971). rafinesque must have examined the g. biflorus specimen nuttall sent to collins between 1820 and 1833. apparently having realized that the name for this species had not been published, rafinesque seized the opportunity. later workers have variably indicated either nuttall or rafinesque as the author of g. biflorus. it is not clear whether crediting nuttall as the author was a repeated accident or an intentional snub toward rafinesque. eight years after rafinesque’s publication of g. biflorus, decaisne (1844) included the species in his newly described genus, chthamalia decne., citing nuttall as the author of the basionym. asa gray also cited the species with nuttall as the author in his synoptical flora of north america (1878). more than 120 years after nuttall’s specimen was first collected, milkweed specialist robert everard woodson, jr. lumped chthamalia, including chthamalia biflora, into the genus matelea, along with over 100 species in more than 20 genera (woodson 1941). currently, floras and databases indicate the authorship of this species as either m. biflora (nutt.) woodson or m. biflora (raf.) woodson. however, because nuttall did not validly publish gonolobus biflorus, and because rafinesque, when validly publishing g. biflorus had 72 oklahoma native plant record volume 14, december 2014 angela mcdonnell apparently taken up the name suggested by nuttall, the proper authorship is g. biflorus nutt. ex raf. and in matelea, m. biflora (nutt. ex raf.) woodson. species description plants prostrate, usually with 5–20+ stems from a thickened taproot, stem length in flower 10–50 cm, lengthening in fruit, malodorous throughout; the largest leaves with petioles 0.7–2.5 cm long, blades broadly lanceolate to broadly ovate or nearly triangular, 1.5–5.0 cm long and 1.0–3.2 cm wide, bases deeply to shallowly cordate, apices acute, youngest leaf bases with a pair of rounded colleters; inflorescences of axillary pairs or solitary flowers; peduncles 0–10 mm; pedicels 0.2–1.1 cm; calyx lobes ovate to triangular, 2.0–3.5 mm long; corolla shallowly campanulate-rotate usually with spreading lobes, maroon to dark brown, 8– 13 mm in diameter, deeply 5-lobed; lobes elliptic to narrowly deltoid, margins often reflexed at maturity, densely pilose adaxially and sparsely pilose abaxially; corona consisting of a fleshy disk arising at the junction of the gynostegial column and the corolla, with 5 fleshy, incurved lobes, maroon to dark brown, approximately triangular in cross section, incumbent on anthers; anthers with entire, white, membranous, apical appendages; fruit a muricate, ellipsoid follicle, 5–10 cm long, protuberances numerous (≥5 per 5 cm of follicle length). distribution and ecology matelea biflora has been found most commonly on or adjacent to the edwards plateau in texas. the range extends north to the glass (gloss) mountains in major county, oklahoma. the easternmost collection was made near idabel in mccurtain county, oklahoma. the western edge of its range is near the texas-new mexico state line, where two specimens have been collected from lea county, new mexico (see fig. 2). in oklahoma, m. biflora is most commonly found south of i-40 in the southern tier of counties, particularly in areas with shale, dolomite, gypsum, limestone, or sandstone substrates (usgs 2005). it is also found west of oklahoma city in comanche, caddo, canadian, and major counties on sandstone, shale and limestone. to the southeast of oklahoma city, it has been collected in murray, pontotoc, johnston, and carter counties on limestone, shale and conglomerates. in the proximity of the ouachita mountains, it has been collected on shale and limestone. matelea biflora is generally found on hillsides or plains, in intact or disturbed prairies, pastures, ditches, or roadsides, where the soils generally include clay, rocks and sand. due to its prostrate, highly branched growth form, m. biflora tolerates mowing quite well and is often locally common when found in mown habitats. among the level iii ecoregions of texas and oklahoma (griffith et al. 2004; woods et al. 2005), this species has been collected in parts of the high plains, the central great plains, and the cross timbers. it is also found throughout the edwards plateau ecoregion of texas (griffith et al. 2004). within oklahoma, m. biflora is also found within the south central plains ecoregion (woods et al. 2005). few collectors have noted associated species; however, available data suggest that these are numerous and diverse. they include graminoids in the genera aristida, bothriochloa, bouteloua, bromus, carex, dicanthelium, erioneuron, and poa. other herbaceous associates include species of aphanostephanus, asclepias, ambrosia, artemisia, atriplex, centaurea, callirhoe, calylophus, chrysopsis, croton, cuscuta, dalea, desmanthus, euphorbia, gaillardia, grindelia, hedeoma, hedyotis, hymenoxys, krameria, lesquerella, linum, melampodium, opuntia, plantago, ruellia, solanum, salvia, stillingia, teucrium, thamnosma, thelesperma, and tragia. oklahoma native plant record 73 volume 14, december 2014 angela mcdonnell woody associates include species of juniperus, prosopis, quercus, and ziziphus. though almost nothing is known about faunal interactions with m. biflora, including potential pollinators, i have observed dung beetles in the genus euphoria on flowers twice, but with no pollinia attached (these have also been observed by mark fishbein, pers. comm.). near fort worth, texas, i have observed blister beetles from the family meloidae on the foliage. additionally, i’ve seen a variety of ants and flies on and around flowers. matelea cynanchoides (engelm. & a. gray) woodson gonolobus cynanchoides engelm. & a. gray vincetoxicum cynanchoides (engelm. & a. gray) a. heller prairie milkvine taxonomic history matelea cynanchoides (see figs. 1b, 1d) was first described as gonolobus cynanchoides by george engelmann and asa gray in 1845. ferdinand lindheimer collected the type specimen during his second collecting trip in texas in 1844. the holotype is held at mo. there are also four duplicates: one at k, two at gh, and one at university of michigan (mich). according to the accompanying label, the specimen was collected in “sandy soil, in open woods, near industry. april-june”. lindheimer was contracted by engelmann and gray to collect specimens in texas, and many new species discovered by lindheimer were described by these two leading botanists of their time (blankinship 1907). on the 1844 collecting trip, lindheimer traveled from the brazos river, near san felipe, to industry and then west to the colorado river. industry, where the specimen was collected, is a small community in austin county between the cities of austin and houston. in the introductory remarks to engelmann and gray’s (1845) published enumeration of lindheimer’s collections, they noted this region had rocks of secondary sandstone, cacti, and prairies with large numbers of anthills. the morphology of g. cynanchoides was described by engelmann and gray as follows: “stems 6 to 15 inches high, diffuse; leaves 1-2 inches long, cordate, with an open sinus, the uppermost sometimes almost truncate at the base. corolla greenish purple, about two lines [i.e., 0.2 in] in diameter”. they also described the coronal structure and pollinia characters in some detail. interestingly, they concluded that this taxon is a likely congener of decaisne’s chthamalia biflora (=matelea biflora, see above). gonolobus cynanchoides was differentiated primarily by its glabrous corolla. engelmann and gray did not take up decaisne’s (1844) generic name, chthamalia, published the previous year, because they argued that the characters possessed by g. cynanchoides were accommodated by the range of variation in gonolobus, as understood by botanists of that time, including decaisne. thus, they rejected decaisne’s concept of chthamalia as a genus (decaisne 1844; engelmann and gray 1845) and maintained the morphological diversity housed within gonolobus. after the initial description of g. cynanchoides, amos arthur heller transferred the species to the genus vincetoxicum (heller 1900). in doing so, he adopted a then current taxonomic opinion that vincetoxicum was the correct generic name for gonolobus, but this opinion was overturned a few decades later. just under 100 years after the first g. cynanchoides specimens were collected, woodson (1941) placed g. cynanchoides into matelea (along with many other species, including m. biflora). species description plants erect, decumbent or prostrate, usually with 3–10+ stems from a thickened 74 oklahoma native plant record volume 14, december 2014 angela mcdonnell taproot, stem length in flower 20–40 cm, lengthening in fruit, malodorous throughout; the largest leaves with petioles 0.7–1.3 cm long, blades broadly ovate to deltoid, 1.5–4 cm long and 1.5–3.2 cm wide, bases truncate to deeply cordate or sagittate, apices acute to rounded, youngest leaf bases with 2–4 elongated, pointed colleters; inflorescences of axillary (sometimes appearing terminal) fascicles or shortly pedunculate umbels; peduncles 0–13 mm; pedicels 3–6 mm long; calyx lobes ovate to elliptic, 2–3 mm long; corolla shallowly campanulate-rotate, usually with ascending lobes, green to maroon or dark brown, 6-9 mm in diameter, 5-lobed; lobes ovate to deltoid, margins not reflexed at maturity, glabrous to sparsely pilose adaxially and glabrous abaxially; corona consisting of a fleshy disk arising at the junction of the gynostegial column and the corolla, with 5 fleshy incurved lobes, green, yellow, or maroon, approximately rhombic in cross section, incumbent on anthers, anthers with lobed, white, membranous, apical appendages; fruit a sparsely muricate, broadly ellipsoid follicle, 7–10 cm long, protuberances few (≤3 per 5 cm of follicle length). distribution and ecology matelea cynanchoides is most commonly found along the gulf coastal plain in texas. the distribution extends northward to oklahoma and is strongly associated with quaternary dunes and alluvial deposits, especially those near the red, canadian, and north canadian rivers (usgs 2005). to the east, the range of m. cynanchoides extends to miller county in the southwest corner of arkansas and caddo parish in the northwest corner of louisiana. to the west, this species largely circumvents the edwards plateau in central texas, but does reach isolated outposts in kent county in northcentral texas, where a specimen was collected from a sand sheet deposit. it has also been found at an isolated site in greer county, oklahoma, where it is associated with terraces of the north fork of the red river, near lake altus-lugert (see fig. 2). along both sides of the red river, m. cynanchoides populations are found on alluvial deposits (mostly cretaceous sands) intercalated between m. biflora populations that occur along upland bluffs on sedimentary substrates. populations in southern and eastern texas are found on various sandy deposits that include queen city sand, carrizo sand, the lissie formation, the willis formation, and the catahoula formation as well as mudstone, sandstone, siltstone, and alluvium. matelea cynanchoides is typically found in openings in cross timbers and pine-oak forests and in prairies. it is strongly associated with stabilized dune systems. this species tolerates disturbance and is regularly found in weedy sites along roads, in pastures, and other deforested areas. unlike its congener, this species is decumbent-upright, but it seems to recover well from the effects of mowing by producing branches from the base or from low axillary buds. in texas and oklahoma, m. cynanchoides has been well collected from two level iii ecoregions (griffith et al. 2004; woods et al. 2005): the south central plains and the east central texas plains. the westernmost collection of m. cynanchoides is from a sand sheet near the lubbock area, in the high plains ecoregion. there are also many collections from within the western gulf coastal plains ecoregion of texas (griffith et al. 2004). in oklahoma, m. cynanchoides also occurs in the central great plains and the cross timbers (woods et al. 2005). though few specimens record associated species, available data suggest that the associated species are numerous and diverse. these include graminoids in the genera aristida, cenchrus, dichanthelium, digitaria, eragrostis, eustachys, panicum, paspalum, and sporobolus. other herbaceous associates include species of acalypha, oklahoma native plant record 75 volume 14, december 2014 angela mcdonnell aristolochia, asclepias, berlandiera, chenopodium, cnidoscolus, croton, commelina, dalea, diodia, ditaxis, erigeron, eriogonum, eupatorium, gaillardia, galactia, helenium, helianthus, heliotropium, hymenopappus, hypericum, indigofera, lantana, lepidium, mimosa, monarda, opuntia, phyllanthus, physalis, richarida, rudbeckia, sida, sphaeralcea, stillingia, tetragonotheca, triodanis, verbena, vernonia, and yucca. woody associates include species of callicarpa, carya, celtis, diospyros, juniperus, pinus, prosopis, prunus, quercus, rhus, vaccinium, and vitis. there are no known pollinators or other faunal interactions for m. cynanchoides, but there has been one observation (fishbein, pers. comm.) of a small, unidentified weevil (curculionidae) visiting the flowers, apparently feeding on nectar. acknowledgements funding from the oklahoma state university department of botany mcpherson fund, the oklahoma state university botanical society, the society of systematic biologists, and the systematics research fund of the linnaean society has made this research possible. i thank dr. mark fishbein, whose expertise and dedication to milkweed research have been invaluable in all aspects of this project. i thank ben haack, kevin mcdonnell, bob o’kennon, and lindsey worcester, who each provided exceptional field support and assistance. i also thank an anonymous reviewer for comments and suggestions that greatly improved this manuscript. finally, i thank the collections managers and curators of each herbarium that i have worked with: ariz, asu, brit, cas, gh, ksc, lsu, mo, ny, ocla, okl, okla, srsc, tex/ll, unm, and us. references blankinship, j.w. 1907. plantae lindheimerianae. part iii. missouri botanical garden annual report 1907. pp. 123-223. boewe, c. 2003. profiles of rafinesque. knoxville: the university of tennessee press. collins, z. 1805-1827. correspondence with various botanists. philadelphia academy of natural sciences. coll. 129. correll, d.s. and m.c. johnston. 1970. manual of the vascular plants of texas. renner (tx): texas research foundation. decaisne, j. 1844. asclepiadeae. in a. p. de candolle (ed.). prodromus systematis naturalis regni vegetabilis.. vol. 8. pp. 490-665. paris: masson. engelmann, g. and a. gray. 1845. plantae lindheimerianae: an enumeration of f. lindheimer's collection of texan plants. with remarks, and descriptions of new species, etc: freeman and bolles. gray, a. 1878. synoptical flora of north america. vol. ii part i. gamopetalae after compositae. new york: ivison, blakeman, taylor, & co. griffith, g.e., s.a. bryce, j.m. omernik, j.a. comstock, a.c. rogers, b. harrison, and d. bezanson. 2004. ecoregions of texas (color poster with map, descriptive text, and photographs). reston (va): u.s. geological survey. heller, a.a. 1900. some changes in nomenclature. muhlenbergia 1(1): 1-8. krings, a., d.t. thomas, and q.-y. xiang. 2008. on the generic circumscription of gonolobus (apocynaceae, asclepiadoideae): evidence from molecules and morphology. systematic botany 33:403-415. lawson, r.m. 2004. the land between the rivers: thomas nuttall's ascent of the arkansas, 1819. ann arbor: the university of michigan press. lottinville, s. 1980. editor's introduction a jounal of travels into the arkansas territory during the year 1819. (pp. ix-xxiv). norman (ok): university of oklahoma press. 76 oklahoma native plant record volume 14, december 2014 angela mcdonnell mclean, e.p. 1980. asclepiadaceae of thomas nuttall at the academy of natural sciences of philadelphia. bartonia 47:31-35. nuttall, t. 1837. collections towards a flora of the territory of arkansas. transactions of the american philosophical society. pp. 139-203. parks, m. 2008. phylogeny of new world milkweed vines (apocynaceae, gonolobinae) [master’s thesis]. portland (or): portland state university. rafinesque, c.s. 1836. new flora of north america. vol. iv: neobotanon. philadelphia. redfield, mr. 1876. botanical correspondence of zaccheus collins. proceedings of the academy of natural sciences of philadelpia 28:81-82. stuckey, r.l. 1966. thomas nuttall's 1816 ohio valley plant collections described in his "genera" of 1818. castanea 187198. stuckey, r.l. 1971. the first public auction of an american herbarium including an account of the fate of the baldwin, collins, and rafinesque herbaria. taxon 20(4):443-459. thiers, b. 2014. index herbariorum: a global directory of public herbaria and associated staff. new york botanical garden's virtual herbarium. http://sweetgum.nybg.org/ih/. retrieved august 2014. torrey, j. 1859. botany of the boundary. in w. h. emory (ed.). report on the united states and mexican boundary survey. vol. 2. washington dc: u.s. govt. tyrl, r.j. and p.a. shryock. 2014. a cavalcade of field botanists in oklahoma–contributors to our knowledge of the flora of oklahoma. oklahoma native plant record 13:55-100. tyrl, r.j., s.c. barber, p. buck, w.j. elisens, j.r. estes, p. folley, l.k. magrath, c.l. murray, a.k. ryburn, b.a. smith, c.e.s. taylor, r.a. thompson, j.b. walker, l.e. watson. (in prep). flora of oklahoma: keys and descriptions. noble (ok): flora oklahoma inc. usgs (cartographer). 2005. preliminary integrated geologic map databases for the united states: central states: montana, wyoming, colorado, new mexico, north dakota, south dakota, nebraska, kansas, oklahoma, texas, iowa, missouri, arkansas, and louisiana. http://pubs.usgs.gov/of/2005/1351/. warren, l. 2004. constantine samuel rafinesque: a voice in the american wilderness: lexington (ky): university press of kentucky. woods, a.j., j.m. omernik, d.r. butler, j.g. ford, j.e. henley, b.w. hoagland, b.c. moran, 2005. ecoregions of oklahoma (color poster with map, descriptive text, summary tables, and photographs). reston (va): u.s. geological survey. woodson, r.e., jr. 1941. the north american asclepiadaceae i. perspective of the genera. annals of the missouri botanical garden 28:193-244. oklahoma native plant record 77 volume 14, december 2014 angela mcdonnell key to the species of matelea in oklahoma the following key includes the four species of matelea native to oklahoma and a version will appear in the next edition of the flora of oklahoma: keys and descriptions (tyrl et al., in prep). gonolobus is included in the key to aid in distinguishing g. suberosus, which has sometimes been placed in matelea as m. gonocarpa. 1. flowers with dorsal anther appendages. follicles thick-walled, winged. ...................... gonolobus 1. flowers lacking dorsal anther appendages. follicles thin-walled, smooth or muricate, but not winged. ..................................................................................... matelea 2. plants non-twining herbs with multiple prostrate to ascending stems from the base, 10–50 cm long. leaf blades 1–6 cm long, conspicuously and generally pubescent. corolla rotate-campanulate with lanceolate to deltoid lobes. 3. stems nearly to fully prostrate. corolla lobes narrowly deltoid to lanceolate, usually spreading, margins reflexed at maturity. adaxial surface of calyx and corolla with dense, thick hairs. ........................................... m. biflora 3. stems decumbent, ascending, or nearly erect. corolla lobes deltoid, usually ascending, margins planar. adaxial surface of calyx and corolla glabrous. ............................................................................................... m. cynanchoides 2. plants vines with 1-few stems from the base, 100–300 cm long. leaf blades 6–18 cm long, inconspicuously puberulent with hairs mostly limited to veins. corolla campanulate with narrowly lanceolate to linear, twisted lobes. 4. corolla white or cream. .................................................................................. m. baldwyniana 4. corolla maroon or brown-purple. ...................................................................... m. decipiens 78 oklahoma native plant record volume 14, december 2014 angela mcdonnell appendix specimens of matelea biflora and m. cynanchoides that were examined are listed below. data are presented in the following format: taxon: provenance, voucher (acronym of herbarium deposition). specimens collected by more than one person are listed here by the first name on the label. matelea biflora (nutt. ex raf.) woodson u.s.a., new mexico: lea co.: hutchins 9411 (nmu), sivinski 8456 (nmu) u.s.a., oklahoma: bryan co.: blain 131 (us), taylor 608 (okl), taylor 1413 (okl), taylor 24871 (okl), caddo co.: magrath 9764 (ocla, 2 sheets), nighswonger 1375 (okl), hoagland 2909 (okl), hoagland 2433 (okl), carter co.: fryxell 1367 (ny), goodman 7841 (okl), choctaw co.: leavenworth s.n. (ny, 2 sheets), magrath 16036 (ocla), comanche co.: thompson s0377 (okl), cotton co.: waterfall 7275 (okl), harmon co.: stevens 1169 (gh, ny), waterfall 7784 (okl), jackson co.: buthod ab-7372 (okl), buthod ab-10028 (okl), johnston co.: taylor 528 (okl), love co.: taylor 3605 (okl), major co.: rein 41 (okla), fishbein 6593 (okla), mccurtain co.: waterfall 17257 (gh, cas), demaree 12644 (okl), buthod ab-7197 (okl), murray co.: johnson 67 (okl), pontotoc co.: goodman 5454 (okl), waterfall 11425 (okl), johnson pon0154 (okl), stephens co.: magrath 16541 (ocla), tillman co.: smith 54 (okl), county uncertain: nuttall s.n. (ny, type), merrill 301 (us) u.s.a., texas: bastrop co.: tharp 1697 (us), bell co.: nesom 6432, bexar co.: harvard 30 (us, gh), schulz 512 (us), jermy s.n. (us), blanco co.: prinzie 221 (mo, okla), brown co.: carr 12728 (tex), rein 40 (okla), comanche co.: lehto l25070 (asu), concho co.: dorr 1563 (tex), dallas co.: lehto l25114 (asu), reverchon 2310 (gh, ny, 4 sheets, us), bush 623 (gh, ny, 2 sheets, us), reverchon s.n. (ny, 2 sheets), bodin 234 (us), reverchon 619 (us), denton co.: lundell 8423 (gh), cory 53715 (ny, us), eastland co.: johnston 73 (asu), edwards co.: cory 39042 (gh), erath co.: hoisington 244 (okl), garza co.: hutchins 546 (tex), wooton s.n. (us), hamilton co.: tharp, s.n. (gh), holland 10093 (ksc), hardeman co.: ball 1121 (us), hockley co.: thurow s.n. (us), johnson co.: lehto l25208 (asu), kerr co.: heller 1681 (gh, ny, us), llano co.: bray 336 (us), lubbock co.: wooton s.n. (us), demaree 7717 (us), demaree 7699 (us), mclennan co.: smith 535 (ny), prinzie 229 (mo, okla), massey 940 (okl), menard co.: mcvaugh 8291 (gh), rein 38 (okla), mitchell co.: tracy 7974 (us, gh, ny, 2 sheets), schleicher co.: turner 21-840 (tex), rein 39 (okla), sutton co.: cory 39624 (gh), tarrant co.: correll 32752 (gh), ruth 93 (ksc, us, 2 specimens, gh, ny, 3 specimens), cory 54532 (tex), carr 12833 (tex), pond s.n. (us, ny), killian 6799 (us), taylor co.: williams s.n. (gh), tom green co.: tweedy s.n. (ny, us), travis co.: poud s.n. (us), young s.n. (gh), carr 11100 (tex), tharp 1691 (us), tharp 1329 (us), wichita co.: whitehouse 10883 (ny), williamson co.: baird 3796 (ny), wise co.: mcdonnell 150 (okla), mcdonnell 172 (okla), bridges 13625 (tex), young co.: vollum s.n. (us, 2 sheets), cory 13144 (gh), county uncertain: wright s.n. (gh, 2 sheets, ny), lindheimer s.n. (gh), wright 545 (gh, 2 sheets), degener 5050 (ny), hayes s.n. (ny), stanfield s.n. (ny), ward s.n. (ny, us), stevenson s.n. (us), bebb 2394 (okl), bebb 2508 (okl) oklahoma native plant record 79 volume 14, december 2014 angela mcdonnell matelea cynanchoides (engelm. & a. gray) woodson u.s.a., arkansas: miller co.: thomas 134244 (ksc, lsu), kral 65495 (tex) u.s.a., louisiana: caddo parish: macroberts 88691 & 6891 (tex, lsu, ny, us), reid 5569 (lsu), reid 5578 (lsu), parish uncertain: leavenworth s.n. (ny) u.s.a., oklahoma: atoka co.: fishbein 6775 (okla), lewallen 2636 (okl), rein 56 (okla), rein 57 (okla), blaine co.: rein 42 (okla), waterfall 7071 (okl), bryan co.: taylor 1654 (okl), taylor 2294 (okl), caddo co.: pettijohn 139 (ocla), bittle 160 (okl), canadian co.: goodman 5846 (gh), goodman 7523 (okl), choctaw co.: waterfall, 16031 (ksc), hoagland hugo396 (okl), cleveland co.: stevens 1569 (gh), jeffs s.n. (okl), barkley s.n. (okl), smith 604 (okl), hawk 3 (okl), pusonett 16 (okl), custer co.: waterfall 2226 (okl), waterfall 7347 (okl), grady co.: mcdonnell 195 (okla), pettijohn 217 (ocla), bowers 224 (ocla), goin 6 (ocla), rein 43 (okla), rein 44 (okla), greer co.: joseph s.n. (okl), jefferson co.: goodman 7198 (okl), taylor 3632 (okl), kingfisher co.: bollenbach 47 (okl), folley 330 (okl), logan co.: carleton 154 (us, ksc), smith 539 (okl), smith 393 (okl), marshall co.: goodman 5926 (okl), mccurtain co.: schwenn 105 (ocla), payne co.: stratton 3046 (okl), pushmataha co.: magrath 11930 (ocla), magrath 15549 (ocla), magrath 11254 (ocla), tillman co.: johnson hf0071 (okl), county uncertain: carleton s.n. (ksu), palmer 182 (ny, us, 2 sheets) u.s.a., texas: anderson co.: rein 107 (okla), angelina co.: rein 105 (okla), aransas co.: berlandier 561 (gh), atascosa co.: orzell 6696 (tex), austin co.: lindheimer 273 (gh, 2 sheets, type), bastrop co.: lott 5093 (tex), lott 4497 (tex), bee co.: carr 24543 (tex), bexar co.: thurber 185 (gh), brazos co.: fryxell 2380 (ny), burnet co.: wolff 1551 (us), cooke co.: lusk s.n. (nmu), de witt co.: drushel 10771 (us), franklin co.: worcester 164 (okla), freestone co.: thomas 133705 (ny), frio co.: palmer 33883 (ny), gonzales co.: cory 5781 (gh), cory 8348 (gh), bogusch 1873 (us), goodman 6215 (okl), warnock 164 (tex), guadalupe co.: rein 54 (okla), harris co.: hall 520 (gh, us, ny, 2 sheets), thuron s.n. (us), henderson co.: correll 22110 (ny), jasper co.: orzell 11045 (tex), kent co.: correll 22110 (ny), leon co.: kral 67245 (gh), palmer 13418 (us), limestone co.: holmes 7116 (tex), navarro co.: joor 96 (us), newton co.: allen 22175 (lsu, 2 sheets), parker co.: quayle 566 (tex), refugio co.: hill 10613 (gh, ny), shelby co.: thomas, 129199 (ny), tyler co.: prinzie 225 (mo, okla), upshur co.: holmes 9964 (tex), van zandt co.: rein 108 (okla), correll 16211 (gh), wilbarger co.: correll 16211 (gh), county uncertain: tharp s.n. (gh), tharp 566 (ny), hayes s.n. (ny), bigelow s.n. (ny), drummond 203 (ny), wright s.n. (ny), wright 545 (gh, 2 sheets) non-twining milkweed vines of oklahoma: an overview ofmatelea biflora and matelea cynanchoides (apocynaceae) by ms. angela mcdonnell journal of the oklahoma native plantsociety, volume 3, number 1, december 2003 oklahoma native plant record 73 volume 3, number 1, december 2003 caddell, g. m. https://doi.org/10.22488/okstate.17.100024 critic’s choice essay take time to watch, not just smell the wildflowers! gloria m. caddell although plant-pollinator interactions between orchids and bees in the tropics may seem more interesting than those closer to home, oklahoma is full of fascinating plantpollinator interactions and mechanisms. the most important pollination agents in oklahoma are wind and insects. wind is particularly effective where many plants of the same species grow close together. prairie grasses, and dominant trees of our forests and woodlands, e.g. post oak and blackjack oak, are wind-pollinated. there are few things more beautiful than anthers dangling from a grass spikelet along with feathery stigmas that trap wind-borne pollen! in spring, male flowers of oaks are borne on pendulous catkins, releasing pollen that catches on stigmas of tiny female flowers held close to the branch. insects are major pollinators of our prairie forbs, and their flowers are visited by a variety of insects, including butterflies, moths, beetles, flies, bees, and wasps. of these, bees are most important. you can observe bumblebees with glistening, saddlebag-shaped pollinia of green milkweed (asclepias viridis) on their legs. you can hear their buzzing as bees clasp the anther cone of western horsenettle (solanum dimidiatum) and use their flight muscles to vibrate pollen out through pores at the top of the anthers. flower characters such as color, shape, size, and amount of nectar can sometimes be used to predict major pollinator(s) of a species. but it takes many hours of observing and collecting insect visitors to see if they are carrying pollen, to determine which are actual pollinators. my students and i have observed over 20 families of insect visitors to a single species, but find that only two or three effectively transfer pollen. differences in flowers among species are often clearly related to pollination, but differences among flowers within a single species may also be related to pollination and are equally intriguing. within a population you find sometimes subtle, and at other times obvious, differences between flowers at different stages. for example, when a pink gentian (sabatia campestris) flower opens its anthers are bright yellow and release pollen, but its style branches are green, coiled together, and lay flat against the petals. as the anthers wither the style branches uncoil, become erect, turn bright yellow, and their stigmas become receptive to pollen. in any population and even on the same plant, you can find flowers with style branches in various stages of uncoiling. difference in timing between pollen release and stigma receptivity is a mechanism to promote cross-pollination. when flowers of fog fruit (phyla) open, they have a yellow spot near the corolla tube opening (the “throat”). later in the day the spot turns a rosy-lavender color, less visible to bees. older flowers remain on the inflorescence as new flowers open, but in many cases such as this, newer, more attractive flowers offer a greater reward, e.g. more nectar. in prairie bluet (hedyotis nigricans) some plants bear flowers with long styles and short stamens. others bear flowers with short styles and long stamens with clearly visible blue anthers. insects that contact anthers of long stamens will likely transfer that pollen to a long-styled flower on another plant. so this mechanism also promotes cross-pollination. details of flowering stages and plantpollinator interactions of many oklahoma plants have not been well-documented. i encourage you to stop, sit, and not only take the time to “smell” the wildflowers, but to watch them as well. you will surely see things that have never been observed before! bacf oklahoma native plant record, volume 15, number 1, december 2015 oklahoma native plant record 105 volume 15, december 2015 paul buck https://doi.org/10.22488/okstate.17.100117 critic’s choice essay mistletoe, phoradendron serotinum (raf.) johnston reprinted from gaillardia, spring 1993 paul buck, deceased professor emeritus department of biological science university of tulsa every oklahoma child quickly becomes familiar with the common mistletoe, the green leaved growth on naked branches of large trees in mid-winter. this native plant occurs over most of the state and is particularly popular as one of the year-end holiday decorations. we all know it is permissible to steal a kiss from someone standing “under the mistletoe.” although considered by many to be a parasite, in reality the plant is only semiparasitic. it does obtain water, minerals, and perhaps some proteins from the host, but it is able to carry out photosynthesis and therefore produce most of its own food. in spite of the plant invading its tissue, the host is seldom harmed, unless of course there is a very heavy infestation. just 100 years ago in february 1893, mistletoe became the floral emblem of the territory of oklahoma. in 1909, the second state legislature conferred the same designation for the state of oklahoma. the following explanation for its selection appeared in the chronicles of oklahoma, the publication of the oklahoma historical society. tradition has it that the first grave made in oklahoma country in the winter after the opening of 1889 was covered with mistletoe since there were no other floral offerings in the new country except the green of the mistletoe with its white berries growing in great clusters on the elms along the dry creek beds and branches. all through the winter, the green bank of the lonely grave could be seen far across the prairie against the sere brown grass or the melting snow of early spring. thus, the mistletoe became associated with sacred thoughts among the pioneer settlers. in oklahoma, mistletoe is most commonly associated with ulmus americana (american elm), a species which has been badly ravaged by dutch elm disease, a fungus with tissue choking the water translocating tissues. mistletoe may also be found on hackberries, oaks, maples, ash, sycamore, and other native deciduous trees. this is fortunate; otherwise, the species might well become a candidate for rare or endangered status. the plants are dioecious (unisexual: staminate and pistillate flowers on different individuals). flowers are about 2 mm across, without petals, and borne on spike-like stalks from the bases of the leaves. the fruit, which are readily consumed by birds, are whitish, mucilaginous, one-seeded drupes, appearing during the winter. it has been suggested that dispersal takes place when the sticky seeds are “glued” to a twig as a bird wipes its bill, or the ingested, but unharmed, seeds are deposited on a limb with fecal material. 106 oklahoma native plant record volume 15, december 2015 paul buck used as a medicinal plant by indians and pioneers, a tea was prepared to relax nervous tension and muscle irritability and to increase blood pressure. other uses were to lessen bleeding, promote clotting, stimulate uterine contraction, and arrest postpartum hemorrhage. however, caution is advisable. like virtually all medications, mistletoe can be poisonous under certain conditions such as improper dosage levels, sensitive individuals, or with the very young, elderly, or feeble. there is no reliable information on safe dosages. although consumption of the fruit is harmless to pigs, 13 hereford cattle, forced to consume the plant when their pasture was reduced, died within 10 hours after the onset of symptoms. death was due to collapse of the cardiovascular system. several deaths among children, having consumed the fruit, have been documented. such is the state’s floral emblem, the oklahoma mistletoe, phoraendron serotinum — an interesting, beneficial, and potentially dangerous member of our native flora. bacf critic’s choice essay: mistletoe, phoradendron serotinum (raf.) johnston by dr. paul buck journal of the oklahoma native plant society, volume 8, number 1, december 2008 37 oklahoma native plant record volume 8, number 1, december 2008 barber, s.c. https://doi.org/10.22488/okstate.17.100060 updated list of taxa for vascular plants of the gypsum hills and redbed plains area of southwestern oklahoma susan c. barber associate provost and professor of biology oklahoma city university 2501 n. blackwelder, oklahoma city, ok 73106-1493 email: sbarber@okcu.edu the following is a list of vascular plants of the redbed plains and gypsum areas of southwestern oklahoma based on specimens collected by the author and deposited in the oklahoma state herbarium and the bebb herbarium of the university of oklahoma. in addition, 26 taxa collected by previous workers and four observed, but not collected, are included and so indicated. each taxon is listed alphabetically within its family and families are listed in order according to the engler-prantl classification scheme. nomenclature originally followed that of correll and johnston (1970) and waterfall (1969), but has been updated by bruce hoagland of the oklahoma biological survey according to the national plant data center, baton rouge, la <http://plants.usda.gov> accessed january 2009. marsileaceae marsilea vestita hook. & grev. (syn. = marsilea mucronata a. braun) polypodiaceae pellaea atropurpurea (l.) link var. atropurpurea cupressaceae juniperus pinchoti sudw.; u.t. waterfall(11261) april 4, 1953. j. virginiana l. ephedraceae ( = gnetaceae) ephedra antisyphilitica berl. ex c.a. mey. typhaceae typha angustifolia l. gramineae (poaceae) andropogon gerardii vitman var. gerardii a. hallii hack. aristida oligantha michx. a. purpurea nutt. a. purpurea nutt. var. fendleriana (steud.) vasey (syn. = a. fendleriana steud.) a. purpurea nutt. var. longiseta (steud.) vasey (syn. = a. longiseta steud.) arundo donax l. avena sativa l. bothriochloa ischaemum (l.) keng (syn.= andropogon ischaemum l.) b. saccharoides (sw.)rydb. (syn.= andropogon saccharoides sw.) bouteloua barbata lag.; u.t. waterfall (8729) august 26, 1948. b. curtipendula (michx.) torr. b. dactyloides (nutt.) j.t. columbus (syn. = buchloe dactyloides (nutt.) engelm.) b. gracilis (willd. ex kunth) lag. ex griffiths b. hirsuta lag. bromus arvensis l. (syn. = b. japonicus thunb.) b. catharticus vahl (syn. = b. unioloides kunth, bromus willdenowii) b. tectorum l. 38 oklahoma native plant record volume 8, number 1, december 2008 barber, s.c. calamovilfa gigantea (nutt.) scribn. & merr. cenchrus spinifex cav. chloris cucullata bisch. c. verticillata nutt. cynodon dactylon (l.) pers. digitaria sanguinalis (l.) scop distichlis spicata (l.) greene (syn. = distichlis spicata l. var. stricta (torr.) scribn.) echinochloa crus-galli (l.) p. beauv. elymus canadensis l. e. elymoides (raf.) swezey ssp. elymoides (syn. = sitanion hystrix (nutt.) j.g. sm.); j.g. smith (199) june 8, 1931; u.t. waterfall (8954) june 14, 1949. e. virginicus l. eragrostis barrelieri daveau e. cilianensis (all.) vign. ex janchen e. curvula (schrad.) nees (observed only) eragrostis secundiflora j. presl ssp. oxylepis (torr.) s.d. koch (syn. = e. oxylepis (torr.) torr. var. oxylepis) e. trichodes (nutt.) alph. wood (syn. = e. trichodes (nutt.) alph. wood var. pilifera (scheele) fernald) erioneuron pilosum (buckley) nash hordeum pusillum nutt. muhlenbergia arenicola buckley g.w. stevens (1111) june 21, 1913. m. asperifolia (nees & meyen ex trin.) parodi; u.t. waterfall august 26, 1948. panicum capillare l. var. capillare p. virgatum l. pascopyrum smithii (rydb.) a. löve (syn. = agropyron smithii (rydb. var. smithii) phalaris caroliniana walter pleuraphis mutica buckley poa arachnifera torr. polypogon monspeliensis (l.) desf. schedonnardus paniculatus (nutt.) trel. schizachyrium scoparium (michx.) nash (syn. = andropogon scoparius michx.) setaria leucophila (schribn. & merr.) k. schum. setaria pumila (poir.) roem. & schult. ssp. pumila (syn. = setaria lutescens (wiegel) f.t. hubb.) setaria reverchonii (vasey) pilg. ssp. reverchonii (syn. = p. reverchonii vasey); u.t. waterfall (7774) june 3, 1948 (7802) june 5, 1948. s. virdis (l.) p. beauv. sorghastrum nutans (l.) nash sorghum halepense (l.) pers. sporobolus airoides (torr.) torr. s. compositus (poir.) merr. var. compositus (syn. = s. asper (michx.) kunth). s. cryptandrus (torr.) a. gray s. giganteus nash; r.j. tyrl (883) & s.c. barber september 28, 1974. tridens albescens (vasey) woot. & standl. t. flavus (l.) hitchc. tridens muticus (torr.) nash var. elongatus (buckley) shinners (syn. = t. elongatus (buckley) nash) urochloa texana (buckley) r. webster (syn. = panicum texanum buckley) cyperaceae cyperus retroflexus buckley (syn. = c. uniflorus torr. & hook.) eleocharis compressa sull. e. macrostachya britton schoenoplectus americanus(pers.) volkart ex schinz & r. keller (syn. = scirpus americanus pers. var. americanus) commelinaceae commelina erecta l. var. erecta tradescantia occidentalis (britton) smyth t. ohiensis raf. forma ohiensis liliaceae allium drummondii regel androstephium coeruleum (scheele) greene forma coeruleum nothoscordum bivalve (l.) britton yucca glauca nutt. var. glauca iridaceae sisyrinchium angustifolium mill. 39 oklahoma native plant record volume 8, number 1, december 2008 barber, s.c. salicaceae populus deltoides bartram ex marsh. salix nigra marsh. fagaceae quercus havardii rydb. ulmaceae celtis laevigata willd. celtis laevigata willd. var. reticulata (torr.) l.d. benson (syn. = c. reticulata torr.) c. occidentalis l. moraceae maclura pomifera (raf.) c.k. schneid. morus alba l. polygonaceae eriogonum annuum nutt. e. longifolium nutt. var. longifolium polygonum lapathifolium l. p. pensylvanicum l. (syn. = polygonum bicorne raf.) rumex altissimus alph. wood r. crispus l. r. hymenosepalus torr. chenopodiaceae atriplex argentea nutt.; u.t. waterfall (8733) august 25, 1948. a. canescens (push) nutt. chenopodium album l. c. incanum (s. watson) a. heller u.t. waterfall (9084) june 14, 1949. kochia scoparia (l.) a.j. scott salsola tragus l. (syn. = salsola kali l. ssp. tenuifolia moq.) suaeda calceoliformis (hook.) moq. (syn. = suaeda depressa (pursh) s. watson) amaranthaceae amaranthus palmeri s. watson tidestromia lanuginosa (nutt.) standl. nyctaginaceae abronia fragrans nutt. ex hook. r.j. tyrl (855), c. mcdonald & p. risk may 15, 1974. portulacaceae portulaca pilosa l. caryophyllaceae arenaria serpyllifolia l. cerastium brachypodum engelm. ex a. gray) b.l. rob. paronychia jamesii torr. & a. gray (formerly in the illecebraceae) silene antirrhina l. forma antirrhina ranunculaceae anemone caroliniana walter forma violacea clute delphinium carolinianum walter ssp. virescens (nutt.) r.e. brooks (syn. = delphinium virescens nutt. var. pernardii (hutt.) l.m. perry) myosurus minimus l. (syn. = m. minimus l. var. interior boivin) papaveraceae argemone polyanthemos (fedde) g.b. ownbey cruciferae (brassicaceae) camelina microcarpa andrz. ex dc. capsella bursa-pastoris (l.) medik. descurainia pinnata (walter) britton ssp. halictorum (cockerell) detling (syn. = d. pinnata (walt.) britt. var. osmiarum (cockerell) shinners) d. sophia (l.) webb ex prantl dimorphocarpa candicans (raf.) rollins (syn. = dithyrea wislizenii engelm. var. palmeri payson) draba brachycarpa nutt. ex torr. & a. gray erysimum repandum l. lepidium austrinum small l. virginicum l. var. medium (greene) c.l. hitchc. lesquerella gordonii (a. gray) watson sibara virginica (l.) rollins 40 oklahoma native plant record volume 8, number 1, december 2008 barber, s.c. rosaceae prunus angustifolia marsh. leguminosae (fabaceae) acacia angustissima (mill.) kuntze var. hirta (nutt.) b.l. rob. (syn. = acacia hirta nutt.) astragalus lindheimeri englem. ex a. gray a. lotiflorus hook. a. missouriensis nutt. a. mollissimus torr. a. nuttallianus dc. var. nuttallianus a. plattensis nutt. a. racemosus prush caesalpinia gilliesii (wall. ex hook.) wall. ex d. dietr. chamaecrista fasciculata (michx.) greene var. fasciculata (syn. = cassia fasciculata michx.) dalea aurea nutt. ex pursh d. candida michx. ex willd. var. oligophylla (torr.) shinners (syn. = petalostemon candidus michx. var. oligophyllus (torr.) f.j. herm.) d. enneandra nutt. d. villosa (nutt.) spreng (syn. = petalostemon villosum nutt.) desmanthus illinoensis(michx.) macmill. ex b.l. rob. & fernald gleditsia triacanthos l. hoffmannseggia glauca (ortega) eifert (syn. = hoffmannseggia densiflora benth.) indigofera miniata ortega (i. miniata ortega var. leptosepala (nutt.) turner) medicago minima (l.) l. mimosa borealis a. gray m. microphylla dryand. (syn. = schrankia uncinata willd.) pediomelum cuspidatum (pursh) rydb. (syn. = psoralea cuspidata pursh) pomaria jamesii (torr. & a. gray) walp. (syn. = hoffmannseggia jamesii torr. & a. gray) psoralidium tenuiflorum (pursh) rydb. (syn. = psoralea tenuiflora pursh) prosopis glandulosa torr. var. glandulosa strophostyles leiosperma (torr. & a. gray.) piper vicia ludoviciana nutt. krameriaceae krameria lanceolata torr. linaceae linum pratense (norton) small (syn. = linum lewisii pursh var. pratense norton) l. rigidum pursh var. rigidum oxalidaceae oxalis corniculata l. o. dillenii jacq. geraniaceae erodium cicutarium (l.) l’hér. ex aiton e. texanum a. gray geranium carolinianum l. zygophyllaceae kallstroemia parviflora j.b.s. norton (syn. = k. intermedia rydb.) tribulus terrestris l. polygalaceae polygala alba nutt. euphorbiaceae chamaesyce albomarginata (torr. & a. gray) small (syn.= euphorbia albomarginata torr. & a. gray) c. glyptosperma (engelm.) small (syn. = euphorbia glyptosperma engelm.) c. lata (engelm.) small (syn. = euphorbia lata engelm.) c. missurica (raf.) shinners (syn. = euphorbia missurica raf.) cnidoscolus texanus (müll. arg.) small croton texensis (klotzsch) müll. arg. euphorbia cuphosperma (engelm.) boiss. (syn. = e. dentata michx. var. cuphosperma (engelm.) fern.) e. hexagona nutt. ex spreng. e. marginata pursh e. spathulata lam. reverchonia arenaria a. gray; u.t. waterfall (8340) july 21, 1948. stillingia sylvatica l. anacardiaceae rhus trilobata nutt. var. trilobata (syn. = r. aromatica 41 oklahoma native plant record volume 8, number 1, december 2008 barber, s.c. aiton var. flabelliformis shinners) r. microphylla engelm. ex a. gray u.t. waterfall (8447) may 13, 1950. toxicodendron radicans (l.) kuntze ssp. radicans (syn. = rhuss radicans l. var. radicans) sapindaceae sapindus saponaria l. var. drummondii (hook. & arn.) l.d. benson (syn. = sapindus drummondii hook. & arn.) rhamnaceae ziziphus obtusifolia (hook. ex torr. & a. gray) a. gray (syn. = condalia obtusifolia (hook. ex torr. & a. gray) weberb.) vitaceae vitis acerifolia raf. malvaceae callirhoe involucrata (torr. & a. gray) a. gray var. involucrata malvella leprosa (ortega) krapov. (syn. = sida leprosa (ortega) k. schum. var. hederaceae (douglas ex hook.) k. schum.) u.t. waterfall (9016) june 16, 1949. rhynchosida physocalyx (a. gray) fryxell (syn. = sida physocalyx a. gray); u.t. waterfall (8996) june 15, 1949. sphaeralcea coccinea (nutt.) rydb. tamaricaceae tamarix gallica l. loasaceae mentzelia decapetala (pursh ex sims) urb. & gilg ex gilg m. nuda (pursh) torr. & a. gray m. nuda (pursh) torr. & a. gray var. stricta (osterh.) harrington (syn. = m. stricta (osterhout) greene) m. oligosperma nutt. ex sims cactaceae cylindropuntia davisii (engelm. & bigelow) f.m. knuth (syn. = opuntia davisii engelm. & bigelow) c. leptocaulis (dc.) f.m. knuth (syn. = o. leptocaulis dc.) echinocactus texensis hopffer echinocereus reichenbachii (terscheck ex walp.) hort ex haage opuntia humifusa (raf.) raf. (syn. = opuntia compressa auct. non j.f. macbr.) lythraceae ammannia coccinea rottb. onagraceae calylophus hartwegii (benth.) p.h. raven ssp. fendleri (a. gray) towner & p.h. raven c. hartwegii (benth.) p.h. raven ssp. pubescens (a. gray) towner & p.h. raven (syn. = c. hartwegii (benth.) p.h. raven var. pubescens (a. gray) shinners) c. serrulatus (nutt.) p.h. raven gaura longiflora spach (syn. = gaura filiformis small) g. mollis james (syn. = g. parviflora douglas ex lehm.) g. sinuata nutt. ex ser. g. suffulta engelm. ex a. gray g. villosa torr. ssp. villosa oenothera grandis (britton) smyth (syn. = o. laciniata hill var. grandiflora (s. watson) b.l. rob.) o. rhombipetala nutt. ex torr. & a. gray o. speciosa nutt. o. triloba nutt. stenosiphon linifolius (nutt. ex james) heynh. umbelliferae (apiaceae) ammoselinum popei torr. & a. gray cymopterus macrorhizus buckley daucus pusillus michx. eurytaenia texana torr.& a. gray waterfall (11981) june 4, 1954. lomatium foeniculaceum (nutt.) j.m. coult. & rose ssp. daucifolium (torr. & a. gray) w.l. theobald (syn. = l. daucifolium (torr. & a. gray) j. m. coult. & rose) torilis arvensis (huds.) link 42 oklahoma native plant record volume 8, number 1, december 2008 barber, s.c. primulaceae androsace occidentalis pursh samolus ebracteatus kunth plumbaginaceae limonium limbatum small; u.t. waterfall (8319) july 21, 1948. oleaceae fraxinus pennsylvanica marsh. asclepiadaceae asclepias asperula (decne.) woodson ssp. capricornu (woodson) woodson (syn. = a. asperula (decne.) woodson var. decumbens (nutt.) shinners) a. arenaria torr. a. engelmanniana woodson cynanchum laeve (michx.) pers. matelea biflora (raf.) woodson convolvulaceae convolvulus arvensis l. cressa truxillensis kunth; u.t. waterfall (9423) may 13, 1950. cuscuta sp. observed only. evolvulus nuttallianus schult. polemoniaceae ipomopsis longiflora (torr.) v.e. grant hydrophyllaceae nama hispidum a. gray n. stevensii c.l. hitchc. phacelia integrifolia torr. boraginaceae cryptantha minima rydb. lappula occidentalis (s. watson) greene var. occidentalis (syn. = l. redowskii (hornem.) greene var. occidentalis (s. watson) rydb.) l. occidentalis (s. watson) greene var. cupulata (a. gray) higgins (syn. = l. texana (scheele) britton) lithospermum incisum lehm. verbenaceae glandularia canadensis (l.) nutt. (syn. = verbena canadensis (l.) britton) g. pumila (rydb.) umber (syn. = v. pumila rydb.) verbena bracteata cav. ex lag. & rodr. v. halei small v. plicata greene labiatae (lamiaceae) hedeoma drummondii benth. lamium amplexicaule l. forma amplexicaule monarda citriodora cerv. ex lag. m. punctata l. ssp. punctata var. occidentalis (epling) palmer & steyerm. (syn. = m. punctata l. ssp. occidentalis epling) salvia azurea michx. ex lam. var. grandiflora benth. scutellaria drummondii benth. s. wrightii a. gray forma wrightii teucrium canadense l. var. canadense (syn. = teucrium canadense l. var. virginicum (l.) eaton) t. laciniatum torr. solanaceae chamaesaracha coniodes (moric. ex dunal) britton datura wrightii regel (syn. = d. meteloides auct. non dunal. p.p.) lycium berlandieri dunal; u.t. waterfall (8994) june 15, 1949. nicotiana obtusifolia m. martens & galeotti var. obtusifolia (syn. = n. trigonophylla dunal); u.t. waterfall (7801) june 5, 1948. quincula lobata (torr.) raf. (syn. = physalis lobata torr. var. lobata) solanum dimidiatum raf. (syn. = s. torreyi a. gray forma torreyi) s. elaeagnifolium cav. s. rostratum dunal s. triflorum nutt.; g.w. stevens (1096) june 23, 1913. scrophulariaceae castilleja purpurea (nutt.) g. don var. citrina (pennell) shinners (syn. = c. citrina pennell) penstemon albidus nutt. p. cobaea nutt. p. fendleri torr.& a. gray veronica arvensis l. v. peregrina l. ssp. xalapensis (kunth) pennell 43 oklahoma native plant record volume 8, number 1, december 2008 barber, s.c. bignoniaceae catalpa bignonioides walter chilopsis linearis (cav.) sweet. martyniaceae proboscidea louisianica (mill.) thell. observed only. plantaginaceae plantago patagonica jacq. (syn. = p. purshii roem. & schult. var. spinulosa (decne.) shinners) p. rhodosperma decne. p. virginica l. p. wrightiana decne. rubiaceae houstonia humifusa (a. gray) a. gray (syn. = hedyotis humifusa a. gray) stenaria nigricans (lam.) terrell (syn. = h. nigricans (lam.) fernald) cucurbitaceae cucurbita foetidissima kunth ibervillea lindheimeri (a. gray) greene; u.t. waterfall (9406) may 13, 1950. campanulaceae triodanis holzingeri mcvaugh t. perfoliata (l.) nieuwl. compositae (asteraceae) achillea millefolium l. ambrosia psilostachya dc. var. lindheimeriana (scheele) blankenship a. trifida l. var. texana scheele amphiachyris dracunculoides (dc.) nutt. (syn. = gutierrezia dracunculoides (dc.) s.f. blake) aphanostephus pilosus buckley a. ramosissimus dc. a. skirrhobasis (dc.) trel. artemisia filifolia torr. a. ludoviciana nutt. ssp. ludoviciana baccharis salicina torr. & a. gray b. texana (torr.& a. gray) a. gray; u.t. waterfall (8361) july 23, 1948. berlandiera lyrata benth. var. lyrata centaurea americana nutt. chaetopappa ericoides (torr.) g.l. nesom (syn. = aster leucelene s.f. blake) cirsium texanum buckley conyza canadensis (l.) conq. var. glabrata (a. gray) conq. croptilon hookerianum (torr.& a. gray) house var. hookerianum (syn. = haplopappus divaricatus (nutt.) a. gray var. hookerianus (torr. & a. gray) waterf.) engelmannia peristenia (rafr.) goodman & c.a. lawson (syn. = e. pinnatifida a. gray ex nutt.) evax verna raf. flaveria campestris j.r. johnst. u.t. waterfall (8735) august 25, 1948. gaillardia pinnatifida torr. g. pulchella fouq. g. suavis (a. gray & engelm) britton & rusby grindelia nuda alph. wood var. nuda (syn. = g. squarrosa (pursh) dunal var. nuda (alph. wood) a. gray) g. papposa g.l. nesom & suh (syn = haplopappus ciliatus (nutt.) dc.) gutierrezia sarothrae (pursh) britton & rusby haploesthes greggii a. gray var. texana (j.m. coul.) i.m. johnst. helenium microcephalum dc. helianthus annuus l. h. petiolaris nutt. heterotheca canescens (dc.) shinners (syn. = chrysopsis villosa (pursh) nutt. ex dc. var. canescens a. gray) h. stenophylla (a. gray) shinners var. stenophylla (syn. = chrysopsis villosa (pursh) nutt. ex dc. var. stenophylla (a. gray) a. gray) h. subaxillaris (lam.) britton & rusby (syn. = heterotheca latifolia buckley) hymenopappusi scabiosaeus l’her var. corymbosus (torr. & a. gray) b.l. turner h. tenuifolius pursh; u.t. waterfall (7307) june 28, 1947. hymenoxys odorata dc. 44 oklahoma native plant record volume 8, number 1, december 2008 barber, s.c. iva annua l. var. annua (syn. = iva ciliata willd.) liatris punctata hook. var. nebraskensis gasier l. punctata hook. var. punctata lindheimera texana a. gray & engelm. lygodesmia texana (torr. & a. gray) greene (syn. = lygodesmia aphylla (nutt.) dc. var. texana torr & a. gray.) machaeranthera pinnatifida (hook.) shinners ssp. pinnatifida var. pinnatifida (syn. = haplopappus spinulosis (pursh) dc.) palafoxia sphacelata (nutt. ex torr.) cory pluchea odorata (l.) cass. var. odorata (syn. = p. purpurascens (sw.) dc.) psilostrophe tagetina (nutt.) greene var. cerifera (a. nelson) b.l. turner (syn. = p. villosa rydb.) pyrrhopappus grandiflorus (nutt.) nutt. p. pauciflorus (d. don) dc. (syn. = p. multicaulis dc. var. geiseri (shinners) northington) ratibida columnifera (nutt.) woot. & stand. forma columnifera r. tagetes (james) barnhart; g.w. stevens (1080) june 21, 1913. rudbeckia hirta l. var. pulcherrima farw. senecio riddellii torr. & a. gray silphium laciniatum torr. var. laciniatum solidago gigantea aiton (syn. = s. gigantea aiton var. leiophylla fernald) s. missouriensis nutt. var. faciculata holz. symphyotrichum divaricatum (nutt.) g.l. nesom (syn. = aster subulatus michx. var. ligulatus shinners) s. ericoides (l.) g.l. nesom (syn. = aster ericoides l.) s. oblongifolium (nutt.) g.l. nesom (syn. = aster oblongifolius nutt.) tetraneuris scaposa (dc.) greene (syn. = hymenoxys scaposa (dc.) k.f. parker var. scaposa) thelesperma filifolium (hook.) a. gray t. megapotamicum (spreng.) kuntze tragopogon dubius scop. (syn. = t. major jacq.) verbesina encelioides (cav.) benth. & hook. f. ex a. gray vernonia baldwinii torr. var. interior (small) faust v. marginata (torr.) raf.; bruce harkins (91) october 17, 1970. xanthisma texanum dc. ssp. drummondii (torr. & a. gray) semple (syn. = x. texanum dc. var. drummondii (torr. & a. gray) a. gray) xanthium strumarium l. var. canadense (mill.) torr.& a. gray x. strumarium l. var. glabratum (dc.) cronquist. zinnia grandiflora nutt. journal of the oklahoma native plantsociety, volume 3, number 1, december 2003 oklahoma native plant record 19 volume 3, number 1, december 2003 additions to black mesa flora study patricia folley bebb herbarium university of oklahoma many changes have taken place in far western oklahoma since jim mcpherson’s untimely death in 1994. then the mesa lobe containing the highest point in oklahoma and its surrounding slopes was owned and managed by the oklahoma chapter of the nature conservancy, but it soon became part of black mesa state park. private lands still separate the two portions of the park, but they are now connected by a public road. since that time and during the course of several weekend outings with the oklahoma academy of science and the oklahoma native plant society, i have been adding to mcpherson’s list, as many plant species as could be found in bloom or fruit. these species are not already listed by mcpherson as present in the mesa area. voucher specimens are housed in the robert bebb herbarium (okl) at the university of oklahoma. most visitors to black mesa camp in the original park area near lake etling. in this list plants collected at that site are noted as “park”. plants found along the roads leading to the mesa or to the outlying canyons are noted as “roadside”. a few plants were found only at privately owned, tessequite canyon, one of the many side canyons leading down from the mesa. those are identified with the name “tessequite”. “mesa” denotes plants from black mesa or its slopes. mcpherson’s collections were solely from the mesa. through the years black mesa state park has been studied by several botanists, including c.m. rogers and u.t. waterfall, as well as mcpherson. it is hoped that their work, along with this list will serve as a basis for the initiation of future explorations in that geographic region. the approximate gps location of black mesa state park is between latitudes 36.833 and 36.861 and longitudes 102.862 and 102.900 the elevation of the mesa ranges from 4960 ft (1512 m) to 4973 ft (1516 m). it is now contained within black mesa state park which contains approximately 349 acres of land. references correll, d.s. and m.c. johnston. 1970. manual of the vascular plants of texas, renner, tx: texas research foundation. taylor, j.r. and c.e.s. taylor. 1991. an annotated list of the ferns, fern allies, gymnosperms and flowering plants of oklahoma. [published by the authors at southeastern oklahoma state university]. tyrl, r.j., susan barber, paul buck, wayne elisens, james estes, patricia folley, lawrence magrath, constance taylor, and rahmona thompson. the flora of oklahoma. the flora of oklahoma editorial board. forthcoming. usda-nrcs 2003. the plants database. (http://plants.usda.gov/plants.) folley, p.a. https://doi.org/10.22488/okstate.17.100019 20 oklahoma native plant record volume 3, number 1, december 2003 additional plant list for black mesa 2003 common name site status longhorn milkweed park native nodding tickseed park introduced fetid marigold park native wright’s cudweed park native rayed palafoxia roadside native paperflower park native groundsel mesa native prairie ironweed park native bindweed heliotrope park native many-flowered puccoon park native wallflower park native hedgehog cactus mesa native brittle prickly pear park native white-spine prickly pear park native brownspine prickly pear park native cardinal flower park native rocky mountain beeplant park native james' clammyweed park native nailwort roadside native 4-wing saltbush park native family/species family asclepiadaceae asclepias macrotis family asteraceae bidens cernua dyssodia papposa gnaphalium wrightii palafoxia sphacelata psilostrophe villosa senecio flaccidus vernonia fasciculata family boraginaceae heliotropium convolvulaceum lithospermum multiflorum family brassicaceae erysimum asperum family cactaceae echinocereus viridiflora opuntia fragilis opuntia macrorhiza opuntia phaeacantha family campanulaceae lobelia cardinalis family capparidaceae cleome serrulata polanisia jamesii family caryophyllaceae paronychia sessiliflora family chenopodiaceae atriplex canescens chenopodium leptophyllum folley, p. a. narrowleaf goosefoot park native oklahoma native plant record 21 volume 3, number 1, december 2003 folley, p.a. kochia scoparia tumbleweed roadside introduced family cyperaceae carex brevior sedge park native cyperus globulosus park native scirpus acutus hardstem bulrush park native scirpus atrovirens darkgreen bulrush park native family dryopteridaceae cystopteris fragilis brittle fern park native family fabaceae astragalus ceramicus painted milkvetch park native colutea arborescens roadside introduced dalea tenuifolia slimleaf prairie clover park native family fagaceae quercus mohriana shin oak tessequite native family lamiaceae teucrium laciniatum cutleaf germander park native family liliaceae nolina texana beargrass tessequite native yucca harrimaniae new mexico yucca park native family loasaceae mentzelia oligosperma stickleaf park native family nyctaginaceae mirabilis albida white 4 o'clock park native mirabilis nyctaginea wild 4 o'clock park native family onagraceae gaura villosa wooly gaura roadside native oenothera engelmannii engelmann’s eve. primrose roadside native oenothera latifolia mountain eve. primrose park native family poaceae andropogon virginicus sand sedge park native chloris virgata park native distichlis spicata saltgrass park native oklahoma native plant record volume 3, number 1, december 2003 22 marsh muhly park native switchgrass park native rabbit-foot grass park native prickleaf gilia base of mesa native no common name park & mesa native water knotweed park native tall dock park native small-flowered buttercup lake native western wood-rose park native wafer ash mesa native water speedwell park introduced cutleaf verbena park native muhlenbergia racemosa panicum virgatum polypogon monspeliensis family polemoniaceae gilia rigidula family polygonaceae eriogonum tenellum polygonum amphibium rumex altissimus family ranunculaceae ranunculus abortivus family rosaceae rosa woodsii family rutaceae ptelia trifoliata family scrophulariaceae veronica anagalis-aquatica family verbenaceae glandularia bipinnatifida family vitaceae vitis acerifolia folley, p.a. no common name park native oklahoma native plant record, volume 15, number 1, december 2015 6 oklahoma native plant record volume 15, december 2015 ben osborn https://doi.org/10.22488/okstate.17.100111 first flowering dates for central oklahoma unpublished report bebb herbarium university of oklahoma may 1, 1934 ben osborn keywords: blooming, climate change, historical, phenology introduction the following list of native and cultivated plants is arranged according to the first recorded date of blooming for each species as observed in oklahoma county or any of the counties contiguous to it. most of the records are from norman in cleveland county, with a few from oklahoma city in oklahoma county and shawnee in pottawatomie county. where more than one has been recorded, the average is given as the arithmetic average of the dates. dates of observations of plants which had attained full bloom, where known, have been eliminated, except where they were the earliest recorded dates for the species. such dates have been starred (*) in the list. a plant is considered in bloom when as many as one flower is open and having either stamens shedding pollen or the stigma ready to receive pollen, as indicated by pollen grains adhering to it or the obviously mature condition of the stigma. exceptionally early bloomings within the shelter of buildings or other barriers are included, but those resulting from artificial heat have been eliminated. oklahoma native plant record 7 volume 15, december 2015 ben osborn first flowering dates editor’s note: where nomenclature has been updated using itis–integrated taxonomic information service (http://www.itis.gov), the original name is in brackets [ ]. there are no voucher specimens for this work; therefore, species identifications are provisional. the dates of observations of plants which have attained full bloom are marked with a single asterisk (*). observations added by fred barkley on 6 june 1934, are marked with a double asterisk (**). january 1 january stellaria media. average 30 january. earliest, oklahoma city, 1933, osborn.** 7 january common dandelion, taraxacum officinale. average 25 january. earliest, norman, 1929, gould. 19 january shepherd’s purse, capsella bursa-pastoris. average 7 february. earliest, norman, 1927, gould. 21 january chinese elm, ulmus pumila. earliest, oklahoma city, 1933, osborn. 23 january american elm, ulmus americana. average 10 february. earliest, oklahoma city, 1933, osborn.** 28 january silver maple, acer saccharinum. average 8 february. earliest, oklahoma city, 1934, osborn. 30 january arbor vitae, thuja occidentalis. oklahoma city, 1933, osborn. february 1 february common violet, viola sororia var. sororia [=viola papilionacea]. oklahoma city, 1933, osborn. 3 february spring beauty, claytonia virginica. average 18 february. earliest, norman, 1927, gould. 5 february bush honeysuckle, lonicera fragrantissima. average 6 february. earliest, norman, 1927, gould. 6 february wild pansy, viola bicolor [=viola rafinesquii]. average 24 february. earliest, norman, 1928, gould. virginia rock-cress, planodes virginica [=arabis virginica]. average 26 february. earliest, norman, 1928, gould. http://www.itis.gov/ 8 oklahoma native plant record volume 15, december 2015 ben osborn 7 february jasminum nudiflorum [=jasminum nudum]. norman, 1927, gould. 8 february least bluet, houstonia pusilla [=houstonia minima]. average 27 february. earliest, norman, 1927, gould. 14 february goldenbells, forsythia suspensa. norman, 1927, gould. 24 february carolina whitlow-grass, draba reptans [=draba caroliniana]. average 2 march. earliest, norman, 1927, gould. midland dogtooth-violet, erythronium mesochoreum. average 6 march. earliest, norman, 1927, gould. anemone caroliniana. average 14 march. earliest, norman, 1927, gould. 25 february shortpod whitlow-grass, draba brachycarpa. average 7 march. earliest, norman, 1927, gould. 28 february red cedar, juniperus virginiana. average 6 march. earliest, norman, 1927, gould. thunberg spirea, spiraea thunbergii. average 7 march. earliest, norman, 1927, gould. march 3 march hoary tansy-mustard, descurainia incana [=sisymbrium canescens]. average 29 march. earliest, norman, 1928, gould. 4 march slippery elm, ulmus rubra [=ulmus fulva]. average 8 march. earliest, norman, 1927, gould.** 5 march chaenomeles japonica. average 14 march. earliest, norman, 1928, gould. 7 march peach, prunus persica. average 16 march. earliest, norman, 1927, gould. androsace occidentalis. average 12 march. earliest, norman, 1927, gould. 10 march pear, pyrus communis. norman, 1927, gould. shortstalk chickweed, cerastium brachypodum. average 10 march. earliest, norman, 1928, gould. common henbit, lamium amplexicaule. (open flowers). average 12 march. earliest, norman, 1928, gould. plantainleaf cat’s-foot, antennaria plantaginifolia. average 24 march. earliest, norman, 1927, gould. chickasaw plum, prunus angustifolia. average 17 march. earliest, norman, 1927, gould.** oklahoma native plant record 9 volume 15, december 2015 © ben osborn journal compilation © 2015 oklahoma native plant society 11 march yellow false-garlic, nothoscordum bivalve. average 19 march. earliest, oklahoma city, 1924, osborn. spreading pearlwort, sagina decumbens. average 26 march. earliest, norman, 1928, gould. 12 march box-elder, acer negundo. average 19 march. earliest, norman, 1928, gould. 15 march prunus triloba. norman, 1927, gould. apricot, prunus armeniaca. norman, 1927, gould. yellow woodsorrel, oxalis stricta. average 31 march. earliest, norman, 1927, gould. 16 march missouri buffalo currant, ribes aureum. average 19 march. earliest, norman, 1927, gould. 18 march redbud, cercis canadensis. average 24 march. earliest, norman, 1927, gould. tulip, tulipa sp. oklahoma city, 1934, osborn. 19 march low plum, prunus gracilis. average 23 march. earliest, norman, 1928, gould.** carrotleaf parsley, lomatium foeniculaceum ssp. daucifolium [=lomatium daucifolium]. average 24 march. earliest, norman, 1928, gould. missouri violet, viola sororia var. missouriensis [=viola missouriensis]. average 24 march. earliest, norman, 1928, gould. (reported as v. papilionacea.) 20 march berberis aquifolium [=mahonia aquifolium]. norman, 1927, gould. 22 march cottonwood, populus deltoides. average 26 march. earliest, norman, 1928, gould. 24 march lilac, syringa x persica [=syringa persica]. norman, 1927, gould. common pansy, viola tricolor. norman, 1927, gould. american plum, prunus americana. norman, 1929, gould. ground-plum, astragalus crassicarpus var. crassicarpus [=astragalus caryocarpus]. average 28 march. earliest, norman, 1929, gould. 25 march narrowleaf puccoon, lithospermum incisum [=lithospermum angustifolium]. average 27 march. earliest, norman, 1928, gould. 27 march lonicera tatarica. norman, 1927, gould. lonicera morrowii. norman, 1927, gould. 10 oklahoma native plant record volume 15, december 2015 © ben osborn journal compilation © 2015 oklahoma native plant society 28 march ash, fraxinus sp. oklahoma city, 1933, osburn. 29 march eastern hackberry, celtis occidentalis. norman, 1928, gould. maclura pomifera. norman, 1928, gould. krigia caespitosa [=scrinia oppositifolia]. average 9 april. earliest, norman, 1929, gould. purple poppy-mallow, callirhoe involucrata. average 13 april. earliest, norman, 1928, gould. 30 march white mulberry, morus alba. norman, 1928, gould. purslane speedwell, veronica peregrina. average 2 april. earliest, norman, 1928, gould. androstephium coeruleum. average 5 april. earliest, norman, 1928, gould. red mulberry, morus rubra. average 9 april. earliest, norman, 1928, gould. 31 march chaenomeles speciosa [=chaenomeles lagenaria]. norman, 1927, gould. prairie false-dandelion, nothocalais cuspidata [=agoseris cuspidata]. average 2 april. earliest, norman, 1927, gould. april 1 april apple, malus pumila [=pyrus malus]. norman, 1927, gould. prunus cerasus. norman, 1927, gould. scarlet strawberry, fragaria virginiana. norman, 1927, gould. thunberg barberry, berberis thunbergii. norman, 1927, gould. common lilac, syringa vulgaris. average 2 april. earliest, norman, 1927, gould. stout blue-eyed grass, sisyrinchium angustifolium [=sisyrinchium gramineum]. average 3 april. earliest, norman, 1928, gould. tall dock, rumex altissimus. average 13 april. earliest, oklahoma city, 1933, osborn. 2 april caragana arborescens. norman, 1927, gould. colutea arborescens. norman, 1927, gould. white ash, fraximus americana. average 6 april. earliest, norman, 1927, gould. philadelphia fleabane, erigeron philadelphicus. average 3 april. earliest, norman, 1928, gould. blackjack oak, quercus marilandica. average 4 april. earliest, norman, 1928, gould. prairie ragwort, packera plattensis [=senecio plattensis]. average 8 april. earliest, norman, 1928, gould. violet wood-sorrel, oxalis violacea. average 9 april. earliest, oklahoma city, 1933, osborn. cutleaf evening-primrose, oenothera laciniata. average 9 april. earliest, norman, 1929, gould. roundleaf moneyflower, mimulus glabratus var. jamesii. average 21 april. earliest, norman, 1928, gould. 3 april comptonia peregrina [=myrica asplenifolia]. norman, 1927, gould. oklahoma native plant record 11 volume 15, december 2015 © ben osborn journal compilation © 2015 oklahoma native plant society 4 april ovalleaf bladderpod, physaria ovalifolia ssp. ovalifolia [=lesquerella ovalifolia]. norman, 1929, gould. spurge, euphorbia sp. oklahoma city, 1933*, osborn. 5 april ellisia nyctelea. norman, 1927, gould. smooth yellow violet, viola pubescens var. scabriuscula [=viola scabriuscula]. norman, 1927, gould. virginia pepper-grass, lepidium virginicum. norman, 1929, gould. poncirus trifoliata. norman, 1927, gould. juglans cinera. norman, 1927, gould. celtis laevigata [=celtis mississippiensis]. norman, 1927, gould. mousetail, myosurus minimus. average 8 april. earliest, norman, 1929, gould. dewberry, rubus flagellaris [=rubus villosus]. average 14 april. earliest, norman, 1927, gould. spiny sow-thistle, sonchus asper. average 15 april. earliest, norman, 1928, gould. 6 april sycamore, platanus occidentalis. norman, 1929, gould. 7 april broussonetia papyrifera. norman, 1929, gould. slender bladderpod, physaria gracilis [=lesquerella gracilis]. norman, 1929, gould. 8 april plantago elongata. norman, 1927, gould. wild columbine, aquilegia canadensis. norman, 1927, gould. spiraea vanhoutei. norman, 1927, gould. tamarisk, tamarix gallica. average 9 april. earliest, norman, 1927, gould. large wild-onion, allium canadense var. mobilense [=allium mutabile]. average 12 april. earliest, norman, 1927, gould. blue toadflax, nuttallanthus canadensis [=linaria canadensis]. average 16 april. earliest, norman, 1927, gould. wooly yarrow, achillea millefolium [=achillea lanulosa]. average 23 april. earliest, norman, 1927, gould. meadow garlic, allium canadense. average 23 april. earliest, oklahoma city, 1933, osborn. 9 april corn speedwell, veronica arvensis. norman, 1929, gould. lonicera flava. norman, 1927, gould. black willow, salix nigra. average 12 april. earliest, oklahoma city, 1933, osborn. bracted false-indigo, baptisia bracteata. average 14 april. earliest, oklahoma city, 1933, osborn. western plantain, plantago virginica. average 14 april. earliest, norman, 1927, gould. 10 april black-haw, viburnum plicatum [=viburnum tomentosum]. norman, 1927, gould. western crab-apple, malus ioensis [=pyrus ioensis]. norman, 1927, gould. aquilegia coerulea. norman, 1927, gould. 12 oklahoma native plant record volume 15, december 2015 ben osborn spreading chervil, chaerophyllum procumbens. average 13 april. earliest, oklahoma city, 1933, osborn. goose-grass, galium aparine. average 14 april. earliest, oklahoma city, 1933, osborn. 11 april tropaeolum majus. norman, 1927, gould. robinia hispida. norman, 1927, gould. salix alba [=salix vitellina]. norman, 1927, gould. western tansy-mustard, descurainia incisa ssp. incisa [=sisymbrium incisum]. norman, 1928, gould. vaillant’s goose-grass, galium aparine [=galium aparine baillantii]. average 12 april. earliest, norman, 1928, gould. tetraneuris linearifolia [=actinea linearifolia]. average 14 april. earliest, norman, 1928, gould. 12 april viburnum opulus [=viburnum opulus sterile]. norman, 1927, gould. black medick, medicago lupulina. average 6 may. earliest, norman, 1927, gould. 13 april fragrant sumac, rhus aromatica [=rhus canadensis]. norman, 1927, gould. kerria japonica. norman, 1927, gould. frost grape, vitis vulpina [=vitis cordifolia]. average 10 april. norman, 1928, gould. 14 april alyssum alyssoides. norman, 1927, gould. crataegus calpodendron [=crataegus globosa]. norman, 1927, gould. black-haw, virburnum prunifolium. norman, 1927, gould. leafystem false-dandelion, pyrrhopappus carolinianus. norman, 1927, gould. corydalis aurea ssp. occidentalis. average 19 april. earliest, norman, 1927, gould. black locust, robinia pseudoacacia. average 20 april. earliest, norman, 1927, gould. beaked corn-salad, valerianella radiata. average 22 april. earliest, norman, 1927, gould. pecan, carya illinoiensis. average 27 april. earliest, norman, 1927, gould. 15 april calycanthus floridus. norman, 1927, gould. pepper-grass, lepidium apetalum. average 15 april. earliest, norman, 1929, gould, and oklahoma city, 1933, osborn. silverleaf nightshade, solanum elaeagnifolium. average 29 april. earliest, oklahoma city, 1933, osborn. 16 april western crab apple, pyrus ioensis. norman, 1927, gould.** cryptanthe fendleri. average 20 april. earliest, norman, 1927, gould. post oak, quercus stellata. average 17 april. earliest, norman, 1927, gould. western daisy, astranthium integrifolium ssp. integrifolium [=bellis integrifolia]. average 20 april. earliest, norman, 1927, gould. red oak, quercus rubra [=quercus borealis]. norman, 1929, gould. 13 oklahoma native plant record volume 15, december 2015 17 april rhodotypos scandens [=rhodotypos kerrioides]. norman, 1927, gould. yellow oak, quercus muhlenbergii. norman, 1929, gould. cotoneaster gaballei. norman, 1927, gould. cotoneaster horizonatlis. norman, 1927, gould. southern black-haw, viburnum rufidulum. average 19 april. earliest, norman, 1929, gould. burr oak, quercus macrocarpa. average 23 april. earliest, norman, 1929, gould. 18 april nandina domestica. norman, 1927, gould. aronia melanocarpa. norman, 1927, gould. carolina cranebill, geranium carolinianum. average 27 april. earliest, norman, 1928, gould. 19 april phacelia hirsuta. norman, 1927, gould. eleagnus angustifolia. norman, 1927, gould. deutzia scabra. norman, 1927, gould. stemless loco-weed, oxytropis lambertii. average 24 april. earliest, norman, 1927, gould.** small skullcap, scutellaria parvula. average 25 april. earliest, norman, 1927, gould. 20 april rosa rubiqinosa [=rosa eglanteria]. norman, 1927, gould. 21 april hoary puccoon, lithospermum canescens. norman, 1929, gould. light poppy-mallow, callirhoe alcaeoides. oklahoma city, 1934, osborn. 22 april white clover, trifolium repens. oklahoma city, 1934, osborn. nuttall onion, allium drummondii [=allium nuttallii]. norman, 1928, gould. true water-cress, nasturtium officinale [=radicula nasturium-aquaticum]. norman, 1929, gould. western spiderwort, tradescantia occidentalis. average 24 april. earliest, norman, 1929, gould. poison oak, toxicodendron pubescens [=rhus toxicodendron]. average 29 april. earliest, norman, 1929, gould. horsenettle, solanum carolinense. average 3 may. earliest, 1929, gould. 23 april smallflower verbena, glandularia bipinnatifida var. bipinnatifida [=verbena bipinnatifida]. norman, 1938, gould. rock sandwort, minuartia tenella [=arenaria stricta]. norman, 1928, gould. petioled wild four-o-clock, mirabilis nyctaginea [=oxybaphus nyctagineus]. average 1 may. earliest, norman, 1929, gould. 24 april alfalfa, medicago sativa. average 1 may. earliest, norman, 1929, gould. curly dock, rumex crispus. average 3 may. earliest, norman, 1929, gould. ben osborn 14 oklahoma native plant record volume 15, december 2015 © ben osborn journal compilation © 2015 oklahoma native plant society 25 april red-haw, crataegus sp. average 25 april. norman, 1928–29, gould. rough false-dandelion, pyrrhopappus grandiflorus [=pyrrhopappus scaposus]. average 26 april. earliest, norman, 1929, gould. blue false-indigo, baptisia australis. average 27 april. earliest, norman, 1929, gould. chaetopappa asteroides. average 29 april. earliest, norman, 1928, gould. black walnut, juglans nigra. average 30 april. earliest, norman, 1928, gould. wild parsnip, pastinaca nativa. average 1 may. earliest, norman, 1928, gould. hairy puccoon, lithospermum caroliniense [=lithospermum gmelini]. average 3 may. earliest, norman, 1928, gould. 26 april vetch, vicia sp. (cultivated). oklahoma city, 1934, osborn. basket oak, quercus michauxii. norman, 1928, gould. 27 april common greenbriar, smilax rotundifolia. norman, 1929, gould. virginia willow, itea virginica. norman, 1929, gould. yellow sweet-clover, melilotus officinalis. average 4 may. earliest, oklahoma city, 1934*, osborn. 28 april spreading fleabane, erigeron divergens. average 3 may. earliest, norman, 1927, gould. serrateleaf evening-primrose, oenothera serrulata. average 1 may. earliest, norman, 1927, gould. 29 april skunk bush, rhus aromatica [=rhus canadensis trilobatus]. norman, 1928, gould. purple milkweed, asclepias purpurascens. norman, 1928, gould. bluntleaf milkweed, asclepias amplexicaulis. average 5 may. earliest, norman, 1929, gould. wavyleaf gaura, oenothera sinuosa [=gaura sinuata]. average 2 may. earliest, norman, 1929, gould.** 30 april reflexed spiderwort, tradescantia ohiensis [=tradescantia reflexa]. norman, 1927, gould. mexican sandbur, tribulus terrestris. average 30 april. earliest, norman, 1928, gould. cutleaf bayless gaillardia, gaillardia suavis. average 5 may. earliest, oklahoma city, 1933, osborn. oblongleaf milkweed, asclepias viridis [=asclepiodora viridis]. average 6 may. earliest, oklahoma city, 1933, osborn. may 1 may vetch, vicia tetrasperma. norman, 1928, gould. small venus’-lookingglass, triodanis perfoliata ssp. biflora [=specularia biflora]. norman, 1928, gould. prairie larkspur, delphinium carolinianum [=delphinium penardi]. norman, 1928, gould. sand grape, vitis rupestris. norman, 1928, gould. american mistletoe, phoradendron serotinum ssp. serotinum [=phoradendron flavescens]. norman, 1928, gould. oklahoma native plant record 15 volume 15, december 2015 ben osborn 2 may sleepy catchfly, silene antirrhina. average 6 may. earliest, norman, 1929, gould. venus’-lookingglass, specularia perfoliata. average 12 may. earliest, norman, 1928, gould. 3 may salsify, tragopogon porrifolius. norman, 1928, gould. river locust, amorpha fruticosa. average 5 may. earliest, norman, 1927, gould. scarlet gaura, oenothera suffrutescens [=gaura coccinea]. norman, 1927, gould. downy phlox, phlox pilosa. average 4 may. norman, 1929, gould. calystegia sepium ssp. angulata [=convolvulus repens]. average 6 may. earliest, norman, 1929, gould. smooth soapweed, yucca glauca. average 7 may. earliest, norman, 1929, gould. large beardtongue, penstemon cobaea. average 7 may. earliest, norman, 1927, gould. slender beardtongue, penstemon gracilis. average 2 may. earliest, norman, 1929, gould. missouri evening-primrose, oenothera macrocarpa ssp. macrocarpa [=oenothera missouriensis]. average 8 may. earliest, norman, 1929, gould. whorled tickseed, coreopsis verticillata. average 8 may. earliest, norman, 1927, gould. spermolepis echinata. average 12 may. earliest, norman, 1927, gould. spurge nettle, cnidoscolus urens var. stimulosus [=jatropha stimulosa]. average 12 may. earliest, norman, 1929, gould. hymenopappus scabiosaeus [=hymenopappus carolinensis]. average 13 may. earliest, norman, 1929, gould. 4 may hairy bedstraw, galium pilosum. norman, 1929, gould. delphinium carolinianum [=delphinium virescens]. norman, 1929, gould. psoralidium tenuiflorum [=psoralea tenuiflora]. norman, 1929, gould. 5 may galium tricornutum [=galium tricorne]. norman, 1928, gould. spinyleaf catbriar, smilax bona-nox. average 9 may. earliest, norman, 1927, gould. showy evening-primrose, oenothera speciosa. average 6 may. earliest, norman, 1927, gould. largeflower flax, linum rigidum. average 7 may. earliest, norman, 1929, gould. sweet-scented grape, vitus vulpina. average 8 may. earliest, norman, 1927, gould. sensitive-briar, mimosa microphylla [=schrankia uncinata]. average 11 may. earliest, norman, 1927, gould. 6 may longstalk green-briar, smilax pseudo-china. norman, 1928, gould. pediomelum digitatum [=psoralea digitata]. norman, 1928, gould. kentucky coffee-tree, gymnocladus dioica. norman, 1928, gould. 7 may polygala senega. average 15 may. earliest, norman, 1927, gould. virginia ground-cherry, physalis virginiana. norman, 1927, gould. dwarf morning-glory, evolvulus nuttallianus [=evolvulus argenteus]. average 10 may. earliest, norman, 1927, gould. 16 oklahoma native plant record volume 15, december 2015 © ben osborn journal compilation © 2015 oklahoma native plant society 8 may bracted plantain, plantago aristata. norman, 1928, gould. honey locust, gledtisia triacanthos. norman, 1928, gould. rib-grass, plantago lanceolata. norman, 1928, gould. 9 may western catalpa, catalpa speciosa. norman, 1929, gould. catalpa, catalpa bignonioides [=catalpa catalpa]. norman, 1927, gould. american vetch, vicia americana. norman, 1927, gould.** american elder, sambucus nigra ssp. canadensis [=sambucus canadensis]. average 13 may. earliest, norman, 1929, gould. white sweet-clover, melilotus albus. average 14 may. earliest, norman, 1927, gould. roughleaf dogwood, cornus asperifolia. average 16 may. earliest, norman, 1927, gould. 12 may potentilla norvegica [=potentilla monspeliensis]. norman, 1927, gould. purple lemon-mint, monarda citriodora [=monarda dispersa]. average 23 may. earliest, norman, 1927, gould. 13 may queen’s delight, stillingia sylvatica. norman, 1928, gould. rabbit tobacco, diaperia prolifera [=evax prolifera]. norman, 1928, gould. canada moonseed, menispermum candense. norman, 1928, gould. bristly greenbriar, smilax tamnoides [=smilax hispida]. norman, 1928, gould. 14 may low ground-cherry, physalis pumila. norman, 1928, gould. low hairy ground-cherry, physalis pubescens. norman, 1928, gould. largeflower tickseed, coreopsis grandiflora. oklahoma city, 1933*, osborn. pediomelum cuspidatum [=psoralea cuspidata]. average 17 may. earliest, norman, 1927, gould. bank-bur, krameria lanceolata. average 21 may. earliest, norman, 1927, gould. 15 may dogbane, apocynum cannabinum. norman, 1929, gould. showy gaillardia, gaillardia puchella. norman, 1928, gould. green dragon, arisaema dracontium. average 17 may. earliest, norman, 1929, gould. 16 may dwarf verbena, glandularia pumila [=verbena pumila]. norman, 1928, gould. 17 may ground ivy, nepeta cataria. norman, 1928, gould. purple cone-flower, echinacea purpurea [=brauneria purpurea]. average 21 may. earliest, norman, 1928, gould. field bindweed, convolvulus arvensis. norman, 1928, gould. oklahoma native plant record 17 volume 15, december 2015 © ben osborn journal compilation © 2015 oklahoma native plant society 19 may decumbent milkweed, asclepias asperula [=asclepiodora decumbens]. norman, 1928, gould. tumble mustard, sisymbrium altissimum. norman, 1929, gould. engelmannia peristenia [=engelmannia pinnatifida]. norman, 1929, gould. persimmon, disopyros virginiana. norman, 1929, gould. 20 may phlox maculata. norman, 1927, gould. leafy white prickly-poppy, argemone polyanthemos [=argemone intermedia]. average 22 may. earliest, norman, 1927, gould. 21 may smooth solomon’s seal, polygonatum biflorum [=polygonatum commutatum]. norman, 1928, gould. bluntleaf spurge, euphorbia spathulata [=euphorbia obtusata]. norman, 1928, gould. low dwarf mallow, malva neglecta [=malva rotundifolia]. norman, 1928. climbing bittersweet, celastrus scandens. norman, 1928, gould. prairie sunflower, helianthus petiolaris. average 24 may. earliest, norman, 1928. 22 may spermolepis inermis [=spermolepis patens]. norman, 1928, gould. 23 may longhead coneflower, ratibida columnifera [=lepachys columnaris]. average 29 may. earliest, norman, 1927, gould. 24 may wild sweet-pea, tephrosia virginiana. average 25 may. earliest, norman, 1928, gould. 25 may ruellia strepens. average 26 may. earliest, norman, 1929, gould. 26 may intermediate bush-clover, lespedeza simulata. norman, 1927, gould. snow-on-the-mountain, euphorbia marginata. norman, 1928, gould. lead plant, amorpha canescens. norman, 1927, gould. claspingleaf coneflower, rudbeckia amplexicaulis. norman, 1927, gould. smooth sumac, rhus glabra. average 28 may. earliest, norman, 1927, gould. black-eyed susan, rudbeckia hirta. earliest, norman, 1927, gould. 27 may denseflower water-willow, justicia americana [=dianthera americana]. norman, 1929, gould.** 28 may lateflower talinum, phemeranthus calycinus [=talinum calycinum]. norman, 1928, gould. smallflower talinum, phemeranthus parviforus [=talinum parviflora]. norman, 1928, gould buffalo burr, solanum rostratum. norman, 1928, gould. 18 oklahoma native plant record volume 15, december 2015 ben osborn 30 may butterfly weed, asclepias tuberosa. norman, 1929, gould. 31 may sabatia campestris. norman, 1927, gould. yellow gaillardia, gaillardia pinnatifida. norman, 1928, gould. june 1 june gold tickseed, coreopsis tinctoria. norman, 1927, gould. first flowering dates for central oklahoma by mr. ben osborn oklahoma native plant record, volume 16, number 1, december 2016 1 oklahoma native plant r ecord journal of the okla hom a native plant society p. o. box 14274 tulsa, oklahoma 74159-1274 volume 16, december 2016 issn 1536-7738 http://ojs.library.okstate.edu/osu/ managing editor: sheila strawn production editor: paula shryock electronic production editor: sandy graue manuscript editor: mark fishbein technical advisor: erica corbett the purpose of onps is to encourage the study, protection, propagation, appreciation, and use of the native plants of oklahoma. membership in onps is open to any person who supports the aims of the society. onps offers individual, student, family, and life memberships. 2016 officers and board members president: joe roberts vice-president: sheila strawn secretary: sandy graue treasurer: mary korthase historian: adam ryburn past president: adam ryburn board members: mike dunn pearl garrison elaine lynch jay pruett bruce smith sara souza chapter chairs: central: patrick bell cross timbers: vacant northeast: connie murray southwest: helen riley publicity/merchandise chair: alicia nelson conservation chair: chadwick cox tulsa garden club liaison: sue amstutz awards chair: sue amstutz membership database: tina julich photo contest chair: lynn michael mailings/printings chair: karen haworth gaillardia editor: marilyn stewart website manager: adam ryburn http://www.oknativeplants.org cover photo: tradescantia ohiensis raf. (spiderwort) at wichita mountains wildlife refuge by lisa roberts, medicine park, ok articles (c) the authors journal compilation (c) oklahoma native plant society except where otherwise noted, this work is licensed under a creative commons attributionnoncommercial-sharealike4.0 international license, https://creativecommons.org/licenses/ by-nc-sa/4.0/, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly attributed, not used for commercial purposes, and, if transformed, the resulting work is redistributed under the same or similar license to this one. https://doi.org/10.22488/okstate.17.100118 http://ojs.library.okstate.edu/osu/ http://www.oknativeplants.org/ 2 oklahoma native plant record volume 16, december 2016 oklahoma native plant record volum e 16 table of contents foreword .................................................................................................................................................... 3 pollination ecology of sabatia campestris nutt. (gentianaceae) .......................................................... 4 dr. constance e. taylor the structure of the gynostegium, breeding system, and pollination ecology of spider milkweed, asclepias viridis walter (apocynaceae). m. s. thesis ........................................ 10 mr. shang-wen liaw a floristic inventory of the university of oklahoma’s kessler atmospheric and ecological field station, mcclain county, oklahoma .............................................................. 45 ms. amy k. buthod and dr. bruce w. hoagland effects of fire severity on habitat recovery in a mixed grass prairie ecosystem ................................................................................................................................... 64 ms. laura e. jardine, dr. adam k. ryburn, and dr. anthony j. stancampiano critic’s choice essay: a conversation with a small beetle. gaillardia, fall 2000 .............................. 79 dr. paul buck editorial policies and procedures ......................................................................................................... 81 five year index to oklahoma native plant record ............................................... inside back cover oklahoma native plant record, journal of the oklahoma native plant society, volume 16, december 2016 title page table of contents foreword journal of the oklahoma native plant society, volume 2, number 1, december 2002 oklahom a n ative plant record journal of the oklahoma native plant society vol. 2 no. 1 december 2002 issn 1536-7738 managing editor, sheila a. strawn technical editor, patricia folley technical advisor, bruce hoagland cd-rom producer, chadwick cox website: http://www.usao.edu/~onps/ the purpose of the onps is to encourage the study, protection, propagation, appreciation and use of the native plants of oklahoma. membership in onps shall be open to any person who supports the aims of the society. onps offers individual, student, family, and life membership. officers and board members president: patricia folley vice-president: chadwick cox conservation chair: open secretary: maurita nations publicity co-chairs: treasurer: mary korthase ruth boyd & betty culpepper board members: marketing chair: lawrence magrath berlin heck photo contest: paul reimer iris mcpherson ann long award chair: paul reimer sue amstutz harriet barclay award chair: james elder constance taylor paul reimer onps service award chair: sue amstutz lawrence magrath newsletter editor: chadwick cox librarian: bonnie winchester northeast chapter chair: james elder website manager: chadwick cox central chapter chair: judy jordan cross-timbers chapter chair: ronald tyrl cover: gaillardia pulchella (indian blanket) historian: lynn allen photo by paul buck, april 25, 1981. winner of the first onps photo contest in 1988. articles (c) the authors journal compilation (c) oklahoma native plant society except where otherwise noted, this work is licensed under a creative commons attributionnoncommercial-sharealike4.0 international license, https://creativecommons.org/licenses/ by-nc-sa/4.0/, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly attributed, not used for commercial purposes, and, if transformed, the resulting work is redistributed under the same or similar license to this one. https://doi.org/10.22488/okstate.17.100009 2 oklahoma native plant record volume 2, number 1, december 2002 oklahoma native plant record volume 2, number 1 table of contents forward ...................................................................................................... 3 ms. patricia folley, onps president vascular plants of the wichita mountains ............................................ 4 dr. paul buck floristic list for comanche county, oklahoma ................................ 22 dr. bruce w. hoagland schoenoplectus hallii and s. saximontanus ................................................... 54 wichita mountains wildlife refuge survey 2000 dr. lawrence k. magrath pontotoc ridge floristic survey: 1999 ................................................. 64 dr. forrest l. johnson ms. patricia folley (ed.) onps editorial ......................................................................................... 82 water, soil, and plant diversity in oklahoma dr. sheila a. strawn journal of the oklahoma native plant society, volume 2, number 1, december 2002 82 oklahoma native plant record volume 2, number 1, december 2002 editorial water, soil, and plant diversity in oklahoma sheila strawn in 1963 john shed and william penfound did a study on legume distribution. they found a general relationship between soils and vegetation types. sand deposits generally became forests while limestone and clay deposits generally became prairie (proc. okla. acad. sci. 44:2-6). but soil-to-plant relationships are not always that simple. tropographic effects such as slope and solar and wind exposure as well as precipitation rates can override the characteristics of the various soils making it difficult to determine specific soil types based on vegetation. in 1964 dr. paul buck wrote "relationships of the woody vegetation of the wichita mountains wildlife refuge to geological formations and soil types (ecology 45:2). he had found that when it rains, soil type and topographic features, together, control the movement of water. granite formations at the refuge erode to make a cobbly soil at the foot of the mountains that drains quickly, making it difficult for some plants to become established. different amounts and patterns of precipitation can also interact with soil type producing gradients of moisture availability for plants with different moisture requirements. some species can live only at certain positions across the moisture gradient. others are more tolerant of variable moisture availability. sugar maple is able to dominate some of the more mesic, cobbly patches near streams, while black-jack oak dominates the less mesic, cobbly upland patches in this newly formed soil. in places like the carolinas bountiful precipitation on limestone soils can actually make soil very poor in nutrients. organic acids from the abundant vegetation react with calcium, sodium, and magnesium ions in the soil the abundant rainfall then leaches them from the soil and washes them into the groundwater below the reach of roots. however, this is not usually the case in oklahoma. oklahoma actually loses more water to evaporation than it gets in precipitation. rivers and groundwater from the rocky mountains make up the difference. here, precipitation is not sufficient to leach all the minerals from our limestone soils. dense stands of forbs and grasses grow in the tallgrass prairie preserve where these mineral-rich formations are not leached by excess precipitation. southwest of the refuge the precipitation rate is so low that much of the water evaporates before it gets very deep into the soil. in those soils, the calcium ions accumulate at the lowest level of water penetration, sometimes, just inches from the surface and form a calcium carbonate. known as "caliche", this layer restricts root growth, so fewer species grow on caliche. gypswn, mostly calcium sulfate, was formed long ago in a similar manner when the seawater evaporated from the plains. in oklahoma mesquite and mixed grass are associated with “gyp soil”. soil and water relations alone do not explain why plants grow where they do in oklahoma. other variables like humus, micro-organisms, nematodes, grazing animals, fire, ice, windstorms, and human influence all work on a smaller scale and are responsible for much of the physical heterogeneity and resultant biodiversity. this small-scale heterogeneity is what actually accounts for our having more species than other continental united states, except california, florida, and texas. ironically, plant diversity in oklahoma has been woefully understudied. with the promise of improved medical treatments based on genetic engineering, we need to know where species are in oklahoma that might be beneficial. we are, therefore, obliged to preserve as much of our diversity as we can. but we will need to save it on the landscape level, because it is our diverse landscape that supports our diverse biota. indeed, researching and preservmg oklahoma's landscape-to-species relationships is worth researching and preserving twice as much elsewhere. onps strawn, s. https://doi.org/10.22488/okstate.17.100015 oklahoma native plant record, volume 17, number 1, december 2017 oklahoma native plant record 37 volume 17, december 2017 erica a. corbett and andrea lashley https://doi.org/10.22488/okstate.18.100005 laboratory studies of allelopathic effects of jun iper us vir gin iana l. on five species of native plants erica a. corbett andrea lashley department of biological sciences southeastern oklahoma state university durant, ok 74701 ecorbett@se.edu keywords: allelopathy, prairie restoration, eastern redcedar, g ermination, encroaching abstract juniperus virginiana l. (eastern redcedar) is known as an encroaching native plant species. it poses particular problems in the great plains, where fire suppression in the 20th century has led to the expansion of its population and the area it affects. there is some evidence that the genus juniperus contains members that are allelopathic; work with western species of juniper have demonstrated negative effects of litter on seedling growth. we established laboratory experiments to test the effect of a water leachate of eastern redcedar litter (100 g litter per liter di water; steeped 8 h) and eastern redcedar litter on the growth and germination of five native herbaceous species. we saw no clear negative effect of leachate or litter; in fact, there is limited evidence that the leachate increased percent germination, and the presence of litter may have led to greater height growth in inland sea-oats. there was no evidence of the litter or leachate acidifying the soil, at least over the short course of the experiment. it is possible the main negative effect of the presence of eastern redcedar on herbaceous species is through light or nutrient competition by mature trees. we are repeating this study in a field setting. introduction juniperus virginiana l. (eastern redcedar), while a native plant in the u.s., has encroached throughout much of the great plains region following fire suppression and extensive land-use changes in the 20th century (van els et al. 2010; linneman et al. 2011). eastern redcedar had invaded 600,000 acres in oklahoma by 1950; that number had increased to 1,400,000 acres in 1985 (engle et al. 1987). it can change a prairie into a closed woodland in less than 100 years (limb et al. 2010) and is considered a weed tree throughout much of its range, including oklahoma. eastern redcedar tends to invade both abandoned land and high-diversity, non-degraded native grassland because of high seed production and rapid seed dispersal (holthuizen and sharik 1984; linneman et al. 2011). juniperus species in general are a common woody invader in many grassland ecosystems (limb et al. 2010; alford et al. 2012). eastern redcedar can also affect soil properties, raising ph and leading to an increase in calcium content (pierce and reich 2010). when junipers come to dominate a site, they greatly reduce the ground cover layer (horman and anderson 2003). the reduction of ground cover is evidenced by the bare patches that often develop below its crown and dripline; there are five possible contributing factors (not mailto:ecorbett@se.edu 38 oklahoma native plant record volume 17, december 2017 erica a. corbett and andrea lashley necessarily mutually exclusive) for this, listed by horman and anderson (2003): 1. the litter (bark, needles, shed branches, cones) of the cedar trees change soil ph and change its properties to prevent germination or growth of seeds. 2. the shade cast by the crown of the tree is deep enough to hinder survival of herbaceous plants and prevents germination. 3. the tree competes strongly for water with other species. 4. the depth of the litter smothers or prevents water from reaching herbaceous plants and seedlings. 5. the tree itself (roots or litter) is allelopathic and hinders seedling germination and/or survival. many western juniper species show this sort of reduction: horman and anderson (2003) demonstrated that utah juniper (juniperus osteosperma (torr.) little) litter did seem to have a negative effect on seedling growth and survival but more from the standpoint of drying than of allelopathy (i.e., the litter prevented the seeds from properly imbibing water in order to germinate). schott and pieper (1985) found that shading by one-seeded juniper (juniperus monosperma (engelm.) sarg.) in new mexico had a major effect on grass growth, and that litter and allelopathy had secondary effects. ashe juniper (juniperus ashei j. buchholz) also has a zone of reduced understory growth beneath its dripline, and furthermore, growth of vegetation may be stunted in areas where junipers were present but had been removed (yager and smeins 1999). however, engle et al. (1987) observed that the zonation noted around the base of western juniper species was not as clear around eastern redcedar. they examined the effect of eastern redcedar on herbage standing crop in tallgrass prairie. their measurements were taken 1 m and 3 m beyond the dripline of trees. they demonstrated an effect of proximity to the tree; biomass of herbaceous vegetation was reduced close to the dripline of the cedar. they concluded this was a result of shading or possibly water competition rather than allelopathy. also, in general, the effect on herbaceous vegetation under eastern redcedar in oklahoma was less than the effect under other juniperus species in the arid southwest (engle et al. 1987). alford et al. (2012) demonstrated that removal of eastern redcedar from grassland areas in oklahoma increased herbaceous plant species diversity and biomass, likely because of reduced competition for light. the effect was stronger the more heavily-invaded the area had been. van els et al. (2010) proposed possible changes j. virginiana could cause in grassland: increased litter depth, increased soil ph, changes in soil n and c balance, and decreased light availability. apparently redcedars increase, rather than decrease, soil ph, unlike the litter of some other conifers. van els et al. (2010) also proposed that the effects of redcedar encroachment would be different on prairie than in forest. smith and stubbendieck (1990) determined that in tallgrass prairie, grass biomass was reduced underneath redcedar canopy, and that water content under the tree canopy was reduced. they concluded competition for soil water was the most important factor. linnemann et al. (2011) demonstrated in a field experiment that removal of redcedar litter and trees increased herbaceous cover and species diversity, with removal of the trees having a bigger effect on forb and prairie grass cover. in a laboratory experiment, stipe and bragg (1989) demonstrated that only one prairie species among those studied (coreopsis palmata nutt.) showed a statistically significant decrease in germination when grown in soil from underneath redcedar. however, the seeds were watered with tapwater, so there was no further influence of litter after its removal. our objectives for this study were to determine to what extent eastern redcedar oklahoma native plant record 39 volume 17, december 2017 erica a. corbett and andrea lashley affected selected prairie species. in particular, we were interested in the possible effects of remaining eastern redcedar litter in areas where prairie restorations might be attempted. we examined litter and leachate; future studies will examine the effects of dripline, proximity, and shading on plant growth. in this study, we focus on determining whether leachate from litter reduces prairie plant germination and growth. we hypothesize that there will be reductions in germination and/or growth of the selected plant species (two prairie grasses, two prairie forbs, one woodland grass) when watered with leachate from redcedar leaves and that the presence of litter will make that effect stronger. methods and materials in early october 2016, approximately 30 pounds of juniperus virginiana branches were collected from pastureland just outside of durant, oklahoma. these bags were transported to southeastern oklahoma state university and maintained at roughly 20oc. on october 8, leachate was prepared: redcedar needles were picked clean of branches and woody material. one hundred grams of this “cedar litter” were steeped for 8 h in 1000 ml deionized water (similar to the 10% leachate as prepared by horman and anderson 2003). at the end of the time period, the leachate was filtered through a 50 mesh sieve, bottled, and frozen at 0oc until needed. natural soil was collected from a disturbed grassland on lake texoma, bryan county, oklahoma (33.999687˚n, -96.587678˚w). the site was dominated by several species of panic-grasses (dichanthelium), crowngrasses (paspalum), and with forbs, lespedeza cuneata (dum. cours.) g. don (sericea lespedeza) and rudbeckia hirta l. (black-eyed susan). there was a stand of ulmus alata michx. (winged elm) within 15 m of where the soil was collected. the soil is in the boxville fine sandy loam series (usda 1978). approximately the top 20 cm of soil was collected and transported to southeastern oklahoma state university. cone-tainers size sc-10 (21 cm deep and roughly 4 cm in diameter) (ray leach company) were set up in racks. there was a total of four treatments and five species, and nine replicates of each species by treatment combination, resulting in 180 cone-tainers. each cone-tainer was filled with soil, and a standard-sized marble (cardinal industries’ “marble bonanza”) was placed in the bottom of each conetainer to allow for drainage and prevent excessive leakage of soil. each cone-tainer was planted with 2–3 seeds of one of the focal species. as the seeds germinated, excess individuals were removed to leave one plant per cone-tainer. the focal species were: little bluestem (schizachyrium scoparium (michx.) nash), indian grass (sorghastrum nutans (l.) nash), inland sea-oats (chasmanthium latifolium (michx.) h.o. yates), partridge-pea (chamaecrista nictitans (l.) moench), and black-eyed susan. seeds were purchased from native american seeds in junction, texas. the racks of cone-tainers were set under a fluorescent-light fixture having six fluorescent tubes (ge plant and aquarium 40 watt t12 warm linear fluorescent tubes). these light fixtures were suspended from pvc frames so that the tubes were approximately 20 cm above the tops of the cone-tainers. the lights were set on a timer to give a 14 h daylength. two identical fluorescent fixtures were used because there was not enough room for both racks under a single fixture. four treatments were applied: control (10 ml of deionized water once a week, no litter), leachate only (10 ml of eastern redcedar leachate once a week), litter only (~ 3 cm of eastern redcedar litter on top of soil surface), and litter plus leachate (~3 cm of litter plus 10 ml leachate once a week). between treatments, all cone-tainers were watered every second or third day (as 40 oklahoma native plant record volume 17, december 2017 erica a. corbett and andrea lashley needed, from examining the soil surface). precise amounts of water given to each of the 180 cone-tainers were not measured in these waterings; however, it was approximately the same quantity to each. we monitored time-to-germination and percent germination of each species. at the end of the first run of the experiment (5 december 2016), we measured the height of each germinated grass individual or leaflength of the largest leaf for the forbs. we measured leaf growth because individual plants were too small to be weighed. in january 2017, we began a second run of the experiment with a few changes. because of high dormancy of inland seaoats, the seeds were subjected to 30 d of cold-wet stratification (between paper towels in a 5o c refrigerator) before planting. additionally, partridge-pea seeds were scarified with 100 grit sandpaper before planting. fresh soil and cedar branches were collected from the same location as for the first experiment. in contrast to the first experiment, the soil was sieved prior to planting to give a more homogeneous substrate. seeds were planted on 25 january 2017. other procedures were the same as for the first experiment. the second experiment concluded on 11 april 2017, and the height of each grass and length of the longest leaf of each forb individual measured. the soil from replicates of each species and treatment combination was pooled, and a 20 g sample was analyzed for ph. we conducted a third study in spring 2017 examining the effects of cedar leachate on seed germination. the same five species were included in this study. two treatments were applied: control (distilled water only) and leachate. ten seeds from each of the five species were placed into separate petridishes between layers of filter paper. for each species, treatments were applied to six petri-dishes: three received distilled water and three received leachate. there were three replicates (of ten seeds each) for each species and each treatment. petri dishes were maintained at room temperature and were watered as needed: control dishes were watered with deionized water, and treatment dishes with leachate. both the distilled water and eastern redcedar leachate were stored at approximately 4o c and were applied to the seeds at this temperature. germination percentage was recorded after 30 d, and shoot extension and radicle length were measured. inland sea-oats was dropped from further analysis because of low germination. analysis of the germination data was challenging because germination was typically low (table 1). we used likelihoodratio chi-square analysis (g-test) based on number of seeds germinating. we tested the fall 2016 and spring 2017 experiments separately. for this test and all other statistical tests, we used an alpha level of 0.05 for significance. we also grouped all species for each treatment separately for the two experiments for analysis of overall effects across species. for the combined data, we tested the data for normality using a shapiro-wilk test. where data were normal, we performed a one-way analysis of variance (anova) on germination by treatment. when data were not normal, we performed kruskal-wallis analysis. we originally planned to use two-way anova to test for treatment by species interactions in germination and growth. however, because germination was low overall and the species differed in growth form (grasses vs. forbs, one forb had compound leaves and the other had simple leaves), we chose instead to do a series of one-way anovas. the soil ph data (2017 experiment only) were analyzed with a one-way anova following a test for normality. for the petriplate experiment, we used chi-square and likelihood-ratio chi-square tests of germination percentages and mann-whitney u-tests on growth of individual species. statistical analyses, including chi-square oklahoma native plant record 41 volume 17, december 2017 erica a. corbett and andrea lashley table 1 germination results for prairie plants in cone-tainers subjected to leachate and litter treatments in 2016 and 2017. control = no litter, no leachate. leachate = leachate only. litter = litter only. both = litter plus leachate. lb = little bluestem, ig = indian grass, so = inland sea-oats, pp = partridge-pea, and bs = black-eyed susan. for all treatments, n = 9. experiment 1: fall 2016 treatment species percent germination control lb 33.3 control ig 55.6 control so 22.2 control pp 0.0 control bs 33.3 leachate lb 22.2 leachate ig 55.6 leachate so 22.2 leachate pp 0.0 leachate bs 77.8 litter lb 66.7 litter ig 66.7 litter so 11.1 litter pp 11.1 litter bs 22.2 both lb 77.8 both ig 77.8 both so 33.3 both pp 0.0 both bs 66.7 42 oklahoma native plant record volume 17, december 2017 erica a. corbett and andrea lashley table 1 continued experiment 2: spring 2017 treatment species percent germination control lb 11.1 control ig 66.7 control so 77.8 control pp 11.1 control bs 44.4 leachate lb 55.6 leachate ig 77.8 leachate so 88.9 leachate pp 11.1 leachate bs 88.9 litter lb 22.2 litter ig 33.3 litter so 100.0 litter pp 0.0 litter bs 33.3 both lb 22.2 both ig 55.6 both so 100.0 both pp 0.0 both bs 88.9 oklahoma native plant record 43 volume 17, december 2017 erica a. corbett and andrea lashley table 2 growth of four species under four treatments. growth data is plant height (grasses) or length of longest leaf (forbs) at the end of the growing period. control = no litter, no leachate. leachate = leachate only. litter = litter only. both = litter plus leachate. n = sample size. partridge-pea is not included as germination was too low (zero for three treatments). fall 2016 treatment growth (cm ± se) n little bluestem control 10.6 + 2.6 3 leachate 8.0 + 0.6 2 litter 13.8 + 2. 5 6 both 10.2 + 0.4 7 indian grass control 12.7 + 2.0 4 leachate 13.8 + 1. 7 5 litter 21.3 + 4.5 6 both 23.6 + 1.3 7 inland sea-oats control 13.9 + 6.4 2 leachate 9.5 + 1.0 2 litter 6.0 + 0.0 1 both 10.6 + 2.0 3 black-eyed susan control 2.8 + 0.7 3 leachate 1.8 + 0.2 7 litter 2.6 + 0.4 2 both 3.5 + 0.1 6 44 oklahoma native plant record volume 17, december 2017 erica a. corbett and andrea lashley table 2 continued spring 2017 treatment growth (cm ± se) n little bluestem control only one observation, no mean 1 leachate 13.3 + 1.9 5 litter 17.7 + 4.0 3 both 16.5 + 0.1 2 indian grass control 22.7 + 3.2 6 leachate 18.8 + 2.6 7 litter 24.5 + 3.9 3 both 20.8 + 1.7 5 inland sea-oats control 9.2 + 0.9 7 leachate 7.7 + 0.8 8 litter 11.3 + 0.7 9 both 12.0 + 0.8 9 black-eyed susan control 3.3 + 0.3 2* leachate 2.4 + 0.4 5** litter 2.4 + 0.6 3 both 3.0 + 0.3 8 *one individual germinated (total germinating = 3) but died before maturity **three individuals germinated (total germinating = 8) but died before maturity oklahoma native plant record 45 volume 17, december 2017 erica a. corbett and andrea lashley figure 1 average height of indian grass for four eastern redcedar treatments in fall 2016. sample sizes: control, n = 4; leachate, n= 5; litter, n=6; litter plus leachate, n=7. treatment significantly affected height, but no pairwise comparisons were significantly different in post hoc tests. 46 oklahoma native plant record volume 17, december 2017 erica a. corbett and andrea lashley figure 2 average leaf length of black-eyed susan by treatment for fall 2016 data. control, n = 3; leachate treatment, n= 7; litter treatment, n=29; litter plus leachate treatment, n=6. oklahoma native plant record 47 volume 17, december 2017 erica a. corbett and andrea lashley tests, anovas, and kruskal-wallis tests, were performed using spss (ibm 2011). results fall 2016 g ermination a nd g rowth (cone-tainer experiment 1) germination percentages are shown in table 1. species-treatment combinations did not differ in germination percentages (g = 9.161, df = 12, p=0.689). to test treatment effects across species, we grouped the data for all species within each treatment. the data were not normal (shapiro-wilk test, w(20) = 0.899, p = 0.040). there was no significant difference in germination among treatments for all species grouped together (kruskalwallis test: χ2 = 5.077, df = 3, p=0.166). growth measures (i.e., the longest leaf height on the grasses or the longest leaf on the forbs) are presented in table 2. partridge-pea had too few germinating individuals to analyze. we analyzed each species separately because of differences in growth form. little bluestem data were not normal (w(18)=0.853, p=0.009) and showed no significant effect of treatment on growth (g= 4.879, df =3, p = 0.181). normality of indian grass data could not be rejected (w(22)= 0.981, p =0.927), and indian grass showed a significant effect of treatment (anova, f(3, 18)=3.598, p=0.034). there is a weak trend for litter application to result in greater height (figure 1). however, this was not statistically significant in post-hoc tests. normality of inland sea-oats data could not be rejected (w(8)=0.878, p=0.180), and inland sea-oats did not show a significant effect of treatment (anova, f (3, 4) =0.557, p=0.671), perhaps because sample sizes were low due to poor germination. normality of black-eyed susan data could not be rejected (w(18)=0.962, p = 0.650), and black-eyed susan showed a significant effect of treatment (anova, f(3, 4)=7.63, p=0.003), but post-hoc tests did not find pairwise differences between treatments (figure 2) spring 2017 g ermination and g rowth (cone-tainer experiment 2) for the spring germination results (see table 1), again there was no significant relationship between treatment, species, and germination (g = 6.190, df =12, p = 0.906). after grouping species within treatment, differences among were not significant (χ2= 1.702, df=3, p=0.637). for spring growth measures (table 2), normality of little bluestem data could not be rejected (w(11) =0.915, p = 0.277), and little bluestem showed no significant effect of treatment on growth (anova, f(3, 7)=1.478, p=0.301). normality of indian grass data could not be rejected (w(21) = 0.979, p=0.908), and indian grass did not show a significant effect of treatment (anova, f(3, 7)= 0.687, p=0.572). normality of inland sea-oats data could not be rejected (w(34)= 0.972, p=0.518), and inland sea-oats growth was affected significantly by treatment (anova, f(3, 30) = 6.211, p=0.002), with individuals in treatment 4 (litter plus leachate) growing larger (student-newmankeuls test) than individuals in treatment 2 (leachate alone; figure 3). normality of black-eyed susan data could not be rejected (w(18) = 0.945, p=0.346), and growth of black-eyed susan did not show a significant effect of treatment (anova, f(3, 14)=0.811, p=0.509). normality of treatment ph (table 3) could not be rejected (w(20)= 0.934, p=0.188). treatments did not affect soil ph (anova, f(3, 16)=1.355, p=0.262). 48 oklahoma native plant record volume 17, december 2017 erica a. corbett and andrea lashley figure 3 average height of inland sea-oats by treatment for spring 2017 data. control, n = 8; leachate treatment, n= 8; litter treatment, n=9; litter plus leachate treatment, n=9. oklahoma native plant record 49 volume 17, december 2017 erica a. corbett and andrea lashley table 3 soil ph for five species under five treatments in spring 2017 and means across species within treatment. control = no litter, no leachate. leachate = leachate only. litter = litter only. both = litter plus leachate. lb = little bluestem, ig = indian grass, so = inland sea-oats, pp = partridge-pea, and bs = black-eyed susan. treatment species ph control lb 6.4 control ig 6.2 control so 6.3 control pp 6.1 control bs 5.7 mean (± se) 6.1 + 0.13 leachate lb 6.0 leachate ig 6.0 leachate so 6.0 leachate pp 6.1 leachate bs 6.2 mean (± se) 6.1 + 0.04 litter lb 6.2 litter ig 5.9 litter so 5.8 litter pp 6.1 litter bs 5.8 mean (± se) 6.0 + 0.08 both lb 6.0 both ig 6.2 both so 6.2 both pp 6.2 both bs 6.2 mean (± se) 6.2 + 0.04 50 oklahoma native plant record volume 17, december 2017 erica a. corbett and andrea lashley germination in petri-dishes species and treatment combinations differed in germination percentage (table 4, chi-square = 42.897, df=4, p < 0.001). this result was likely influenced by differences in germination of indian grass (ca 27% for control vs. ca 37% for added leachate) and little bluestem (ca 10% vs. 30% ). in both cases, the presence of the cedar leachate seemed to increase germination. for little bluestem, there was no treatment effect of leachate on radicle length (mann-whitney test: u=5.0, n=12, p=0.145) or shoot length (u=7.5, n=12, p=0.282). for indian grass, there was a decrease in radicle length in response to leachate (u=11.50, n=19, p=0.005). however, there was no effect of treatment on shoot length (u=41.5. n=19, p=0.840). black-eyed susan showed no effect of treatment on radicle length (u=211.5, n=42, p=0.828), but had a marginally significant effect of treatment on shoot length (u=145.5, n=42, p=0.052). table 4 percent germination in petri dishes under two treatments. see text for chi-square tests of treatment by species. treatment species average percent germination control little bluestem 10.0 ± 0.0 control indian grass 26.7 ± 8.8 control inland sea-oats 3.3 ± 3.3 control partridge-pea 3.3 ±3.3 control black-eyed susan 66.7 ± 8.8 leachate little bluestem 30.0 ± 0.0 leachate indian grass 36.7 ± 6.7 leachate inland sea-oats 0.0 ± 0.0 leachate partridge-pea 10.0 ± 5.8 leachate black-eyed susan 73.3 ± 6.7 discussion in general, there were few effects of the application of redcedar leachate and/or litter. treatment with leachate and/or litter did not hamper germination, and there was no clear effect of treatment on growth in the cone-tainer experiments. there were weak trends suggesting in some cases that application of litter plus leachate might increase growth, but trends were weak. a negative effect of redcedar was not demonstrated over the months-long course of these experiments (56 d for fall 2016 and 77 d for spring 2017). there is some evidence from the petri-dish germination oklahoma native plant record 51 volume 17, december 2017 erica a. corbett and andrea lashley experiment that the redcedar leachate may stimulate germination rate (at least in indian grass and little bluestem) but reduce growth of seedling root in indian grass and seedling shoot in black-eyed susan, but it is possible that the growth reductions are a short-lived effect. anecdotally, we observed that the treatment receiving the cedar leachate required more frequent watering in order to maintain the same moist environment as those receiving the water only. further study of the rate of evaporation of the water and cedar leachate may provide additional insight regarding whether water resources may be affected by an eastern redcedar population. an additional anecdotal observation was that partridge-pea was susceptible to mold growth that was possibly inhibited by the redcedar leachate. a study of the allelopathic effects of cedar leachate on mold growth may provide additional data to test this observation. we speculate that in soils with long exposure to redcedar litter (years rather than months), perhaps effects would be greater, or there might be a negative effect. we are considering future experiments planting seeds or seedlings within the driplines of existing cedars and comparing their growth to the growth of individuals away from the dripline. it is also possible that juniperus virginiana may lack the same allelopathic compounds found in the western junipers (e.g., juniperus monosperma). it would be informative to repeat the experiment on a larger scale, comparing the effects of eastern redcedar, one-seeded juniper, and utah juniper. it is also possible that a harmful effect of redcedar is produced by shade; future research could include planting individuals at varying distances from the trunk of the tree to determine whether shading has an effect, or if the dripline of the tree has an effect. in future work, we plan to examine the ph of soils within the dripline and 3 and 5 m beyond the dripline of redcedars. smith and stubbendeick (1990) suggested that the effects of red-cedar on herbaceous species in the field is mainly mediated through light and water competition from the mature trees; they demonstrated reduction in biomass for prairie species grown inside the dripline of cedar trees. it is also possible that water competition is the mechanism of limitation; many of the studies showing reduced herbaceous growth under junipers (e.g., schott and pieper 1985; yager and smeins 1999) were conducted in climates drier than that of south-central oklahoma and certainly under more water-limited conditions than our lab experiment. however, in northeastern oklahoma, engle et al. (1987) did demonstrate reduction in herbaceous standing crop within the dripline of redcedar trees. van els et al. (2010) demonstrated reduced species richness in forest understory under juniperus trees in oklahoma; they attributed these changes to increased litter depth but did not separate what chemical or physical characteristic of the litter served as a barrier to plant germination and growth. finally, we plan to conduct a field experiment to determine whether the greater stress of growth outdoors or over a longer time is sufficient to show effects. acknowledgments we thank the oklahoma native plant society for providing a small grant to e.a.c. to purchase seeds and equipment for this study. we thank tom and charlene tucker for allowing us to cut redcedar branches on their pasture land. we thank matt spears for his assistance in setting up the second run of this experiment. we also thank two anonymous reviewers for comments that led to improvement of this paper. 52 oklahoma native plant record volume 17, december 2017 erica a. corbett and andrea lashley literature cited alford, a.l., e.c. hellgren, r. limb, and d.m. engle. 2012. experimental tree removal in tallgrass prairie: variable responses of flora and fauna along a woody cover gradient. ecological applications 22:947–958. engle, d.m., j.f. stritzke, and p.l. claypool. 1987. herbage standing crop around eastern redcedar trees. journal of range management 48:100–107. holthuizen, a.m.a. and t.l. sharik. 1984. seed longevity and mechanism of regeneration of eastern red cedar (juniperus virginiana l.). journal of the torrey botanical society 111:153–158. horman, c.s. and v.j. anderson. 2003. understory response to utah juniper litter. journal of range management 56:68– 71. ibm corp. released 2011. ibm spss statistics for windows, version 20.0. armonk, ny: ibm corp. limb, r.f., d.m. engle, a.l. alford, and e.c. hellgren. 2010. tallgrass prairie plant community dynamics along a canopy cover gradient of eastern redcedar (juniperus virginiana l.). rangeland ecology and management 63:638– 644. linneman, j.s., m.s. allen, and m.w. palmer. 2011. the effects of removal of juniperus virginiana l. trees and litter from a central oklahoma grassland. oklahoma native plant record 11:43–60. pierce, a.m and p.b. reich. 2010. the effects of eastern red cedar (juniperus virginianus) invasion and removal on a dry bluff prairie ecosystem. biological invasions 12:241–252. schott, m.r. and r.d. pieper. 1985. influence of canopy characteristics of one-seed juniper on understory grasses. journal of range management 38:328–331. smith, s.d. and j. stubbendieck. 1990. production of tall-grass prairie herbs below eastern redcedar. prairie naturalist 22:12–18. stipe, d.j. and t.b. bragg. 1989. effect of eastern red cedar on seedling establishment of prairie plants. proceedings of the eleventh north american prairie conference. pp. 101–102. [usda, scs] united states department of agriculture, soil conservation service. 1978. soil survey of bryan county, oklahoma. in cooperation with the oklahoma agricultural experiment station. van els, p., r.e. will, m.w. palmer, and k.r. hickman. 2010. changes in forest understory associated with juniperus encroachment in oklahoma, usa. applied vegetation science 12:346–368. yager, l.y. and f.e. smeins. 1999. ashe juniper canopy and litter effects on understory vegetation in a juniper-oak savannah. southwestern naturalist 44:6–16. laboratory studies of allelopathic effects of juniperus virginiana l. on five species of native plants by erica a. corbett and andrea lashley oklahoma native plant record, volume 15, number 1, december 2015 oklahoma native plant record 49 volume 15, december 2015 amy k. buthod and bruce w. hoagland https://doi.org/10.22488/okstate.17.100114 contributions to the flora of cimarron county and the black mesa area amy k. buthod bruce w. hoagland oklahoma biological survey oklahoma biological survey university of oklahoma department of geography and norman, ok 73019 environmental sustainablity amybuthod@ou.edu university of oklahoma norman, ok 73019 keywords: flora, cimarron county, black mesa, vascular plants, rare plants abstract this paper reports the results of recent collection activities in cimarron county, including the black mesa area, in the state of oklahoma. a total of 331 taxa in 60 families were collected. two-hundred and six genera, 279 species and 52 infraspecific taxa were identified. the largest families were the poaceae with 72 taxa and the asteraceae with 63. thirty-six exotic taxa were collected (10.9 % of the flora), including two species new to oklahoma: scorzonera laciniata and ranunculus testiculatus. forty-six taxa tracked by the oklahoma natural heritage inventory were found. introduction cimarron county has long been recognized as a botanically significant region in oklahoma. a total of 95 vascular plants tracked by the oklahoma natural heritage inventory (onhi) occur in the county (oklahoma natural heritage inventory 2013). included among these is asclepias uncialis greene, which, prior to 1996, was listed as a likely candidate for federal listing as threatened or endangered by the u.s. fish and wildlife service (united states department of the interior 1993). before this survey, nineteen of the tracked taxa had an onhi ranking of sh, meaning that reports of occurrences are older than twenty years (oklahoma natural heritage inventory 2013; natureserve 2015). the number of taxa in cimarron county that are rare at the state level is due in part to the presence of black mesa, an extension of the mesa de maya, which extends for 72 km from east of raton, new mexico, though colorado and into northwestern cimarron county. the eastern-most extension of the rocky mountain foothill vegetation is present in the area; rogers (1953) found it to be “an excellent example of the intergradation of the flora of the great plains with that of the rocky mountain foothills”. our intent for this work was to relocate the rare taxa, update their onhi ranks, and, hopefully, expand our knowledge of the area’s current flora. the earliest botanical collections from the black mesa region were made in 1820 by edwin james, botanist for major stephen long’s expedition to the rocky mountains. eighty-four years later, per axel rydberg, author of flora of colorado (1906) and flora of the rocky mountains and adjacent plains (1917), botanized in the area. the first thorough botanical inventory of the mesa de maya was completed by rogers (1953). from 1947 and 1949, he collected along the mesa in colorado, new mexico and oklahoma, as well as from some of the secondary mesas in the area (rogers 1953). according mailto:amybuthod@ou.edu 50 oklahoma native plant record volume 15, december 2015 amy k. buthod and bruce w. hoagland to a list published in 1953, rogers collected 267 taxa from 51 families in oklahoma, but in a later work (1954) he notes that “approximately five-hundred were found, or could be found”. u. t. waterfall collected at black mesa and in cimarron county within the same time period, adding approximately 30 taxa to the state’s flora (waterfall 1949, 1950a, b). james k. mcpherson completed an inventory with the sole focus of black mesa in the early 1990s, reporting 236 taxa from 58 families (2003a, b). his collecting activities were confined to the areas of the mesa on the property belonging to the state of oklahoma (township 6n, range 1e, sections 28–33 and township 5n, range 1e, section 6). patricia folley (2003) supplemented the mcpherson list with collections made from 1994 through 2003. folley collected over a wider area than mcpherson, surveying the state park around lake etling, the roadsides leading to the park and mesa, and some private lands, including tesequite canyon (folley 2003). she found an additional 49 taxa from 25 families. other botanists have contributed to the knowledge of the black mesa/cimarron county flora over the years, including delzie demaree, who worked in the area in the 1930s, george goodman (from the late 1930s through the early 1970s), john and connie taylor (1960s and 1970s), and larry magrath in the 1980s (oklahoma vascular plants database 2015). study site cimarron county falls within the high plains and the cimarron river valley geomorphic provinces (curtis et al. 2008). the high plains province consists of flat uplands over tertiary-era dakota sandstones and is found throughout most of the county (rogers 1953). the cimarron river valley is found in the northeastern part of the county and is distinguished by dissected valleys of mesozoic-era shale and sandstone. the black mesa, the flat, eroded remnant of a tertiary-era lava flow, is located in this area (curtis et al. 2008). the highest point in oklahoma, at 1515 m, is on the mesa. rolling, low hills and canyons surround the mesa. four soil associations occur within cimarron county. travessilla-kim soils are only found in the northeastern corner of the county. they consist of “loam, calcareous, and humus-poor soils on steep slopes” (carter and gregory 2008). dalhart-vona soils are found primarily in the southern half of the county; these are “very deep loamy soils on gentle slopes” (carter and gregory 2008). sherm-ulysses type soils dominate the eastern half of the county. these soils are “very deep, silty and clayey, humus-rich soils on gentle slopes” (carter and gregory 2008). conlen-pastura-plack soils are the least common soil type in the county; they consist of “loamy and calcareous soils on moderately steep slopes” (carter and gregory 2008). potential vegetation types in cimarron county include shortgrass high plains, sandsage grassland, piñon pine/juniper mesa, and bottomland forest (duck and fletcher 1943; hoagland 2008). cimarron county has a dry climate, falling within trewartha’s steppe or semiarid type (1968). average annual precipitation ranges from 38–50 cm, with most falling from may through august. thunderstorms occur in the spring and summer. average temperature is 13–14°c. the average high (in july) is 34°c, and the average low (in january) is -7°c. southsouthwesterly winds are dominant and relative humidity ranges from 29–84%. over 70% of days are sunny (oklahoma climatological survey 2015). methods plants were collected at 100 sites throughout cimarron county (fig. 1; table 1). collection sites were chosen based on location information from the oklahoma natural heritage inventory database and oklahoma native plant record 51 volume 15, december 2015 amy k. buthod and bruce w. hoagland figure 1 cimarron county, oklahoma. dots indicate collection sites. shaded areas indicate black mesa nature preserve lands. map by todd fagin, oklahoma biological survey. the oklahoma vascular plants database. additional collections were also made opportunistically. coordinates of each site were collected using a garmin gpsmap 76cx unit. sites were located between latitudes n36.98989 and n36.62313 and longitudes w102.67913 and w102.68063. elevations ranged from 1118 m to 1513 m. field work began in march of 2013, with subsequent monthly trips until september. an additional trip was made in may of 2014. one example of each taxon encountered was collected and processed according to standard herbarium protocols. specimens were deposited at the robert bebb herbarium (okl) at the university of oklahoma. manuals used to identify plants included great plains flora association (1986), tyrl et al. (2010) and allred and ivey (2012); the collections of the robert bebb herbarium were also used to verify identifications. taxonomy follows the integrated taxonomic information system (2015). duration and nativity to oklahoma were determined using the plants database (usda-nrsc 2015); if the information from plants was ambiguous, taylor and taylor (1991) was consulted. vegetation classifications were assigned based on hoagland (2000). results and discussion three-hundred and thirty-one taxa in 60 families were collected in this study (appendix a). two-hundred and six genera, 279 species, and 52 infraspecific taxa were identified. two-hundred thirty-one taxa were perennials; there were 96 annuals and four biennials. thirty-six taxa were nonnative to oklahoma, including two species new to the state (scorzonera laciniata in the asteraceae and ranunculus testiculatus in the 52 oklahoma native plant record volume 15, december 2015 amy k. buthod and bruce w. hoagland ranunculaceae); non-native taxa accounted for 10.9% of the total flora. the poaceae had the greatest number of exotic taxa with 11; the brassicaceae had five. the largest families were the poaceae with 72 taxa and the asteraceae with 63. forty-six taxa tracked by the oklahoma natural heritage inventory were found (table 2). asclepias uncialis, the former candidate for federal listing, was not located. vegetation classes encountered in this study included the artemisia filifolia/sporobolus cryptandrus-schizachyrium scoparium shrubland association. it is found on sandy soils and stabilized dunes in the northwestern and central portions of the study site. associated taxa included andropogon gerardii ssp. hallii, abronia fragrans, and eriogonum annuum (duck and fletcher 1943; hoagland 2000). two intergrading variations of shortgrass prairie were noted. the bouteloua curtipendula-b. gracilis-b. dactyloides herbaceous association is found on rocky slopes and well-drained soils in the southern part of the study area (duck and fletcher 1943; hoagland 2000). plants found here included muhlenbergia torreyi, ratibida columnifera, and sphaeraclea coccinea. the bouteloua gracilishilaria jamesii herbaceous association is found in northwestern cimarron county on slopes and uplands (hoagland 2000). plants found in this type included cylindropuntia imbricata, melampodium leucanthum, and zinnia grandiflora. the bouteloua gracilis-hilaria jamesii herbaceous association intergrades with the fourth vegetation type, the juniperus monosperma woodland alliance. this alliance includes the juniperus monosperma/bouteloua curtipendula woodland association and the juniperus monosperma-pinus edulis/bouteloua curtipendula woodland association and is found in northwestern cimarron county. plants from this type included bouteloua gracilis, cercocarpus montanus, and prunus virginiana (hoagland 2000). herbaceous wetland vegetation was found at only a few sites, including those with seeps, lakes, and intermittently flowing streams and rivers. plants found in this vegetation type included polypogon monspeliensis, populus deltoides, salix exigua, and tamarix chinensis. vegetation of disturbed areas includes taxa found around lawns, stock tanks, campgrounds, parking lots, and gravel pits. plants in this vegetation type included conyza canadensis, descurainia sophia, kochia scoparia, and malva neglecta. table 1 collection sites in cimarron county latitude longitude township, range, and section 36.623130 -102.68063 sec. 24-t2n-r3e 36.690420 -102.95001 sec. 33-t3n-r1e 36.698390 -102.9484 sec. 28-t3n-r1e 36.719380 -102.89576 sec. 13-t3n-r1e 36.719660 -103.00208 sec. 18-t3n-r1e 36.722180 -102.877 sec. 18-t3n-r2e oklahoma native plant record 53 volume 15, december 2015 amy k. buthod and bruce w. hoagland 36.733790 -102.7183 sec. 15-t3n-r3e 36.733800 -102.76698 sec. 17-t3n-r3e 36.733920 -102.74949 sec. 16-t3n-r3e 36.739900 -102.51231 sec. 10-t3n-r5e 36.741100 -102.51344 sec. 10-t3n-r5e 36.753940 -102.96656 sec. 4-t3n-r1e 36.756350 -102.96655 sec. 4-t3n-r1e 36.765380 -102.96653 sec. 32-t4n-r1e 36.772640 -102.96652 sec. 32-t4n-r1e 36.780300 -102.87736 sec. 30-t4n-r2e 36.790710 -102.96668 sec. 29-t4n-r1e 36.804340 -102.97145 sec. 20-t4n-r1e 36.806220 -102.37202 sec. 13-t4n-r6e 36.817480 -102.80509 sec. 13-t4n-r2e 36.829480 -102.87738 sec. 7-t4n-r2e 36.832480 -102.65052 sec. 8-t4n-r4e 36.835430 -102.96116 sec. 9-t4n-r1e 36.836080 -102.88737 sec. 6-t4n-r2e 36.840080 -102.88219 sec. 6-t4n-r2e 36.845780 -102.87656 sec. 5-t4n-r2e 36.846420 -102.88263 sec. 6-t4n-r2e 36.848380 -102.62216 sec. 3-t4n-r4e 36.849240 -102.88435 sec. 6-t4n-r2e 36.850360 -102.87642 sec. 31-t5n-r2e 36.850360 -102.87642 sec. 31-t5n-r2e 54 oklahoma native plant record volume 15, december 2015 amy k. buthod and bruce w. hoagland 36.851790 -102.86967 sec. 32-t5n-r2e 36.853670 -102.87138 sec. 32-t5n-r2e 36.854090 -102.88454 sec. 31-t5n-r2e 36.856980 -102.94078 sec. 34-t5n-r1e 36.859200 -102.38917 sec. 34-t5n-r6e 36.881280 -102.88344 sec. 19-t5n-r2e 36.882050 -102.97772 sec. 20-t5n-r1e 36.883330 -102.97295 sec. 20-t5n-r1e 36.886460 -102.97238 sec. 20-t5n-r1e 36.887550 -102.97424 sec. 20-t5n-r1e 36.889260 -102.96963 sec. 20-t5n-r1e 36.891690 -102.96015 sec. 21-t5n-r1e 36.892660 -102.98643 sec. 19-t5n-r1e 36.893120 -102.82283 sec. 22-t5n-r2e 36.893790 -102.95947 sec. 16-t5n-r1e 36.895370 -102.9677 sec. 17-t5n-r1e 36.895760 -102.98476 sec. 18-t5n-r1e 36.895960 -102.98691 sec. 18-t5n-r1e 36.897520 -102.91134 sec. 13-t5n-r1e 36.897950 -102.96324 sec. 16-t5n-r1e 36.898540 -102.98034 sec. 17-t5n-r1e 36.899190 -102.97931 sec. 17-t5n-r1e 36.899370 -102.8527 sec. 16-t5n-r2e 36.899560 -102.84465 sec. 16-t5n-r2e 36.899830 -102.82454 sec. 15-t5n-r2e oklahoma native plant record 55 volume 15, december 2015 amy k. buthod and bruce w. hoagland 36.900190 -102.96891 sec. 17-t5n-r1e 36.900640 -102.97209 sec. 17-t5n-r1e 36.901170 -102.96404 sec. 16-t5n-r1e 36.901410 -102.95449 sec. 16-t5n-r1e 36.903700 -102.94789 sec. 16-t5n-r1e 36.904800 -102.93303 sec. 15-t5n-r1e 36.907530 -102.44369 sec. 7-t5n-r6e 36.908160 -102.45214 sec. 7-t5n-r6e 36.910440 -102.92143 sec. 11-t5n-r1e 36.912730 -102.82081 sec. 11-t5n-r2e 36.913450 -102.97624 sec. 8-t5n-r1e 36.914270 -102.96875 sec. 8-t5n-r1e 36.919640 -102.4009 sec. 10-t5n-r6e 36.920710 -102.51988 sec. 9-t5n-r5e 36.921370 -102.60638 sec. 10-t5n-r4e 36.921370 -102.60638 sec. 10-t5n-r4e 36.929980 -102.58279 sec. 1-t5n-r4e 36.931820 -102.99784 sec. 6-t5n-r1e 36.934710 -102.9383 sec. 3-t5n-r1e 36.934710 -102.93839 sec. 3-t5n-r1e 36.934850 -102.57666 sec. 1-t5n-r4e 36.936330 -102.55646 sec. 21-t6n-r5e 36.936870 -102.52358 sec. 33-t6n-r5e 36.936880 -102.47233 sec. 36-t6n-r5e 36.937150 -103.0018 sec. 6-t5n-r1e 56 oklahoma native plant record volume 15, december 2015 amy k. buthod and bruce w. hoagland 36.938120 -103.00098 sec. 31-t6n-r1e 36.938800 -103.00023 sec. 31-t6n-r1e 36.939080 -102.99954 sec. 31-t6n-r1e 36.940380 -102.98649 sec. 31-t6n-r1e 36.943380 -102.95534 sec. 33-t6n-r1e 36.944330 -102.95544 sec. 33-t6n-r1e 36.945420 -102.618 sec. 34-t6n-r4e 36.945760 -102.97118 sec.32-t6n-r1e 36.947060 -102.97128 sec. 32-t6n-r1e 36.947930 -102.96566 sec. 33-t6n-r1e 36.948080 -102.45784 sec. 31-t6n-r6e 36.952680 -102.96242 sec. 28-t6n-r1e 36.955610 -102.72656 sec. 27-t6n-r3e 36.960120 -102.71428 sec. 27-t6n-r3e 36.962150 -102.80867 sec. 26-t6n-r2e 36.964600 -102.62363 sec. 28-t6n-r4e 36.967830 -102.71885 sec. 22-t6n-r3e 36.982940 -102.24962 sec. 13-t6n-r7e 36.989890 -102.67913 sec. 13-t6n-r3e oklahoma native plant record 57 volume 15, december 2015 amy k. buthod and bruce w. hoagland table 2 taxa located during this study that are tracked by the oklahoma natural heritage inventory (oklahoma natural heritage inventory 2013; natureserve explorer 2015). status ranks are on a 1–5 scale, with a 1 indicating the taxa is critically imperiled. g ranks are at the global level and s ranks are at the subnational or state level. infraspecific taxa are assigned a t rank. a taxon with nr indicates that it has not been ranked at the global level (natureserve 2015). highlighted taxa were re-ranked as a result of this survey. family taxon ranking amaranthaceae krascheninnikovia lanata (pursh) a. meeuse & a. smit s1g5 apocynaceae asclepias macrotis torr. s1g4 asteraceae ambrosia confertiflora dc. s1g5 asteraceae artemisia carruthii alph. wood ex carruth. s2g4? asteraceae brickellia brachyphylla (a. gray) a. gray s1g5 asteraceae brickellia californica (torr. & a. gray) a. gray s1g5 asteraceae brickellia eupatorioides (l.) shinners var. s1g5t5 asteraceae ericameria nauseosa (pall. ex pursh) g.l. nesom & s1g5t5 asteraceae picradeniopsis woodhousei (a. gray) rydb. s2g4g5 asteraceae solidago velutina dc. ssp. sparsiflora (a. gray) semple s1g5?tnr boraginaceae cryptantha cinerea (greene) cronquist var. s2g5t5? boraginaceae cryptantha thyrsiflora (greene) payson s2g4 cactaceae cylindropuntia imbricata (haw.) f.m. knuth s2g5 cactaceae echinocereus reichenbachii (terscheck ex walp.) j.n. s3g5 cactaceae echinocereus viridiflorus engelm. s1g5 cactaceae escobaria vivipara (nutt.) buxb. s1g5 cactaceae opuntia polyacantha haw. var. polyacantha s2g5t5 convolvulaceae cuscuta umbellata kunth s1g5 crossomataceae glossopetalon spinescens a. gray var. s1g5tnr cupressaceae juniperus monosperma (engelm.) sarg. s2g4g5 fabaceae dalea formosa torr. s2g5 58 oklahoma native plant record volume 15, december 2015 amy k. buthod and bruce w. hoagland fabaceae dalea jamesii (torr.) torr. & a. gray s1g5 fabaceae desmanthus cooleyi (eaton) trel. s2g5 fabaceae hoffmannseggia drepanocarpa a. gray s2g5 fabaceae lupinus plattensis s. watson s1g4 grossulariaceae ribes cereum douglas s1g5 malvaceae sphaeralcea angustifolia (cav.) g. don s2g5 nyctaginaceae abronia fragrans nutt. ex hook. s2g5 papaveraceae argemone squarrosa greene s1g4 pinaceae pinus edulis engelm. s1g5 plantaginaceae penstemon fendleri torr. & a. gray s1g5t4? poaceae aristida arizonica vasey s1g4 poaceae bouteloua barbata lag. s1g5 poaceae bouteloua eriopoda (torr.) torr. s1g5 poaceae hesperostipa neomexicana (thurb.) barkworth s1g4g5 poaceae hilaria jamesii (torr.) benth. s1g5 poaceae muhlenbergia phleoides (kunth) columbus s1g5 poaceae muhlenbergia porteri scribn. ex beal s1g5 poaceae muhlenbergia torreyi (kunth) hitchc. ex bush s1g4 poaceae piptatherum micranthum (trin. & rupr.) barkworth s1g5 polygonaceae eriogonum jamesii benth. s1g5 polygonaceae eriogonum lachnogynum torr. ex benth. s1g4? polygonaceae eriogonum tenellum torr. s1g5 rosaceae cercocarpus montanus raf. s1g5 rosaceae rubus deliciosus torr. s1g4? selaginellaceae selaginella underwoodii hieron. s1g5? oklahoma native plant record 59 volume 15, december 2015 amy k. buthod and bruce w. hoagland discussion one-hundred sixty taxa from 46 families reported in the rogers, mcpherson, and folley studies were not found (appendix b), and only 46 of the 95 taxa tracked by the oklahoma natural heritage inventory were located. one explanation for this difference is land access. for instance, we were not able to collect in tesequite canyon, which is known to have populations of tracked taxa (oklahoma natural heritage inventory 2015), as was done in the folley study. we were uncomfortable botanizing along some of the public roads, as well. another explanation could be that vegetation changes have occurred in the area. vegetation analysis by graham et al. (unpubl. data) indicates a decrease in the amount of grassland/herbaceous vegetation and an increase in forest/shrubland since 1992. this is most probably due to the increased amount of cholla (cylindropuntia imbricata) in the area. the most likely explanation for our results, however, is drought. cimarron county is considered to be the epicenter of the exceptional drought experienced by the high plains regions of northern texas, southwestern kansas, northeastern new mexico, southeastern colorado, and the northwestern oklahoma panhandle (lindsey 2008; south central climate science center 2013). throughout the survey period, western cimarron county experienced exceptional, extreme, or extreme/severe drought (national oceanic and atmospheric administration et al. 2015). rogers (1953) stated that the “severe drouth of the 1930s had a disturbing effect on the vegetation”, but noted a “great recovery” in the following decade. although the national weather service predicts that the drought status for the area will likely be removed, another “great recovery” is unlikely (u. s. geological survey 2014). the area could be as much as 5°c hotter by the end of the century, and decreases in precipitation, runoff, and amounts of soil water storage are also likely (u. s. geological survey 2014). acknowledgements this work was supported by a grant from the department of interior, u. s. fish and wildlife service. the authors wish to thank todd fagin (oklahoma biological survey/department of geography and environmental sustainability, university of oklahoma) for assistance with map preparation. literature cited allred, k.w. and r.d. ivey. 2012. flora neomexicana iii: an illustrated identification manual. self-published. carter, b.j. and m.s. gregory. 2008. soil map of oklahoma. in: earth sciences and mineral resources of oklahoma. johnson, k.s. and k.v. luza (eds.). norman (ok): oklahoma geological survey. curtis, n.m., w.e. ham, and k.s. johnson. 2008. geomorphic provinces of oklahoma. in: earth sciences and mineral resources of oklahoma. johnson, k.s. and k.v. luza (eds.) norman (ok): oklahoma geological survey. duck, l.g. and j.b. fletcher. 1943. a survey of the game and fur-bearing animals of oklahoma. oklahoma city (ok): oklahoma department of wildlife conservation. folley, p.a. 2003. additions to black mesa flora study. oklahoma native plant record 3:19–22. great plains flora association. 1986. flora of the great plains. lawrence (ks): university of kansas press. hess, w.j. 2002. nolina. pp. 415–422. in: flora of north america north of mexico. vol. 26. flora of north american editorial committee (eds.). new york and oxford. 60 oklahoma native plant record volume 15, december 2015 amy k. buthod and bruce w. hoagland hoagland, b.w. 2000. the vegetation of oklahoma: a classification for landscape mapping and conservation planning. the southwestern naturalist 45:385–420. hoagland, b.w. 2008. vegetation of oklahoma. in: earth sciences and mineral resources of oklahoma. johnson, k.s. and k.v. luza (eds.). norman (ok): oklahoma geological survey. integrated taxonomic information system. 2015. http://www.itis.gov (12 february 2015). lindsey, r. 2008. devastating drought settles on the high plains. http://earthobservatory.nasa.gov (17 march 2015). mcpherson, j.k. 2003a. black mesa flora study. oklahoma native plant record 3:4–7. mcpherson, j.k. 2003b. black mesa flora study: year two supplement. oklahoma native plant record 3:8–18. national drought mitigation center. 2015. http://droughtmonitor.unl.edu (17 march 2015). national oceanic and atmospheric administration, u.s. department of agriculture, and national drought mitigation center. 2015. u.s. drought monitor. http://droughtmonitor.unl.edu/home.aspx (9 march 2015). natureserve. 2015. natureserve explorer. http://www.natureserve.org/explorer (12 february 2015). noaa center for weather and climate prediction. 2015. http://www.cpc.noaa.gov (23 march 2015). oklahoma climatological survey. 2015. the climate of cimarron county. http://www.ocs.ou.edu (11 march 2015). oklahoma natural heritage inventory. 2013. plant tracking list. http://www.oknaturalheritage.ou.edu (15 january 2013). oklahoma natural heritage inventory. 2015. heritage database. http://www.oknaturalheritage.ou.edu (17 march 2015). oklahoma vascular plants database. 2015. http://www.oklahomaplantdatabase.org (4 november 2015). rogers, c.m. 1953. the vegetation of the mesa de maya region of colorado, new mexico and oklahoma. lloydia 16:257–290. rogers, c.m. 1954. some botanical studies in the black mesa region of oklahoma. rhodora 56: 205–212. south central climate science center. 2013. drought history for the oklahoma panhandle. http://www.southcentralclimate.org (17 march 2015). taylor, r.j. and c.e.s. taylor. 1991. an annotated list of the ferns, fern allies, gymnosperms and flowering plants of oklahoma. self-published. trewartha, g.t. 1968. an introduction to climate. new york: mcgraw-hill. tyrl, r.j., s.c. barber, p. buck, w.j. elisens, j.r. estes, p. folley, l.k. magrath, c.l. murray, a.k. ryburn, b.a. smith, c.e.s. taylor, r.a. thompson, j.b. walker, and l.e. watson. 2010. keys and descriptions for the vascular plants of oklahoma. noble (ok): flora oklahoma incorporated. united states department of the interior, fish and wildlife service. 1993. federal register, part iv 58(188):51160. united states geological survey. 2014. summary of cimarron county. http://regclim.coas.oregonstate.edu/n exdcp30_app/data/counties_201403 31/summaries/40025/40025.pdf (23 march 2015). usda, nrcs. 2015. the plants database. http://plants.usda.gov/plants (9 march 2015). waterfall, u.t. 1949. some results of a summer’s botanizing in oklahoma. rhodora 51:18–28. http://www.itis.gov/ http://earthobservatory.nasa.gov/ http://droughtmonitor.unl.edu/ http://droughtmonitor.unl.edu/home.aspx http://droughtmonitor.unl.edu/home.aspx http://www.natureserve.org/explorer http://www.cpc.noaa.gov/ http://www.ocs.ou.edu/ http://www.oknaturalheritage.ou.edu/ http://www.oknaturalheritage.ou.edu/ http://www.oklahomaplantdatabase.org/ http://www.southcentralclimate.org/ http://regclim.coas.oregonstate.edu/nexdcp30_app/data/counties_20140331/summaries/40025/40025.pdf http://regclim.coas.oregonstate.edu/nexdcp30_app/data/counties_20140331/summaries/40025/40025.pdf http://regclim.coas.oregonstate.edu/nexdcp30_app/data/counties_20140331/summaries/40025/40025.pdf http://plants.usda.gov/plants oklahoma native plant record 61 volume 15, december 2015 amy k. buthod and bruce w. hoagland waterfall, u.t. 1950a. some additions to the oklahoma flora. rhodora 52:19–24, 35– 41. waterfall, u.t. 1950b. some results of a third summer’s botanizing in oklahoma. rhodora 52:165–175. 62 oklahoma native plant record volume 15, december 2015 amy k. buthod and bruce w. hoagland appendix a list of plant taxa in cimarron county and black mesa, oklahoma taxa list with duration, vegetation type, and nativity. a=annual, b=biennial, p=perennial; afsa=artemisia filifolia shrubland association, bcbgbd=bouteloua curtipendula-bouteloua gracilisbouteloua dactyloides herbaceous association, bghj=bouteloua gracilis-hilaria jamesii herbaceous association, daof=disturbed area/old field vegetation, hwv=herbaceous wetland vegetation, jmwa=juniperus monosperma woodland alliance. an asterisk (*) indicates a taxon that is non-native to the united states. a dagger (†) indicates a tracked taxon. taxonomy follows the integrated taxonomic information system (2015). duration and nativity to oklahoma were determined using the plants database (usda-nrsc 2015); if the information from plants was ambiguous, taylor and taylor (1991) was consulted. vegetation classifications were based on hoagland (2000). alismataceae alisma subcordatum raf., p, hwv amaranthaceae amaranthus palmeri s. watson, a, afsa amaranthus tuberculatus (moq.) j.d. sauer, a, afsa atriplex canescens (pursh) nutt., p, bghj *chenopodium album l., a, bghj chenopodium berlandieri moq., a, daof chenopodium incanum (s. watson) a. heller, a, bghj chenopodium leptophyllum (moq.) nutt. ex s. watson, a , daof chenopodium pratericola rydb., a, bghj chenopodium simplex (torr.) raf., a, bcbgbd chenopodium standleyanum aellen, a, jmwa froelichia floridana (nutt.) moq., a, jmwa *kochia scoparia ssp. scoparia (l.) schrad., a, daof †krascheninnikovia lanata (pursh) a. meeuse & a. smit, p, jmwa monolepis nuttalliana (schult.) green, a, daof *salsola tragus l., a, bcbgbd tidestromia lanuginosa (nutt.) standl., a, afsa amaryllidaceae allium drummondii regel, p, bghj anacardiaceae rhus aromatica aiton var. pilosissima (engl.) shinners, p, bghj toxicodendron rydbergii (small ex rydb.) greene, p, jmwa apiaceae cymopterus montanus nutt. ex torr. & a. gray, p, jmwa oklahoma native plant record 63 volume 15, december 2015 amy k. buthod and bruce w. hoagland apocynaceae apocynum androsaemifolium l., p, jmwa asclepias asperula (decne.) woodson ssp. capricornu (woodson) woodson, p, jmwa asclepias engelmanniana woodson, p, afsa asclepias latifolia (torr.) raf., p, afsa †asclepias macrotis torr., p, jmwa asclepias subverticillata (a. gray) vail, p, afsa asclepias viridiflora raf., p, bcbgbd asparagaceae *asparagus officinalis l., p, bcbgbd yucca glauca nutt., p, afsa asteraceae †ambrosia confertiflora dc., p, afsa ambrosia grayi (a. nelson) shinners, p, daof ambrosia psilostachya dc., p, daof ambrosia trifida l., a, bghj amphiachyris dracunculoides (dc.) nutt., a, afsa †artemisia carruthii alph. wood ex carruth., p, bcbgbd artemisia filifolia torr., p, afsa artemisia ludoviciana nutt., p, bghj baccharis salicina torr. & a. gray, p, hwv berlandiera lyrata benth., p, afsa †brickellia brachyphylla (a. gray) a. gray, p, bghj †brickellia californica (torr. & a. gray) a. gray, p, bghj †brickellia eupatorioides (l.) shinners var. chlorolepis (woot. & standl.) b.l. turner, p, bghj cirsium ochrocentrum a. gray ssp. ochrocentrum, p, bghj cirsium undulatum (nutt.) spreng., p, bghj conyza canadensis (l.) cronquist, a, daof diaperia prolifera (nutt. ex dc.) nutt., a, bghj dyssodia papposa (vent.) hitchc., a, jmwa engelmannia peristenia (raf.) goodman & c.a. lawson, p, bghj †ericameria nauseosa (pall. ex pursh) g.l. nesom & baird var. graveolens (nutt.) reveal & schuyler, p, jmwa erigeron bellidiastrum nutt., afsa, a erigeron flagellaris a. gray, b, afsa gaillardia pinnatifida torr., p, bghj gaillardia pulchella foug., a, bghj grindelia squarrosa (pursh) dunal, p, bghj gutierrezia sarothrae (pursh) britton & rusby, p, bghj helianthus annuus l., a, bghj helianthus ciliaris dc., p, bcbgbd helianthus petiolaris nutt., a, daof heterotheca stenophylla (gray) shinners var. angustifolia (rydb.) semple, p, jmwa heterotheca subaxillaris (lam.) britton & rusby spp. latifolia (buckley) semple, a, bghj 64 oklahoma native plant record volume 15, december 2015 amy k. buthod and bruce w. hoagland heterotheca villosa (pursh) shinners var. villosa, p, jmwa hymenopappus flavescens a. gray, b, afsa hymenopappus tenuifolius pursh, b, bghj *lactuca serriola l., a, daof liatris punctata hook. var. punctata, p, afsa lygodesmia juncea (pursh) d. don ex hook., p, jmwa machaeranthera tanacetifolia (kunth) nees, a, jmwa melampodium leucanthum torr. & a. gray, p, bghj packera plattensis (nutt.) w.a. weber & a. löve, p, bghj palafoxia sphacelata (nutt. ex torr.) cory, a, bcbgbd †picradeniopsis woodhousei (a. gray) rydb., p, bghj pseudognaphalium canescens (dc.) w.a. weber ssp. canescens, b, bghj ratibida columnifera (nutt.) woot. & standl., p, bcbgbd ratibida tagetes (james) barnhart, p, daof *scorzonera laciniata l., p, daof senecio flaccidus less. var. flaccidus, p, bghj senecio riddellii torr. & a. gray, p, jmwa solidago gigantea aiton, p, daof †solidago velutina dc. ssp. sparsiflora (a. gray) semple, p, bghj symphyotrichum subulatum (michx.) g.l. nesom, a, hwv *taraxacum officinale f.h. wigg., p, daof tetraneuris acaulis (pursh) greene var. acaulis, p, jmwa tetraneuris scaposa (dc.) greene var. scaposa, p, bghj thelesperma ambiguum a. gray, p, afsa thelesperma filifolium (hook.) a. gray, p, bghj thelesperma megapotamicum (spreng.) kuntze, p, bghj townsendia exscapa (richardson) porter, p, bghj *tragopogon dubius scop., a, jmwa vernonia marginata (torr.) raf., p, jmwa xanthisma spinulosum (pursh) d.r. morgan & r.l. hartm. var. spinulosum, p, bghj xanthium strumarium l., a, hwv zinnia grandiflora nutt., p, bghj boraginaceae †cryptantha cinerea (greene) cronquist var. jamesii (torr.) cronquist, p, afsa, cryptantha minima rydb., a, afsa cryptantha thyrsiflora (greene) payson, p, bghj lappula occidentalis (s. watson) greene var. cupulata (a. gray) higgins, a, daof lappula occidentalis (s. watson) greene var. occidentalis, a, daof lithospermum incisum lehm., p, bghj onosmodium bejariense dc. ex a. dc. var. occidentale (mack.) b.l. turner, p, jmwa brassicaceae *camelina microcarpa dc., a, bcbgbd descurainia pinnata (walter) britton ssp. brachycarpa (richardson) detling, a, jmwa *descurainia sophia (l.) webb ex prantl, a, daof erysimum asperum (nutt.) dc., p, bghj oklahoma native plant record 65 volume 15, december 2015 amy k. buthod and bruce w. hoagland erysimum capitatum (douglas ex hook.) greene, p, bghj *erysimum repandum l., a, bghj *lepidium densiflorum schrad., a, daof physaria ovalifolia (rydb.) o'kane & al-shehbaz ssp. ovalifolia, p, jmwa rorippa sinuata (nutt.) hitchc., p, hwv *sisymbrium altissimum l., a, bghj cactaceae †cylindropuntia imbricata (haw.) f.m. knuth, p, bghj †echinocereus reichenbachii (terscheck ex walp.) j.n. haage, p, afsa †echinocereus viridiflorus engelm., p, jmwa †escobaria vivipara (nutt.) buxb., p, jmwa opuntia humifusa (raf.) raf. var. humifusa, p, bghj opuntia macrorhiza engelm., p, jmwa opuntia phaeacantha engelm., p, bghj, †opuntia polyacantha haw.var. polyacantha, p, jmwa cannabaceae celtis reticulata torr., p, bghj caryophyllaceae paronychia jamesii torr. & a. gray, p, bghj paronychia sessiliflora nutt., p, bghj cleomaceae polanisia dodecandra (l.) dc., a, bghj commelinaceae commelina erecta l., p, jmwa tradescantia occidentalis (britton) smyth var. occidentalis, p, bghj convolvulaceae *convolvulus arvensis l., bghj, p convolvulus equitans benth., bghj, p †cuscuta umbellata kunth, a, daof evolvulus nuttallianus schult., p, bghj ipomoea leptophylla torr., p, bghj crossomataceae †glossopetalon spinescens a. gray var. planitierum (ensign) yatsk., p, jmwa, cucurbitaceae cucurbita foetidissima kunth, p, bghj cyclanthera dissecta (torr. & a. gray) arn., a, jmwa cupressaceae †juniperus monosperma (engelm.) sarg., p, jmwa 66 oklahoma native plant record volume 15, december 2015 amy k. buthod and bruce w. hoagland cyperaceae carex gravida l.h. bailey, p, hwv carex muehlenbergii schkuhr ex willd., p, hwv schoenoplectus acutus (muhl. ex bigelow) á. löve & d. löve var. acutus, p, hwv schoenoplectus pungens (vahl) palla var. pungens, p, hwv euphorbiaceae croton texensis (klotzsch) müll. arg., a, bghj ditaxis mercurialina (nutt.) j.m. coult., p, jmwa euphorbia dentata michx., a, afsa euphorbia exstipulata engelm., a, bghj euphorbia fendleri torr. & a. gray, p, jmwa euphorbia glyptosperma engelm., a, afsa euphorbia lata engelm., p, bghj euphorbia marginata pursh, a, bcbgbd euphorbia missurica raf., a, bcbgbd euphorbia serpyllifolia pers. var. serpyllifolia, a, bcbgbd tragia ramosa torr., p, jmwa fabaceae amorpha canescens pursh, p, jmwa astragalus missouriensis nutt., p, bghj astragalus mollissimus torr., p, bghj dalea aurea nutt. ex fraser, p, bghj dalea candida michx. ex. willd var. oligophylla (torr.) shinners, p, jmwa dalea enneandra nutt. ex fraser, p, afsa †dalea formosa torr., p, jmwa †dalea jamesii (torr.) torr. & a. gray, p, bghj dalea lanata spreng., p, bghj dalea tenuifolia (a. gray) shinners, p, bghj dalea villosa (nutt.) spreng., p, daof †desmanthus cooleyi (eaton) trel., p, bghj glycyrrhiza lepidota pursh, p, bghj †hoffmannseggia drepanocarpa a. gray, p, bghj hoffmannseggia glauca (ortega) eifert, p, bcbgbd †lupinus plattensis s. watson, p, afsa *medicago sativa l., p, bghj *melilotus officinalis (l.) lam., a, daof mimosa borealis a. gray, p, bghj oxytropis lambertii pursh, p, afsa pediomelum cuspidatum (pursh) rydb., p, bghj prosopis glandulosa torr. var. glandulosa, p, bghj psoralidium tenuiflorum (pursh) rydb., p, bghj robinia pseudoacacia l., p, daof sophora nuttalliana b.l. turner, p, bghj oklahoma native plant record 67 volume 15, december 2015 amy k. buthod and bruce w. hoagland fagaceae quercus mohriana buckley ex rydb., p, jmwa geraniaceae *erodium cicutarium (l.) l'hér. ex aiton, a, daof grossulariaceae ribes aureum pursh var. villosum dc., p, bcbgbd †ribes cereum douglas, p, jmwa juncaceae juncus interior wiegand, p, hwv juncus torreyi coville, p, hwv krameriaceae krameria lanceolata torr., p, bghj lamiaceae hedeoma drummondii benth., p, bghj *marrubium vulgare l., p, bghj monarda pectinata nutt., a, afsa salvia reflexa hornem., a, jmwa teucrium laciniatum torr., p, jmwa linaceae linum pratense (norton) small, a, bghj linum rigidum pursh var. rigidum, a, bcbgbd loasaceae mentzelia multiflora (nutt.) a. gray, a, afsa mentzelia nuda (pursh) torr. & a. gray, p, afsa mentzelia oligosperma nutt. ex sims, p, bghj malvaceae callirhoe involucrata (torr. & a. gray) a. gray, p, bcbgbd *malva neglecta wallr., a, daof †sphaeralcea angustifolia (cav.) g. don, p, afsa sphaeralcea coccinea (nutt.) rydb., p, bcbgbd martyniaceae proboscidea louisianica (mill.) thell., ssp. louisianica, a, afsa moraceae *morus alba l., p, jmwa nyctaginaceae †abronia fragrans nutt. ex hook., p, afsa 68 oklahoma native plant record volume 15, december 2015 amy k. buthod and bruce w. hoagland mirabilis albida (walter) heimerl, p, jmwa mirabilis linearis (pursh) heimerl var. subhispida (heimerl) spellenb., p, jmwa mirabilis nyctaginea (michx.) macmill., p, jmwa oleaceae forestiera pubescens nutt., p, bghj onagraceae oenothera cespitosa nutt., p, jmwa oenothera cinerea (wooton & standl.) w.l. wagner & hoch ssp. cinerea, p, bcbgbd oenothera curtiflora w.l. wagner & hoch, a, daof oenothera hartwegii benth. ssp. pubescens (a. gray) w.l. wagner & hoch, p, bghj oenothera serrulata nutt., p, bcbgbd oenothera suffrutescens (ser.) w.l. wagner & hoch, p, bghj oenothera triloba nutt., p, bghj orobanchaceae orobanche ludoviciana nutt. ssp. multiflora (nutt.) t.s. collins ex h.l. white & w.c. holmes, a, bghj papaveraceae †argemone squarrosa greene, p, bghj corydalis aurea willd. ssp. occidentalis (engelm. ex a. gray) g.b. ownbey, a, bcbgbd pinaceae †pinus edulis engelm., p, jmwa plantaginaceae penstemon albidus nutt., p, bghj penstemon ambiguus torr., p, bghj †penstemon fendleri torr. & a. gray, p, afsa plantago patagonica jacq., a, bghj veronica anagallis-aquatica l., p, hwv poaceae *aegilops cylindrica host, a, daof andropogon gerardii vitman ssp. hallii (hack.) wipff, p, afsa andropogon gerardii vitman ssp. gerardii, p, bcbgbd aristida adscensionis l., a, bghj †aristida arizonica vasey, p, bghj aristida havardii vasey, p, bcbgbd aristida oligantha michx., a, afsa aristida purpurascens poir., p, bcbgbd aristida purpurea nutt. var. purpurea, p, bghj bothriochloa barbinodis (lag.) herter, p, bghj *bothriochloa ischaemum (l.) keng, p, afsa bothriochloa laguroides (dc.) herter, p, bghj †bouteloua barbata lag., a, jmwa oklahoma native plant record 69 volume 15, december 2015 amy k. buthod and bruce w. hoagland bouteloua curtipendula (michx.) torr., p, afsa bouteloua dactyloides (nutt.) columbus, p, bghj †bouteloua eriopoda (torr.) torr., p, bghj bouteloua gracilis (kunth) lag. ex griffiths, p, bcbgbd bouteloua hirsuta lag. , p, bghj *bromus arvensis l., a, daof *bromus catharticus vahl, a, daof *bromus racemosus l., a, bghj *bromus tectorum l., a, daof calamovilfa gigantea (nutt.) scribn. & merr., p, bcbgbd cenchrus spinifex cav., p, bghj chloris verticillata nutt., p, afsa chloris virgata sw., a, bghj *cynodon dactylon (l.) pers., p, daof distichlis spicata (l.) greene var. stricta (torr.) thorne, p, bghj echinochloa muricata (p. beauv.) fernald, a, daof elymus canadensis l., p, bghj elymus elymoides (raf.) swezey, p, jmwa elymus virginicus l., p, afsa *eragrostis cilianensis (bellardi) vignolo ex janch., a, afsa erioneuron pilosum (buckley) nash, p, jmwa †hesperostipa neomexicana (thurb.) barkworth, p, bghj †hilaria jamesii (torr.) benth., p, bghj hopia obtusa (kunth) zuloaga & morrone, p, afsa hordeum jubatum l., p, daof hordeum pusillum nutt., a, daof leptochloa fusca (l.) kunth spp. fasicularis n.w. snow, a, hwv muhlenbergia asperifolia (nees & meyen ex trin.) parodi, p, afsa muhlenbergia paniculata (nutt.) columbus, p, daof †muhlenbergia phleoides (kunth) columbus, p, bghj †muhlenbergia porteri scribn. ex beal, p, jmwa †muhlenbergia torreyi (kunth) hitchc. ex bush, p, bcbgbd munroa squarrosa (nutt.) torr., a, bghj panicum capillare l., a, daof panicum hallii vasey, p, bghj panicum virgatum l., p, jmwa pascopyrum smithii (rydb.) barkworth & d.r. dewey, p, afsa paspalum setaceum michx. var. stramineum (nash) d.j. banks, p, daof †piptatherum micranthum (trin. & rupr.) barkworth, p, jmwa poa fendleriana (steud.) vasey, p, jmwa *polypogon monspeliensis (l.) desf., a, hwv schizachyrium scoparium (michx.) nash, p, afsa setaria macrostachya kunth, p, daof *setaria viridis (l.) p. beauv., a, daof sorghastrum nutans (l.) nash, p, bghj *sorghum halepense (l.) pers., p, bghj sporobolus airoides (torr.) torr. , p, bghj 70 oklahoma native plant record volume 15, december 2015 amy k. buthod and bruce w. hoagland sporobolus cryptandrus (torr.) a. gray, p, afsa sporobolus pyramidatus (lam.) hitchc., p, afsa polemoniaceae ipomopsis laxiflora (j.m. coult.) v.e. grant, a, jmwa polygalaceae polygala alba nutt., p, bghj polygonaceae eriogonum annuum nutt., a, afsa †eriogonum jamesii benth., p, bcbgbd †eriogonum lachnogynum torr. ex benth., p, bghj †eriogonum tenellum torr., p, jmwa persicaria amphibia (l.) delarbre, p, hwv persicaria lapathifolia (l.) gray, a, hwv *polygonum aviculare l., a, daof rumex altissimus alph. wood, p, hwv *rumex crispus l., p, hwv rumex venosus pursh, p, daof portulacaceae phemeranthus parviflorus (nutt.) kiger, p, afsa portulaca oleracea l., a, jwma portulaca pilosa l., a, daof potamogetonaceae zannichellia palustris l., p, hwv pteridaceae cheilanthes eatonii baker, p, jmwa notholaena standleyi, p, jmwa ranunculaceae delphinium carolinianum walter ssp. virescens (nutt.) r.e. brooks, p, jmwa ranunculus abortivus l., p, hwv ranunculus sceleratus l., a, hwv *ranunculus testiculatus crantz, a, daof rosaceae †cercocarpus montanus raf., p, jmwa prunus virginiana l. var. demissa (nutt.) torr., p, jmwa †rubus deliciosus torr., p, jmwa rutaceae ptelea trifoliata l., p, jmwa oklahoma native plant record 71 volume 15, december 2015 amy k. buthod and bruce w. hoagland salicaceae populus deltoides w. bartram ex marshall, p, hwv salix amygdaloides andersson, p, hwv salix exigua nutt., p, hwv salix nigra marshall, p, hwv santalaceae comandra umbellata (l.) nutt. ssp. pallida (a. dc.) piehl, p, jmwa sapindaceae sapindus saponaria l. var. drummondii (hook. & arn.) l.d. benson, p, daof selaginellaceae †selaginella underwoodii hieron., p, jmwa solanaceae chamaesaracha coniodes (moric. ex dunal) britton, p, jmwa datura quercifolia kunth, a, daof physalis hederifolia a. gray var. fendleri (a. gray) cronquist, p, jmwa physalis longifolia nutt. var. longifolia, p, afsa quincula lobata (torr.) raf., p, jmwa solanum elaeagnifolium cav., p, daof solanum ptychanthum dunal, a, bcbgbd solanum rostratum dunal, a, afsa solanum triflorum nutt., a, daof tamaricaceae *tamarix chinensis lour., p, hwv verbenaceae glandularia bipinnatifida (nutt.) nutt. var. ciliata (benth.) b.l. turner, a, bghj glandularia canadensis (l.) nutt., p, jmwa glandularia pumila (rydb.) umber, a, bghj phyla cuneifolia (torr.) greene, p, hwv verbena bracteata cav. ex lag. & rodr., a, afsa violaceae hybanthus verticillatus (ortega) baill., p, bghj vitaceae vitis vulpina l., p, jmwa zygophyllaceae kallstroemia parviflora norton, a, afsa *tribulus terrestris l., a, afsa 72 oklahoma native plant record volume 15, december 2015 amy k. buthod and bruce w. hoagland appendix b list of plant taxa in cimarron county and black mesa, oklahoma not found by buthod and hoagland taxa from the published lists of rogers (1953), mcpherson (2003a, b), and folley (2003) that were not found by buthod and hoagland. r=rogers collection, m=mcpherson collection, f=folley collection. taxonomy has been updated and follows the integrated taxonomic information system (2015). amaranthaceae amaranthus retroflexus l., m chenopodium albescens small, r cycloloma atriplicifolium (spreng.) j.m. coult., r froelichia gracilis (hook.) moq., r guilleminea densa (humb. & bonpl. ex schult.) moq. var. densa, r salsola kali l. ssp. tenuifolia moq., m suckleya suckleyana (torr.) rydb., m amaryllidaceae allium canadense l. var. fraseri ownbey, m anacardiaceae rhus aromatica aiton var. simplicifolia (greene) cronquist, r toxicodendron radicans (l.) kuntze, m apiaceae cymopterus glomeratus (nutt.) dc., m apocynaceae asclepias arenaria torr. , m asclepias involucrata engelm. ex torr. , r asclepias pumila (a. gray) vail, r, m asclepias uncialis greene, m funastrum crispum (benth.) schltr., r, m araceae lemna minor l., m asparagaceae nolina texana s. watson, f (collections are actually nolina greenei s. watson ex trel.; hess 2002)) yucca harrimaniae trel., f aspleniaceae asplenium septentrionale (l.) hoffm., m asteraceae antennaria parvifolia nutt., r oklahoma native plant record 73 volume 15, december 2015 amy k. buthod and bruce w. hoagland artemisia dracunculus l., r, m baccharis wrightii a. gray, r bidens cernua l., f brickellia eupatorioides (l.) shinners var. corymbulosa (torr. & a. gray) shinners, r chaetopappa ericoides (torr.) g.l. nesom, r, m ericameria nauseosa (pall. ex pursh) g.l. nesom & baird var. nauseosa, r, m erigeron nudiflorus buckley, r erigeron tracyi greene, m nothocalais cuspidata (pursh) greene, m oonopsis foliosa (a. gray) greene var. foliosa, r packera tridenticulata (rydb.) w.a. weber & a. löve, r, m pericome caudata a. gray, r, m, f picradeniopsis oppositifolia (nutt.) rydb. ex britton, r psilostrophe villosa rydb., f solidago mollis bartlett , m solidago petiolaris aiton, m stephanomeria pauciflora (torr.) a. nelson, r, m symphyotrichum ericoides (l.) g.l. nesom, r, m symphyotrichum fendleri (a. gray) g.l. nesom, m symphyotrichum oblongifolium (nutt.) g.l. nesom, m verbesina encelioides (cav.) benth. & hook. f. ex a. gray, m vernonia fasciculata michx., f xanthisma spinulosum (pursh) d.r. morgon & r.l. hartm. var. glaberrimum (rydberg) d.r. morgan & r.l. hartm., r boraginaceae cryptantha cinerea (greene) cronquist var. cinerea, r cryptantha crassisepala (torr. & a. gray) greene, r euploca convolvulacea nutt., f lithospermum multiflorum torr. ex a. gray, f brassicaceae boechera fendleri (s. watson) w.a. weber, m cactaceae opuntia fragilis (nutt.) haw., f campanulaceae lobelia cardinalis l., f cleomaceae peritoma serrulata (pursh) dc., r, f polanisia jamesii (torr. & a. gray) iltis, f cupressaceae juniperus scopulorum sarg., m 74 oklahoma native plant record volume 15, december 2015 amy k. buthod and bruce w. hoagland cyperaceae carex brevior (dewey) mack. , f cyperus croceus vahl, f cyperus schweinitzii torr., r, m schoenoplectus tabernaemontani (c.c. gmel.) palla, m scirpus atrovirens willd., f scirpus pallidus (britton) fernald, r cystopteridaceae cystopteris fragilis (l.) bernh., f equisetaceae equisetum laevigatum a. br., r euphorbiaceae ditaxis humilis (engelm. & a. gray) pax, r, m euphorbia geyeri engelm., r euphorbia spathulata lam., r fabaceae astragalus ceramicus e. sheld., f astragalus crassicarpus nutt., r astragalus crassicarpus nutt. var. paysonii (e.h. kelso) barneby, m astragalus gracilis nutt., r astragalus hallii a. gray, r astragalus lotiflorus hook. , r, m astragalus puniceus osterh., m colutea arborescens l., f dalea candida michx. ex willd var. candida, r dalea compacta spreng. var. compacta, r dalea nana torr. ex a. gray, r dalea purpurea vent. var. purpurea, r hedysarum boreale nutt., r melilotus albus medik., r pediomelum argophyllum (pursh) j.w. grimes, m pediomelum hypogaeum (nutt.) rydb. var. hypogaeum, r pomaria jamesii (torr. & a. gray) walp., r, m vicia americana muhl. ex willd. , m vicia ludoviciana nutt. ex torr. & a. gray var. leavenworthii (nutt. ex torr. & a. gray) broich, r fagaceae quercus gambelii nutt., r quercus grisea liebm., r quercus x undulata torr., r lamiaceae salvia azurea michx. ex lam. var. grandiflora benth., m oklahoma native plant record 75 volume 15, december 2015 amy k. buthod and bruce w. hoagland linaceae linum lewisii pursh , r, m loasaceae mentzelia decapetala (pursh ex sims) urb. & gilg, r, m lythraceae lythrum alatum pursh, r nyctaginaceae mirabilis glabra (s. watson) standl., r, m mirabilis linearis (pursh) heimerl var. linearis, r onagraceae oenothera albicaulis pursh, r oenothera engelmannii (small) munz, r, f oenothera lavandulifolia torr. & a. gray, m oenothera pallida lindl. ssp. latifolia (rydb.) munz, f orobanchaceae castilleja sessiliflora pursh, r, m papaveraceae argemone polyanthemos (fedde) g.b. ownbey, r plantaginaceae penstemon angustifolius nutt. ex pursh var. caudatus (a. heller) rydb., r poaceae achnatherum hymenoides (roem. & schult.) barkworth, r, m achnatherum scribneri (vasey) barkworth, r, m andropogon virginicus l., f aristida barbata e. fourn., r aristida divaricata humb. & bonpl. ex willd., r aristida purpurea nutt. var. fendleriana (steud.) vasey, r aristida purpurea nutt. var. longiseta (steud.) vasey, r aristida purpurea nutt. var. wrightii (nash) allred, r, m bothriochloa saccharoides (sw.) rydb., m bouteloua hirsuta lag. var. hirsuta, m bromus japonicus thunb. ex murray, r bromus lanatipes (shear) rydb., r, m cenchrus incertus m.a. curtis, r cenchrus longispinus (hack.) fernald, m dichanthelium oligosanthes (schult.) gould, r digitaria californica (benth.) henrard, r digitaria cognata (schult.) pilg., r echinochloa crus-galli (l.) p. beauv., m 76 oklahoma native plant record volume 15, december 2015 amy k. buthod and bruce w. hoagland enneapogon desvauxii p. beauv., r eragrostis curtipedicellata buckley, r eragrostis intermedia hitchc., r eragrostis secundiflora j. presl, r eragrostis sessilispica buckley, r eragrostis trichodes (nutt.) alph. wood, m hesperostipa comata (trin. & rupr.) barkworth, r, m leptochloa dubia (kunth) nees, r muhlenbergia arenicola buckley, r muhlenbergia racemosa (michx.) britton, sterns & poggenb., r, f phalaris caroliniana walter, r phragmites australis (cav.) trin. ex steud., r poa nemoralis l., r poa pratensis l., r setaria leucopila (scribn. & merr.) k. schum., m sphenopholis obtusata (michx.) scribn., r tridens muticus (torr.) nash var. elongatus (buckley) shinners, r triplasis purpurea (walter) chapm., r vulpia octoflora (walter) rydb., r, m polemoniaceae giliastrum rigidulum (benth.) rydb., f polygonaceae polygonum ramosissimum michx. , m pteridaceae astrolepis sinuata (lag. ex sw.) d.m. benham & windham ssp. sinuata, r cheilanthes feei t. moore, r, m cheilanthes lanosa (michx.) d.c. eaton, m pellaea atropurpurea (l.) link, r, m ranunculaceae clematis hirsutissima pursh var. scottii (porter) r.o. erickson, m ranunculus cymbalaria pursh, r rhamnaceae ceanothus herbaceus raf., r rosaceae fallugia paradoxa (d. don) endl. ex torr., r physocarpus monogynus (torr.) j.m. coult., r, m prunus americana marshall , m rosa woodsii lindl., f rubiaceae galium texense a. gray, m oklahoma native plant record 77 volume 15, december 2015 amy k. buthod and bruce w. hoagland salicaceae salix interior rowlee, m selaginellaceae selaginella densa rydb., r solanaceae solanum nigrum l., r tamaricaceae tamarix gallica l., r, m urticaceae parietaria pensylvanica muhl. ex willd., m verbenaceae verbena plicata greene, r vitaceae parthenocissus quinquefolia (l.) planch., m vitis acerifolia raf., r, f, woodsiaceae woodsia oregana d.c. eaton, r, m contributions to the flora of cimarron county and the black mesa area by ms. amy k. buthod and dr. bruce w. hoagland journal of the oklahoma native plant society volume 13, december 2013 oklahoma native plant record 3 volume 13, december 2013 foreword this year we have a special critic’s choice essay, “a cavalcade of field botanists in oklahoma” by ron tyrl and paula shryock. it is a tribute to all those botanists who have contributed to the database of native plant species in oklahoma. many are native oklahomans. many have been members of the oklahoma native plant society. all have been dedicated to documenting the state’s botanical diversity. they have been compiling biographical and professional information to write this article for us for several years. we are excited to share it with you in volume 13. one of those botanists ron tyrl writes about is connie taylor, author of our historical article in this volume. a professor emeritus at southeastern oklahoma state university, she submitted her master’s thesis to the university of oklahoma in 1961, based on research done in the water canyon complex. it is a taxonomic comparison of the canyons, based on their ecology; their geology, microclimate, and habitat. she has revised her thesis and updated her species list so that we can make the data globally available. her figures and detailed descriptions of the canyons help the reader picture the landscape and hypothesize about the causes and effects of the environmental conditions with which the species interact. data from this study could provide an important building block for long term ecological studies and future research in climate change effects. while many researchers have reported on the effects of fire, a common environmental factor in oklahoma, stan rice and sonya ross have been looking at several different effects of fire, including the effects of chemicals in smoke on plant establishment after a fire. it is a preliminary report that looks at germination rates of phacelia strictifloria seeds that have been watered with smoke-produced chemicals dissolved in water. also in this issue, amy buthod, of the oklahoma biological survey and oklahoma natural heritage inventory, provides us with another valuable flora; that of the oxley nature center in mohawk park in northwestern tulsa county. this study has provided voucher specimens for oxley personnel to use in education and outreach, as well as a species list for their use and for your enjoyment when you visit there. open access journals published by non-profit or not-for-profit organizations can provide a quick way to share data and bring valuable feedback to the author. they are a valid way of ensuring conflict of interest is not the issue that it can be when publication is profit driven. however, it depends heavily upon authors to review each other’s articles and upon volunteer editorial staff to proof-read and format them. please volunteer to write, review, or serve on the editorial board of the oklahoma native plant record. we need your help to get more of oklahoma’s native plant research out to the world. remember to tell everyone that all volumes of the record are now available online through oklahoma state university’s edmon low library as an e-journal publication. our abstracts are indexed by the centre for agricultural biosciences international (cabi). the journal is listed in the directory of open access journals (doaj). it can be accessed globally at http://ojs.library.okstate.edu/osu/ sheila strawn managing editor http://ojs.library.okstate.edu/osu/ journal of the oklahoma native plant society, volume 2, number 1, december 2002 54 oklahoma native plant record volume 2, number 1, december 2002 schoenoplectus hallii and s. saximontanus 2000 wichita mountain wildlife refuge survey dr. lawrence k. magrath curator-usao (ocla) herbarium chickasha, ok 73018-5358 a survey to determine locations of populations of schoenoplectus hallii and s. saximontanus was conducted at wichita mountains wildlife refuge in august and september 2000. one or both species were found at 20 of the 134 locations surveyed. a distinctive terminal achene character was found specifically that the transverse ridges of s. hallii appeared to be rounded and s. saximontanus appeared to be rounded with a projecting narrow wing. basal macroachenes have not yet been properly described but are borne singly at the base of each culm and are about 3-4 times larger than the terminal achenes. it is speculated that amphicarpy may be related to grazing pressure, the basal macroachene being produced even if the upper portion is consumed, as a response to grazing. both species are grazed/disturbed by bison, elk, and longhorns on the refuge. introduction a survey to determine locations of populations of schoenoplectus hallii (a. gray) s.g. smith (hall’s bulrush) and s. saximontanus (fernald) j. raynal (rocky mountain bulrush) was conducted on the wichita mountains wildlife refuge during late august through september 2000. the survey team members were myself, sam waldstein, refuge manager; chip kimball, range biologist; and bob timberman, biology technician. sites selected for observation were areas similar to the jed johnson dam habitat, which is the site of the original collection of s. hallii made in 1987. selection of sites to be surveyed was done by wmwr team members. the three sundays spent in the field were some of the hottest of the season with temperatures ranging from 100o to 110of. general observations the presence of schoenoplectus halllii and s. saximontanus at the various sites sampled. represent, for the most part, a response to the drawdown of water levels in the various lakes and ponds. the plants seem to occur mainly on magrath, l.k. https://doi.org/10.22488/okstate.17.100013 the drawdown mud, sand, or gravel flats. however in some places they occur in shallow water up to a depth of about a foot [30.5cm]. they seem to compete with perennial emergent plants and with most emergent annuals. in addition to the 36 sites that i personally examined, wmwr staff examined an additional 98 sites with similar habitat and found no schoenoplectus present. schoenoplectus occurred in only 20 of the 134 sites (14.93%). of those 134 sites, 4 had both species present (3%), 14 had s. halllii present (10.45%) and 10 had s. saximontanus present (7.46%) (see table). based on the wmwr observations, it is strongly advised that the adjacent area of fort sill should be inventoried since these two species are, most likely, present there. however, rahmona thompson who has conducted plant surveys at fort sill has not found either species at this time (pers. comm. 2002). at several of the sites plants had been uprooted as a result of trampling by bison and other animals in the mud flat areas. the number of uprooted plants ranged between 0.0% and 1.0% at those oklahoma native plant record 55 volume 2, number 1, december 2002 magrath, l.k. locations, but in general was less than .05%. while this obviously does cause some damage to populations in localized areas the damage appears to be negligible and it is even possible that it may be an important transport method of matured achenes to previously uncolonized areas. this could readily be accomplished by mud containing seeds or inflorescences with seeds adhering to the hooves, hair, or skin of the lower part of the animals’ legs. there appear to be three useful characters associated with the achenes that can be used to separate the two species: style branching, achene cross section, and transverse ridges. s. hallii s. saximontanus style branches mostly bifid trifid achene cross section unequally biconvex (1 of 2 sides may be flat) trigonus (3-sided) transverse ridges rounded mostly rounded with narrow wing to my knowledge, “transverse ridges” has not been mentioned in the literature on these two taxa. mckenzie (1988) does not mention this character in his status assessment report on s. hallii, nor does yatskievych (1999) in steyermark’s flora of missouri. dr. marian smith at southern illinois university is working on both terminal achenes and basal macroachenes using scanning electron microscopy and more precise measuring techniques (pers. comm. 2001). so this may be a new useful character to separate these two taxa. (see figures). mckenzie (1998) reports that “heavy grazing has been noted at sites in kansas, missouri, and wisconsin but it is not known whether this disturbance negatively impacts the species…” it is my hypothesis that amphicarpy may well be a response to heavy grazing pressure disturbance by native grazing animals such as bison, elk, and deer prior to the introduction of exotic grazers such as cattle, horses and sheep by european settlers. it would be a way that the plant could insure seed production even if it is heavily grazed and the terminal achenes damaged or destroyed. the observation that the plant produces only a very limited number of basal macroachenes (one per culm) with significantly more food reserves per achene would seem to support this interpretation. the smaller, more numerous terminal achenes would offer a relatively easy method for seeds to be transported farther distances and to new sites while the macroachenes would provide an excellent way to persist in presently occupied sites. the number and identification of the two species at sites where both occur could be somewhat problematic. s.g. smith and a. e. schuyler independently identified a hybrid from wmwr that was collected by m. smith and mckenzie july 28, 2002 as the first documented, putative hybrid of the two species ever recorded. [usa oklahoma: comanche co., wmwr, 28 july 02, p. mckenzie 2028 pers., wis, mo] (pers. comm. mckenzie 2002; smith 2002). there obviously needs to be further research done to confirm or deny this hypothesis as well as possible hybridization between the two species at the refuge. voucher specimens of schoenoplectus hallii and s. saximontanus resulting from this study are kept in the usao herbarium (ocla) at the university of science and arts of oklahoma in chickasha, oklahoma. 56 oklahoma native plant record volume 2, number 1, december 2002 magrath, l.k. fig. (a) and (b) fruit and inflorescence of schoenoplectus saximontanus (steyermark’s flora of missouri, 1999. used by permission). fig. (e), (f), and (g) fruit, inflorescence, and habit of schoenoplectus hallii (steyermark’s flora of missouri, 1999. used by permission). oklahoma native plant record 57 volume 2, number 1, december 2002 magrath, l.k. table schoenoplectus occurrence in thirty-six survey sites august and september 2000 (based on achene characteristics of the two species) site # description & date abundance/# collected/voucher# s. hallii s. saximontanu 1-pond 27-aug draw-down mudflat around pond some plants grazed, some uprooted some dried out on shore few floating in water still alive 0 scat/8/21254 2-pond 27-aug draw-down mudflat 0 0 3-pond 27 aug draw-down mudflat 0 0 4-corral area 27 aug stream with draw-down mudflat utricularia observered 0 0 5-kiowa lake 27 aug draw-down mud flat around pond some plants grazed, some uprooted some dried out on shore few floating in water still alive 0 scattered/4/21259 6-wing pasture west of creek 27 aug draw-down mud flat around pond some plants grazed 0 rare/12/21261 7-buford lake 27 aug draw-down mud flats utricularia and nelumbo abundant 0 0 58 oklahoma native plant record volume 2, number 1, december 2002 magrath, l.k. 8-quanah parker lake 27 aug near nature center draw-down mud flat around pond scat/6/21263 0 9-new pond by crater lake 27 aug south of visitor center draw-down mud flat 0 0 10-pond at sulphur trap corrals 27 aug east of visitor center draw-down mud flat 0 0 11-pond in sulphur trap 27 aug north of visitor center draw-down mud flat around pond 0 0 12-jed johnson lake 18 aug 8 identifiable plants. not collected plants just coming into bloom original site for the original collections for wmwr rare/0/0 27 aug 8 identifiable plants no collections made rare/0/0 13-crater lake 27 aug draw-down mud flat apparently too much perennial vegetation present 0 0 14-west gate pond 3 sep draw-down mud flat apparently too much perennial vegetation present 0 0 15-comanche lake 18 aug draw-down mud flat scat/2/21237 0 oklahoma native plant record 59 volume 2, number 1, december 2002 magrath, l.k. 3 sep draw-down mud flat mixed populations scat-loc com/24/21272 scat-locom/83/27271 16-grama lake near dam & gram flat 18 aug draw-down mud flat several hundred plants numerous basal rosettes in shallow water (20-30 cm) 0 scat/3/21236 3 sep mixed populations on draw-down several hundred plants numerous submerged basal rosettes in shallow water (20-30 cm) scat-loc com/48/21274 scat-loc com/15/21273 10 sep revisit & complete walk-around both present on draw-down mud flat several thousand plants dominant plant in a few places numerous basal rosettes present in shallows in several areas scat-loc abd/211/21216 scat/99/21322 17-hollis lake 3 sep apparently too much perennial vegetation in draw-down zone 0 0 18-pond 0.5 mile west of hollis 3 sep com-loc abd/300+/21278 0 19-pond 3 sep draw-down mudflat 0 0 20-pond 3 sep draw-down mudflat 0 0 21-boggy flats 18 august 2000 draw-down mudflat around pond some plants grazed or uprooted some dried out on shore few floating in water still alive first located by sam waldenstein on aug 17, 2000. 0 loc abd/x/21231 60 oklahoma native plant record volume 2, number 1, december 2002 magrath, l.k. sun 3 september 2000 draw-down mud flats on ponds 0 loc abd/5/21280 22-pond southwest of grace mountain 3 sep draw-down mud flat water-clover fern present 0 0 23-cut throat lake 3 sep draw-down mud flat water-clover fern present beautiful clear water and bass 0 0 24-northwest corner of pinchot loop 3 sep draw-down mud flat around pond 0 0 25-barow pit east side of pinchot loop 3 sep draw-down mud flat around pond 0 0 26-straight east of site 25 3 sep draw-down mud flat around pond bacopa present scat/8/21287 0 27-medicine tank 3 sep draw-down mud flat around pond colonial bryozoan pectinatella magnifica leidy in shallow water by dam identification by dr. mike mather, usao common/12/21290 scat/3/21289 28-west gate road, buffalo gap 10 sep draw-down mud flat around pond bacopa present 0 scat/11/21303 29-winter valley at end of wing fence at hot trap 10 sep draw-down mud flat around pond mixed collection not discovered until laboratory observations were made. rare/3/21304 in part* rare/6/21304 in part* oklahoma native plant record 61 volume 2, number 1, december 2002 magrath, l.k. 30-winter valley southeast fo road 10 sep draw-down mudflat around pond bacopa present scat/2/21305 0 31-north end of research 10 sep draw-down mud flat around pond bacopa present 0 scat/14/21306 32-pond in exhibition pasture 10 sep pond in exhibition pasture draw-down mud flat one plant/0/0 0 33 ingram house pond 10 sep draw-down mud flat several hundred plants bacopa present scat-loc com/19/21309 0 34-quanah parker 10 sep near dam draw-down mud flat several thousand? plants scat-loc com/54/21210 0 35-quanah parker 10 sep se of environmental center draw-down mud flat several hundred plants scat-loc com/7/21313 scat-loc com/25/21311 36-elmer thomas lake 10 sep draw-down mud flat along north shore scat-loc abd/46/21314 0 key to abundance descriptors rare = fewer than 10 plants at site scat = scattered, a few plants occurring over several square meters common = many plants occurring over several square meters abundant = large numbers of plants (often the local dominant plant) loc com = many plants in a small area, but may be scattered over a large area loc abd = large numbers of plants in small area, but may be scattered over a large area 62 oklahoma native plant record volume 2, number 1, december 2002 magrath, l.k. references mckenzie, paul m. 1998. hall’s bulrush (schoenoplectus hallii) status assessment. u.s. fish and wildlife service. columbia, mo. yatskievych, george. 1999. steyermark’s flora of missouri, vol 1, rev. ed. missouri dept. of conservation in cooperation with the missouri botanical garden press. st. louis, mo. editor’s note: exact locations of individual sites were determined by gps and are recorded in the wmwr database. however, that information and access to the sites is strictly limited and permission must be requested from the refuge manager. oklahoma native plant record 63 volume 2, number 1, december 2002 magrath, l.k. wichita mountains wildlife refuge, comanche county echinocerus baileyi on elk mountain trail (photos by sheila strawn) oklahoma native plant record 3 volume 1, number 1, december 2001 waterfall, u.t. https://doi.org/10.22488/okstate.17.100003 the spermatophyta of oklahoma county, oklahoma exclusive of the grasses, sedges and rushes a thesis approved for the department of botany and bacteriology university of oklahoma graduate school by u. t. waterfall norman, oklahoma, 1942 chapter i introduction this paper represents a preliminary taxonomic study of the flowering plants indigenous to oklahoma county. collections during the springs, summers and falls of 1939, 1940, and 1941 and also during the spring of 1942. after the first general but extensive collections were made a number of special stations of widely varying ecological structures were selected. collections were made from these at regular intervals of about two weeks throughout the growing season, or at a corresponding time during the next year. in addition a search was made for stations containing different, ecological elements. thus the finding of a maximum number of species over a limited period of time was assured by a combination of extensive and intensive methods of collection. the specimens were pressed in the standard way used in the leading herbaria. duplicates were obtained in nearly every case and were deposited in the bebb herbarium of the university of oklahoma. among the most outstanding of the recent investigations, which may be applied to the flora of this region, are fernald's series of "virginia" papers published annually in rhodora since 1935. fernald reported1 that a number of wide-ranging continental plants were first collected in virginia by john clayton. they were described by the italian botanist gronovius in his flora virginica (1739), and later given binomial designation by linneas in the species plantarum. thus the type locality for these linnean species, which are based on clayton's material, is in southeastern virginia. collections from that region were often found to differ from the wider-ranging inland plants referred erroneously, by most botanists, to the linnean species. fernald’s restudy of many of these types has shown that the variety occurring in a restricted range along the coast is usually the typical one, i.e., the variety which gronovius had before him when writing the description upon which linneas based his generic and specific name, while the wide-ranging plant of the interior must, in the large majority of cases, be given a new varietal name. a similar situation has been found to be true for plants collected along the coast and named by other botanists. this will help to account for the appearance of many of the varietal designations in this paper that are not found in the existing floras and manuals pertaining to oklahoma. 1ferna1d and griscom, three days of botanizing in southwestern virginia, rhodora 37, pp. 129-13l, 1935. 4 oklahoma native plant record volume 1, number 1, december 2001 waterfall, u.t. chapter ii history one of the first americans to traverse what is now oklahoma county was washington irving, who with charles latrobe and his fellow travelers, made a trip through this region in the fall of 1832. irving, in his tour on the prairies, recorded the events and his impressions of this trip. his companion, latrobe1, may have had some botanical training as he mentioned various genera of plants seen on the journey. the party approached the present site of edmond on the 23rd of october. their line of search took them past the sites arcadia, spencer, oklahoma city and, on the 28th, over what is now the southern boundary of oklahoma county in the direction of moore2. irving gave a good description of the post oak-blackjack woods, even mentioning the dwarf oak, quercus prinoides, although not by name3, and of the prairies he saw when emerging from the woodland near oklahoma city4. he also tells of the cottonwoods, sycamores and willows found along the streams5. josiah gregg, a santa fe trader, in commerce of the prairies (1844), told of eight expeditions across the prairies. two of these were along the course of the canadian river, hence probably through oklahoma county. he also described the "cross timbers" (post oak-blackjack associes), dwarf oaks and prairie fires.6 [see editor’s note at end.] sitgraves and woodruff, with s. w. wodehouse as naturalist, surveyed the northern boundary of the creek indian country in 1849 and 1850, returning to ft. gibson by way of the north canadian.7 bigelow (1856) discussed the vegetation of oklahoma as seen in traveling from east to west. he also mentioned briefly the "cross timbers."8 a large number of plants from oklahoma county have undoubtedly been collected by thomas r. stemen and w. stanley meyers in the course of their investigations on which the oklahoma flora9 is based. these have not been available for study by the author. 1charlee latrobe, rambler in north america, (excerpts in irving's tour on the prairies, edited by joseph b. thoburn and george c. wells. xxv. harlow pub1ishing company, oklahoma city, oklahoma, 1930). 2washington irving, tour on the prairies, (1.c.), pp. 240-243. 3ibid., p 145. 4ibid., p. 173. 5ibid., p. 151. 6w. e. bruner, the vegetation of oklahoma, ecological monographs vol. 1., no. 2, p. 128, april, 1931. 7ibid. 8ibid. 9thomas r. stemen and w. stanley myers, oklahoma flora, harlow publishing corporation, oklahoma city, oklahoma, 1937. chapter iii physical features location and size oklahoma county is in central oklahoma, being a part of the region known before the run as "old oklahoma". it is bounded on the north by logan county, on the east by lincoln and pottawatomie counties, on the south by cleveland county and on the west by canadian county. it is rectangular in shape, extending thirty miles from east to west, and twenty-four miles from north to south. it covers an area of 720 square miles. the total population of the county is 224,159. oklahoma city has a population of 204,424. oklahoma native plant record 5 volume 1, number 1, december 2001 waterfall, u.t. edmond is next with 4,002, while bethany has 2,590 and britton 2,239. other towns in the county, all under 1,000 in population, are harrah, arcadia, luther, nicoma park, newalla and marion. topography the county is drained chiefly by the north canadian river and its tributaries. the majority of the creeks, especially the small ones in this drainage system have water running in them only during the spring and after rains during the rest of the year. especially in the hot summer and early fall months one is apt to find them dried up. a tier of sections along the southern boundary south and southwest of oklahoma city are in the watershed of the south canadian river. the north canadian enters the county west of oklahoma city. here it has been dammed to form lake overholser, which furnishes the city's water supply. it runs through the southern part of oklahoma city, then swings northeast, through spencer into the central part of the county. it then bends southeast, to leave the county near harrah, l8 miles east of oklahoma city, having curved 10 miles north between these two places. the eastern part of the county is made up of sandy oak-covered hills and small prairies, together with outcroppings of red sandstone. the western townships are, for the most part, rolling prairie. near bethany there is a region of aeolian sand hills1, which support a vegetation similar to that found on the sandy soils in the eastern part of the county. geology and soils oklahoma county is in the permian system of rocks,2 which has been called the permian redbeds. the western half of the county is in the lower part of the enid formation of the permian system.3 this system consists of layers of thin red sandstones and soft red shales. the soil in the western part of the county is a prairyerth4, which is a mature soil composed mostly of clay, but containing some sand. near bethany and the northern part of lake overholser there is a small area of aeolian sandhills.5 the eastern part of the county is covered with a residual sandy soil.6 running through the prairyerths and the sandy soils is another type of transported soil. this is alluvial soil7 found chiefly along the north canadian river and its tributary creeks. 1c. e. thornwaite, map of soils, university of oklahoma (unpublished). 2hugh d. miser, geologic map of oklahoma, u.s. geologic survey, 1926. 3ibid. 4c. w. thornwaite, op. cit. 5ibid. 6ibid. 7ibid. chapter iv climate the climate of oklahoma county is of the continental type modified to some extent by winds from the gulf of mexico. the annual range in temperature is, therefore, rather marked. the summer temperatures are quite high, while in winter there are often cold spells when the thermometer hovers near zero for several days. in summer there are often droughts of several weeks duration. 6 oklahoma native plant record volume 1, number 1, december 2001 waterfall, u.t. the prevailing winds are from the south with an average velocity at oklahoma city of 11.3 miles per hour.1 the monthly average at this station varies from slightly more than 9 miles per hour in august to nearly 14 miles per hour in march and april. the highest wind velocity recorded here for a five-minute period was 57 miles per hour on june 24, 1915, and again on june 29, 1918. temperature records have been kept in oklahoma city since 1891. between that year and 1941 inclusive, the average temperature for january was found to be 37.5 degrees f. for july it was 81.3 degrees. the maximum temperature recorded over this fifty-one year period was 113 degrees on august 11, 1936, and the minimum was -17 on february l2, 1899. the average date of the last, killing frost is march 29, while in the fall the average date of the first killing frost is november 5, giving a growing season of 221 days. the average annual precipitation is 31.37 inches. february is the driest month of the year, having an average precipitation of 1.13 inches, while may is the wettest month with an average of 4.89 inches. the largest total monthly precipitation was in june 1932, when 14.12 inches were recorded. this was 10.40 inches more than the average for this month, and 2.13 inches more than may 1902, the wettest month previous to this time. the wettest year was 1902 when there were 52.03 inches of precipitation. 1annual meteorological summary with comparative data, 1941, compiled under the direction of h.f. walgren, weather bureau office, oklahoma city. published in oklahoma city, 1942. chapter v ecology oklahoma county has two principal vegetational regions. the eastern three-fifths of the county is occupied by a post oak-black jack post [sic] climax, while the western part supports a mixed grass formation. in addition there is to be found a flood-plain forest of a distinct nature running along the streams through both the prairie and the savanna. the range of the latter two is determined by the type and texture, as well as by the ph of the soil. the oak savanna is found in sandy soil, which would show a high ph value, while the mixed-grass prairie begins abruptly in the finer-textured clay soils of a lower ph value. since these soils occur in intermixed spots, zones and belts where they merge together, their resulting vegetational expressions are similarly interrupted, although separated from one another. the dominants in the savanna are quercus marilandica and q. stellata. there is not much hickory associated with these two oaks although some plants of carya buckleyi var. arkansana may befound. the association of grasses in the true prairie in the western part of the county has been called by bruner 1 the stipa-koeleria association after the grasses that are dominant in the prairie states farther north. in our area, however, their places have been taken to a large extent by the bluestems of southern origin.2 the chief dominants are species of andropogon and bouteloua. the flood-plain forests are characterized by a populus-salix associes*. in association with these is often found cephalanthus occidentalis, while farther from the water ulmus americana and u. fulva always occur, often with celtis laevigata and rather scattered specimens of prunus mexicana. oklahoma native plant record 7 volume 1, number 1, december 2001 waterfall, u.t. the forests often merge into the prairie with a narrow band of chaparral consisting of characteristic shrubs. between the flood-plain forests and the grassland these are usually rhus glabra, r. copallina var. latifolia, diospyros virginiana, symphoriocarpos orbiculatus, and prunus augustifolia if any sand is present. the ecotone between the post oak-black jack associes and the prairie is characterized by quercus prinoides, symphoriocarpos orbiculatus, rhus copallina var. latifolia, rhus glabra, and prunus angustifolia var. watsoni. some of these shrubs are common to both ecotones, but sambucus canadensis is characteristic of the transition from flood-plain forest to prairie, while quercus prinoides is found only in the ecotone between the post oak-black jack associes and the prairie. there are two common disclimaxes, or disturbance climaxes, present. one is made up of cultivated crops.3 here man determines what the climax vegetation shall be. the second consists of overgrazed pasturelands. where this condition exists in the prairies the taller grasses are replaced by bouteloua hirsuta and b. gracilis associated with buchloe dactyloides. thus the pasture assumes the aspect of the short grass plains farther west. between the two disclimaxes there is often very little of the original vegetation left. the botanist is often confined to following railroad tracks or searching for out-of-the-way corners if he is to find much of interest. even in the post oak-black jack woods overgrazing has played a destructive part. in some cases about all that remains is buck brush. the overgrazed prairies are characterized not only by the short grass species already mentioned, but also by such weedy inedible species as vernonia baldwinii var. interior, achillea lanulosa, gutierrezia dracunculoides, artemesia gnaphalodes, and cirsium undulatum. in fact these species always serve as indicators of overgrazing. their prominence in a pasture or field should be a warning to the farmer or cattleman to decrease the number of cattle pastured in a given area, or to change pastures long enough to allow the original vegetation to assume its normal dominance. it is interesting to note that different species of the same genus may be used as indicators of soil types. thus tradescantia occidentalis is found, in the prairyerths while t. canaliculata grows in the sands. liatrus punctata may grow in clay soil, but l. squarrosa var. intermedia is found in sandy soil or on sandstone outcroppings of soils that may contain some clay. lithospermum incisum is found in clay, but l. caroliniense grows only in sandy soil in the post oak-black jack associes. other species, characteristic of sandy soil, are psoralea cuspidata, p. villosum, ipomoea leptophyllum, and penstemon laxiflorus. prevernal societies on the prairies include anemone caroliniana, claytonia virginica, houstonia minima, lithospermum incisum, draba brachycarpa, d., reptans, northoscordum bivalve, and androsace occidentalis. forming a succession on previously cultivated soil one finds stel1aria media, viola kitabeliana var. rafinesquii, capsella bursa-pastoris, taraxacum laevigatom, and lamium amplexicaule. in the post oak-black jack region the common plants of this society are antennaria fallax, sagina decumbens, and a small sedge, carex microrynchia. scattered 8 oklahoma native plant record volume 1, number 1, december 2001 waterfall, u.t. individuals of viola papilionacea grow along the creeks through such environments but there is not the abundance of forbs which may be found in the prairie. some of the conspicuous components of the vernal societies on the prairies are sisyrinchium bushii, baptisia leucophaea, b.australis var. minor, senecio plattensis, valerianella stenocarpa var. parviflora, tradescantia occidentalis, callirhoe involucrata, specularia biflora, linum lewisii var. pratense, and achillea lanulosa. forbs now form a more conspicuous component of the postoak-black jack flora. they include lithospermum caroliniense and astranthium integrifolium var. ciliatum. in more open spots and along fields and roadways coreopsis grandiflora, schrankia unciniata, penstemon laxiflorus and tradescantia canaliculata are often found abundance. in succession on disturbed soils often occur linaria canadensis var. texana, lepidium densiflorum, l. virginicum, silene antirrhina, descurainia pinnata var. brachycarpa, and chaerophyllum tainturieri var. floridanum. prairie aestival societies include petalostemum purpureum, p. candidum, psoralea floribunda, sabatia campestris, coreopsis tinctoria, rudbeckia hirta var. sericia, rudbeckia amplexicaulis, oenothera serrulata, ruellia caro1iniensis, krameria secudiflora, amorpha canescens, acacia angustissima var. hirta, ratibida columnifera,4 asclepiodora decumbens, thelesperma trifidum, physalis mollis, solanum eleaegnifolium and solanum torreyi. in the oak postclimax are found petalostemum villosum, psoralea cuspidata, galium pilosum var. puncticulosum, and ruellia caroliniensis. in succession on disturbed areas are found helianthus annuus, h. petiolaris, croton monanthogynous, c. capitatus, c. texense and several species of polygonum including p. punctatum, p. opelousanum, and p. muhlenbergii. cardiospermum halicacibum is also abundant here. several species of vitis in combination with ampelopsis cordata, and parthenocissus quinquefolia form lianas. commelina erecta var. typica is a species tolerant of shade, which can be found under those layers. some of the serotinal prairie dominants are euphorbia marginata, gutierrezia dracunculoides, liatrus punctata, chrysopsis berlandieri, aster ericoides, aster oblongifolius var. rigidulus, vernonia baldwinii var. interior, solidago radula, artemisia gnaphalodes, ambrosia coronopifolia, helianthus maximi1ianus and heterotheca aubaxillaria. growing in the post oak-black jack associes one finds desmondium marilandicum, d. paniculatum, aster patens var. gracilis and acalypha gracilens. common along the wooded creek sides are acalypha rhomboidea, ambrosia trifida var. texana,aster drummondii, aster exilis, verbesina virginica, solidago petiolaris, irensine rhizomatosa and euphorbia heterophylla. in summing up the ecological aspects of the county one finds that it lies in a climate favorable to the development of a grassland formation, but due to the presence of sand the eastern three-fifths of the area is largely covered by a post oak-black jackpost climax. a second post climax is the floodplain forest found along the north canadian river and its tributaries. two disclimaxes are present, one caused by overgrazing, the other by cultivation. oklahoma native plant record 9 volume 1, number 1, december 2001 waterfall, u.t. 1w. e. bruner, the vegetation of oklahoma, ecological monographs, vol. 1, no. 2, pp. 110-111, april, 1931. 2w.e. bruner, the vegetation of oklahoma, ecological monographs, vol. 1, no. 2, l.c. 3weaver and clements, plant ecology, pp. 86-89, mcgraw-hill book company, inc, new york, 1938. 4w. w. fernald, new species, varieties and transfers, rhodora 40: 353, 1938. chapter vi range extensions and plants new to the county in the course of the investigations on which this study is based several plants were collected which have been previously unrecorded from the state. these include typha truxillensis,1 medicago minima, gaura filiformis var. typica2, achillea lanulosa forma rubicunda and tragopogon major.3 the latter has since been found in several sections of the state. the pink-rayed form of achillea lanulosa is fairly common, but apparently has escaped previous notice. eloecharis parvula, var. anachaeta was collected near oklahoma city, definitely establishing its occurrence within the state. in his monograph4 svenson included oklahoma in the mapped range of var. anachaeta (map 3, page 387) but no specimens we recited from our area. this leads one to conclude that svenson assumed the presence of the variety in oklahoma, but had no actual specimens from the state. herbarium sheets were cited by him from iowa, colorado, new mexico and texas, but from kansas, nebraska, missouri and oklahoma he had seen no material. cyperus rivularis was found in the eastern portion of the county. it seems to be a rarely collected species. dr. f. j. hermann of the u.s. department of agriculture has seen no material from oklahoma. dr. hugh o'neill of the catholic university of washington, d.c., writes5 that he has seen only two sheets from the state, both of which are in the gray herbarium of harvard university. this station is west of the range as given in all the published floras and manuals. xyris torta was an unusual "find". my station in the southeastern part of the county seems to be the identical one from which dr. milton hopkins of the university of oklahoma collected this species two years earlier. at any rate this appears to be the most northwestern station in the state. acer negundo var. interior, previously unrecorded from the state was found along the north canadian river in the extreme eastern part of the county near harrah. bergia texana, collected north of oklahoma city, is neither listed by jeffs and little in their check list, nor by stemen and myers in the oklahoma flora. however, its occurrence was to be expected as it falls within the range as given by rydberg's flora. professor m. l. fernald wrote6 that there is a sheet in the gray herbarium "from arkansas, indian territory, september 28, 1894, b.f. bush, no. 33". in an investigation of ambrosia aptera and ambrosia trifida7 the author came to the conclusion that all of our specimens should be reduced to varietal rank. professor fernald8 agreed that this entity should be accorded varietal status as ambrosia trifida var. texana scheele, the first available varietal designation. 10 oklahoma native plant record volume 1, number 1, december 2001 waterfall, u.t. some highly localized species were found in the southeastern part of the county. one station where several were found was in marshy springy soil surrounding a small lake about three miles south of harrah. the lake had been made by damming a small creek, but presumably the spring and marsh, and hence the species characterizing them, were inexistence previously. here were found cyperus rivularia, agrimonia parviflora, rotala ramosier var. interior, prunella vulgaris var. lanceolata, mimulus ringens, and mimulus glabratus var. oklahomensis. growing in alluvial soil in the wooded valley of the north canadian river about a mile south of harrah were found acer negundo var. interior, ampelopsis arborea, polymnia uvedalia var. densipilis, pluchea purpurascens, and again prunella vulgaris var. lanceolata. most of these were probably at the western limit of their range. mimulus glabratus var. oklahomensis however is found farther west9, but this is the only station the author has found in the county. pluchea purpurascens may be found elsewhere. in their extreme forms, it and p. camphorata (p. petiolata) seem distinct, but there are several sheets in the bebb herbarium of the university of oklahoma which appear difficult positively to assign to either species. 1m. l. fernald, midsummer vascular plants of virginia. rhodora 37: 385-387, 1935. 2u. t. waterfall, interesting plants of oklahoma. rhodora 42: 499-502, 1940. 3ibid. 4h. k. svenson, monographic studies in eleocharis iii. rhodora 36: 386-389, 1934. 5correspondence with the author. 6correspondence with dr. milton hopkins. 7u. t. waterfall. interesting plants of oklahoma, l. c 8m. l. fernald, as editor of rhodora, in editor’s footnote to waterfall's paper. 9norman c. fassett, notes from the herbarium of the university of wisconsin xvii. rhodora, 41: 525, 1939 *ed. note: according to j.e. clements and f.e. weaver, plant ecology (p46) mcgraw hill 1929; the term associes is “the developmental equivalent of the association … used where the community is not permanent but is replaced by another in the process of development of succession”. [b.h.] oklahoma native plant record 11 volume 1, number 1, december 2001 waterfall, u.t. chapter vii spermatophyta of oklahoma county, oklahoma exclusive of the grasses, sedges and rushes (based on collections of the author) angiospermae monocotyledonae typhaceae typha truxillensis hbk typha latifolia l. alismaceae echinodorus cordifolius (l.) griseb. forma lanceolatus engelm.) fernald. xyridaceae xyris torta j. e. smith commelinaceae commelina communis l. var. ludens (miquel) clark. commelina diffusa burm. f. (c.nudiflora of authors, c. longicaulis jacq.). commelina erecta l. var. typica fern. commelina erecta l. var. typical fern., forma intercursa fern. commelina erecta l. var. angustifolia (michx.)fern. forma crispa (wooton) fern pontederiaceae heteranthera limosa (sw.) willd. liliaceae allium canadense l. allium mutabile michx. allium nuttallii wats. androstephium coeruleus (scheele) greene. asparagus officinalis l. nothoscordium bivalve (l.) britton smilax bona-nox l. smilax hispida muhl. yucca glauca nutt. amaryllidaceae cooperia drumondii herb. iridaceae sisyrinchium bushii bickn. sisyrinchium campestre bickn. sisyrinchium graminoides bickn. sisyrinchium varians bickn. orchidaceae spiranthea cernuus l. dicotyledoneae salicaceae populus deltoides marsh. salix interior rowlee. salix interior rowlee var. wheeleri rowlee. salix nigra marsh. juglandaceae carya buckleyi durand var. arkansana sarg. carya pecan (marsh) engler and graebner. juglans nigra l. fagaceae quercus bicolor willd. quercus macrocarpa michx. quercus marilandica moench. quercus prinoides willd. quercus stellata wang. urticaceae boehmeria cylindrica (l.) sw. var. drummondiana weddell. celtis laevigata willd. celtis reticulata torr. maclura pomifera (ref.) schneider. morus alba l., var. tatarica (l.) loud. morus rubra l. parietaria pennsylvanica muhl. ulmus americana l. ulmus fulva michx. loranthaceae phoradendron flavescens (pursh) nutt. polygonaceae eriogonum annuum nutt. eriogonum longifolium nutt. polygonum buxiforme small. polygonum convolvulus l. polygonum cristatum engelm. & gray. polygonum dumetorum l. 12 oklahoma native plant record volume 1, number 1, december 2001 waterfall, u.t. polygonum exsertum small. polygonum lapathifolium l. polygonum longistylus small. polygonum muhlenbergii (meisn.) wats. polygonum opelousanum riddell. polygonum pennsylvanicus l. var. laevigatum fernald. polygonum punctatum ell. polygonum scandens l. polygonum tenue michx. tovaria virginiana (l.) adams. chenopodiaceae atriplex argentea nutt. chenopodium ambrosioides l., ss. eu-ambrosioides aellen. chenopodium gigantospermum aellen. (c. hybridum of am. authors). chenopodium pratericola rydb. (c. leptophyllum of most authors). chenopodium standleyanum aellen. (c. boscianum moq.in part). cycloloma atriplicifolium (spreng.) coult. monolepis nuttalliana (r. & s.) wats. salsola kali l. var. tenuifolia g.f.w. mey saueda linearis (ell.) moq. amaranthacaeae acnida tamariscina (nutt.) wood. amaranthus blitoides wats. amaranthus graeciszans l. amaranthus palmeri s. wats. amaranthus spinosus l. amaranthus torreyi (gray) benth. froelichia floridana (nutt.) moq. var. campestris (small) fern. froelichia gracillis moq. iresine rhizomatosa standley. phytolaccaceae phytolacca americana l. nyctaginaceae oxybaphus albidus (walt.) sweet. oxybaphus floribundus chois. oxybaphus hirsutus (pursh.) robinson. illecebraceae paronychia jamesii t. & g. paronychia wardii small. aizoaceae mollugo verticillata l. caryophyllaceae cerastium brachypodum (engelm.) robinson cerastium nutans raf. sagina decumbens (ell.) t. & g. silene antirrhina l. silene antirrhina l. var. divericata robinson. stellaria media (l.) cyrill. portulacaceae claytonia virginica l. claytonia virginica l. forma robusta (somes) palmer & steyermark. portulaca oleraceae l. portulaca parvula gray. (p. pilosa) talinum parviflorum nutt. nymphaeaceae castalia odorata (ait.) woodville & wood. nelumbo pentapetala (walt.) fernald. ranunculaceae anemone caroliniana walt. clematis pitcheri t. & g. delphinum virescens nutt. var. camporum (greene) martin. myosurus minimus l. ranunculus pusillus poir. ranunculus sceleratus l. menispermaceae cocculus carolinus (l.) dc. menispermum canadense l. papaveraceae argemone intermedia sweet. fumariaceae corydalis aurea willd. var. ocidentalis engelm. corydalis campestris (britton) buckholz & plamer. oklahoma native plant record 13 volume 1, number 1, december 2001 waterfall, u.t. cruciferae arabis virginica (l.) poir. brassica campestrus l. camelina microcarpa andrz. capsella bursa-pastoris (l.) medic. cardamine parviflora l. var. arenicola (britt.) o.e. schultz. chorispora tenella dc. descursinia pinnata (walt.) britton. var. brachycarpa (richardson) fern. draba brachycarpa nutt. draba cuneifolia nutt. var. helleri (small) o. e. schultz. draba reptans (lam.) fernald. erysimum repandum l. lepidium densiflorum schrad. (l. apetalum) lepidium oblongum small. lepidium virginicum l. rorippa islandica (oeder ex murr) borbas. rorippa sessiliflora (nutt.) hitchc. rorippa sinuata (nutta.) greene. sisymbrium altissima l. sisymbrium officinale scop. streptanthus hyacinthoides hook. thlaspi arvense l. capparidaceae cleomella angustifolia torr. polansia trachysperma t. & g. saxifragaceae ribes odoratum wendl. platanaceae platanus occidentalis l. rosaceae agrimonia parviflora ait. crataegus sp. fragaria virginiana duchesne, var. illinoensis (prince) gray. geum canadense jacq. prunus angustifolia marsh. var. watsoni (sarg.) waugh. prunus gracilis engelm. gray. prunus mexicana wats. rosa foliosa nutt. rubus sp. sanguicorba annua nutt. leguminosae acacia angustissima (will.) kuntze. var. hirta (nutt.)robinson. amorpha canescens pursh. amorpha fruticosa l. var. angustifolia pursh. apios americana medic. astragalus canadensis l. astragalus nuttallianus dc. astragalus plattensis nutt. baptisia australis (l.) r. br. var. minor (lehm.) fernald. baptisia leucantha t. & g. baptisia leucophaea nutt. (b. bracteata) cassia fasciculata michx. (c. chamascrista) cassia marilandica l. (c. medsgeri) cercis canadensis l. desmanthus illinoenis (michx.) macm. desmodium ciliare dc. (d. obtusum). desmodium dillenii darl. desmodium illinoense gray. desmodium paniculatum (l.) dc. var. pubens t. & g. desmodium sessilifolium (torrey) t. & g. galactia volubillis (l.) britton. var. mississippiensis vail. gleditsia tricanthos l. gleditsia tricanthos l.forma inermis c.k. schneider glycyrrhiza lepidota (nutt.) pursh. gymnocladus diocica (l.) koch. hosackia americana (nutt.) piper. indigofera leptosepala nutt. krameria lancolata torr. lespedeza capitata michx. lespedeza intermedia (l.) britton. lespedeza intermedia (l.) britton, forma hahnii (blake) hopkins. lespedeza procumbens michx. lespedeza repens (l.) barr. lespedeza striata (thub.) h. & a. lespedeza stuevei nutt. lespedeza stuevei nutt, forma augustifolia (britt.) hopkins. medicago lupulina l. medicago minima l. medicago sativa l. 14 oklahoma native plant record volume 1, number 1, december 2001 waterfall, u.t. melilotus alba desv. melilotus officinalis (l.) lam. neptunea lutea (leavenw.) benth. oxytropus lambertii pursh. parosela aurea (nutt.) britton. parosela enneandra (nutt.) britton. petalostemum candidum michx. petalostemum occidentale (gray) fernald. petalostemum purpureum (vent.) rydb. petalostemum purpureum (vent.) rydb. forma pubescens fassett. petalostemum villosum nutt. psoralea digitata nutt. psoralea floribunda nutt. rhynchosia latifolia nutt. robinia pseudo-acacia l. schrankia nuttallii (dc.) standley. strophostyles helvola (l.) britton strophostyles pauciflora (benth.) wats. stylosanthes biflora (l.) bsp. var. hispidissma (michx.) pollard & ball. tephrosia virginiana (l.) pers. tephrosia virginiana (l.) pers. var. holosericia (nutt.) t. & g. trifolium carolinianum michx. trifolium pratense l. vicia caroliniana walt. vicia ludoviciana nutt. vicia villosa roth. linaceae linum berlandieri hook. linum lewisii pursh. var. pratense norton. linum rigidum pursh. linum sulcatum riddell. oxalidaceae oxalis stricta l. oxalis violaceae l. geraniaceae geranium carolinianum l. zygophyllaceae kallstroemia intermedia rydb. tribulus terrestris l. rutaceae ptelea trifoliate l. zanthoxylum americanum mill. polygalaceae polygala alba nutt. polygala incarnata l. polygala verticillata l. var. isocycla fernald. euphorbiaceae acalypha gracilens gray acalypha ostryaefolia ridd. acalypha rhomboidea raf. (a. virginica l.) croton capitatus michx. croton glandulosis l. var. septentrionalis muell. arg. croton lindheimerianus scheele. croton monanthogyhous michx. croton texenis (klotzsh) muell. arg. euphorbia arkansana engelm. & gray. euphorbia chamaesyche l. (e. malaca (small) little). euphorbia corollata l. euphorbia corollata l. var. mollis millsp. euphorbia geyeri engelm. euphorbia heterophylla l. euphorbia hexagona nutt. euphorbia humistrata engelm. ex. gray. euphorbia maculata l. (e. preslii guss.) euphorbia marginata pursh. euphorbia missurica raf. (e. zygophylloides boiss.) euphorbia missurica raf. var. intermedia (engelm.) l. c.wheeler. (e. petaloidea (engelm.) l. c. euphorbia obtusata pursh. euphorbia serpens hbk. euphorbia strictospora engelm. euphorbia supine raf. (e. maculata l.) jatropha texana muell arg. stillingia sylvatica l. anacardiaceae rhus copallina l. var. latifolia engelm. rhus glabra l. rhus radicans l. oklahoma native plant record 15 volume 1, number 1, december 2001 waterfall, u.t. celastraceae celastrus scandens l. evonymis atropurpureus jacq. aceraceae acer negundo l. acer negundo l. var. interior (britton) sarg. acer negundo l. var. texanum pax. sapindaceae cardiospermum hallicacabum l. sapindus drummondii h. & a. rhamnaceae ceanothus evatus desf. var. pubescens wats. vitaceae ampelopsis arborea (l.) rusby. ampelopsis cordata michx. parthenocissus quinquefolia (l.) planchl. vitis cinerea engelm. vitis palmata vahl. vitis riparia michx. (v. vulpina auth.) vitis vulpina l. (v. cordifolia michx.) malvaceae callirhoe alcaeoides (michx.)gray. callirhoe involucrate (t.& g.) gray. hibiscus trioneum l. sida spinosa l. sphaeralcea coccinea (pursh.) rydb. hypericaceae ascyrum hypericoides l. var. multicaule (michx.) fern. hypericum multilum l. hypericum punctatum lam.var. pseudomaculatum (bush) fern. elatinaceae bergia texana (hook.) seubert. cistaceae lechea tenuifolia michx. var. occidentalis hodgdon. lechea villosa ell. violaceae viola kataibeliana roem. & schultes, var. rafinesquii greene) fern. viola missourienses greene. viola papilionaceae pursh. viola primulifolia l. var. villosa a. eaton. passifloraceae passiflora incarnata l. loasaceae mentzelia oligosperma nutt. cactaceae mamillaria similes engelm. opuntia humifusa raf. lythraceae ammannia coccines rottb. lythrum lanceolatum ell. rotala ramosior (l.) koehne. onagraceae gaura filiformis small. var. typica munz. gaura parviflora dougl. var. typica munz. garua parfiflora dougl. var typica munz. forma glabra munz. gaura tripetala cav. var. triangulata (buckl.) munz. jussiaea diffusa forsakal. ludwigia alternifolia l. ludwigia palustris (l.) ell. var. americana (dc.) fern. & grisc. oenothera canovirens steele (oe. biennis in part). oenothera lacinata hill. oenothera lacinata hill, var. grandiflora (walt.) robinson. oenothera linifolia nutt. var. typica munz. oenothera missourienses sims. var. oklahomensis (norton) munz. oenothera rhombipetala nutt. oenothera serrulata nutt. var. typica munz. oenothera serrulata nutt. var. drummondii t.& g. forma flava munz. oenothera speciosa nutt. 16 oklahoma native plant record volume 1, number 1, december 2001 waterfall, u.t. oenothera triloba nutt. stenosiphon linofolium (nutt.) britton. umbelliferae ammoselinum butleri (engelm.) coult & rose. chaerophyllum tainturieri hook. var. floridanum coult. & rose. chaerophyllum texanum coult. & rose cicuta maculata l. daucus pusillus michx. lomatium daucifolium (nutt.) coutl. & rose. lomatium foeniculaceum (nutt.) coult. & rose. pastinaca sativa l. polytaenia nuttallii (dc.) britton. ptilimnium capillaceum (michx.) raf. sanicula canadensis l. spermolepis divericata (watt.) britton. spermolepis echinata (nutt.) heller. spermolepis inermis (nutt.) mathias & constance (s. patens). torilis japonicus (houtt.) dc. (t. anthriscus (l.) bernh.) cornaceae cornus drummondii meyer. (c. asperifolia of authors). primulaceae androsace occidentalis pursh. samolus pauciflorus raf. (s. floribundus hbk.) sapotaceae bumelia lanuginose (michx.) pers. ebenaceae diospyros virginiana l. diospyros virginiana l. var. platycarpa sarg. oleaceae fraxinus pennsylvanica marsh. var. americana (borkh.) sarg. loganiaceae polypremum procumbens l. gentianaceae sabatia angularis (l.) bursh. sabatia campestris nutt. apocynaceae apocynum cannabium l. var. glaberrimum a. dc. apocynum cannabium l. var. pubescens (r. br.) a. dc. apocynum sibiricum jacq. apocynum sibiricum jacq. var. farwellii (greene) woodson. asclepiadaceae acerates auriculata engelm. acerates viridiflora (rar.)eaton. ampelamus albidus (nutt.) britt. (gonolobus laevis sensu vail). asclepias amplexicaulis j.e. smith. asclepias galioides hbk. asclepias incarnata l. asclepias speciosa torr. asclepias stenophylla gray. asclepias tuberosa l. asclepias tuberosa l. forma lutea clute. asclepiodora decumbens (nutt.) gray. asclepiodora viridis (walt.) gray. gonolobus gonocarpos (walt.) perry. convolvulaceae convolvulus ambigens house. convolvulus sepium l. cuscuta arvensis bevrich. evolvulus nuttallianus schultze. (e. pilosus nutt.) ipomoea lacunosa l. ipomoea leptophylla torr. ipomoea longifolia benth. hydrophyllaceae ellisia nyctelea (l.) nemophila phacelioides nutt. phacelia hirsute nutt. boraginaceae hackelia virginiana (l.) i. m. johnston. (lappula virginiana (l.) greene. heliotropum tenellum (nutt.) torr. lappula texana (scheele) britton. lithospermum arvense l. oklahoma native plant record 17 volume 1, number 1, december 2001 waterfall, u.t. lithospermum caroliniense (walt.) macm. (l. gmeleni in part). lithospermum incisum lehm. (l. angustifolium michx.) verbenaceae lippia cuneifolia (torr.) steud. lippia lanceolata michx. var. recognita fern. & grisc. verbena bipinnatifida nutt. verbena bracteata lag. & rodr. verbena canadensis (l.) britton. verbena hastate l. verbena pumila rydb. verbena stricta vent. verbena urticaefolia l. labiateae hedeoma camporum rydb. hedeoma hispida pursh. lamium amplexicaule l. lycopsus americanus muhl. mondarda clinopodioides gray. monarda mollis l. prunella vulgaris l. var. lanceolata (barton) fern. salvia pitcheri torr. scutellaria lateriflorus l. scutellaria parvula michx. var. australis fassett. teucrium canadense l. var. virginicum (l.) eaton. solanaceae datura stramonium l. physalis heterophylla nees. physalis ixiocarpa brot. physalis lobata torr. physalis macrophysa rydb. physalis mollis nutt. physalis pendula rydb. physalis pumila nutt. physalis subglabrata mack. & bush. solanum carolinense l. solanum elaeagnifolium cav. solanum nigrum l. solanum rostratum dunal. solanum torreyi gray. scrophulariaceae buchnera americana l. gerardia densiflora benth. gerardia grandiflora benth. var. serrata (torr.) robinson. gerardia heterophylla nutt. llysanthes anagallidea (michx.) robinson. leucospora multifida (michx.) nutt. (conobea multifida). linaria canadensis (l.) dumont. var. texana pennell. mimulus glabratus hbk. var. oklahomensis fassett. mimulus ringens l. penstemon cobaea nutt. penstemon laxiflorus pennell. penstemon oklahomensis pennell. verbascum thapsus l. veronica arvensis l. veronica peregrina l. var. xalapensis (hbk.) pennell. bignoniaceae catalpa speciosa warder acanthaceae dicliptera brachiata (pursh) spreng. ruellia caroliniensis (walt.) steud. (r. ciliosa pursh.) ruellia strepens l. plantaginaceae plantago aristata michx. plantago purshii r. & s. plantago pusilla nutt. plantago rhodosperma dcne. plantago rugelii dcne. plantago virginica l. rubiaceae cephalanthus occidentalis l. cephalanthus occidentalis l. var. pubescens raf. diodia teres walt. var. setifera fern. & grisc. galium aparine l. var. vaillantii (dc.) koch. galium circaezans michx. var. hypomalacum fern. galium obtusum bigel. galium pilosum ait. var. puncticulosum (michx.) t. & g. galium virgatum nutt. houstonia nigricans (lam.) fern. h. angustifolia michx.) houstonia minima beck. 18 oklahoma native plant record volume 1, number 1, december 2001 waterfall, u.t. caprifoliaceae sambucus canadensis l. symphoriocarpos orbiculatus moench. virburnum rufidulum raf. valerianaceae valerianella amarelle (lindl.) krok. valerianella radiata (l.) dufr. valerianella stenocarpa (engelm) (krok.) var. parviflora dyall. cucurbitaceae cucurbita foetidissima hbk. melothria pendula l. campanulaceae specularia biflora (r.& p.) f. & m. specularia leptocarpa (nutt.) gray. specularia perfoliata (l.) a. dc. lobeliaceae lobelia splendens willd. compositae achillea lanulosa nutt. achillea lanulosa nutt. forma rubicunda farwell. actinea linearifolia (hook.) kuntze. actinomeris alternifolia (l.) dc. agoseris cuspidata (pursh) steud. ambrosia artemesifolia l. var. elatior (l.) descourtils. ambrosia coronopifolia t. & g. ambrosia trifida l. var. texana scheele (a. aptera dc.) antennaria fallax greene. anthemis cotula l. aphanostephus skirrobasis (dc.) trel. aplopappus ciliatus (nutt.) dc. aplopapous divericatus (nutt.) gray artemisia gnaphalodes nutt. aster azureus lindl. aster drummondii lindl. aster ericoides l. (a. multiflorus) aster exilis ell. aster oblongifolius nutt. var. rigidulus gray. aster patens ait. var. gracilis hook. aster praealtus poir. (a. salicifolius) astranthium integrifolium (michx.) nutt. var. ciliatum larsen. baccharis salicina t. & g. bidens bipinnata l. bidens cernua l. bidens involucrate (nutt.)britton. bidens vulgata greene. chaetopappa asteroids dc. chrysopsis berlandieri greene. chrysopsis pilosa nutt. cirsium undulatum (nutt.) spreng. cirsium virginianum (l.) michx. coreopsis cardaminefolia (dc.) t.& g. coreopsis grandiflora hogg. ex sweet. coreopsis tinctoria nutt. echinacea angustifolia dc. eclipta alba (l.) hassk. elephantopus carolinianus raeuschel. erigeron canadensis l. erigeron diverticatus michx. erigeron philadelphicus l. erigeron ramosus (walt.) bsp. eupatroium coelestinum l. eupatorium perfoliatum l. eupatorium serotinum michx. evax multicaulis dc. gaillardia lanceolata michx. gaillardia suavis (gray) britt. & rusby. gaillardia trinervata small gnaphalium obtusifolium l. gnaphalium purpureum l. gutierrezia dracunculoides (dc.) blake. amphiachyris dracunculoides). helenium tenuifolium nutt. helianthus annus l. helianthus hirsutus raf. helianthus maximiliani schrad. helianthus mollis lam. helianthus petiolaris nutt. helianthus tuberosus l. heterotheca subaxillaris (lam.) britt. & rusby. hieracium gronovii l. hieracium longipilum torr. hymenopappus tenuifolius pursh. oklahoma native plant record 19 volume 1, number 1, december 2001 waterfall, u.t. iva ciliata willd. krigia occidentalis nutt. kuhnia eupatorioides l. kuhnia eupatorioides l. var. corymbulosa t. & g. lactuca campestris greene. lactuca canadensis l. var. latifolia o. ktze. lactuca canadensis l. var. longifolia (michx.) farwell. lactuca floridana (l.) gaertn. lactuca scariola l. liatrus acidota engelm. & gray. liatrus punctata hook. liatrus squarrosa willd. var. intermedia (lindl.) dc. matricaria matricarioides (less.) porter. parthenium hysterophorus l. pluchea marilandica (michx.) cass. pulchea purpurascens (sw.) dc. polymnia uvedalia l. var. densipilis blake. polypteris macrolepis (rydb.) pyrrhopappus carolinianus (walt.) dc. pyrrhopappus scaposus dc. ratibida columnifera (nutt.) woot. & standl. ratibida columnifera (nutt.) woot. & standl. forma pulcherrima (dc.) fern. & standl. forma pulcherrima (dc.) fern. rudbeckia hirta l. var. sericea (t.v. moore) fernald. senecio glabellus poir. senecio plattensis nutt. serinia oppositifolia (raf.) kuntze. silphium asperrimum hook. silphium laciniatum l. solidago canadensis l. solidago hellari small. solidago leptocephala t. & g. solidago petiolaris ait. solidago radula nutt. solidago rigida l. solidago serotina ait. solidago ulmifolia muhl. sonchus asper (l.) hill. taraxacum laevigatum (willd.) dc. taraxacum palustre (lyons) lam. & dc. var. vulgare (lam.) fern. thelesperma trifidum (poir) britton. tragopogon major jacq. verbesina encelioides (cav.) gray. verbesina virginica l. vernonia baldwinii torr. var. interior (small) schuberr. xanthisma texanum dc. xanthium italicum moretti. 20 oklahoma native plant record volume 1, number 1, december 2001 waterfall, u.t. chapter viii tabular view of the families of the spermatophyta classes, families, etc. genera species/ var-form monocotyledonae 25. ranunculaceae 5 5/1 1. typhaceae 1 2 26. menispermaceae 2 2/1 2. alismaceae 1 1 27. papaveraceae 1 1 3. xyridaceae 1 1 28. fumariaceae 1 1/1 4. commelinaceae 2 5/3 29. crucifera 14 18/3 5. pontederiaceae 1 1 30. capparidaceae 2 2 6. liliaceae 6 9 31. saxifragaceae 1 1 7. amaryllidaceae 1 1 32. platanaceae 1 1 8. iridaceae 1 4 33. rosaceae 8 8/2 9. orchidaceae 1 1 34. leguminosae 31 52/12 dicotyledonae 35. linaceae 1 3/1 10. salicaceae 2 3/1 36. oxalidaceae 1 2 11. juglandaceae 2 3 37. geraniaceae 1 1 12. fagaceae 1 5 38. zygophyllaceae 2 2 13. urticaceae 6 7/2 39. rutaceae 2 2 14. loranthaceae 1 1 40. polygalaceae 1 2/1 15. polygonaceae 3 15/1 41. euphorbiaceae 5 23/3 16. chenopodiaceae 6 9/1 42. anacardiaceae 1 2/1 17. amaranthaceae 4 9/1 43. celastraceae 2 2 18. phytolaccaceae 1 1 44. aceraceae 1 1/2 19. nyctaginaceae 1 3 45. sapindaceae 2 2 20. illecebraceae 1 2 46. rhamnaceae 1 1/1 21. aizoaceae 1 1 47. vitaceae 3 7 22. caryophyllaceae 4 5/1 48. malvaceae 4 5 23. portulacaceae 3 4/1 49. tamaricaceae 1 1 24. nymphaceae 2 2 50. hypericaceae 2 1/2 oklahoma native plant record 21 volume 1, number 1, december 2001 waterfall, u.t. 51. elatinaceae 1 1 74. solanaceae 3 14 52. cistaceae 1 1/2 75. scrophulariaceae 9 11/4 53. violaceae 1 2/2 76. bignoniaceae 1 1 54. passifloraceae 1 1 77. acanthaceae 2 3 55. loasaceae 1 1 78. plantaginaceae 1 6 56. cactaceae 2 2 79. rubiaceae 4 5/5 57. lythraceae 3 3 80. caprifoliaceae 3 3 58. onagraceae 5 12/6 81. valerianaceae 1 2/1 59. umbelliferae 11 14/1 82. cucurbitaceae 2 2 60. cornaceae 1 1 83. campanulaceae 1 3 61. primulaceae 2 2 84. lobeliaceae 1 1 62. sapotaceae 1 1 85. compositae 60 95/15 63. ebenaceae 1 1/1 total** monocotyledonae 15 26/3 64. oleaceae 1 1/1 total** dicotyledonae 283 449/84 65. loganiaceae 1 1 total** angiospermae 298 475/87 66. gentianaceae 1 2 total** spermatophyta 298 475/87 67. apocynaceae 1 1/3 68. asclepidaceae 4 12/1 69. convolvulaceae 5 7 **ed. note: while numbering of species in the chart has been edited, errors in totals have not been corrected. editor counts 25 species of monocots and 456 species of dicots making angiosperm and spermatophyte totals of 481. also, a total of 85 varieties or forms of dicots and 3 monocot varieties total 88 varieties and forms. author did not have the benefit of an electronic calculator. [s.s.] 70. hydrophyllaceae 3 3 71. boraginaceae 4 6 72. verbenaceae 2 8/1 73. labiateae 8 8/3 22 oklahoma native plant record volume 1, number 1, december 2001 waterfall, u.t. acknowledgements the author is deeply grateful to dr. milton hopkins, associate professor of botany in the university of oklahoma, under whose supervision this work was done. his inspiration and encouragement, his suggestions and guidance have been of great assistance. he is also indebted to the curators of several herbaria, and to various specialists for identifications of difficult, or critical specimens. these include dr. m. l. fernald and other members of the staff of the gray herbarium of harvard university, dr. e. j. alexander of the new york botanical gardens, dr. p.a. munz of pomona college, dr.francis r. pennell of the academy of natural sciences of philadelphia, and dr. s. f. blake of the bureau of plant industry of the united states department of agriculture. bibliography anderson, edgar, and r. e. woodson, jr. the species of tradescantia indigenous to the united states. contr. arnold arboretum of harvard university. 9: 1-132, 1935. britton, n. l., and a. brown. an illustrated flora of the northern united states, canada, and the british possessions. three vols. 2nd ed. revised and enlarged. lancaster press, inc., lancaster, pennsylvania. deam, charles c. flora of indiana. department of conservation, division of forestry. indianapolis, indiana, 1940. dyal, sarah c. key to the species of oaks of eastern north america based on foliage and twig characteristics. rhodora 38: 53-63, 1936. dyal, sarah c, valerianella in north america. rhodora 40: 185212, 1938. fassett, roman c. notes from the herbarium of the university of wisconsin. xiii. rhodora 38: 94-97, 1936. fassett, norman c. notes from the herbarium of the university of wisconsin. xv. rhodora 39: 377-379, 1937. fassett, norman c. notes from the herbarium of the university of wisconsin. xviii. rhodora 41: 524-529, 1939. fernald, m. l. midsummer vascular plants of southeastern virginia. rhodora 37: 378-413, 423-455,1935. fernald, m. l. contributions from the gray herbarium. xiii. rhodora 38: 165-l82, 261-239 [sic],1936. fernald, m. l. plants from the outer coastal plain of virginia. rhodora 38: 376-404, 414-452, 1936. fernald, m. l. petalostemum occidentae. rhodora 39: 28, 1937. fernald, m. l. nomenclatural transfers and new varieties and forms. rhodora 39: 309-320, 1937. fernald, m. l. local plants of the inner coastal plain of virginia. rhodora 39: 321-366, 379-415, 433-459, 465-491, 1937. fernald, m. l. noteworthy plants of southeastern virginia. rhodora 40: 364-424, 434-459, 467-485, 1935. fernald, m. l. new species varieties and transfers. rhodora 40: 331-358, 1938. fernald, m. l. new species varieties and transfers, rhodora 41: 423-461, 1939. fernald, m. l. last survivors in the flora of tidewater virginia. rhodora 4l: oklahoma native plant record 23 volume 1, number 1, december 2001 waterfall, u.t. 465-504, 529-558, 564-574, 1939. fernald, m. l. campestrian variety of froelichia floridana. rhodora 43: 336, 1941. fernald, m. l. another century of additions to the flora of virginia. rhodora 43: 485553, 559-630, 635-657, 1941. fernald, m. l. some forms in the aliasmaceae. rhodora 38: 73, 1938. fernald, m.l., and ludlow griscom. three days botaniing in southeastern virginia. rhodora 37: 129-157, 167-189, 1935. fernald, m. l., and ludlow griscom. notes on diodia. rhodora 39: 306-308, 1937. freeman, florence l. variations of psoralea psoralioides. rhodora 39: 425-428. 1937. henson, dorthy h. the genus monarda in oklahoma. american midland naturalist 25: 358360, 1941. hodgdon, albion r. taxonomic study of lechea. rhodora 40: 29-69, 87-131, 1938. hopkins, milton. notes on lespedeza. rhodora 37: 265266, 1935. hopkins, milton. arabis in eastern and central north america. rhodora 39: 63-98, 106-148, 155-186, 1937. hopkins, milton. cercis in north america. rhodora 44: 193-211, 1942. larisey, mary m. monograph of the genus baptisia. annals of the missouri botanical gardens. 27: 119-244, 1940. larsen, esther l. astranthium and related genera. annals of the missouri botanical gardens. 20: 23-44, 1933. leonard, emery c. the north american species of scutellaria, contrib. from the u.s. nat. herb. vol. 22, part, 10, govmt. printing office, washington, d. c., 1927. leyendecker, jordon phillip jr. a taxonomic study of the genus galium in iowa. proc. iowa acad. sci. xlvii. 101-114, 1940. mcvaugh, rogers. studies in the taxonomy and distribution of the eastern north american species of lobelia. rhodora 38: 214-263, 276-298, 305-329, 346-362, 1936 mathias, mildred e., and lincoln constance. new combinations and new names in the umbelliferae. bul. torre bot. club. 68: 128124 [sic]. 1942. merrill, e. d. on houttuyn's overlooked binomials for native or introduced plants in eastern north america. rhodora 40: 228293, 1938. munz, philip a. studies in onagracea. iv. a revision of the subgenera salpingia and calylophis of the genus oenothera. amer. journ. bot. 16: 702-715, 1929. munz, philip a. studies in onagraceae xi. a revision of the genus gaura. bul. torr. bot. club. 65: 105-122, 211-228, 1938. pennell, francis w. commelina in the united states. bull. torr. bot. club. 43: 96-111, 1916. pennell, francis w. scrophulariaceae of eastern temperate north america. acad. nat. sciences of philadelphia, monograph i. 1935. pennell, francis w. commelina communis in eastern united states. bartonia, no. 19: 19-22, 1937. pennell, francis w. a supplemental note concerning commelina nudiflora. proc. acad. sciences of philadelphia. 40: 39, 1938 perry, lily m. evolvulus pilosus an invalid name. rhodora 37: 63, 1935. perry, lily m. a revision of the north american species of verbena. annais mo. bot. gard. 20: 239-352, 1933. 24 oklahoma native plant record volume 1, number 1, december 2001 waterfall, u.t. perry, lily m. gonolobus within the gray's manual range. rhodora 40: 281-287, 1938. rickett, h. w. cornus asperifolia and its relatives. robinson, b. l., and m. l. fernald.gray's new manual of botany. amer. book co., new york. 1908. rydberg, p. a. flora of the prairies and plains of central north america. new york bot. gard. new york, 1932. sargent, c. s. manual of the trees of north america. riverside press. cambridge, mass. 3rd. ed., 1933. schubert, bernice c. notes on vernonia. rhodora 38: 369372. 1936. small, j. k. flora of the southeastern united states. pub. by j. k. small. new york. 2nd ed., 1913. small, j. k. manual of the southeastern flora. pub. by j. e. small. new york, 1933. stemen, thomas r., and w. stanley myers. oklahoma flora. harlow pub. corp., oklahoma city. 1937. stevens, g. l. the flora of oklahoma. unpublished original deposited in the widener library of harvard university, 1916. steyermark, julian a. spring flora of missouri. mo. bot. gard. and field mus. nat. history. chicago, 1940. waterfall, u. t. interesting plants of oklahoma. rhodora 42: 499 502, 1940. weatherby, c. a. typification of acalypha virginica. rhodora 39: 14-16, 1937. wheeler, l. c. euphorbia subgenus chamaesyche in canada and the united states exclusive of southern florida. rhodora 43: 97-154, 168-206, 231-236, 1941. woodson, robert e. apocynaceae. n. am. flora 29, part 2: 188-192, 1938. *ed. note: for historical purposes much of waterfall’s original format has been retained. species epithets derived from a person’s name are capitalized and margins are left-justified only. a similar font has been used. however, we have edited the thesis for readability. footnotes have been moved from the bottom of each page to the end of each chapter. italics have been substituted for underscoring of scientific names and the text has been formatted in two columns. [s.s.] journal of the oklahoma native plant society, volume 8, number 1, december 2008 67 oklahoma native plant record volume 8, number 1, december 2008 murray, c.s. https://doi.org/10.22488/okstate.17.100064 tribute to paul buck paul buck passed away on january 16, 2008. respected and admired by his colleagues and friends, he left a legacy of lifelong commitment to ecology and botany. paul was a founding member of the board of directors of the oklahoma native plant society in 1987, and, after serving on its board for several subsequent terms, he was the second recipient of the onps service award. in the early days of onps he was an active leader and participant in society field excursions throughout the state. paul was at the organizing core of the onps color oklahoma committee, serving on its first board from 2003 through 2006. for more than ten years he wrote a column entitled “botany bay” for the onps quarterly newsletter, gaillardia. in each issue he presented a puzzling or intriguing botanical problem and then decoded it in colorful terms, accessible to both amateur and professional botanists. most knew paul as professor of botany at the university of tulsa. he was there from 1964 to 1987, teaching and inspiring students, majors and non-majors alike. for decades he transported students all over the state to oklahoma academy of science (oas) field meetings and technical meetings, to southwest association of naturalists meetings, and on spring break excursions to mexico and new mexico he encouraged students to attend the university of oklahoma biological station, which he had attended as a student, and rocky mountain biological station, at which he taught during the summer. paul was the faculty advisor of the tu student chapter of zero population growth. with his colleague, estelle levetin, he established the longest pollen record in the u.s., also one of the longest in the world. after paul retired from the university of tulsa he continued to curate the harriet g. barclay herbarium there and to teach and guide students. in the mid-1980s paul was a founding member of the flora of oklahoma committee, a group of oklahoma botanists dedicated to writing the flora of oklahoma. this has been and continues to be a monumental work to write and update keys and descriptions of all the vascular plant species in oklahoma, replacing the keys of u.t. waterfall. paul actively participated in the flora of oklahoma board of directors until retiring in the spring of 2006 as its treasurer. paul was elected president of the oklahoma academy of science in 1971 and served as executive secretary-treasurer in the late 1980s and early 1990s. he was recognized by oas for his meritorious service to oklahoma scientists with the tenure service award in 1991, the education service award in 1994, and the oas lifetime achievement award presented at a botanical symposium in his honor in 2006. oas continues to publish paul’s distribution and identification of woody plants of oklahoma in the winter condition (1983). paul was founder of the mary kay oxley nature center, a natural area along bird creek at the edge of mohawk park in tulsa. along with harriet barclay, he was instrumental in encouraging the nature conservancy to purchase both red bud valley nature preserve in 1970 and the tall grass prairie preserve in 1989. he served on the boards of all three of these organizations for many years until they were well established. he also served on the board of the oklahoma nature conservancy. many are surprised to learn that paul buck was not a native oklahoman. he was born in lansing, michigan, september 9, 1927. at 17 he joined the us navy and was stationed in norman, oklahoma. on leave 68 oklahoma native plant record volume 8, number 1, december 2008 murray, c.s. in tulsa, he met lou ann clark, whom he later married. in tulsa he served on the tulsa police force as a “beat cop” in oakhurst in west tulsa. working the night shift enabled him to attend classes at the university of tulsa during the day. there he was inspired by harriet barclay and ralph kelting to pursue a career in botany. after his bs and ms at ut, he attended the university of oklahoma where he worked with elroy rice. his dissertation, relationships of woody vegetation of the wichita mountains wildlife refuge to geological formations and soil types, was among the first ecological studies of the wichita refuge. in 2002 oklahoma native plant record published “vascular plants of the wichita mountains”, from an informational pamphlet previously used by refuge biologists, which was based on that work. paul’s commitment to botany, ecology, and environmental conservation extended beyond formal and academic venues. he involved neighborhood children in observing the natural world. paul was active in community and student efforts to start recycling programs. he led boy scout trips to philmont in new mexico and spoke to citizen groups at the tulsa library. he rode his bicycle to campus for years, attired in a tuxedo. paul even rode his bicycle to his daughter’s wedding. he lived his commitment to conservation and never lost his joy and wonder at the beauty and complexity of the natural world. nothing in this formal description can convey the serene and honorable way in which paul conducted his life. as he mentored and encouraged students, he persisted in his community activism and cared for his mentor and colleague, harriet barclay in her later years. he did so with kindness and good humor. nor can it convey the remarkable grace with which he accepted the unfairness and disability of parkinson’s disease as it limited his field experiences in his own later years. paul was remembered in a memorial service may 3, 2008 at the harriet g. barclay nature center at red bud valley. it was a glorious spring afternoon, sun shining, gentle breeze – oklahoma at its best. family and friends, colleagues and former students, neighbors and community activists were all in attendance, remembering the life of the man who had formed, shared, or changed their lives and left this world a better place for his having been here. paul is survived by lou ann clark buck, his wife of more than fifty years, and by his children, paul buck iii of gunnison, colorado and dana buck of atlanta, georgia. intellectually and inspirationally, he is survived by us all. paul’s knowledge of the natural world and his tireless pursuit of its further understanding inspired students, future scientists, and laymen for more than five decades. his gentle manner, his patience, his persistence, and his kindness, even in the face of personal, professional, political, and environmental adversity, make him a model for each of us as we continue his commitment to the botany and ecology of oklahoma. constance murray, 1 june 2008 onps journal of the oklahoma native plant society, volume 7, number 1, december 2007 54 oklahoma native plant record volume 7, number 1, december 2007 the vascular flora of the oklahoma centennial botanical garden site osage county, oklahoma bruce w. hoagland amy buthod oklahoma biological survey oklahoma biological survey department of geography university of oklahoma university of oklahoma norman, ok 73019-0575 norman, ok 73019-0575 * e-mail: bhoagland@ou.edu this paper is a report on the results of an inventory of the vascular plants at the future site of the oklahoma centennial botanical garden in osage county, oklahoma. we collected a total of 293 taxa in 208 genera and 68 families. the families poaceae and asteraceae had the greatest number of species with 50 and 44 species respectively. fortyone species of woody plants were present. forty-four non-native species were present, representing 15% of the flora. no species tracked by the oklahoma natural heritage inventory were present. introduction the objective of this study was to complete a floristic inventory at the future site of the oklahoma centennial botanical garden (ocbg) in southeast osage county (36.2017°n to 36.2109°n and 96.0555°w and 96.0678°w). construction of the ocbg is scheduled to begin in late 2007 on 87 hectares (215 acres). the master plan, developed by marshall tyler rausch of pittsburgh, pennsylvania, includes a mexican garden, oklahoma wildflower garden, cross timbers prairie and woodland, folk garden, horticultural therapy garden, children’s garden, demonstration gardens, and others. in addition, a 17-acre lake, an amphitheater, a visitor center, education buildings, and a conservatory will be constructed (oklahoma centennial botanical garden 2007). the ocbg site is located in the claremore cuesta plains geomorphic province of southeastern osage county (curtis and ham 1979). surface geology is predominantly pennsylvanian sandstone and shale (branson and johnson 1979). soils belong to the niotaze-darnell association, described as moderately deep and shallow, hoagland, b.w. https://doi.org/10.22488/okstate.17.100053 gently sloping to steep, loamy soils over shale and sandstone (bourlier et al. 1979). the climate is subtropical humid (cf) (trewartha 1968). summers are warm and humid. mean july temperature is 27.5oc (81.5of). winters are relatively short and mild with a mean january temperature of 1.5oc (34.7of). mean annual precipitation is 111.7 cm (43.8 in) (oklahoma climatological survey, 2007). elevation ranges from 259 to 302 m (849.5 to 990.6 ft). potential natural vegetation at ocbg is post oak-blackjack forest and tallgrass prairie (duck and fletcher 1943). historical land use of the site has included livestock grazing and oil exploration. methods three collection sites were visited monthly for floristic sampling. the predominant vegetation association at these sites was classified according to hoagland (2000). additional collections were also made opportunistically throughout the ocbg. collecting began in july of 2006 and continued through july of 2007. vouchers for non-native species were made from naturalized populations only, thus 55 oklahoma native plantrecord volume 7,number 1,december 2007 hoagland & buthod excluding cultivated and ornamental plants. specimens were processed following standard procedures and deposited at the robert bebb herbarium at the university of oklahoma (okl). manuals used for specimen identification included waterfall (1973) and steyermark (1963). origin, either native or introduced to north america, was determined using the plants database (usda-nrcs, 2007). nomenclature follows the united states department of agriculture-natural resources conservation service (usda-nrcs 2007). results and discussion a total of 293 taxa of vascular plants in 68 families and 208 genera were collected at the ocbg (appendix). of the angiosperms, 92 species were liliopsida and 199 were magnoliopsida (table). there was one species of pteridophyta and one of coniferophyta. forty-one species were trees, shrubs, and woody vines. the poaceae with 50 taxa, and the asteraceae with 44 taxa, were the largest families. the genera symphyotrichum (formerly aster) and cyperus had the most species, with seven and six species respectively. one hundred and seven taxa were annuals, 2 were biennials and 184 were perennials. forty-four species (15% of the flora) in 25 families were non-native to oklahoma. the percentage of non-native species at the ocbg is high when compared to other floristic surveys from oklahoma, which range from 6.6%-15% (hoagland and buthod 2004; hoagland and johnson 2005). the greatest numbers of non-native species occurred in the poaceae, with eleven and fabaceae, with eight. no species tracked by the oklahoma natural heritage inventory (2007) were encountered. collection sites selected at ocbg occurred within four vegetation associations. a description of each vegetation category follows: 1. quercus stellata-quercus marilandica forest association [qsqm] this vegetation association occupied a small percentage of the ocbg. common associated species included amelanchier arborea, antennaria plantaginifolia, baptisia bracteata var. leucophaea, danthonia spicata, helianthus hirsutus, hypericum hypericoides, symphyotrichum patens, myosotis verna, opuntia humifusa, sideroxylon lanuginosum, smilax rotundifolia, ulmus alata, and viburnum rufidulum. 2. schizachyrium scoparium-sorghastrum nutans [ssn] this herbaceous grassland vegetation association occupied the greatest area at the ocbg. soils were typically shallow with exposed cobble. associated species included amorpha canescens , arnoglossum plantagineum, callirhoe alcaeoides , coreopsis grandiflora, cyperus echinatus, echinacea atrorubens, krameria lanceolata, lespedeza cuneata, minuartia drummondii, and pediomelum linearifolium. 3. wetland and aquatic vegetation [wetl] wetland vegetation was restricted to a small stream bisecting the site and its associated beaver pond. common associates included alisma subcordatum, ammannia auriculata, callitriche heterophylla, cephalanthus occidentalis, eclipta prostrata, fimbristylis autumnalis, juncus brachycarpus, ludwigia palustris, nelumbo lutea, polygonum pensylvanicum, sagittaria ambigua, and samolus ebracteatus. 4. disturbed areas and old-field vegetation [daof] disturbed areas coincided with roadways and oil extraction sites. common associated species included achillea millefolium, aegilops cylindrica, capsella bursapastoris, carduus nutans, convolvulus arvensis, daucus pusillus, juniperus virginiana, lamium amplexicaule, rhus copallinum, r. glabra, and torilis arvensis. 56 oklahomanative plantrecord volume 7,number 1,december 2007 hoagland & buthod literature cited bourlier, b.g., j.d. nichols, w.j. ringwold, p.j. workman, and s. clemmons. 1979. osage county soil survey. united states department of agriculture, washington, dc. branson, c.c., and k.s. johnson. 1979. generalized geologic map of oklahoma. in: johnson, k.s., c.c. branson, n.m. curtis, w.e. ham, w.e. harrison, m.v. marcher, and j.f. roberts (eds.). geology and earth resources of oklahoma. oklahoma geological survey, norman, oklahoma. curtis, n.m. and w.e. ham. 1979. geomorphic provinces of oklahoma. in: johnson, k.s., c.c. branson, n.m. curtis, w.e. ham, w.e. harrison, m.v. marcher, and j.f. roberts (eds.). geology and earth resources of oklahoma. oklahoma geological survey, norman, oklahoma. duck, l.g., and j.b. fletcher. 1943. a game type map of oklahoma. in: duck, l.g., and j.b. fletcher (eds.). a survey of the game and furbearing animals of oklahoma. oklahoma department of wildlife conservation, oklahoma city, oklahoma. hoagland, b.w. 2000. the vegetation of oklahoma: a classification of landscape mapping and conservation planning. southwest naturalist 45: 385-420. hoagland, b.w. and f.l. johnson. 2004. vascular flora of red slough and grassy slough wildlife management areas, gulf coastal plain, mccurtain county, oklahoma. castanea 69: 284296. hoagland, b.w. and f.l. johnson. 2005. vascular flora of the deep fork river in okmulgee, creek, and okfuskee counties. publications of the oklahoma biological survey 6: 15-29. oklahoma climatological survey. 2007. oklahoma climatological data. university of oklahoma, norman. (www.ocs.ou.edu accessed 1 august 2007). oklahoma centennial botanical garden. 2007. oklahoma centennial botanical garden research and education center. tulsa, oklahoma. www.oklahomacentennialbotanicalgar den.com accessed 1 september 2007). oklahoma natural heritage inventory. 2007. oklahoma natural heritage inventory working list of rare oklahoma plants. university of oklahoma, norman. (www.biosurvey.ou.edu/publicat.html accessed 1 august 2007). palmer, m.w., g.l. wade, and p. neal. 1995. standards for the writing of floras. bioscience 45: 339-345. steyermark, j.a. 1963. flora of missouri. iowa state university press. ames, iowa. trewartha, g.t. 1968. an introduction to climate. mcgraw-hill, new york, new york. usda-nrcs. 2007. the plants database. national plant data center, baton rouge, la 70874-4490 usa. (http://plants.usda.gov accessed 1 may 2007). waterfall, u.t. 1973. keys to the flora of oklahoma. published by the author, stillwater, oklahoma. 57 oklahoma native plantrecord volume 7,number 1,december 2007 hoagland & buthod table summary of floristic collections from the oklahoma centennial botanical garden site, osage county, oklahoma. format follows palmer et al. (1995). figure oklahoma centennial botanical garden site, osage county, oklahoma. taxonomic group taxa native nonnative pteridophyta 1 1 0 coniferophyta 1 1 0 magnoliophyta 291 247 44 magnoliopsida 199 165 34 liliopsida 92 82 10 total 293 249 44 58 oklahomanative plantrecord volume 7,number 1,december 2007 hoagland & buthod appendix annotated species list for the oklahoma centennial botanical garden, osage county, oklahoma. the first entry is habitat (qsqm=quercus stellata-quercus marilandica forest association, sssn=schizachyrium scoparium-sorghastrum nutans grassland association, wetl=wetland and aquatic vegetation, daof=disturbed areas and old-field vegetation); followed by the life history (a=annual, b=biennial, p=perennial); habit (t=tree, s=shrub, v=woody vine, h=herb, g=graminoid); and collection number. exotic species are denoted with an asterisk. voucher specimens were deposited at the robert bebb herbarium of the university of oklahoma (okl). pteridophyta aspleniaceae asplenium platyneuron (l.) b.s.p. (ebony spleenwort) qsqm; p; h; obg-152 coniferophyta cupressaceae juniperus virginiana l. (eastern red cedar) qsqm; p; t; obg-231 magnoliophyta magnoliopsida acanthaceae ruellia humilis nutt. (fringeleaf wild petunia) sssn; p; h; obg-285 r. strepens l. (limestone wild petunia) qsqm; p; h; obg-153 amaranthaceae amaranthus albus l. (prostrate pigweed) daof; a; h; obg-012 anacardiaceae rhus aromatica ait. (fragrant sumac) qsqm; p; s; obg-184 r. copallinum l. (flameleaf sumac) sssn; p; s; obg-247 r. glabra l. (smooth sumac) sssn; p; s; obg-255 toxicodendron radicans l. (kuntze) (poison ivy) qsqm; p; s; obg-334 apiaceae chaerophyllum tainturieri hook. (hairyfruit chervil) daof, qsqm, sssn; a; h; obg198 daucus carota* (queen anne’s lace) daof; b; h; obg-296 d. pusillus michx. (american wild carrot) daof, sssn; a; h; obg-254 ptilimnium nuttallii (dc.) britt. (laceflower) sssn; a; h; obg-304 spermolepis divaricata (walt.) raf. ex ser. (roughfruit scaleseed) sssn; a; h; obg305 torilis arvensis* (huds.) link (spreading hedgeparsley) daof; a; h; obg-256 aquifoliaceae ilex decidua walt. ( possumhaw) qsqm; p; s; obg-144 asclepiadaceae asclepias viridis walt. (green antelopehorn) sssn; p; h; obg-032 asteraceae achillea millefolium l. (common yarrow) daof, sssn; p; h; obg-219 ambrosia psilostachya dc. (cuman ragweed) daof; p; h; obg-079 amphiachyris dracunculoides (dc.) nutt. (prairie broomweed) daof, sssn; a; h; obg-097 antennaria plantaginifolia (l.) richards. (woman’s tobacco) qsqm; p; h; obg-187 arnoglossum plantagineum raf. (groovestem indian plantain) sssn; p; h; obg-221 carduus nutans* l. (nodding plumeless thistle) daof; b; h; obg-208 cirsium altissimum (l.) hill (tall thistle) sssn; p; h; obg-114 59 oklahoma native plantrecord volume 7,number 1,december 2007 hoagland & buthod c. undulatum (nutt.) spreng. (wavyleaf thistle) daof, sssn; p; h; obg-279 conyza canadensis (l.) cronq. (canadian horseweed) daof; a; h; obg-091 c. ramosissima cronq. (dwarf horseweed) daof; a; h; obg-089 coreopsis grandiflora hogg ex sweet (largeflower tickseed) sssn; p; h; obg300 c. tinctoria nutt. (golden tickseed) daof, sssn; a; h; obg-059 echinacea atrorubens nutt. (topeka purple coneflower) sssn; p; h; obg-284 eclipta prostrata (l.) l. (false daisy) wetl; a; h; obg-073 erigeron strigosus muhl. ex willd. (prairie fleabane) daof, sssn; a; h; obg-015 eupatorium serotinum michx. (lateflowering thoroughwort) qsqm; p; h; obg-062 euthamia gymnospermoides greene (texas goldentop) sssn; p; h; obg-148 gamochaeta purpurea (l.) cabrera (spoonleaf purple everlasting) -qsqm; p; h; obg-237 grindelia lanceolata nutt. (narrowleaf gumweed) daof, sssn; p; h; obg048 g. papposa nesom & suh (spanish gold) daof, sssn; a; h; obg-020 helenium amarum (raf.) h.rock (yellowdicks) daof; a; h; obg-106 helianthus hirsutus raf. (hairy sunflower) qsqm; p; h; obg-121 iva angustifolia nutt. ex dc. (narrowleaf marshelder) wetl; a; h; obg-125 krigia caespitosa (raf.) chambers (weedy dwarfdandelion) qsqm; a; h; obg-240 packera plattensis (nutt.) w.a. weber & a. löve (prairie groundsel) qsqm; p; h; obg-192 pluchea camphorata (l.) dc. (camphor pluchea) wetl; p; h; obg-011 pseudognaphalium obtusifolium (l.) hilliard & burtt (rabbit tobacco) daof, sssn; a; h; obg-002 pyrrhopappus carolinianus (walt.) dc. (carolina desert chicory) daof, sssn; a; h; obg-280 rudbeckia hirta l. (blackeyed susan) sssn; a; h; obg-251 solidago speciosa nutt. (showy goldenrod) sssn; p; h; obg-132 s. ulmifolia muhl. ex willd. (elmleaf goldenrod) qmqv; p; h; obg-049 sonchus asper* (l.) hill (spiny sowthistle) daof; a; h; obg-185 symphyotrichum cordifolium (l.) nesom (common blue wood aster) qsqm; p; h; obg-130 s. ericoides (l.) nesom var. ericoides (white heath aster) daof, sssn; p; h; obg-154 s. lanceolatum (willd.) nesom var. lanceolatum (white panicle aster) qsqm, sssn; p; h; obg-133 s. oblongifolium (nutt.) nesom (aromatic aster) sssn; p; h; obg-129 s. patens (ait.) nesom (late purple aster) qsqm, sssn; p; h; obg-100 s. praealtum (poir.) nesom var. praealtum (willowleaf aster) sssn; p; h; obg-134 s. subulatum (michx.) nesom (eastern annual saltmarsh aster) daof, sssn, wetl; a; h; obg-105 taraxacum officinale* g.h. weber ex wiggers (common dandelion) daof; p; h; obg180 verbesina virginica l. (white crownbeard) qsqm; p; h; obg-096 vernonia arkansana dc. (arkansas ironweed) qsqm; p; h; obg-007 v. baldwinii torr. (baldwin’s ironweed) daof, sssn; p; h; obg-024 xanthium strumarium l. (rough cocklebur) wetl; a; h; obg-064 boraginaceae myosotis verna nutt. (spring forget-me-not) qsqm; a; h; obg-267 brassicaceae brassica nigra* (l.) w.d.j. koch (black mustard) daof; a; h; obg-195 capsella bursa-pastoris* (l.) medik. (shepherd’s purse) daof; a; h; obg-327 60 oklahomanative plantrecord volume 7,number 1,december 2007 hoagland & buthod cardamine parviflora l. var. arenicola (britt.) o.e. schulz (sand bittercress) qsqm; a; h; obg-194 erysimum repandum* l. (spreading wallflower) daof; a; h; obg-197 lepidium densiflorum schrad. (common pepperweed) daof, sssn; a; h; obg190 cactaceae opuntia humifusa (raf.) raf. (devil’s tongue) qsqm, sssn; p; s; obg-169 callitrichaceae callitriche heterophylla pursh (twoheaded water-starwort) wetl; a; h; obg-216 campanulaceae triodanis biflora (ruiz & pavón) greene (clasping venus’ looking-glass) sssn; a; h; obg-241 caprifoliaceae lonicera japonica* thunb. (japanese honeysuckle) qsqm; p; v; obg-214 sambucus nigra l. ssp. canadensis (l.) r. bolli (common elderberry) qsqm; p; s; obg265 symphoricarpos orbiculatus moench (coralberry) qsqm; p; s; obg-082 viburnum rufidulum raf. (rusty blackhaw) qsqm; p; s; obg-189 caryophyllaceae arenaria serpyllifolia* l. (thymeleaf sandwort)daof; a; h; obg-242 cerastium pumilum* w. curtis (european chickweed) daof; a; h; obg-191 dianthus armeria* l. (deptford pink) daof, sssn; a; h; obg-235 minuartia drummondii (shinners) mcneill (drummond’s stitchwort) sssn; a; h; obg-273 stellaria media* (l.) vill. (common chickweed) daof; a; h; obg-176 clusiaceae hypericum hypericoides (l.) crantz (st. andrew’s cross) qsqm; p; h; obg-098 h. punctatum lam. (spotted st. johnswort) qsqm; p; h; obg-295 convolvulaceae convolvulus arvensis* l. (field bindweed) daof; p; h; obg-253 cornaceae cornus drummondii c.a. mey. (roughleaf dogwood) qsqm; p; t; obg-232 cuscutaceae cuscuta cuspidata engelm. (cusp dodder) daof; p; h; obg-131 ebenaceae diospyros virginiana l. (common persimmon) qsqm; p; t; obg-213 euphorbiaceae acalphya monococca (engelm. ex gray) l. mill. & gandhi (slender threeseed mercury) sssn; a; h; obg-281 a. virginica l. (virginia threeseed mercury) qsqm; a; h; obg-119 chamaesyce maculata (l.) small (spotted sandmat) daof; a; h; obg-023 croton capitatus michx. (hogwort) daof, sssn; a; h; obg-084 c. monanthogynus michx. (prairie tea) daof, sssn; a; h; obg-146 c. willdenowii g. l. webster (willdenow’s croton) sssn; a; h; obg-043a euphorbia dentata michx. (toothed spurge) qsqm; a; h; obg-102 e. heterophylla l. (mexican fireplant) qsqm; a; h; obg-081 e. spathulata lam. (warty spurge) sssn; a; h; obg-277 phyllanthus caroliniensis walt. (carolina leafflower) daof; a; h; obg-072 61 oklahoma native plantrecord volume 7,number 1,december 2007 hoagland & buthod fabaceae amorpha canescens pursh (leadplant) sssn; p; s; obg-150 a. fruticosa l. (desert false indigo) sssn, wetl; p; s; obg-076 baptisia bracteata muhl. ex ell. var. leucophaea (nutt.) kartesz & gandhi (longbract wild indigo) sssn; p; h; obg193 cercis canadensis l. (eastern redbud) qsqm; p; t; obg-208 chamaecrista nictitans (l.) moench (partridge pea) daof, sssn; a; h; obg-087 crotalaria sagittalis l. (arrowhead rattlebox) sssn; p; h; obg-043 dalea purpurea vent. (violet prairie clover) sssn; p; h; obg-258 galactia volubilis (l.) britt. (downy milkpea) qsqm; p; h; obg-103 gleditsia triacanthos l. (honeylocust) qsqm; p; t; obg-115 kummerowia stipulacea* (maxin.) makino (korean clover) daof; a; h; obg-090 lathyrus hirsutus* l. (caley pea) daof; a; h; obg-282 lespedeza cuneata* (dum.-cours.) g. don (chinese lespedeza) daof, sssn; p; h; obg-107 l. repens (l.) w. bart. (creeping lespedeza) qsqm; p; h: obg-156 l. violacea (l.) pers. (violet lespedeza) qsqm; p; h; obg-035 l. virginica (l.) britt. (slender lespedeza) qsqm; p; h; obg-022 medicago lupulina* l. (black medick) daof; a; h; obg-172 melilotus officinalis* (l.) lam. (yellow sweetclover) daof; a; h; obg-060 mimosa nuttallii (dc.) b.l. turner (nuttall’s sensitive-briar) sssn; p; h; obg-264 pediomelum linearifolium (torr. & gray) j. grimes (narrowleaf indian breadfruit) sssn; p; h; obg-299 stylosanthes biflora (l.) b.s.p. (sidebeak pencilflower) sssn; p; h ; obg-252 trifolium dubium* sibthorp (suckling clover) daof; a; h; obg-243 t. pratense* l. (red clover) daof; a; h; obg290 vicia villosa* roth (winter vetch) daof; a; h; obg-288 fagaceae quercus muehlenbergii engelm. (chinkapin oak) qsqm; p; t; obg-145 q. shumardii buckl. (shumard’s oak) qsqm; p; t; obg-139 q. stellata wangenh. (post oak) qsqm; p; t; obg-083 q. velutina lam. (blackjack oak) qsqm; p; t; obg-335 gentianaceae sabatia campestris nutt. (texas star) sssn; a; h; obg-283 geraniaceae geranium carolinianum l. (carolina geranium) daof, sssn; a; h; obg-275 haloragaceae myriophyllum aquaticum* (vell.) verdc. (parrot feather watermilfoil) wetl; p; h; obg-202 juglandaceae carya illinoinensis (wangenh.) k. koch (pecan) qsqm; p; t; obg-135 c. texana buckl. (black hickory) qsqm; p; t; obg-128 krameriaceae krameria lanceolata torr. (trailing krameria) sssn; p; h; obg-250 lamiaceae hedeoma drummondii benth. (drummond’s false pennyroyal) sssn; p; h; obg-276 lamium amplexicaule* l. (henbit deadnettle) daof; a; h; obg-181 l. purpureum* l. (purple deadnettle) daof; a; h; obg-182 monarda fistulosa l. (wild bergamot) qsqm; p; h; obg-260 62 oklahomanative plantrecord volume 7,number 1,december 2007 hoagland & buthod prunella vulgaris l. (common selfheal) qsqm; p; h; obg-257 pycnanthemum tenuifolium schrad. (narrowleaf mountain mint) qsqm; p; h; obg-249 salvia azurea michx. ex lam. (azure blue sage) sssn; p; h; obg-093 scutellaria parvula michx. (small skullcap) sssn; p; h; obg-302 teucrium candense l. (canada germander) wetl; p; h; obg-101 linaceae linum medium (planch.) britt. (stiff yellow flax) sssn; a; h; obg-031 l. sulcatum riddell (grooved flax) sssn; a; h; obg-030 lythraceae ammannia auriculata willd. (eared redstem) wetl; a; h; obg-013 cuphea viscosissima jacq. (blue waxweed) wetl; a; h; obg-058 malvaceae callirhoe alcaeoides (michx.) gray (light poppymallow) sssn; p; h; obg-270 menispermaceae cocculus carolinus (l.) dc. (carolina coralbead) qsqm; p; h; obg-226 nelumbonaceae nelumbo lutea willd. (american lotus) wetl; p; h; obg-287 oleaceae fraxinus americana l. (white ash) qsqm; p; t; obg-155 ligustrum quihoui* carr. (waxyleaf privet) qsqm; p; s; obg-143 l. sinense* lour. (chinese privet) qsqm; p; s; obg-046 onagraceae gaura villosa torr. (wooly beeblossom) sssn; p; h; obg-010 ludwigia palustris (l.) ell. (marsh seedbox) wetl; p; h; obg-001 l. peploides (kunth) raven (floating primrosewillow) wetl; p; h; obg-067 oenothera linifolia nutt. (threadleaf eveningprimrose) sssn; a; h; obg-220 oxalidaceae oxalis stricta l. (common yellow oxalis) daof, sssn; p; h; obg-095 o. violacea (violet woodsorrel) qsqm, sssn; p; h; obg-199 passifloraceae passiflora incarnata l. (purple passionflower) sssn; p; h; obg-248 plantaginaceae plantago aristata michx. (largebraced plantain) qsqm; a; h; obg-225 p. virginica l. (virginia plantain) sssn; a; h; obg-271 polygalaceae polygala incarnata l. (procession flower) sssn; a; h; obg-274 polygonaceae polygonum pensylvanicum l. (pennsylvania smartweed) wetl; a; h; obg-298 p. punctatum ell. (dotted smartweed) wetl; a; h; obg-071 rumex crispus* l. (curly dock) daof, wetl; p; h; obg-209 portulacaceae phemeranthus parviflorum (nutt.) kiger (sunbright) sssn; p; h; obg-033 portulaca oleracea* l. (little hogweed) daof; a; h; obg-108 primulaceae samolus ebracteatus kunth (limewater brookweed) wetl; p; h; obg-137 63 oklahoma native plantrecord volume 7,number 1,december 2007 hoagland & buthod ranunculaceae delphinium carolinianum walt. (carolina larkspur) sssn; p; h; obg-229 rosaceae amelanchier arborea (michx. f.) fern. (common serviceberry) qsqm; p; t; obg-183 crataegus mollis scheele (arnold hawthorn) qsqm; p; s; obg-167 c. viridis l. (green hawthorn) qsqm; p; s; obg-186 prunus mexicana s. wats. (mexican plum) qsqm; p; t; obg-110 p. serotina ehrh. (black cherry) qsqm; p; t; obg-045 rosa multiflora* thunb. ex murr. (multiflora rose) qsqm; p; v; obg-263 r. setigera michx. (climbing rose) sssn; p; v; obg-292 rubus sp. (blackberry) daof, qsqm; p; v; obg-204 rubiaceae cephalanthus occidentalis l. (common buttonbush) wetl; p; s; obg-050 diodia teres walt. (poorjoe) sssn; a; h; obg070 galium aparine l. (stickywilly) daof, qsqm; a; h; obg-233 g. pilosum ait. var. puncticulosum (michx.) torr. & gray (hairy bedstraw) qsqm; p; h; obg-061 g. virgatum nutt. (southwestern bedstraw) sssn; a; h; obg-233 houstonia pusilla schoepf (tiny bluet) sssn; a; h; obg-174 sherardia arvensis* l. (blue fieldmadder) daof; a; h; obg-177 salicaceae populus deltoides bartr. ex marsh. (eastern cottonwood) wetl; p; t; obg-118 salix nigra marsh. (black willow) wetl; p; t; obg-051 sapotaceae sideroxylon lanuginosum michx. (gum bully) qsqm; p; t; obg-052 scrophulariaceae bacopa rotundifolia (michx.) wettst. (disk waterhyssop) wetl; p; h; obg-109 castilleja indivisa engelm. (entireleaf indian paintbrush) sssn; a; h; obg-200 lindernia dubia (l.) pennell (yellowseed false pimpernel) wetl; a; h; obg-078 nuttallanthus canadensis (l.) d.a. sutton (canada toadflax) sssn; a; h; obg-272 veronica polita* fries (gray field speedwell) daof; a; h; obg-173 solanaceae physalis pubescens l. (husk tomato) sssn; a; h; obg-014 solanum americanum p. mill. (american black nightshade) qsqm; a; h; obg-017 s. carolinense l. (carolina horsenettle) daof, sssn; p; h; obg-075 s. elaeagnifolium cav. (silverleaf nightshade) daof, sssn; p; h; obg-116 s. rostratum dunal (buffalobur nightshade) daof; a; h; obg-075 ulmaceae ulmus alata michx. (winged elm) qsqm; p; t; obg-245 u. rubra muhl. (slippery elm) qsqm; p; t; obg-246 valerianaceae valerianella radiata (l.) dufr. (beaked cornsalad) sssn; a; h; obg-268 verbenaceae verbena bracteata cav. ex lag. & rodr. (bigbract verbena) daof, sssn; a; h; obg-113 v. stricta vent. (hoary verbena) daof; p; h; obg-278 v. urticifolia l. (white vervain) qsqm; p; h; obg-141 64 oklahomanative plantrecord volume 7,number 1,december 2007 hoagland & buthod vitaceae parthenocissus quinquefolia (l.) planch. (virginia creeper) qsqm; p; v; obg-222 vitis cinerea (engelm.) millard (graybark grape) qsqm; p; v; obg-297 zygophyllaceae tribulus terrestris* l. (puncturevine) daof; a; h; obg-019 liliopsida alismataceae alisma subcordatum raf. (american water plantain) wetl; p; h; obg-286 sagittaria ambigua j.g. sm. (kansas arrowhead) -wetl ; p; h; obg-266 s. latifolia willd. (broadleaf arrowhead) -wetl ; p; h; obg-027 commelinaceae commelina erecta l. (whitemouth dayflower) daof, sssn; p; h; obg-028 tradescantia ohiensis raf. (bluejacket) sssn; p; h; obg-236 cyperaceae carex aggregata mackenzie (glomerate sedge) qsqm; p; g; obg-328 c. blanda dewey (eastern woodland sedge) qsqm; p; g; obg-326 c. festucacea schkuhr ex willd. (fescue sedge) qsqm; p; g; obg-330 c. frankii kunth (frank’s sedge) -wetl; p; g; obg-163 c. microdonta torr. & hook. (littletooth sedge) sssn; p; g; obg-203 cyperus croceus vahl (baldwin’s flatsedge) sssn; p; g; obg-166 c. echinatus (l.) wood (globe flatsedge) sssn; p; g; obg-099 c. odoratus l. (fragrant flatsedge) wetl; a; g; obg-164 c. squarrosus l. (bearded flatsedge) daof; a; g; obg-065 c. strigosus l. (strawcolored flatsedge) sssn; p; g; obg-161 c. virens michx. (green flatsedge) wetl; p; g; obg-147 eleocharis lanceolata fern. (daggerleaf spikerush) wetl; a; g; obg-325 e. obtusa (willd.) j.a. schultes (blunt spikerush) wetl; a; g; obg-127 e. palustris (l.) roemer & j.a. schultes (common spikerush) wetl; p; g; obg324 fimbristylis autumnalis (l.) roemer & j.a. schultes (slender fimbry) wetl; a; g; obg-069 f. puberula (michx.) vahl (hairy fimbry) wetl; p; g; obg-322 f. vahlii (lam.) link (vahl’s fimbry) wetl; a; g; obg-063 isolepis carinata hook. & arn. ex. torr. (keeled bulrush) wetl; a; g; obg-239 rhynchospora globularis (chapman) small (globe beaksedge) -wetl; p; g; obg-323 scirpus pendulus muhl. (rufous bulrush) wetl; p; g; obg-244 iridaceae sisyrinchium campestre bickn. (prairie blueeyed grass) sssn; p; h; obg-201 juncaceae juncus brachycarpus engelm. (whiteroot rush) wetl; p; g; obg-318 j. diffusissimus buckl. (slimpod rush) wetl; p; g; obg-055 j. interior wieg. (inland rush) sssn; p; g; obg-321 j. marginatus rostk. (grassleaf rush) wetl; p; g; obg-319 j. tenuis willd. (poverty rush) wetl; p; g; obg-168 lemnaceae lemna minor l. (common duckweed) wetl; p; h; obg-332 wolffia columbiana (columbian watermeal) wetl; p; h; obg-333 65 oklahoma native plantrecord volume 7,number 1,december 2007 hoagland & buthod liliaceae allium canadense (meadow garlic) sssn; p; h; obg-188 erythronium mesochoreum knerr (midland fawnlily) qsqm; p; h; obg-301 hypoxis hirsuta (l.) coville (common goldstar) sssn; p; h; obg-217 nothoscordum bivalve (l.) britt. (crowpoison) sssn; p; h; obg-179 orchidaceae spiranthes vernalis engelm. & gray (spring ladies’-tresses) sssn; p; h: obg-291 poaceae aegilops cylindrica* host (jointed goatgrass) daof; a; g; obg-210 agrostis hyemalis (walt.) b.s.p. (winter bentgrass) sssn; p; g; obg-307 a. perennans (walt.) tuckerman (upland bentgrass) qsqm; p; g; obg-306 andropogon gerardii vitman (big bluestem) sssn; p; g; obg-112 a. virginicus l. (virginia wildrye) sssn; p; g; obg-117 aristida dichotoma michx. (churchmouse threeawn) sssn; a; g; obg-038 bothriochloa laguroides (dc.) herter (silver beardgrass) daof, sssn; p; g; obg-005 bromus catharticus* vahl (rescuegrass) daof; a; g; obg-206 b. tectorum* l. (cheatgrass) daof; a; g; obg-309 buchloe dactyloides (buffalograss) sssn; p; g; obg-218 danthonia spicata (l.) beauv. ex roemer & j.a. schultes (poverty oatgrass) qsqm; p; g; obg-207 dichanthelium acuminatum (sw.) gould & c.a. clark var. fasciculatum (torr.) freckmann (western panicgrass) qsqm; p; g; obg124 d. depauperatum (muhl.) gould (starved panicgrass) qsqm; p; g; obg-311 d. malacophyllum (nash) gould (softleaf rosette grass) qsqm; p; g; obg-120 d. scoparium (lam.) gould (velvet panicum) qsqm; p; g; obg-317 d. villosissimum (nash) freckmann (whitehair rosette grass) qsqm; p; g; obg-310 digitaria cognata (j.a. schultes) pilger (carolina crabgrass) daof; p; g; obg003 d. sanguinalis (l.) scop. (hairy crabgrass) daof; a; g; obg-004 echinochola crus-galli* (l.) beauv. (barnyardgrass) wetl; a; g; obg-025 eleusine indica* (l.) gaertn. (indian goosegrass) daof; a; g; obg-068 elymus virginicus l. (virginia wildrye) qsqm, sssn; p; g; obg-312 eragrostis barrelieri* daveau (mediterranean lovegrass) daof; a; g; obg-008 e. intermedia a.s. hitchc. (plains lovegrass) qsqm; p; g; obg-021 e. spectabilis (pursh) steud. (purple lovegrass) daof, sssn; p; g; obg-018 hordeum pusillum nutt. (little barley) sssn; a; g; obg-211 leersia oryzoides (l.) sw. (rice cutgrass) wetl; p; g; obg-314 l. virginica willd. (whitegrass) -wetl; p; g; obg-140 lolium perenne* l. (perennial ryegrass) daof; p; g; obg-230 muhlenbergia sobolifera (muhl. ex willd.) trin. (rock muhly) qsqm; p; g; obg-029 panicum anceps michx. (beaked panicgrass) qsqm, sssn; p; g; obg-104 p. dichotomiflorum michx. (fall panicgrass) wetl; a; g; obg-122 p. philadelphicum bernh. ex trin. (philadelphia panicgrass) sssn; a; g; obg-123 p. rigidulum bosc ex nees (redtop panicgrass) wetl; p; g; obg-162 p. virgatum l. (switchgrass) sssn; p; g; obg094 paspalum pubiflorum rupr. ex fourn. (hairyseed paspalum) sssn, wetl; p; g; obg-053 phalaris caroliniana walt. (carolina canarygrass) wetl; a; g; obg-228 66 oklahomanative plantrecord volume 7,number 1,december 2007 hoagland & buthod poa annua* l. (annual bluegrass) qsqm; a; g; obg-178 schedonnardus paniculatus (nutt.) trel. (tumblegrass) daof, sssn; p; g; obg057 schedonorus phoenix* (scop.) holub daof; p; g; obg-238 schizachyrium scoparium (michx.) nash (little bluestem) sssn; p; g; obg-149 setaria parviflora (poir.) kerguélen (marsh bristlegrass) wetl; p; g; obg-158 s. pumila* (poir.) roemer & j.a. schultes (yellow foxtail) daof; a; g; obg-009 sorghastrum nutans (l.) nash (indian grass) sssn; p; g; obg-336 sorghum halepense* (l.) pers. (johnsongrass) daof, sssn; p; g; obg-111 sphenopholis obtusata (michx.) scribn. (prairie wedgescale) qsqm; p; g; obg-308 sporobolus cryptandrus (torr.) gray (sand dropseed) sssn; p; g; obg-016 steinchisma hians (ell.) nash (gaping grass) wetl; p; g; obg-262 tridens strictus (nutt.) nash (longspike tridens) daof, sssn; p; g; obg-044 vulpia elliotea (raf.) fern. (squirreltail fescue) sssn; a; g; obg-315 v. octoflora (walt.) rydb. (sixweeks fescue) sssn; a; g; obg-315 potamogetonaceae potamogeton diversifolius raf. (waterthread pondweed) wetl; p; h; obg-160 smilacaceae smilax rotundifolia l. (roundleaf greenbriar) qsqm; p; v; obg-223 s. tamnoides l. (bristly greenbriar) qsqm; p; v; obg-224 typhaceae typha domingensis pers. (southern cattail) wetl; p; h; obg-126 microsoft word 2014onpr_20150103final_txt.docx oklahoma native plant record 3 volume 14, december 2014 foreword we are very excited that such a wide range of contributors, from gardeners and students to professional botanists and ecologists, submitted articles for volume 14. this demonstrates the strength of our membership and helps us, as a society, bring all our interests together in a way that best promotes our goal of encouraging the study, protection, propagation, appreciation, and use of the native plants of oklahoma. our “historic” article this year is about the flora of kiowa county. there is very little historic plant distribution information from that far southwestern part of the state, but we hope that lottie o. baldock’s 1938 master’s thesis will spark interest there. this article will be of special value to today’s botanists and ecologists studying historic species distributions and environmental changes. stan rice and sonya ross have done a small scale study of the different effects our warmer winter temperatures might have on the timing of spring budburst in three native tree species: sycamore (platanus occidentalis), pecan (carya illinoensis), and sweetgum (liquidambar styraciflua). katie keil raises awareness of three invasive species by updating and proposing revisions in the formats of distribution maps for purple loosestrife (lythrum salicaria), japanese honeysuckle (lonicera japonica), and multiflora rose (rosa multiflora). sadie gordon reports on research that she has done regarding the use of native species in historic, domestic gardens in the ne oklahoma, se kansas, sw missouri, and nw arkansas region. all three of these articles will pique the interests of both professional and amateur botanists and gardeners. angela mcdonnell’s article will be valuable to both professional and amateur field biologists. she describes the characteristic features and distributions of two milkweed vines, matelea biflora and m. cynanchoides, and provides a valuable key for the species in that genus that, until now, have been difficult to discern. educators will be inspired by gloria caddell’s critic’s choice essay. as professor of botany at the university of central oklahoma, she describes pollination studies done by undergraduate students at uco’s field site at lake arcadia east of edmond as well as the arcadia conservation education area. as we continue to develop the quality of the journal and its usefulness for botanists, researchers, enthusiasts, and gardeners, the global footprint of the society grows. statistics show that, in addition to the hundreds of printed volumes sold, valuable information from the oklahoma native plant record has been accessed thousands of times from oklahoma state university’s ejournals digital collections. the oklahoma native plant record is listed in the “directory of open access journals”, and our abstracts are indexed in the “centre for agricultural bioscience international”, which is based in the u.k. our editorial board has included many society members over the years, and the record could not have reached those milestones without their help. we are especially grateful to paula shryock, who has been our valuable, multi-talented production editor in this process since 2008. sandy graue has updated our previous electronic versions, produced between 2001 and 2010, and reformatted them for upload into the osu digital collections website. she has been our electronic production editor since she joined us in 2010, and she now uploads each new volume of the record. we thank both of them for the time and work they put into getting our journal out each year. we also appreciate the many members and colleagues who have authored and reviewed articles, as well as the members who have served on our editorial board as technical assistants and proof-readers. we thank them all for their support. sheila strawn, managing editor oklahoma native plant record, volume 11, number 1, december 2011 oklahoma native plant record volume 11, december 2011 clark, l. g. 4 survey of the vascular flora of the boehler seeps and sandhills preserve submitted to the department of botany of oklahoma state university in partial fulfillment of the requirements for the degree of doctor of philosophy july 1997 linda gatti clark e-mail: gatti.clark@gmail.com located in atoka county of southcentral oklahoma, the nature conservancy’s boehler seeps and sandhills preserve comprises sandhills, acidic hillside seeps, marshes, intermittent and permanent streams, and shallow lakes. the sandhills are the site of the highest quality, old-growth vegetation of the western gulf coastal plains (s. orzell, pers. comm. to ian butler). the flora is a unique assemblage of plants that is present nowhere else in the state and considered globally rare. approximately 400 species are believed to be present (jones 1993). more than 20 rare species have been reported to occur in the area, including some that are globally rare (oklahoma natural heritage inventory 1997). eriocaulon kornickianum, for example, is designated g2 and s1. other rare species in the preserve include penstemon murrayanus (g4, s1s3), polygonella americana (g5, s1s2), and paronychia drummondii (g4g5, s1s2). prior to this study, our knowledge of the vascular plant species in the preserve was incomplete. although several partial lists of its flora had been compiled, a systematic survey of the area to inventory all of the plants had not been conducted. such information is essential for understanding the ecology of the site and making decisions about its management. this study was undertaken to provide this information. specific objectives were to: (1) compile a list of the terrestrial and aquatic vascular plant species present and (2) prepare a set of herbarium specimens to document the preserve’s flora. this note summarizes my findings and provides a reference to the information compiled in gatti clark (1997). boehler seeps and sandhills preserve the preserve is a 235 ha site located in southern atoka county, oklahoma, approximately 11 miles north of boswell (s25 & 26, t4s, r13w; boswell nw quad). it comprises two tracts bisected by a paved country road and is located in the watersheds of muddy boggy and clear boggy creeks. situated in the dissected coastal plain geomorphic province and western coastal plain land resource area (johnson et al. 1979, usda natural resources conservation service 1992), its underlying strata are cretaceous in age. also underlying the preserve is the antlers sandstone aquifer that is composed primarily of non-marine sand and clay, and marine limestone and clay up to 915 m thick and is saturated with water that has a moderate to high mineral content (johnson et al. 1979). the water table is generally within 1-1.3 m of the surface, with seeps occurring where it reaches the surface (jones 1993, pers. comm.). soil series of the site are the bernow-romia complex, 8-12% slopes; boggy fine sandy loam; and larue loamy fine sand, 0-8% slopes (shingleton and watterson 1979). all are susceptible to erosion by both water and wind. precipitation in the area of the preserve https://doi.org/10.22488/okstate.17.100081 oklahoma native plant record 5 volume 11, december 2011 clark, l. g. occurs primarily in the spring and summer, and averages 119 cm per year (ruffner 1980). the average growing season is 255 days; mean maximum annual temperature is 24.5˚ c and mean minimum is 11˚ c; the average number of days below 0˚ c is 52 (ruffner 1980). vegetation of the area is oak-hickory forest (duck and fletcher 1943) with several communities present. quercus stellata, carya texana, and sideroxylon lanuginosum spp. lanuginosum (=bumelia lanuginose) dominate and form an almost continuous canopy in the drier areas. typically a thick leaf layer is present on the ground, and understory vegetation is sparse. juniperus virginiana and pinus echinata, while not dominant, can be found scattered throughout the preserve. common woody understory species are nyssa sylvatica, vaccinium arboreum, berchemia scandens, and vitis rotundifolia. herbaceous understory taxa include galium arkansanum, g. obtusum, tephrosia virginia, carex spp., cyperus spp., and juncus spp. scattered throughout the preserve and most conspicuous are glades dominated by aristida desmantha and selaginella rupestris. other glades are present and are dominated by various grasses, such as panicum spp., mosses, and forbs, such as gaillardia aestivalis and hieracium longipilum. at the glade edges, trees other than the forest dominants are encountered, in particular quercus incana. its saplings are occasionally found in the centers of the glades. hassel and boehler lakes are small, shallow bodies of water maintained by beaver dams. both have dense stands of emergent and floating-leaved species at their edges and open water in their centers. dominant taxa include typha angustifolia, nuphar lutea, and nymphaea odorata. often quite abundant, free-floating species are azolla caroliniana and utricularia biflora. two types of seeps are present in the preserve. one has water percolating slowly to the surface and accumulating in one area because of the topography. the ground surface has a spongy feel because of the thick carpet of vegetation, primarily mosses; sphagnum lescurii and polytrichum commune in particular dominate. occupying natural drainage ways, the second type is characterized by water flowing away from the seepage point. ferns, sedges, and rushes typically are in abundance along these watercourses. between the lakes and the seeps are marshes dominated by osmunda regalis, o. cinnamonea, cephalanthus occidentalis, scirpus spp., rhynchospora spp., and cyperus spp. associated with boehler lake are rather deep drainages that resemble sloughs. they have less vegetation and are often banked by large trees such as quercus falcata, q. nigra, and q. phellos. method of survey a systematic collection of the terrestrial and aquatic vascular plants occurring in the preserve was conducted during the 1994 and 1995 growing seasons. the area was divided into three survey units using roads and fences as boundaries. each unit was traversed on foot several times during the growing season. plants were collected in both the flowering and fruiting stages, and prepared using standard herbarium techniques (radford et al. 1974). one set of 440 voucher herbarium specimens (appendix h) was prepared and deposited in the oklahoma state university herbarium (okla). specimens previously collected by conservancy personnel were identified and included in the inventory (gatti clark 1997, appendices i and j). identification was accomplished using the resources of the herbarium. nomenclature used was based primarily on that of waterfall (1969), correll and johnston (1979), and gray’s manual of botany (fernald 1950). common names were taken from correll and johnston (1979) and taylor and taylor (1994). oklahoma native plant record volume 11, december 2011 clark, l. g. 6 flora of the preserve three hundred forty-five species in 225 genera and 84 families were encountered in this survey or by previous workers (gatti clark 1997, appendices h, i, and j). three families, asteraceae (56 taxa), poaceae (41 taxa), and cyperaceae (35 taxa), composed 38% of the preserve’s vascular flora. other large families were the fabaceae (21 taxa), apiaceae (11 taxa), lamiaceae (10 taxa), and scrophulariaceae (10 taxa). the largest genera present were carex, represented by 17 species, and panicum, represented by 12 species. species designated by the u.s. fish and wildlife service (1996) as endangered, threatened, or candidate (formally category 1) were not encountered. species ranked by the onhi (1997) as s1 or s2 and present in the preserve included the previously mentioned eriocaulon kornickianum (g2, s1), penstemon murrayanus (g4, s1s3), polygonella americana (g5, s1s2), and paronychia drummondii (g4g5, s1s2). other rare species are listed in the table. although demonstrably secure globally and ranked g4 or g5 by onhi, several species of interest were found in the preserve. the insectivorous drosera brevifolia and the mycotrophic/parasitic monotropa hypopithys were encountered. lycopodiella appressa (=lycopodium appressum) is reported for the site but was not seen in this study or in collections of the nature conservancy personnel (l. k. magrath, pers. comm.). with 22 rare taxa reported for the site, monitoring of the bluejack oak sandhills and seep communities should continue. the communities and its assemblage of plants at boehler seeps and sandhills preserve are rare within the state and deserve continued study. literature cited correll, d. s. and m. s. johnston. 1970. manual of the vascular plants of texas. texas research foundation, renner, tx. duck, l. g. and j. b. fletcher. 1943. a game type map of oklahoma. state of oklahoma game and fish department, division of wildlife restoration. oklahoma biological survey, norman, ok. fernald, m. l. 1950. gray’s manual of botany. 8th ed. american book company, new york. gatti clark, l. c. 1997. floristic and biosystematic investigations in plant taxonomy. ph.d. dissertation. oklahoma state university, stillwater, ok. jones, n. 1993. a baseline study of the water quality, vegetative gradients, and hydrology of boehler seeps and sandhills preserve. report. the nature conservancy, tulsa, ok. johnston, k. s., c. c. branson, n. m. curtis, jr., w. e. ham, w. e. harrison, m. v. marcher, and j. f. roberts. 1979. geology and earth resources of oklahoma. educational publication 1. oklahoma geological survey, norman, ok. oklahoma natural heritage inventory. 1997. short working list of rare plants. version of 22 january 1997. oklahoma biological survey, norman, ok. radford, a. e., w. c. dickison, j. r. massey, and c. r. bell. 1974. vascular plant systematics. harper and row, new york. ruffner, j. a. 1980. climate of the united states: national oceanic and atmospheric administration narrative summaries, tables, and maps for each state with overview of state climatologist programs. 2nd ed. volume 2. gale research company, detroit, mi. oklahoma native plant record 7 volume 11, december 2011 clark, l. g. shingleton, l. c. and a. watterson, jr. 1979. soil survey of atoka county, oklahoma. usda soil conservation service, stillwater, ok. taylor, r. j. and c. e. s. taylor. 1994. an annotated list of the ferns, fern allies, gymnosperms, and flowering plants of oklahoma. 3rd ed. published by authors, durant, ok. usda natural resources conservation service. 1992. map of major land resource areas of oklahoma. stillwater, ok. us fish and wildlife service. 1996. endangered and threatened wildlife and plants. october 31, 1996. 50 cfr 17.11-17.12. washington, d. c. waterfall, u. t. 1969. keys to the flora of oklahoma. 4th ed. published by author, stillwater, ok. oklahoma native plant record volume 11, december 2011 clark, l. g. 8 table taxa of boehler seeps and sandhills preserve that are ranked as rare by the oklahoma natural heritage inventory (onhi) onhi rankings* scientific name common name global state agalinis tenuifolia (vahl.) raf. slender leaved agalinis g5 s2s3 aristolochia reticulata jacq. netleaved snakeroot g4 s2 azolla caroliniana willd. mosquito fern g5 s2 brasenia schreberi j. f. gmel. water-shield g5 s1 carex hyalina boott whitesheath sedge g5 s1 carex swanii (fernald) mack. swan sedge g5 s1 carya myristiciformis (michx. f.) nutt. nutmeg hickory g5 s2s3 drosera brevifolia pursh sundew g5 s2s3 dulichium arundinaceum (l.) britton threeway sedge g5 s1 eriocaulon kornickianum van heurch and müll.arg. small pipewort g2 s1 galium arkansanum a. gray arkansas bedstraw g5 s1s2 houstonia micrantha (shinners) terrell (=hedyotis australis w.h. lewis & d.m. moore) bluet g4g5 s1s2 iris virginica l. southern blue flag g5 s2? monotropa hypopithys l. pinesap g5 s1 paronychia drummondii torr. & a. gray drummond's nailwort g4g5 s1s2 penstemon murrayanus hook. cupleaf beardtongue g4 s1s3 platanthera flava (l.) lindl. pale green orchid g4 s1 polygonella americana (fisch. & c.a. mey.) small southern jointweed g5 s1s2 quercus incana bartram bluejack oak g5 s1s2 rhynchospora caduca elliott anglestem beakrush g5 s1 saccharum giganteum (walter) pers. (=erianthus giganteus (walter) p. beauv.) giant plumegrass g5 s1s2 sacciolepis striata (l.) nash american cupscale g5 s2 *onhi global rankings: g2 imperiled globally because of its rarity (6 to 20 occurrences or few remaining individuals or acres) or because of other factors demonstrably making it vulnerable to extinction throughout its range. g4 apparently secure globally, though it may be quite rare in parts of its range, especially at the periphery. g5 demonstrably secure globally though it may be quite rare in parts of its range, especially at the periphery. s1 critically imperiled in oklahoma because of extreme rarity (5 or fewer occurrences or very few remaining individuals or acres) or because of some factor of its biology making it especially vulnerable to extinction. s2 imperiled in oklahoma because of extreme rarity (6 to 20 occurrences or few remaining individuals or acres) or because of other factors making it very vulnerable to extinction throughout its range. s3 rare and local in oklahoma (thought it may be abundant at some of its locations); in the range of 21-100 occurrences. oklahoma native plant record volume 11, december 2011 clark, l.g. 9 appendix vascular plant collections from boehler seeps and sandhills preserve arranged by family. collections of l. c. gatti clark and the nature conservancy personnel. [ed. notes: all plants are collected by l. gatti clark, unless indicated by an asterisk * for the nature conservancy or a tilde ~ for plants collected by both. nomenclature has been updated using the plants database (plants.usda.gov/plants).] fern allies selaginellaceae – spikemoss family selaginella rupestris (l.) spring rock spikemoss ferns azollaceae – azolla family azolla caroliniana willd. mosquito fern dryopteridaceae – wood fern family onoclea sensibilis l. sensitive fern woodsia obtusa (spreng.) torr. blunt-lobed cliff fern ophioglossaceae – adder’s-tongue family botrychium virginianum (l.) sw. rattlesnake fern osmundaceae – royal fern family osmunda cinnamomea l. cinnamon fern ~ osmunda regalis l. var. spectabilis royal fern (willd.) a. gray gymnosperms cupressaceae – cypress family juniperus virginiana l. eastern redcedar pinaceae – pine family ~ pinus echinata mill. shortleaf pine angiosperms liliopsida – monocots alismataceae – water plantain family alisma subcordatum raf. water plaintain echinodorus tenellus (mart.ex schult. f.) buchenau lanceleaf burweed sagittaria latifolia willd. wapato, duck potato commelinaceae – spiderwort family commelina erecta l. erect day flower tradescantia ohiensis raf. ohio spiderwort oklahoma native plant record volume 11, december 2011 clark, l. g. 10 cyperaceae – sedge family ~ carex bicknellii britton bicknell’s sedge carex blanda dewey loose flowered sedge carex cherokeensis schwein. cherokee sedge * carex complanata torr. & hook. sedge carex crinita lam. fringed sedge ~ carex digitalis willd. sedge * carex frankii kunth frank’s sedge carex granularis muhl. ex willd. meadow sedge carex gravida l.h. bailey heavy sedge carex hyalina boott whitesheath sedge carex lupulina muhl. ex willd. hop sedge * carex muehlenbergii schkuhr ex willd. muhlenberg’s sedge carex normalis mack. sedge carex retroflexa muhl. ex willd. reflexed sedge * carex squarrosa l. sedge carex swanii (fernald) mack. swan sedge carex vulpinoidea michx. fox sedge ~ cyperus echinatus (l.) alph. wood globe flatsedge (=c. ovularis (michx.) torr.) cyperus retroflexus buckley one-flower flatsedge (=c. uniflorus torr. & hook., non thunb. * cyperus strigosus l. false nutgrass cyperus virens michx. green flatsedge dulichium arundinaceum (l.) britton threeway sedge eleocharis acicularis var. acicularis (l.) roem. & schult. needle spikesedge (=e. acicularis (l.) roem. & schult. var. gracilescens) ~ eleocharis compressa sull. flatstem spikesedge eleocharis engelmannii steud. engleman’s spikesedge ~ eleocharis lanceolata fernald blunt spikesedge (=e. obtusa (willd.) schultes var. lanceolata (fernald) gilly eleocharis parvula (roem. & schult.) link ex bluff, nees. dwarf spikesedge & schauer (=e. parvula (roem. & schult.) link var. anachaeta (torr.) svens. eleocharis tenuis (willd.) schult. slender spikesedge var. verrucosa (svens.) svens. ~ isolepis carinata hook. & arn. ex torr. bulrush (=scirpus koilolepis (steud.) gleason * lipocarpha aristulata (coville) g. tucker hemicarpa (=hemicarpha aristulata (coville) smyth rhynchospora caduca elliott anglestem beakrush ~ rhynchospora capitellata (michx.) vahl false bogrush rhynchospora glomerata (l.) vahl clustered beakrush scleria ciliata michx. fringed nutrush scleria triglomerata michx. whip nutrush oklahoma native plant record volume 11, december 2011 clark, l.g. 11 eriocaulaceae – pipewort family ~ eriocaulon kornickianum van heurch & müll. arg. small pipewort iridaceae – iris family iris virginica l. southern blue flag sisyrinchium angustifolium mill. blue-eyed grass juncaceae – rush family juncus acuminatus michx. jointed rush ~ juncus coriaceus mack. leathery rush juncus effusus l. bog rush ~ juncus marginatus rostk. grassleaf rush ~ juncus scirpoides lam. needlepod rush juncus tenuis willd. tender rush luzula bulbosa (alph. wood) smyth & smyth bulb woodrush lemnaceae – duckweed family spirodela polyrrhiza (l.) schleid. duck meat liliaceae – lily family ~ allium canadense l. wild onion ~ hypoxis hirsuta (l.) coville yellow stargrass orchidaceae – orchid family * platanthera flava (l.) lindl. pale green orchid poaceae – grass family agrostis perennans (walter) tuck. autumn bentgrass * andropogon gerardii vitman big bluestem andropogon ternarius michx. splitbeard bluestem aira elegans willd. ex kunth annual silver hairgrass aristida desmantha trin. & rupr. curly threeawn bouteloua hirsuta lag. hairy grama bromus arvensis l. (=b. japonicus thunb.) japanese brome bromus catharticus vahl rescue grass * bromus hordeaceus l. soft chess ~ cenchrus spinifex cav. (=c. incertus m. a. curtis) sandbur ~ chasmanthium latifolium (michx.) yates inland seaoats ~ chasmanthium sessiliflorum (poir.) yates spike-inland seaoats (=c. laxum (l.) yates spp. sessiliflorum (poir.) l. clark) danthonia spicata (l.) p. beauv. ex roem. & schult. poverty oatgrass ~ dichanthelium acuminatum (sw.) gould & c.a. clark wooly panicum var. fasciculatum (torr.) freckmann (=panicum lanuginosum eliott, non bosc ex spreng.) dichanthelium boscii (poir.) gould & c.a. clark bosc panicum (=panicum boscii poir.) oklahoma native plant record volume 11, december 2011 clark, l. g. 12 dichanthelium depauperatum (muhl.) gould slimleaf panicum (=panicum depauperatum muhl.) dichanthelium dichotomum (l.) gould var. dichotomum forked panicum (=panicum dichotomum l.) dichanthelium linearifolium (scribn. ex nash) gould slimleaf panicum (=panicum linearifolium scribn.) ~ dichanthelium oligosanthes (schult.) gould var. oligosanthes small panicgrass (=panicum oligosanthes schult.) dichanthelium ravenelli (scribn. & merr.) gould panicum (=panicum ravenelii scribn. & merr.) dichanthelium sphaerocarpon (elliott) gould var. sphaerocarpon leafy panicum (=panicum sphaerocarpon elliott) ~ elymus virginicus l. virginia wildrye * eragrostis capillaris (l.) nees lacegrass * eragrostis hirsuta (michx.) nees bigtop lovegrass eragrostis secundiflora j. presl red lovegrass eragrostis spectabilis (pursh) steud. purple lovegrass gymnopogon ambiguus (michx.) britton, sterns & poggenb. broadleaf skeletongrass leersia oryzoides (l.) sw. swartz cutgrass ~ panicum anceps michx. beaked panicum panicum dichotomiflorum michx. fall panicum * paspalum laeve michx. field paspalum ~ paspalum setaceum michx. thin paspalum * saccharum giganteum (walter) pers. giant plumegrass (=erianthus giganteus (walter) p. beauv.) sacciolepis striata (l.) nash american cupscale setaria parviflora (poir.) kerguélen knotroot bristlegrass (=s. geniculata (willd.) p. beauv., nom. illeg.) ~ sorghum halepense (l.) pers. johnsongrass sphenopholis obtusata (michx.) scribn. prairie wedgescale ~ steinchisma hians (elliott) nash (=panicum hians elliott) gaping panicum ~ tridens flavus (l.) hitchc. purpletop vulpia octoflora (walter) rydb. sixweeks fescue zizaniopsis miliacea (michx.) döll. & asch. southern wildrice potemogetonaceae – pondweed family potamogeton pulcher tuck. spotted pondweed smilacaceae – catbriar family smilax bona-nox l. greenbrier smilax rotundifolia l. common greenbrier typhaceae – cattail family typha angustifolia l. narrow-leaved cattail oklahoma native plant record volume 11, december 2011 clark, l.g. 13 magnoliopsida dicots acanthaceae – acanthus family ruellia humilis nutt. fringed leaf ruellia amaranthaceae – amaranth family froelichia floridana (nutt.) moq. snake cotton anacardiaceae – sumac family rhus aromatica aiton lemon sumac rhus copallinum l. (=r. copallina l., orth. var.) winged sumac toxicodendron radicans (l.) kuntze poison ivy apiaceae – carrot family * chaerophyllum tainturieri hook. hairy fruit wild chervil ~ daucus pusillus michx. southwestern carrot eryngium prostratum nutt. ex dc. creeping eryngo * hydrocotyle verticillata thunb. whorled pennywort ~ ptilimnium capillaceum (michx.) raf. threadleaf mockbishopweed ~ sanicula canadensis l. black snakeroot sanicula odorata (raf.) k.m. pryer & l.r. phillippe cluster snakeroot (=s. gregaria e.p. bicknell) spermolepis divaricata (walter) raf. ex ser. forked scaleseed spermolepis echinata (nutt. ex dc.) a. heller bristly scaleseed spermolepis inermis (nutt. ex dc.) mathias & constance spreading scaleseed torilis arvensis (huds.) link hedge parsley apocynaceae – dogbane family apocynum cannabinum l. indianhemp aquifoliaceae – holly family ilex decidua walter deciduous holly aristolochiaceae – birthwort family ~ aristolochia reticulata jacq. netleaved snakeroot asclepiadaceae – milkweed family asclepias tuberosa l. butterfly milkweed asclepias verticillata l. whorled milkweed asclepias viridis walter green milkweed matelea biflora (raf.) woodson twoflower milkvine asteraceae – sunflower family * achillea millefolium l. yarrow * ambrosia artemisiifolia l. common ragweed ~ ambrosia bidentata michx. lanceleaf ragweed * ambrosia trifida l. giant ragweed ~ antennaria parlinii fernald plainleaf pussytoes oklahoma native plant record volume 11, december 2011 clark, l. g. 14 astranthium integrifolium (michx.) nutt. western daisy * bidens aristosa (michx.) britton tickseed sunflower centaurea americana nutt. american basket flower * chaetopappa asteroides nutt. ex dc. least daisy ~ chrysopsis pilosa nutt. softhair golden aster * cirsium altissimum (l.) hill tall thistle cirsium horridulum michx. bull thistle * conoclinium coelestinum (l.) dc. blue boneset (=eupatorium coelestinum l.) ~ conyza canadensis (l.) cronquist horseweed ~ coreopsis grandiflora hogg ex sweet bigflowered tickseed ~ croptilon divaricatum (nutt.) raf. scratch daisy (=haplopappus divaricatus (nutt.) a. gray) ~ echinacea pallida (nutt.) nutt. pale coneflower ~ elephantopus carolinianus raeusch. elephant’s foot * erechtites hieracifolia (l.) raf. ex dc. fireweed ~ erigeron strigosus muhl. ex willd. daisy fleabane * eupatorium perfoliatum l. boneset evax prolifera nutt. ex dc. rabbit’s tobacco evax verna raf. var. verna (=e. multicaulis dc.) rabbit’s tobacco gaillardia aestivalis (walter) h. rock prairie gaillardia ~ gamochaeta purpurea (l.) cabrera (=gnaphalium purpureum l.) purple cudweed * helenium amarum (raf.) h. rock sneezeweed * helianthus angustifolius l. narrow-leaf sunflower ~ helianthus hirsutus raf. hairy sunflower heterotheca villosa (pursh) shinners var. villosa roughhair golden aster (=chrysopsis villosa (pursh.) nutt. ex dc.) ~ hieracium gronovii l. hawkweed hieracium longipilum torr. longbeard hawkweed hymenopappus scabiosaeus l’her. old plainsman ~ krigia cespitosa (raf.) k. l. chambers common dwarf dandelion ~ krigia dandelion (l.) nutt. potato dandelion krigia virginica (l.) willd. dwarf dandelion lactuca canadensis l. wild lettuce * lactuca sativa l. prickly lettuce liatris aspera michx. tall gayfeather * liatris elegans (walter) michx. beautiful gayfeather liatris squarrosa (l.) michx. gayfeather * mikania scandens (l.) willd. climbing hempweed packera obovata (muhl. ex willd.) w.a. weber & a. love roundleaf groundsel (=senecio obovatus muhl. ex willd. var. rotundus britton) * pluchea camphorata (l.) dc. camphorweed pseudognaphalium obtusifolium (l.) hilliard & b.l. burtt sweet everlasting ssp. obtusifolium (=gnaphalium obtusifolium l.) ~ pyrrhopappus carolinianus (walter) dc. false dandelion * rudbeckia grandiflora (d. don) j.f. gmel. ex dc. mexican hat ~ rudbeckia hirta l. blackeyed susan oklahoma native plant record volume 11, december 2011 clark, l.g. 15 solidago canadensis l. common prairie goldenrod solidago missouriensis nutt. missouri goldenrod * solidago odora aiton fragrant goldenrod * solidago rugosa mill. rough-leaved goldenrod ~ solidago ulmifolia muhl. ex willd. elmleaf goldenrod * symphyotrichum patens (aiton) g.l. nesom var. patens late purple aster (=aster patens aiton) * symphyotrichum subulatum (michx) g.l. nesom salt marsh aster (=aster subulatus michx.) ~ verbesina helianthoides michx. yellow crownbeard * vernonia baldwinii torr. western ironweed balsaminaceae – touch-me-not family impatiens capensis meerb. spotted touch-me-not berberidaceae – barberry family podophyllum peltatum l. may apple bignoniaceae – trumpet creeper family campsis radicans (l.) seem. ex bureau trumpet creeper boraginaceae – borage family ~ lithospermum caroliniense (walter ex j.f. gmel.) macmill. plains pucoon myosotis verna nutt. early scorpiongrass brassicaceae – mustard family cardamine pensylvanica muhl. ex willd. bitter cress ~ lepidium virginicum l. poorman’s peppergrass buddlejaceae – butterfly-bush family polypremum procumbens l. juniperleaf cabombaceae – water shield family brasenia schreberi j. f. gmel. water shield callitrichaceae – water-starwort family callitriche heterophylla pursh water-starwort campanulaceae – bellflower family ~ triodanis perfoliata (l.) nieuwl. clasping venus looking-glass caprifoliaceae – honeysuckle family lonicera japonica thunb. japanese honeysuckle symphoricarpos orbiculatus moench buckbrush viburnum rufidulum raf. rusty blackhaw oklahoma native plant record volume 11, december 2011 clark, l. g. 16 caryophyllaceae – pink family arenaria serpyllifolia l. thyme-leaved sandwort paronychia drummondii torr. & a. gray drummond’s nailwort stellaria media (l.) vill. chickweed ceratophyllaceae – hornwort family ceratophyllum demersum l. coontail cistaceae – rockrose family lechea villosa elliott pinweed clusiaceae – mangosteen family ~ hypericum drummondii (grev. & hook.) torr. & a. gray nits-and-lice ~ hypericum hypericoides (l.) crantz st. andrew’s cross ~ hypericum prolificum l. (=h. spathulatum (spach.) steud. st. john’s wort convolvulaceae – morning glory family ~ ipomoea pandurata (l.) g. mey. wild potatovine ~ stylisma pickeringii (torr. ex m.a. curtis) a. gray stylisma cornaceae – dogwood family cornus florida l. flowering dogwood nyssa sylvatica marsh. black gum droseraceae – sundew family ~ drosera brevifolia pursh sundew ericaceae – heath family ~ vaccinium arboreum marsh. farkleberry euphorbiaceae – spurge family acalypha rhomboidea raf. rhombic copperleaf acalypha virginica l. three seeded mercury chamaesyce serpens (kunth) small (=euphorbia serpens kunth) round-leaved spurge cnidoscolus texanus (müll. arg.) small texas bullnettle croton capitatus michx. woolly croton croton glandulosus l. sand croton croton willdenowii g.l. webster (=crotonopsis elliptica willd.) rush-foil ~ stillingia sylvatica l. queen’s delight fabaceae – pea family apios americana medik. ground nut astragalus distortus torr. & a. gray bentpod milkvetch * baptisia bracteata muhl. ex elliott plains wild indigo baptisia leucophaea nutt. var. leucophaea (nutt.) white wild indigo kartesz & gandhi (=b. leucophaea nutt.) cercis canadensis l. redbud oklahoma native plant record volume 11, december 2011 clark, l.g. 17 chamaecrista fasciculata (michx.) greene var. fasciculata partridge pea (=cassia fasciculata michx.) chamaecrista nictitans (l.) moench ssp. nictitans var. nictitans sensitive pea (=cassia nictitans l.) clitoria mariana l. butterfly pea dalea phleoides (torr. & a. gray) shinners var. phleoides longbract prairie clover (=petalostemon phleoides torr. & a. gray) desmodium paniculatum (l.) dc. var. paniculatum tall tickclover desmodium sessilifolium (torr.) torr. & a. gray sessile-leaved tickclover desmodium viridiflorum (l.) dc. velvetleaf tickclover galactia regularis (l.) britton, sterns & poggenb. downey milkpea gleditsia triacanthos l. (=caesalpiniaceae family) honey locust lespedeza stuevei nutt. tall lespedeza mimosa nutallii (dc. ex britton & rose) b.l. turner sensitive briar (=schrankia nuttallii (dc. ex britton & rose) standl.) mimosa microphylla dryand. (=schrankia ucinata willd.) catclaw briar ~ orbexilum pendunculatum (mill.) rydb. var. psoralioides sampson’s snakeroot (walter) isely (=psoralea psoraloides (walt.) cory) * orbexilum simplex (nutt. ex torr. & a. gray) rydb. singlestem scurf pea (=psoralea simplex (nutt. ex torr. & a. gray) rydb.) pediomelum digitatum (nutt. ex torr. & a. gray) isely palm-leaved scurf pea (=psoralea digitata nutt. ex torr. & a. gray) * pediomelum hypogaeum (nutt. ex torr. & a. gray) rydb. sara scurf pea var. subulatum (bush) j. grimes (=psoralea subulata bush) rhynchosia latifolia nutt. ex torr. & a. gray broadleaf snoutbean strophostyles helvola (l.) elliott wild bean stylosanthes biflora (l.) britton, sterns & poggenb. pencil-flower ~ tephrosia virginiana (l.) pers. goat’s rue ~ trifolium campestre schreb. low hop clover vicia sativa l. common vetch fagaceae – beech family quercus falcata michx. southern red oak quercus falcata michx southern red oak (=q. falcata michx. var. triloba (michx.) nutt) * quercus incana bartram bluejack oak ~ quercus nigra l. water oak * quercus phellos l. willow oak quercus stellata wangenh. post oak quercus velutina lam. black oak fumariaceae – fumitory family corydalis micrantha (engelm. ex a. gray) a. gray slender fumewort geraniaceae – geranium family geranium carolinianum l. carolina cranesbill oklahoma native plant record volume 11, december 2011 clark, l. g. 18 hydrophyllaceae – waterleaf family * hydrolea ovata nutt. ex choisy hairy hydrolea ~ phacelia strictiflora (engelm. & a. gray) a. gray prairie blue curls var. robbinsii constance juglandaceae – walnut family carya myristiciformis (michx. f.) nutt. nutmeg hickory carya texana buckley black hickory lamiaceae – mint family * lycopus virginicus l. virginia bugleweed ~ monarda punctata l. horsemint monarda russeliana nutt. ex sims. red spotted horsemint ~ prunella vulgaris l. heal-all ~ pycnanthemum albescens torr. & a. gray whiteleaf mountainmint * pycnanthemum tenuifolium schrad. narrowleaf mountainmint ~ salvia lyrata l. lyreleaf age scutellaria elliptica muhl. ex spreng. hairy skullcap * scutellaria laterifolia l. sideflowering skullcap scutellaria parvula michx. var. missouriensis (torr.) skullcap goodman & c.a. lawson (=s. parvula michx. var. leonardii (epling) fernald) lauraceae – laurel family sassafras albidum (nutt.) nees sassafras (=s. albidium (nutt.) nees var. molle (raf.) fernald lentibulariaceae – bladderwort family utricularia gibba l. (=u. biflora lam.) twoflower bladderwort lythraceae – loosestrife family rotala ramosior (l.) koehne toothcup melastomaceae – melastome family rhexia mariana l. meadow beauty menyanthaceae – buckbean famiy nymphoides peltata (s.g. gmel.) kuntze yellow floating heart monotropaceae – indian pipe family ~ monotropa hypopithys l. pinesap nymphaeaceae – water lily family nuphar lutea (l.) sm. yellow pond lily nymphaea odorata aiton american water lily oklahoma native plant record volume 11, december 2011 clark, l.g. 19 onagraceae – evening primrose family ludwigia alternifolia l. bushy seedbox oenothera laciniata hill cutleaf evening primrose oxalidaceae – wood sorrel family oxalis stricta l. yellow wood sorrel oxalis violaceae l. . violet wood sorrel plantaginaceae – plantain family plantago lanceolata l. buckhorn plantain plantago patagonica jacq. wooly plantain plantago virginica l. paleseed plantain plantago wrightiana decne. wright’s plantain polemoniaceae – phlox family phlox glaberrima l. smooth phlox phlox pilosa l. prairie phlox polygalaceae – milkwort family * polygala sanguinea l. blood polygala polygonaceae – buckwheat family ~ eriogonum longifolium nutt. longleaf eriogonum * eriogonum multiflorum benth. heartsepal wild buckwheat ~ polygonella americana (fisch. & c.a. mey.) small southern jointweed ~ polygonum hydropiperoides michx. mild water pepper polygonum persicaria l. lady’s thumb * polygonum sagittatum l. arrowvine ~ rumex hastatulus baldw. heartwing sorrel primulaceae – primrose family ~ hottonia inflata elliott american featherfoil * lysimachia lanceolata walter lanceleaf loosestrife ranunculaceae – buttercup family delphinium carolinianum walter prairie larkspur ranunculus laxicaulis (torr. & a. gray) darby spearwort rhamnaceae – buckthorn family berchemia scandens (hill.) k. koch rattan vine ceanothus americanus l. new jersey tea frangula caroliniana (walter) a. gray buckthorn (=rhamnus caroliniana walter) rosaceae – rose family crataegus spathulata michx. littlehip hawthorn potentilla simplex michx. old-field cinquefoil oklahoma native plant record volume 11, december 2011 clark, l. g. 20 prunus serotina ehrh. black cherry rubus occidentalis l. blackberry rubus ostryifolius rydb. highbush blackberry rubiaceae – madder family cephalanthus occidentalis l. buttonbush diodia teres walter rough buttonweed ~ galium arkansanum a. gray arkansas bedstraw galium circaezans michx. woods bedstraw ~ galium obtusum bigelow bluntleaf bedstraw galium pilosum aiton hairy bedstraw houstonia micrantha (shinners) terrell bluet (=hedyotis australis w.h. lewis & d.m. moore) sapotaceae – sapodilla family sideroxylon lanuginosum michx. ssp. lanuginosum chittamwood (=bumelia lanuginosa (michx.) pers.) scrophulariaceae – figwort family * agalinis tenuifolia (vahl.) raf. slenderleaf agalinus * castilleja coccinea (l.) spreng. indian paintbrush castilleja indivisa engelm. indian paintbrush collinsia violacea nutt. violet collinsia gratiola virginiana l. virginia hedgehyssop nuttallanthus canadensis (l.) d.a. sutton blue toadflax (=linaria canadensis (l.) chaz.) lindernia dubia (l.) pennell yellowseed false pimpernell pedicularis canadensis l. ssp. canadensis common lousewort (=p. canadensis l. var. dobbsii fernald) * penstemon laxiflorus pennell loose flower penstemon penstemon murrayanus hook. cupleaf penstemon solanaceae – potato family physalis heterophylla nees. clammy ground cherry solanum carolinense l. carolina horsenettle ulmaceae – elm family celtis tenuifolia nutt. dwarf hackberry urticaceae – nettle family ~ boehmeria cylindrica (l.) sw. false nettle valerianaceae – valerian family ~ valerianella radiata (l.) dufr. common beaked cornsalad verbenaceae – verbena family callicarpa americana l. american beautyberry oklahoma native plant record volume 11, december 2011 clark, l.g. 21 phryma leptostachya l. lopseed * verbena simplex lehm. narrow-leaved verbena violaceae – violet family viola villosa walter wooly violet vitaceae – grape family ampelopsis arborea (l.) koehne peppervine parthenocissus quinquefolia (l.) planch. virginia creeper vitis aestivalis michx. pigeon grape vitis rotundifolia michx. muscadine survey of the vascular flora of the boehler seeps and sandhills preserve, ph.d. dissertation by dr. linda gattie clark oklahoma native plant record, volume 2, number 1, december 2002 oklahoma native plant record, volume 14, number 1, december 2014 80 oklahoma native plant record volume 14, december 2014 gloria m. caddell https://doi.org/10.22488/okstate.17.100107 critic’s choice essay pollination ecology of our native prairie plants gloria m. caddell department of biology university of central oklahoma the oklahoma prairie in the summer is an ideal place and time to study pollination ecology. with its "cornucopia" pattern of flowering, where many plants flower synchronously, it has many flowers available every day. this past summer at the oklahoma department of wildlife conservation's arcadia conservation education area, dr. rebecca pace, an entomologist, and i taught a course in pollination ecology for the university of central oklahoma. i was glad to once again slow down and really pay attention to our native plants. the goals for each student were to choose an insect-pollinated species and determine its flowering phenology, i.e. the timing of the life cycle, its mating system, attractants, and pollinators; to gain an understanding of diverse pollination strategies; and to learn how synchronously-flowering plants within a community compete for and share pollinators. students often study members of the sunflower family (compositae) because they are so common here. although composites are intimidating because of their tiny flowers that are difficult to manipulate, the students quickly come to appreciate them as they see the diversity of pollinators they attract as well as the intricate details of their phenology. some students, especially those studying winecup (callirhoe involucrata) and trailing ratany (krameria lanceolata), dealt with high levels of herbivory or florivory. although it is frustrating to find buds with holes and extensive damage by insect larvae, this is an important phenomenon that affects fruit and seed set in natural populations and that can have long-term effects on the distribution of plant species. how are such pollination ecology studies conducted? the students first become familiar with their flowers — the numbers and degree of fusion of parts, their symmetry, and whether or not the flowers are aggregated into inflorescences. all these traits influence the orientation and behavior of insect visitors, the placement of pollen on an insect's body, and the subsequent deposition of pollen on stigmas. viewing the petals under high magnification allowed students to determine the type(s) of color-producing pigments. if the cells appear to be filled with colored "water balloons", the pigments are watersoluble and are in the cell's large vacuole. if the color is scattered in "dots" within the cells, the pigments are water-insoluble and are located in tiny cellular structures called plastids. by recording observations each day in the field, students determined their species’ phenological events. they described the sequence in which flowers open throughout the life of their plant or inflorescence and described all flower stages from tight buds to withering. the flowers of some species opened early in the morning, but students studying the lazy daisy (aphanostephus skirrhobasis), sleepy daisy (xanthisma texanum), and passion flower (passiflora incarnata) had to patiently wait for them to "wake up" by midday. by opening at different times of day, flowering species of a community can share pollinators. oklahoma native plant record 81 volume 14, december 2014 gloria m. caddell at close inspection, the differences among flowers become apparent, including size and color of the various parts, and position of parts relative to one another. the position of the anthers and stigmas is of crucial importance, as well as how the anthers release their pollen; different species might share pollinators by placing pollen on different parts of a pollinator's body, so that pollen of each species is transferred to a stigma of a flower of the same species. within a single flower, the anthers sometimes release pollen before the stigma is receptive to it, or vice versa. this difference in timing of the male and female parts of a flower reduces self-pollination. nectar production is often associated with the peak activity time of pollinators, but can be highly variable. tiny capillary tubes can be inserted into nectaries at various stages and times of day to draw out any available nectar. nectaries are often hidden, located within the flowers, or they may be extra-floral. for example, those of the passion flower (passiflora incarnata) are on the leaf stalk where they attract ants that defend it against herbivores. flowers can signal insects that they have pollen and nectar rewards. for example, prairie gaillardia has bright yellow styles and stigmas that contrast with the maroon disk flower petals when rewards for insects are available. as the flowers get older, the styles and stigmas turn maroon. older flowers might help attract pollinators to the inflorescence, but pollinators will visit younger more-rewarding flowers once they land. the flowers of most composites open from the periphery to the center of the inflorescence, so there are often concentric rings of flowers in various stages. students could determine whether their flowers self-pollinated, self-fertilized, or even produced seeds without sex! pollen-producing stamens were removed from some flowers; then, the flower was bagged and later checked to see if seeds were produced. some flowers were pollinated by hand with pollen from another flower on the same plant, while others were cross-pollinated with pollen from different plants. students added pollen to flowers left open to determine whether or not it increased fruit and seed set and to determine if pollinators are sufficient. from dawn to dusk, students recorded insect visitors to their species. to determine whether insects were just "visitors" or effective pollinators, they gathered pollen from flowers, viewed it under a scanning electron microscope, and compared it with the pollen loads on insect visitors to the same plant. this allowed them to determine whether the visitors were able to carry pollen, and whether they had visited flowers of a single species or several species at the same time. bees are generally the most efficient insect pollinators; they are able to carry large amounts of pollen, can learn to tell differences among flowers, can learn to "handle" them, and they show floral constancy by revisiting flowers of the same species. if you would like to delve into and be amazed at what is currently known about pollination biology across the world, i suggest the comprehensive and up-to-date (2011) book pollination and floral ecology by pat willmer, published by the princeton university press. 82 oklahoma native plant record volume 14, december 2014 gloria m. caddell halictid bee visiting passion flower (passiflora incarnata). note the extra-floral nectaries on the leaf stalk. lanceleaf gaillardia (gaillardia aestivalis). note ring of styles emerging from newly-opened flowers. pollinators visit newly-opened flowers of gaillardia aestivalis. bumblebee on dalea candida. all photos by gloria caddell. bacf critic’s choice essay: pollination ecology of our native prairie plants by dr. gloria m. caddell oklahoma native plant record 3 volume 16, december 2016 foreword the effects of climate change on the phenology and pollination of plants is expected to be a major factor in the problem of maintaining biodiversity as well as that of feeding a growing global population. though ecologists and agricultural researchers have historically tried to stay out of each other’s domains, climate change has dealt them a globe of shared problems that will not easily be teased apart. for example, the honey bees that agriculture depends upon to pollinate crops are not native to north america. whether those introduced pollinators help or hinder native species is a complicated problem, but they are certainly a part of the environment that researchers are going to have to study more closely as the world’s population grows. many native species and human-produced cultivars have co-evolved to depend upon or compete with each other in ways that we haven’t yet discovered. we are reaching back in time to pull out some phenology and pollination data that can be compared with current data to study changes in species genomes or gene activity that might be related to climate change. connie taylor wrote “pollination ecology of sabatia campestris” in 1972 based on data she collected while taking a summer course at the university of oklahoma biological station at lake texoma. written as a student research paper, its significance lies not in numerical data, but in her descriptions of pollination processes she observed in the field, which differed from those processes described from research done in green house environments. shang-wen liaw was a graduate student at the university of central oklahoma who studied under gloria caddell. his 1999 master’s thesis went unpublished as he took advantage of an opportunity to go directly into a ph.d. program. we are proud to publish “the structure of the gynostegium, breeding system, and pollination ecology of spider milkweed, asclepias viridis.” if that requires more phenology and pollination terminology than you know, you can flip to the critic’s choice essay in the back where we’ve reprinted paul buck’s botany bay article from the fall 2000 gaillardia, “a conversation with a small beetle.” his explanation of pollination from the standpoint of the pollinator is both entertaining and educating. you can take a break from pollination studies and read amy buthod’s floristic inventory of kessler atmospheric and ecological field station. this site holds great potential for future climate change comparisons using sophisticated environmental monitoring equipment that will enable a coupling of species inventories with climate change. we also have an assessment of a five-year recovery from a burn at wichita mountains wildlife refuge by oklahoma city university authors, laura jardine, adam ryburn, and anthony stancampiano. this is a great piece of local ecological research that can play an important role in predicting dynamics due to fires which may become more frequent due to climate change. again this year, we have something for everyone. if you do research in or about oklahoma’s native plant environments, please consider submitting your own manuscript next year. we want manuscripts based on the newest concepts in research as much as we want manuscripts based on historical data. we want manuscripts written by authors with years of experience, but our editorial staff is also ready to help first-time authors get the experience they need to develop science writing skills. the oklahoma native plant record is a professionally reviewed publication. our abstracts are indexed in the “centre for agricultural bioscience international” that is based in the u. k., and the record is listed in the “directory of open access journals” https://doaj.org. sheila strawn, managing editor https://doaj.org/ oklahoma native plant record, volume 14, number 1, december 2014 4 oklahoma native plant record volume 14, december 2014 lottie opal baldock https://doi.org/10.22488/okstate.17.100102 flora of kiowa county, oklahoma master’s thesis oklahoma agricultural and mechanical college [oklahoma state university] 1938 lottie opal baldock keywords: distribution, ecology, historic, vascular [abstract] this paper presents the results of taxonomic and ecological studies of the plants of kiowa county, oklahoma. the collections were begun in 1933 and continued until the summer of 1938; however, little intensive collecting was done until the spring and summer of 1938. the flora of kiowa county, oklahoma includes six species of pteridophytes, one species of gymnosperms, and 489 species of angiosperms. more than one third of these are in compositae, gramineae, and leguminosae. there are 81 families represented. the 11 largest families, with the number of species are compositae, 86; gramineae, 58; leguminosae, 41; onagraceae, 17; euphorbiaceae, 16; cruciferae, 16; polygonaceae, and solanaceae, 12 each; asclepiadaceae, cyperaceae, and labiatae, 11 each. the three largest families comprise 37.4 per cent of the total number of species. [species names used in the original thesis which appear in brackets have been updated using the usda plants database.] preface the main value of studies such as this is to establish the distribution of species and to observe the varying ecological conditions in which the plants under consideration are growing. plants considered in this study and included in the list are native wild species and mainly indigenous to the county; however, a few species of cultivated plants are listed in cases where they have escaped cultivation and seem to have established themselves in the new habitat. the author does not aim to give a complete list of the vascular plants of the county as the time was limited, and such a survey is a fit subject for more advanced graduate work. introduction this paper presents the results of taxonomic and ecological studies of the plants of kiowa county, oklahoma (figure). the collections were begun in 1933 and continued until the summer of 1938; however, little intensive collecting was done until the spring and summer of 1938. more systematic work has been done with the spring and summer flowering plants than with those flowering in the fall. in most cases the nomenclature is that of gray’s manual (robinson and fernald 1908); however, the oklahoma flora by stemen and meyers (1937) was used as a check, and in some cases plants were listed in the latter publication only. for the grasses, hitchcock (1935) was the final authority. oklahoma native plant record 5 volume 14, december 2014 lottie opal baldock location and size kiowa county is in southwestern oklahoma. it comprises an area of 1,025 square miles, or 656,000 acres [2,655 km2]. the county is bounded by washita on the north, greer and jackson on the west, tillman on the south, and comanche and caddo counties on the east. hobart, the county seat and largest town, is located in the northwestern part, about 136 miles [219 km] southwest of oklahoma city. the elevation at hobart is 1,536 feet [468 m] (wahlgren). history in 1834, a large military expedition was sent out from forts gibson and towson to stop the warfare among the indians and to pay a visit to the wild bands of kiowas, wichitas, and comanches who lived among the wichita mountains. this was probably the first official expedition to reach any part of kiowa county. they explored the region about the wichita mountains going as far west as north fork red river. george catlin, the famous painter of indian pictures, was with this expedition and left many pictures of things he observed. the county was opened to settlement by a proclamation of president mckinley on july 4, 1901. the land was allotted by drawing for a choice. this county was formerly a part of the kiowa, comanche, and apache indian reservation. in 1910, a part of the county was taken with a part of comanche county to form swanson county (goke and holopeter 1931), but the creation of swanson county was declared illegal by a decision of the supreme court of oklahoma, august 9, 1911, and the territory was restored to the former counties. a part was annexed to tillman county. the main industry is farming, with cotton and wheat as the leading crops. quarrying of granite is carried on to a small extent. original dominant vegetation in the county consisted of grasses, a scattered growth of mesquite (prosopis glandulosa torr. var. glandulosa [=prosopis juliflora glandulosa]), and cactus (opuntia humifusa (raf.) raf.). along the streams, cottonwood (populus deltoids w. bartram ex marshall), elm (ulmus americana l.), and pecan (carya illinoiensis (wagenh.) k. koch) were in predominance. during the winter, the grasses in the valley provide the chief source of feed for livestock. before the land was open for settlement, these valleys were highly prized among the indians for grazing purposes. topography the wichita mountains in the south central and eastern portion rise abruptly above the gently rolling plains which are characteristic of the prairies. the mountains are composed of igneous rocks surrounded by sedimentary formations. the igneous rocks are pre-cambrian, but younger than the proterozoic rocks which they have intruded. most of the rock is medium to fine grained pink granite, except those of the northern range in the eastern part of the county which are made up of limestone. the granite mountains are covered with a scrubby growth of oaks, but the limestone hills are comparatively barren. the southward facing escarpment which crosses the northern part of the county shows a distinctly different physical feature. this escarpment is composed of calcareous ledges of the blaine formation (sawyer 1929). streams and drainage north fork red river, bounding the county on the west, and washita river, along the northeastern edge, are the two largest streams. most of the drainage waters flow through them from their several 6 oklahoma native plant record volume 14, december 2014 lottie opal baldock tributaries. east fork of deep red creek and its tributaries with east, west, and middle otter creeks drain the lower southern section. north fork red river with elk creek and their tributaries drain the western section. the northeastern section of the county is drained by washita river and rainy mountain, saddle mountain, and stinking creeks. soil the soils of 95 per cent of the area of kiowa county are heavy in texture either in the surface soil or subsoil, with clay loam mainly in both; the rest, which occur in irregularly shaped areas in different parts of the county, are sandy. the sandy soils are found along the two rivers mentioned, at the western boundary and the northeastern corner. the outstanding difference between the sandy soils and the clay loam soils is that the material of the sandy areas is much more friable throughout the surface soil and subsoil, continuing to a depth of several inches. foard silt loam comprises about 128,896 acres [522 km2] (goke and holopeter 1931) or 19.6 per cent of the total. this type has a dark-brown surface soil that extends to a depth of six inches, where it gradually passes downward to a dark-brown or brown heavy plastic subsoil. the color in this layer gradually changes to a yellowish-brown. at a depth of 18 inches [46 cm], lime is present in sufficient quantitites to effervesce in acid. tillman clay loam is next in importance with 112,064 acres [454 km2] or 17.1 per cent. it has a chocolate-brown friable surface soil that passes at a depth of six inches [15 cm] into a dark-brown friable subsurface soil. at about 12 inches [30 cm], this changes to a chocolate-brown or reddish-brown clay subsoil which is tough and plastic when wet and very hard and dense when dry. at a depth of about 24 inches [60 cm], lime is first reached in the form of hard concretions or in a finely disseminated form. the third important type of soil is vernon clay loam which covers 65,536 acres [265 km2] or 10.0 per cent of the land. the surface soil of vernon clay loam consists of reddish-brown, brown, or chocolate-brown friable material to a depth of four inches. the subsoil is reddish-brown granular clay loam which continues to a depth of about 12 inches [30 cm] where it changes into a reddish-brown clay which is plastic when wet but very hard when dry. this soil is found chiefly on slopes along the valleys and in areas that are cut by many drainage channels. both tillman and vernon clay loams are best suited for pasture (goke and holopeter 1931), as shown by the severely eroded areas over the county where these soils have not been cultivated carefully. in many places, erosion is quite severe although the land has been cultivated little more than 30 years. foard silt loam belongs to the better productive group of soils and is more suited to cultivation. climate the average yearly rainfall for hobart from 1903 to 1930 was 28.13 inches [71 cm]. the months april, may, and june received the most rainfall while december, january, and february proved to be the driest months for those years. the lowest average rainfall came in the year 1910, which was 12.72 inches [32 cm]. the other extreme was 43.33 inches [110 cm] for 1908 (wahlgren); however, the year 1938 proved a record one for moisture. from january to may, the average precipitation was from an inch to an inch and one-half [2.5-3.75 cm] above the average for each month. prevailing winds are from the south in all months except december when they are from the north. the lowest temperature recorded over a period of 28 years is -11ºf [-23.9ºc]; the highest is 114ºf [45.6ºc], with an average minimum temperature of 48ºf oklahoma native plant record 7 volume 14, december 2014 lottie opal baldock [8.9ºc] and an average maximum of 74.6ºf [23.7ºc] for the period. the average date of the last killing frost comes on november 2. there is an average growing season of 213 days. previous collectors dr. g. g. shumard (bull 1932; eskew 1937) was perhaps the first person to make a collection of plants in this vicinity. he was with captain r. b. marcy on his expedition of 1852 to the source of the north fork of the red river. the expedition entered the state near the center of the southern border and passed through the wichita mountains and into the panhandle of texas. about 100 plants were collected within the present boundaries of oklahoma. probably the largest single collection was made by the late dr. g. w. stevens in 1913 while he was preparing his flora of oklahoma. this complete collection is now in the gray herbarium at harvard. oklahoma agricultural and mechanical college has more specimens from the stevens collection than any other herbarium within the state. professor robert stratton of oklahoma agricultural and mechanical college has collected in the vicinity to add to his personal herbarium of leguminosae and for the college herbarium. in 1932, miss rotha bull made a collection of the plants of greer county which is separated from kiowa county on the west by north fork red river. mr. c. t. eskew made a collection of plants in 1937 of the wichita national forest within the boundaries of comanche county which adjoins kiowa county on the east. ecology the most common pre-vernal plants are claytonia virginica l., glandularia canadensis (l.) nutt. [=verbena canadensis], viola bicolor pursh [=viola rafinesquii], v. sororia willd. [=v. papilionacea], erysimum asperum (nutt.) dc., anemone caroliniana walter, a. berlandieri pritz. [=a. decapetala], lithospermum incisum lehm. [=lithospermum angustifolium], cercis canadensis l., glandularia bipinnatifida (nutt.) nutt. [=verbena bipinnatifida], allium canadense l. var. mobilense (regel) ownbey [=allium mutabile], a. drummondii regel [=a. nuttallii], nothascordum bivalve (l.) britton, and quincula lobata (torr.) raf. [=physalis lobata]. all of these were to be found on the streams and ravines. the prairie communities have fewer flowering plants; the outstanding ones are glandularia bipinnatifida, allium drummondii, northascordum bivalve, quincula lobata, anemone, and lepidium [=lepidium apetalum]. liliaceae and violaceae have more representatives at this time than other families. among the spring plants are tradescantia ohiensis raf. [=tradescantia reflexa], t. occidentalis (britton) smyth, baptisia bracteata muhl. ex elliott, b. australis (l.) r. br., corydalis, oxalis, and a great percent of cruciferae, all of which appear on mountains and streams with the budding trees and other woody plants. cruciferae are in more abundance on the prairies along with sphaeralcea coccinea (nutt.) rydb. [=malvastrum coccineum], opuntia humifusa, oenothera laciniata hill, hordeum pusillium nutt., bromus catharticus vahl. [=bromus unioloides ], vulpia octoflora (walter) rydb. [=festuca octoflora], aristida purpurea nutt., yucca glauca nutt., and oxalis. grasses begin flowering in late spring and early summer when they become predominant. leguminosae is another family which flowers mainly in the summer. other plants which become predominant at this time are argemone albiflora hornem. [=argemone alba], a. polyanthemos (fedde) g.b. ownbey [=a. intermedia], cirsium [=cirsium discolor], centaurea americana nutt., gaillardia, tribulus terrestris l., plantago patagonica jacq. [=plantago purshii], krameria lanceolata torr. [=krameria secundiflora], erigeron strigosus muhl. ex willd. 8 oklahoma native plant record volume 14, december 2014 lottie opal baldock [=erigeron ramosus], chloris verticillata nutt., solanum elaeagnifolium cav., s. rostratum dunal, polygonum, salsola tragus l. [=salsola kali], and the greatest percent of euphorbiaceae. during the latter part of the summer the composites begin to gain predominance as for number of species in flower, but the grass family is still the most important as to the amount of space it covers. during the autumn the outstanding plants are helianthus, rhus, vernonia, euphorbia marginata pursh, liatris punctata hook., solidago, aster, ambrosia, xanthium, sorghastrum nutans (l.) nash, and others of the tall grass group. annual and biennial plants on the mountains and streams are not so different from those of the prairies in the summer and fall as they are in the spring. the fall grasses are more adapted to the former habitat. many woody plants are seeding in the autumn, and in the latter part of the year the mountainsides are colorful with the brilliant foliage of the trees and shrubs. along the streams, trees and woody plants are dominant. carya illinoinensis, juglans nigra l., rhus glabra l., ulmus americana, vitis, fraxinus, toxicodendron, sapindus saponaria l. var drummondii (hook. & arn.) l.d. benson [=sapindus drummondii], and salix nigra marshall are the outstanding plants in this type of vegetation. these same genera are found on the mountains, but species of quercus become dominant in the eastern section of the county. other mountain plants are rhus aromatica aiton [=rhus trilobata], ptelea trifoliata l., ribes aureum pursh, baptisia, sedum, ceanothus americanus l., rubus, galium aparine l., and poa arachnifera torr. plants in dry sand and along the rivers form another distinctly different type. artemisia, mentzelia, sporobolus, and species of prunus form the dominant covering on the sand hills, and tamarix [=tamarix gallica] is found in abundance in damp sandy soil. other outstanding plants here are glandularia canadensis, comandra umbellata (l.) nutt. ssp. pallida (a. dc.) piehl [=comandra pallida], lithaspermum incisum, cenchrus, and a species of gaura. the types of vegetation mentioned above are all connected by the prairie type which covers the greatest percent of the area of the county. here is found one main association in the undisturbed pastures. prosopis glandulosa var. glandulosa forms an orchard type of growth, and under the trees the dominant vegetation is bouteloua dactyloides (nutt.) j.t. columbus [=buchloe dactyloides] interspersed with opuntia humifusa. summary the flora of kiowa county, oklahoma includes six species of pteridophytes, one species of gymnosperms, and 489 species of angiosperms. more than one third of these are in compositae, gramineae, and leguminosae. there are 81 families represented. the 11 largest families, with the number of species, are compositae, 86; gramineae, 58; leguminosae, 41; onagraceae, 17; euphorbiaceae, 16; cruciferae, 16; polygonaceae and solanaceae, 12 each; and asclepiadaceae, cyperaceae, and labiatae, 11 each. the three largest families comprise 37.4 per cent of the total number of species. the county lies in the plains region. the prairies are broken by the wichita mountains and a few streams, of these the north fork red river is the largest. the vegetation is mainly that adapted to the prairies. tall grass is found near mountains or streams; it is predominately a short-grass area. trees are to be seen along streams or on mountainsides. the only native trees on the prairies are mesquite (prosopis glandulosa var. glandulosa) which grow in association with cactus (opuntia humifusa) and buffalo grass (bouteloua dactyloides). oklahoma native plant record 9 volume 14, december 2014 lottie opal baldock acknowledgements the writer wishes to express her appreciation to the following people for their services in preparing this paper: dr. h. i. featherly of the oklahoma agricultural and mechnical college, under whose direction this study was made, for constant advice and criticism; dr. k. starr chester, head of the department of botany, and professor r. h. stratton, of the same department, for their aid in securing materials; and dr. elbert l. little, jr., in the united states forest service at flagstaff, arizona, for suggestions. the writer also wishes to express her appreciation to her family for assistance in collecting and preparing specimens. bibliography britton, n.l. and a. brown. 1913. an illustrated flora of the northern united states, canada, and the british possessions. 2nd ed. 3 volumes. new york: [charles scribner and sons]. bull, r.z. 1932. vascular plants of greer county, oklahoma [master’s thesis]. norman (ok): university of oklahoma. eskew, c.t. 1937. flowering plants of the wichita national forest [master’s thesis]. norman (ok): university of oklahoma. featherly, h.i. 1938. grasses of oklahoma. oklahoma agricultural experiment station technical bulletin no. 3. stillwater (ok): oklahoma agricultural and mechanical college. featherly, h.i. and e.e. still. 1934. the ferns of oklahoma. botanical studies no. 1. experiment station circular no. 80. stillwater (ok): oklahoma agricultural and mechanical college. goke, a.w. and c.a. holopeter. 1931. soil survey of kiowa county, oklahoma. united states department of agriculture bulletin no. 14. hitchock, a.s. 1935. manual of the grasses of the united states. misc. pub. no. 200. washington (dc): government printing office. jeffs, r.e. and e.l. little, jr. a preliminary list of the ferns and seed plants of oklahoma. university of oklahoma biological survey, vol. 11, no. 2. little, e.l., jr. flora of muskogee county, oklahoma. 1938. the american midland naturalist 19:369-389. mattoon, w.r. and g.g. phillips. 1936. forest trees of oklahoma. oklahoma forest commission publication no. 1. reprint no. 4. united states department of agriculture. robinson, b.l. and m.l. fernald. 1908. gray’s new manual of botany. 7th ed. new york: american book company. rydberg, p.a. 1932. flora of the prairies and plains of central north america. new york: [new york botanical garden]. sawyer, r.w. 1929. kiowa and washita counties, oklahoma. geological survey bulletin 40hh. small, j.k. 1913. flora of the southeastern united states. 2nd ed. new york: [published by author]. stemen, t.r. and w.s. meyers. 1937. oklahoma flora. oklahoma city: harlow. wahlgren, h.f. [date unknown]. climatological data. united states department of agriculture. weather bureau. oklahoma city. 10 oklahoma native plant record volume 14, december 2014 lottie opal baldock figure map of kiowa county, oklahoma oklahoma native plant record 11 volume 14, december 2014 lottie opal baldock appendix a list of species, kiowa county, ok [nomenclature has been updated using the plants database (http://plants.usda.gov/plants).] pteridophyta dryopteridaceae [polypodiaceae] woodsia obtusa (spreng.) torr. blunt-lobed woodsia mountainsides marsileaceae marsilea vestita hook. & grev. water fern, hairy pepperwort low places, pastures pteridaceae [polypodiaceae] cheilanthes eatonii baker eaton’s lip fern mountainsides cheilanthes lanosa (michx.) d.c. eaton hairy lip fern mountainsides [=cheilanthes lanulosa (michx.) watt] notholaena standleyi maxon standley’s notholaena mountainsides pellaea atropurpurea (l.) link purple cliff brake mountainsides spermatophyta gymnosperms cupressaceae [pinaceae] juniperus virginiana l. red cedar hillsides angiosperms acanthaceae ruellia pedunculata torr. ex a. gray stalked ruellia mountainsides; summer, fall ruellia sp. hairy ruelllia rivers; summer, fall [=ruellia ciliosa pursh, misapplied] agavaceae [liliaceae] yucca glauca nutt. yucca, bear-grass, soap weed pastures, roadsides; spring aizoaceae mollugo verticillata l. carpet-weed pastures, common; summer amaranthaceae amaranthus blitoides s. watson prostrate amaranth fields, pastures, common; summer 12 oklahoma native plant record volume 14, december 2014 lottie opal baldock amaranthus hybridus l. dark green pig-weed pastures; spring to fall amaranthus retroflexus l. red root roadsides; spring, summer amaranthus tuberculatus (moq.) sauer western water-hemp pastures; summer anacardiaceae rhus aromatica aiton fragrant sumac, sumac mountains, streams; [=rhus canadense mill., rhus trilobata nutt.] spring, summer rhus glabra l. smooth upland sumac creeks, hillsides; common; summer toxicodendron radicans (l.) kuntze poison ivy mountainsides, streams; summer apocynaceae amsonia tabernaemontana walter broad-leaved amsonia mountain ravines; spring apocynum cannabinum l. dogbane, indian hemp roadsides; common; summer asclepiadaceae asclepias amplexicaulis sm. milkweed, silkweed near rivers; spring, summer asclepias asperula (decne.) woodson ssp. milkweed mountainsides; capricornu (woodson) woodson spring [=asclepiodora decumbens (nutt.) a. gray] asclepias engelmanniana woodson green milkweed roadsides; summer [=acerates auriculata engelm. ex torr.] asclepias latifolia (torr.) raf. broad-leaved milkweed dry sandy soils; summer asclepias stenophylla a. gray narrow-leaved milkweed prairies; summer [incl. acerates angustifolia (nutt.) decne.] asclepias tuberosa l. butterfly weed sand, near rivers; spring, summer asclepias verticillata l. whorled milkweed mountainsides; spring, summer asclepias viridiflora raf. green milkweed prairies; summer [=acerates viridiflora (raf.) pursh ex eaton] asclepias viridis walter oblong-leaved milkweed prairies; summer [=asclepiodora viridis (walter) a. gray] gonolobus suberosus (l.) r. br. large-leaved angle-pod rivers; early summer [=vincetoxicum gonocarpos walter] boraginaceae heliotropium convolvulaceum (nutt.) a. gray sand heliotrope rivers; late summer heliotropium indicum l. indian heliotrope pastures, creeks; spring to fall oklahoma native plant record 13 volume 14, december 2014 lottie opal baldock lappula occidentalis (s. watson) western stick-weed mountainsides; greene [=lappula redowskii (hornem.) spring greene var. occidentalis (s. watson) rydb.] lithospermum incisum lehm. puccoon dry sandy soils; [=lithospermum angustifolium michx.] spring myosotis verna nutt. spring or early scorpion-grass sand near rivers; [=myosotis virginica (l.) britton, sterns & spring poggenb., misapplied] onosmodium bejariense dc. ex a. dc. western false gromwell prairies; summer [=onosmodium occidentale mack.] cactaceae echinocereus reichenbachii (terscheck ex lace cactus mountainsides walp.) j.n. haage [=echinocereus caespitosus (englem.) engelm. opuntia humifusa (raf.) raf. western prickly-pear pastures; common; spring campanulaceae [incl. lobeliaceae] lobelia cardinalis l. cardinal flower, red lobelia mountain ravines; summer lobelia spicata lam. var. leptostachys spiked lobelia mountainsides; (a. dc.) mack. & bush summer [=lobelia leptostachys a. dc.] triodanis leptocarpa (nutt.) nieuwl. western venus’s looking-glass pastures; spring, [=specularia leptocarpa (nutt.) a. gray] summer triodanis perfoliata (l.) nieuwl. venus’s looking-glass prairies, streams; [=specularia perfoliata (l.) a. dc.] spring, summer capparaceae cleome serrulata pursh pink cleome, stinking clover prairies; summer cleomella angustifolia torr. caprifoliaceae symphoricarpos orbiculatus moench coral-berry, indian currant streams; summer viburnum rufidulum raf. southern black-haw mountainsides; spring caryophyllaceae [incl. illecebraceae] cerastium brachypodum (engelm. ex a. gray) short-stalked chickweed prairies; spring b.l. rob. cerastium nutans raf. long-stalked chickweed mountainsides; spring paronychia jamesii torr. & a. gray james’s whitlow-wort prairies; summer silene antirrhina l. sleepy catchfly roadsides; spring stellaria media (l.) vill. common chickweed, starwort low damp places; early spring 14 oklahoma native plant record volume 14, december 2014 lottie opal baldock chenopodiaceae chenopodium album l. lamb’s quarters roadsides, common; summer cycloloma atriplicifolium (spreng.) j.m. coult. sand tumbleweed rivers; summer monolepis nuttalliana (schult.) greene monolepis common near dwellings; spring salsola tragus l. russian thistle roadsides, cultivated [=salsola kali l., misapplied] soil; summer commelinaceae commelina erecta l. slender day-flower, mountainsides, [incl. commelina crispa woot.] crinkle-leaved day-flower creeks; spring, summer, fall commelina virginica l. virginia day-flower, mountain ravines, [incl. commelina hirtella vahl] bearded day-flower streams; summer tradescantia occidentalis (britton) smyth western spiderwort, trinity mountainsides; spring compositae achillea millefolium l. common yarrow, wooly roadsides, prairies, [incl. achillea lanulosa nutt.] common yarrow creeks; summer ambrosia artemisiifolia l. ragweed ravines; summer, fall ambrosia psilostachya dc. western ragweed roadsides, pastures; fall ambrosia trifida l. great ragsweed creeks; summer, fall amphiachyris dracunculoides (dc.) nutt. august flower kindling-weed roadsides; fall aphanostephus ramosissimus dc. sand-daisy hillsides; summer [=aphanostephus humilis (benth.) a. gray, misapplied] aphanostephus skirrhobasis (dc.) trel. white-flowered sand-daisy rivers; summer artemisia ludoviciana nutt. dark-leaved mugwort dry hillsides near rivers; late summer baccharis salicina torr. & a. gray willow baccharis rivers; early summer berlandiera betonicifolia (hook.) small texas berlandiera streams; summer [=berlandiera texana dc.] brickellia eupatorioides (l.) shinners false boneset prairies; summer, fall centaurea americana nutt. centaurea roadsides; summer chaetopappa asteroides nutt. ex dc. chaetopappa rivers; spring chaetopappa ericoides (torr.) g.l. nesom aster hillsides; summer [=aster ericaefolius rothr.] chrysopsis pilosa nutt. nuttall’s golden aster mountainsides; summer, fall chrysopsis sp. hispid golden aster rivers; summer [=chrysopsis villosa (pursh) nutt. ex dc. var. hispida (hook.) a. gray, misapplied] oklahoma native plant record 15 volume 14, december 2014 lottie opal baldock cirsium ochrocentrum a. gray yellow-spined thistle prairie pastures; summer cirsium sp. field thistle roadsides; common; [=cirsium discolor (muhl. ex willd.) spreng., summer misapplied] conyza canadensis (l.) cronquist horsetail, horse-weed pastures; summer, [=erigeron canadensis l.] fall coreopsis grandiflora hogg ex sweet large-flowered coreopsis rivers; summer coreopsis tinctoria nutt. golden coreopsis, streams; spring, garden tickseed summer coreopsis sp. whorled tickseed rivers; late summer [=coreopsis verticillata l., misapplied] dracopis amplexicaulis (vahl) cass. cone flower streams; late spring [=rudbeckia amplexicaulis vahl] echinacea angustifolia dc. narrow-leaved purple hillsides; spring to [=brauneria angustifolia (dc.) a. heller] cone-flower fall engelmannia peristenia (raf.) goodman & engelmannia hillsides; summer c.a. lawson [=engelmannia pinnatifida a. gray ex. nutt.] erigeron strigosus muhl. ex willd. daisy fleabane pastures; spring [=erigeron ramosus (walter) britton, sterns & poggenb.] euthamia gymnospermoides greene viscid bushy goldenrod prairies; late summer evax prolifera nutt. ex dc. rabbit tobacco roadsides, pastures; spring, summer evax verna raf. rabbit tobacco roadsides, pastures; [=evax multicaulis dc.] common; spring, early summer flaveria campestris j.r. johnst. plains flaveria dry plains; late summer gaillardia pulchella foug. showy gaillardia hillsides; summer, fall gaillardia suavis (a. gray & engelm.) britton cut-leaved rayless prairie roadsides; & rusby three-nerved gaillardia spring gaillardia sp. gaillardia roadsides; common; [=gaillardia aristata pursh, misapplied] summer grindelia papposa g.l. nesom & suh rosin-weed prairie roadsides; [=haplopappus ciliatus (nutt.) dc.] spring to fall grindelia squarrosa (pursh) dunal broad-leaved gum plant, rivers; summer, fall rosin-weed helenium amarum (raf.) h. rock var. amarum fine-leaved sneezeweed creeks; summer [=helenium tenuifolium nutt.] helenium amarum (raf.) h. rock var. badium fine-leaved sneezeweed streams; summer (a. gray ex s. watson) waterf. [=helenium tenuifolium nutt. var. badium a. gray ex s. watson] 16 oklahoma native plant record volume 14, december 2014 lottie opal baldock helenium autumnale l. false or swamp sunflower creeks; late summer helianthus annuus l. common sunflower prairies; common; summer helianthus hirsutus raf. stiff-haired sunflower mountainsides; summer helianthus maximiliani schrad. maximilian’s sunflower prairies; late summer helianthus petiolaris nutt. sunflower roadsides; common; summer heterotheca subaxillaris (lam.) britton & rusby heterotheca hillsides; summer hymenopappus scabiosaeus l’hér. var. corymbed, smooth roadside ditches; corymbosus (torr. & a. gray) b.l. turner hymenopappus summer [=hymenopappus corymbosus torr. & a. gray] hymenopappus scabiosaeus l’hér. var. carolina hymenopappus roadsides; spring scabiosaeus [=hymenopappus carolinensis (lam.) porter] hymenopappus tenuifolius pursh wooly white hymenopappus prairies; spring to fall iva annua l. rough marsh elder creeks; late summer [=iva ciliata willd.] lactuca ludoviciana (nutt.) riddell western lettuce prairies; common; summer to fall lactuca sp. prickly lettuce fields, pastures; late [=lactuca virosa l., misapplied] summer liatris punctata hook. dotted button snakeroot, prairies; late small blazing star summer liatris squarrosa (l.) michx. scaly blazing star roadsides; summer packera plattensis (nutt.) w.a. weber & prairie ragwort pastures, prairies; á. löve [=senecio plattensis nutt.] spring pyrrhopappus carolilnianus (walter) dc. leaf-stemmed false dandelion fields; infrequent; spring pyrrhopappus grandiflorus (nutt.) nutt. rough false dandelion roadsides; prairies; [=pyrrhopappus scaposus dc.] spring ratibida columnifera (nutt.) woot. & standl. lepachys, cone-flower common; late [=lepachys columnaris (pursh) torr. a. gray] spring, summer senecio riddellii torr. & a. gray riddell’s ragwort prairies; late summer silphium integrifolium michx. entire-leaved rosin-weed prairies; late summer silphiium laciniatum l. compass-plant prairies; common; summer solidago altissima l. tall goldenrod hillsides; late summer solidago arguta aiton var. boottii (hook.) boott’s goldenrod, mountains; summer, palmer & steyerm. wreath goldenrod fall [=solidago boottii hook.] oklahoma native plant record 17 volume 14, december 2014 lottie opal baldock solidago gigantea aiton late goldenrod creeks; late summer [=solidago serotina aiton] solidago missouriensis nutt. missouri goldenrod mountainsides; summer solidago petiolaris aiton downy ragged goldenrod fields; fall solidago radula nutt. western rough goldenrod prairies; summer sonchus asper (l.) hill spiny sow-thistle roadsides; spring to fall symphyotrichum divaricatum (nutt.) slim aster creeks; summer g.l. nesom [aster exilis elliott] symphyotrichum ericoides (l.) g.l. nesom many-flowered aster prairies; summer, [=aster multiflorus aiton] fall symphyotrichum falcatum (lindl.) g.l. aster prairies; spring nesom var. commutatum (torr. & a. gray) g.l. nesom [=aster commutatus (torr. & a. gray) a. gray] symphyotrichum fendleri (a. gray) g.l. fendler’s aster pastures; summer nesom [=aster fendleri a. gray] taraxacum officinale f.h. wigg. common dandelion fields, pastures; [=taraxacum vulgare lam.] spring to fall tetraneuris linearifolia (hook.) greene fine-leaved actinea hillsides; summer [=actinea linearifolia (hook.) kuntze] tetraneuris scaposa (dc.) greene narrow-leaved actinea rivers; summer [=actinea scaposa (dc.) kuntze var. linearis (nutt.) b.l. rob.] thelesperma filifolium (hook.) a. gray thelesperma, tickseed prairies, mountains; [=thelesperma trifidum (poir.) britton] common; summer to fall thelesperma megapotamicum (spreng.) rayless thelesperma prairies; summer kuntze [=thelesperma gracile (torr.) a. gray] vernonia baldwinii torr. baldwin’s ironweed rivers; summer vernonia gigantea (walter) trel. tall ironweed streams; summer [=vernonia altissima nutt.] vernonia missurica raf. missouri ironweed prairies; fall xanthisma texanum dc. texas xanthisma, sleepy daisy prairies; summer xanthium strumarium l. cocklebur, great cocklebur roadsides, ravines; [=xanthium pensylvanicum wallr., common; summer, xanthium speciosum kearney] fall convolvulaceae cuscuta cephalanthi engelm. button-bush dodder parasite; summer cuscuta cuspidata engelm. cuspidate dodder pastures; ragweeds; summer cuscuta indecora choisy pretty dodder on composites; summer evolvulus nuttallianus schult. dwarf morning-glory prairies; summer [evolvulus argenteus pursh] 18 oklahoma native plant record volume 14, december 2014 lottie opal baldock ipomoea leptophylla torr. bush morning-glory roadsides; early summer ipomoea pandurata (l.) g. mey. wild potato vine roadside ditches; summer cornaceae cornus drummondii c.a. mey. rough-leaved dogwood streams; spring [=cornus asperifolia michx., misapplied] cornus florida l. flowering dogwood mountainsides; spring crassulaceae sedum nuttallianum raf. nuttall’s stonecrop rocks on mountainsides; spring cruciferae capsella bursa-pastoris (l.) medik. shepherd’s purse fields, meadows; spring descurainia pinnata (walter) britton tansy-mustard hillsides; spring [=sisymbrium canescens nutt.] descurainia sp. western tansy-mustard prairie roadsides; [=sisymbrium incisum englem. ex a. gray, spring, summer misapplied] dimorphocarpa candicans (raf.) rollins spectacle pod dry, sandy soils, [=dithyrea wislizeni engelm.] near rivers; summer draba brachycarpa nutt. ex torr. & a. gray short-fruited whitlow-grass fields, pastures; early spring draba cuneifolia nutt. ex torr. & a. gray wedge-leaved whitlow-grass fields; common; early spring erysimum asperum (nutt.) dc. yellow phlox mountainsides; spring lepidium virginicum l. wild pepper grass abundant; spring lepidium sp. wild pepper grass roadsides; common; [=lepidium apetalum willd., misapplied] spring lesquerella auriculata (engelm. & a. gray) hairy bladder-pod prairies; early spring s. watson lesquerella densiflora (a. gray) s. watson bladder-pod near rivers; spring lesquerella ovalifolia rydb. ex britton slender bladder-pod rocky hillsides; spring nasturtium officinale w.t. aiton water cress streams; spring [=radicula nasturtium-aquaticum (l.) britten & rendle] rorippa sessiliflora (nutt.) hitchc. sessile-flowered cress mountain ravines; [=radicula sessiflora (nutt.) greene] spring sibara virginica (l.) rollins cut-leaved rock-cress mountainsides; [=arabis virginica (l.) poir.] early spring oklahoma native plant record 19 volume 14, december 2014 lottie opal baldock cucurbitaceae cucurbita foetidissima kunth missouri gourd fields, streams; [=pepo foetidissima (kunth) britton] summer cyperaceae carex vulpinoidea michx. fox sedge mountain ravines; summer cyperus echinatus (l.) alph. wood globose cyperus rivers [=cyperus ovularis (michx.) torr.] cyperus esculentus l. yellow nut-grass rivers cyperus odoratus l. coarse cyperus damp soils, [=cyperus ferax rich.] pastures cyperus strigosus l. straw colored cyperus mountain ravines eleocharis compressa sull. flat-stemmed spike-rush ponds; summer fuirena simplex vahl western-umbrella-grass rivers; late summer lipocarpha micrantha (vahl) g. tucker dwarf sedge riversides; summer [=hemicarpha micrantha (vahl) pax] schoenoplectus americanus (pers.) volkart three-cornered bulrush, damp ravines; ex schinz & r. keller sand-bar bulrush summer [=scirpus americanus pers.] scirpus pendulus muhl. reddish bulrush damp ravines; [=scirpus lineatus, misapplied] summer ebenaceae diospyros virginiana l. persimmon mountain ravines; spring euphorbiaceae cnidoscolus texanus (müll. arg.) small spurge nettle rivers; spring, [=jatropha stimulosa michx.] summer croton capitatus michx. goat-weed, hogwort roadsides; spring to fall croton glandulosus l. var. septentrionalis creeks; summer müll. arg. croton lindheimerianus scheele lindheimer’s croton-weed pastures; summer croton texensis (klotzsch) müll. arg. texas croton roadsides; mid summer euphorbia dentata michx. toothed spurge streams; summer euphorbia spathulata lam. reticulate-seeded spurge plains; summer [=euphorbia dictyosperma fisch. & c.a. mey.] euphorbia geyeri engelm. & a. gray geyer’s spurge rivers; summer euphorbia maculata l. spurge prairies; spring to fall euphorbia marginata pursh snow-on-the-mountain hillsides, rivers; summer, fall euphorbia missurica raf. white-flowered spurge prairies; summer [=euphorbia petaloidea engelm.] 20 oklahoma native plant record volume 14, december 2014 lottie opal baldock euphorbia nutans lag. large spotted spurge, mountains, spring to [=euphoribia preslii guss.] upright spotted spurge fall euphorbia serpens kunth round-leaved spreading spurge prairies; spring to fall stillingia sylvatica l. queen’s delight prairies; spring to fall tragia ramosa torr. branching tragia mountainsides; summer tragia sp. catnip-leaved tragia rivers; summer [=tragia nepetifolia cav., misapplied] fagaceae quercus fusiformis small live oak mountains; [=quercus virginiana mill., misapplied] pre-vernal quercus macrocarpa michx. bur oak, mossy-cup oak mountainsides; pre-vernal quercus marilandica münchh. black jack oak mountains; pre-vernal quercus muehlenbergii engelm.? cow oak, swamp oak mountains; spring [=quercus prinus l.] quercus shumardii buckley var. schneckii schneck’s red oak, mountains; spring (britton) sarg. spotted oak quercus stellata wangenh. post oak mountains; pre-vernal fumariaceae corydalis aurea willd. golden corydalis prairies; spring corydalis micrantha (engelm. ex a. gray) plains corydalis creeks, pastures, a. gray [=corydalis campestris (britton) near moisture; j. bucholz & palmer] spring gentianaceae eustoma exaltatum (l.) salisb. ex g. don russell’s eustoma creeks; summer [=eustoma russellianum (hook.) g. don] sabatia angularis (l.) pursh rose pink, bitter bloom creeks; summer sabatia campestris nutt. prairie sabatia prairies; summer geraniaceae geranium carolinianum l. wild geranium mountains, streams; spring gramineae agrostis hyemalis (watt) britton, sterns & ticklegrass mountainsides poggenb. alopecurus geniculatus l. foxtail streams andropogon gerardii vitman forked beard-grass, mountainsides [=andropogon furcatus muhl. ex willd.] big blue-stem oklahoma native plant record 21 volume 14, december 2014 lottie opal baldock aristida dichotoma michx. aristida prairies; summer aristida oligantha michx. few-flowered aristida pastures; summer aristida purpurascens poir. purplish aristida pastures, roadsides aristida purpurea nutt. purple three-awn prairies; spring bothriochloa laguroides (dc.) herter ssp. andropogon creeks; summer torreyana (steud.) allred & gould bouteloua curtipendula (michx.) torr. fall grama-grass prairies; summer bouteloua dactyloides (nutt.) j.t. columbus buffalo grass pastures; summer [=buchlöe dactyloides (nutt.) engelm.] bouteloua gracilis (willd. ex kunth) lag. blue grama-grass pastures; summer, ex. griffiths fall bouteloua hirsuta lag. hairy mesquite-grass pastures; summer bouteloua sp. bouteloua creeks; spring, [=bouteloua breviseta vasey, not in ok] summer bromus arvensis l. field chess roadsides; summer bromus catharticus vahl brome grass pastures, roadsides; [=bromus unioloides kunth] spring bromus racemosus l. brome grass roadsides; common; [=bromus commutatus schrad.] summer cenchrus spinifex cav. field sandbur rivers; summer [=cenchrus pauciflorus benth.] chasmanthium latifolium (michx.) yates broadleaf uniola mountains; autumn [=uniola latifolia michx.] chloris verticillata nutt. windmill grass prairies; common; spring cynodon dactylon (l.) pers. bermuda grass roadsides; common dichanthelium acuminatum (sw.) gould & panicum mountain ravines c.a. clark [=panicum tennesseense ashe] digitaria cognata (schult.) pilg. diffuse crag-grass fields; fall [=leptoloma cognata (schult.) chase] digitaria sanguinalis (l.) scop. large crab-grass fields echinochloa crus-galli (l.) p. beauv. barnyard grass streams eleusine indica (l.) gaertn. goosegrass pastures elymus canadensis l. canada wild-rye streams, ravines elymus glabriflorus (vasey ex l.h. dewey) virginia wild-rye streams scribn. & c.r. ball [=e. virginicus l. var. glabriflorus (vasey) bush] elymus repens (l.) gould couch grass fields; summer [=agropyron repens (l.) p. beauv.] eragrostis capillaris (l.) nees lace-grass prairies; summer eragrostis cilianensis (all.) vign. ex janchen stinkgrass fields, roadsides; summer eragrostis curtipedicellata buckley short-stalked love-grass roadsides, pastures; summer eragrostis secundiflora j. presl love-grass near river; summer eragrostis trichodes (nutt.) alph. wood eragrostis near river; summer 22 oklahoma native plant record volume 14, december 2014 lottie opal baldock erioneuron pilosum (buckley) nash hairy triodia mountainsides; [=triodia pilosa (buckley) merr.] spring hordeum pusillum nutt. little barley prairies; spring melica nitens (scribn.) nutt. ex piper three-flower melic mountainsides; spring panicum anceps michx. panicum rivers panicum capillare l. witch-grass, tumbleweed fields; summer panicum dichotomiflorum michx. fall panicum streams; fall panicum obtusum kunth blunt panic-grass, range-grass rivers panicum rigidulum bosc ex nees panicum streams [=panicum agrostoides spreng.] panicum virgatum l. switch-grass, wild red-top creeks; fall pascopyrum smithii (rydb.) á. löve western wheat-grass prairies; spring, [=agropyron smithii rydb.] summer paspalum setaceum michx. paspalum along rivers [=paspalum pubescens muhl. ex willd.] phalaris caroliniana walter carolina canary-grass moist places, roadsides; spring poa arachnifera torr. texas blue grass highways, hillsides; spring schedonnardus paniculatus (nutt.) trel. texas crab-grass, wire-grass prairies; common; summer schizachyrium scoparium (michx.) nash prairie beard-grass roadsides setaria parviflora (poir.) kerguélen knot-root bristle-grass creeks, roadsides; [=setaria geniculata (willd.) p. beauv.] spring setaria pumila (poir.) roem. & schult. yellow fox-tail roadsides; common; [=setaria lutescens (weigel) f.t. hubbard] late spring setaria viridis (l.) p. beauv. green foxtail-grass fields; summer sorghastrum nutans (l.) nash indian-grass mountains; fall sorghum halepense (l.) pers. johnson grass roadside ditches; summer sphenopholis obtusata (michx.) scribn. prairie wedge grass streams sporobolus cryptandrus (torr.) a. gray sand dropseed rivers; summer, fall tridens flavus (l.) hitchc. purpletop mountains, ravines [=triodia flava (l.) smyth] vulpia octoflora (walter) rydb. six-weeks fescue fields, pastures; [=festuca octoflora] spring grossulariaceae [saxifragaceae] ribes aureum pursh missouri or buffalo currant hillsides, streams; spring hydrophyllaceae phacelia hirsuta nutt. hairy phacelia prairies; spirng phacelia sp. small-flowered phacelia prairies; spring [=phacelia dubia (l.) trel., misapplied] oklahoma native plant record 23 volume 14, december 2014 lottie opal baldock iridaceae sisyrinchium angustifolium mill. blue-eyed grass , creeks, prairies; [incl. sisyrinchium gramineum curtis] stout blue-eyed grass spring sisyrinchium langloisii greene variable blue-eyed grass prairies; spring [=sisyrinchium varians e.p. bicknell] juglandaceae carya illinoinensis (wagenh.) k. koch pecan streams; spring juglans microcarpa berl. little walnut creeks; spring [=juglans rupestris engelm. ex torr.] juglans nigra l. black walnut creeks; spring juncaceae juncus biflorus elliott large grass-leaved rush riversides [=juncus aristulatus michx.] juncus brachycarpus engelm. rush roadside ditches; summer juncus interior wiegand indian rush roadside ditches; summer juncus marginatus rostk. awn-petaled rush rivers; summer juncus torreyi coville torrey’s rush rivers; summer krameriaceae [leguminosae] krameria lanceolata torr. bank-bur prairie roadsides; [=krameria secundiflora dc., misapplied] common; summer labiatae hedeoma hispida pursh rough or little pennyroyal plains; summer lamium amplexicaule l. henbit, dead nettle fields, roadsides; early spring monarda citriodora cerv. ex lag. purple lemon mint praries; summer [=monarda dispersa small] monarda fistulosa l. horse mint, wild bergamot ravines; early summer monarda punctata l. horse mint dry sandy soils; summer salvia azurea michx. ex lam. var. grandiflora tall blue sage plains; spring, benth. summer salvia reflexa hornem. lance-leaved sage creeks; spring, [=salvia lancaefolia poir.] summer scutellaria drummondii benth. drummond’s skullcap roadside ditches; common; spring scutellaria wrightii a. gray wright’s skullcap hillsides; spring teucrium canadense l. germander, wood sage ravines; summer 24 oklahoma native plant record volume 14, december 2014 lottie opal baldock leguminosae acacia angustissima (mill.) kuntze acacia sandy soils, near rivers; summer amopha canescens pursh lead-plant, devil’s shoe-string creeks; summer amorpha fruticosa l. river-locust, false indigo streams; summer astragalus canadensis l. tall astragalus creeks; summer astragalus crassicarpus nutt. ground plum creeks, pastures; [=astragalus caryocarpus ker gawl.] spring astragalus lotiflorus hook. low astragalus prairies; spring astragalus nuttallianus dc. annual astragalus prairies; spring baptisia australis (l.) r. br. blue false indigo mountainsides; spring baptisia bracteata muhl. ex elliott false indigo mountainsides; spring cercis canadensis l. redbud, judas tree creeks, mountain ravines; pre-vernal chamaecrista fasciculata (michx.) greene partridge pea fields, pastures; [=cassia chamaecrista l.] summer dalea aurea nutt. ex pursh golden parosela hillsides; summer [=parosela aurea (nutt. ex pursh) britton] dalea candida michx. ex willd. white prairie clover near rivers, sandy [=petalostemon candidus michx.] soils; summer dalea enneandra nutt. slender parosela rivers; summer [=parosela enneandra (nutt.) britton] dalea multiflora (nutt.) shinners round-headed prairie clover prairies; summer [=petalostemon multiflorus nutt.] dalea purpurea vent. purple prairie-clover prairies; summer [=petalostemon purpureus (vent.) rydb.] dalea sp. slender white prairie clover prairies; summer [=petalostemon gracilis nutt., misapplied] desmanthus illinoensis (michx.) macmill. ex illinois desmanthus rivers; summer b.l. rob. & fernald desmodium cuspidatum (muhl. ex willd.) dc. pointed-leaved tick trefoil, prairies; summer [=desmodium grandiflorum dc.] sticktight desmodium sessilifolium (torr.) torr. & sessile-leaved tick-trefoil mountains; summer a. gray glycyrrhiza lepidota pursh wild liquorice dry sands, roadsides; summer gymnoclados dioicus (l.) k. koch kentucky coffee-tree ravines; spring hoffmannseggia glauca (ortega) eifert blue-weed prairies; spring [=hoffmannseggia falcaria cav.] indigofera miniata ortega western indigo plant prairies; summer to [=indigofera leptosepala nutt. ex torr. & fall a. gray] lathyrus pusillus elliott low wild pea rivers; spring lotus unifoliolatus (hook.) benth. prairie bird’s foot, trefoil prairies; summer [=hosackia americana (nutt.) piper] oklahoma native plant record 25 volume 14, december 2014 lottie opal baldock medicago sativa l. alfalfa escaped cultivation, fields, roadsides; spring, summer melilotus officinalis (l.) lam. yellow meliot, sweet clover roadsides; abundant; summer mimosa microphylla dryand. sensitive brier roadside ditches; [=schrankia angustata torr. & a. gray, common; summer schrankia uncinata willd.] neptunia lutea (leavenworth) benth. neptunia mountain ravines; summer pediomelum cuspidatum (pursh) rydb. large-bracted psoralea prairies; spring [=psoralea cuspidata pursh] pomaria jamesii (torr. & a. gray) walp. james’s hoffmannseggia prairies; early [=hoffmannseggia jamesii torr. & a. gray] summer prosopis glandulosa torr. var. glandulosa prairie mesquite prairies; common; [=prosopis juliflora (sw.) dc. var. glandulosa late spring (torr.) cockerell] psoralidium tenuiflorum (pursh) rydb. few-flowered psoralea prairies; spring to [=psoralea tenuiflora pursh] fall robinia pseudoacacia l. black or yellow locust low waste places, cultivated; summer vicia caroliniana walter pale vetch prairies; spring liliaceae [incl. amaryllidaceae] allium canadense l. var. mobilense (regel) wild onion damp soils, ownbey [=allium mutabile michx.] roadsides; spring allium drummondii regel nuttall’s wild onion roadsides; spring [=allium nuttallii s. watson] allium textile a. nelson & j.f. macbr. wild onion mountain ravines; [=allium reticulatum g. don] late spring androstephium coeruleum (scheele) greene androstephium prairies, rare; early spring camassia scilloides (raf.) cory hyacinth, eastern camas mountain ravines [=camassia esculenta (raf.) cory] cooperia drummondii herb. prairie lily mountain ravines; spring nothoscordum bivalve (l.) britton yellow false garlic pastures, fields; spring, fall polygonatum biflorum (walter) elliott great solomon’s seal damp shady places; summer linaceae linum lewisii pursh lewis’s wild flax roadsides; spring linum rigidum pursh large-flowered yellow flax prairies; spring linum sulcatum riddell prairies; summer 26 oklahoma native plant record volume 14, december 2014 lottie opal baldock loasaceae mentzelia decapetala (pursh ex sims) stick-leaf sand, near rivers; urb. & gilg ex gilg summer mentzelia oligosperma nutt. ex sims stick-leaf, few seeded rivers; summer mentzelia lythraceae ammannia coccinea rottb. long-leaved ammannia mountain ravines; summer malvaceae callirhoe involucrata (torr. & a. gray) a. gray purple poppy mallow roadside ditches; common; spring callirhoe papaver (cav.) a. gray larger purple poppy mallow creeks; spring, summer sphaeralcea coccinea (nutt.) rydb. red false-mallow roadsides; common; [=malvastrum coccineum (nutt.) a. gray] spring martyniaceae proboscidea louisianica (mill.) thell. unicorn plant cultivated soils; [=martynia louisiana mill.] summer menispermaceae cocculus carolinus (l.) dc. carolina moonseed streams; summer moraceae [urticaceae] morus rubra l. red mulberry; wild mulberry creeks; spring nyctaginaceae mirabilis albida (walter) heimerl white oxybaphus roadsides, dry sand; [=oxybaphus albidus (walter) sweet] summer mirabilis hirsuta (pursh) macmill. hairy oxybaphus dry soils, roadsides; [=oxybaphus hirsutus (pursh) sweet] summer mirabilis linearis (pursh) heimerl oxybaphus prairies; summer [=oxybaphus linearis (pursh) b.l. rob.] mirabilis nyctaginea (michx.) macmill. petioled wild four-o’clock creeks; spring [=oxybaphus nyctagineus (michx.) sweet] oleaceae fraxinus americana l. white ash creeks; spring, summer fraxinus pennsylvanica marshall red ash mountain ravines; spring onagraceae ludwigia peploides (kunth) p.h. raven creeping primrose-willow ponds; summer [=jussiaea repens l.] oklahoma native plant record 27 volume 14, december 2014 lottie opal baldock oenothera cinerea (wooton & standl.) w.l. wooly gaura creeks; summer wagner & hoch [=gaura villosa torr.] oenothera curtiflora w.l. wagner & hoch gaura dry sandy soils; [=gaura parviflora douglas ex lehm.] summer oenothera glaucifolia w.l. wagner & hoch flax-leaved stenosiphon sandy soils near [=stenosiphon linifolius (nutt. ex e. james) rivers; summer heynh.] oenothera grandis (britton) smyth evening-primrose hillsides; spring [=oenothera laciniata hill var. grandiflora (s. watson) b.l. rob.] oenothera hartwegii benth. evening-primrose plains; summer oenothera laciniata hill evening-primrose sand, near rivers; summer oenothera macrocarpa nutt. missouri evening-primrose hillsides; summer [=oenothera missouriensis sims] oenothera rhombipetala nutt. ex torr. & evening-primrose near rivers; summer a. gray oenothera serrulata nutt. tooth-leaved primrose pastures, roadsides; summer oenothera sinuosa w.l. wagner & hoch wavy-leaved gaura hillsides; summer [=gaura sinuata nutt. ex ser.] oenothera speciosa nutt. showy evening-primrose prairies; spring oenothera suffrutescens (ser.) w.l. wagner scarlet gaura roadside ditches, & hoch [=gaura coccinea nutt. ex pursh] mountainsides; spring oenothera triloba nutt. three-lobed evening-primrose rivers; summer oenothera sp. biennial gaura rivers; summer [=gaura biennis l., misapplied] oenothera sp. evening-primrose near rivers; summer [=oenothera humifusa nutt., misapplied] oenothera sp. evening-primrose sandy soils, near [=oenothera oakesiana (a. gray) j.w. rivers; summer robbins ex s. watson & j.m. coult., misapplied] oxalidaceae oxalis corniculata l. yellow or procumbent wooddamp soils, sorrel mountainsides; spring oxalis stricta l. upright yellow wood-sorrel damp soils, mountainsides; spring oxalis violacea l. violet wood-sorrel damp soils, creeks, mountainsides; spring 28 oklahoma native plant record volume 14, december 2014 lottie opal baldock papaveraceae argemone albiflora hornem. white prickly poppy roadsides; spring, [=argemone alba lestib. f.] summer argemone polyanthemos (fedde) g.b. prickly poppy roadsides; summer ownbey [=argemone intermedia sweet] phrymaceae phryma leptostachya l. lop-seed mountains; summer phytolaccaceae phytolacca americana l. pokeweed creek banks; summer plantaginaceae plantago aristata michx. ribwort prairies; common; spring plantago patagonica jacq. pursh’s plantain pastures, roadsides; [=plantago purshii roem. & schult.] spring plantago rhodosperma decne. red-seeded plantain rivers; early spring plantago virginica l. dwarf plantain creeks; spring polemoniaceae ipomopsis rubra (l.) wherry red gilia mountainsides; [=gilia rubra (l.) a. heller] summer phlox pilosa l. phlox prairie roadsides; spring, summer polygalaceae polygala alba nutt. white milkwort prairies; summer polygonaceae eriogonum annuum nutt. annual gray-weed prairies; summer eriogonum longifolium nutt. long-leaved gray-weed rivers; summer, fall polygonum aviculare l. joint-weed, pink-weed near dwellings; summer polygonum hydropiper l. common smart-weed lakes; late summer polygonum lapathifolium l. dock-leaved joint-weed ravines; summer polygonum pensylvanicum l. showy joint-weed streams; summer polygonum punctatum elliott water smart-weed, dotted mountain ravines; [=polygonum acre kunth] water pepper summer, fall polygonum ramosissimum michx. bushy joint-weed rivers; summer polygonum tenue michx. slender joint-weed mountains; summer rumex altissimus alph. wood tall dock roadsides; summer rumex crispus l. dock, curly dock damp soils, [incl. rumex elongatus guss.] mountains; spring, summer oklahoma native plant record 29 volume 14, december 2014 lottie opal baldock portulacaceae [incl. caryophyllaceae, in part] claytonia virginica l. spring beauty creeks, pastures; common; early spring primulaceae androsace occidentalis pursh androsace pastures, fields; early spring samolus valerandi l. water pimpernel, brookweed streams; summer [=samolus floribundus kunth] ranunculaceae anemone berlandieri pritz. ten-petaled anemone pastures; spring [=anemone decapetala ard.] anemone caroliniana walter carolina anemone pastures; common; march, april clematis pitcheri torr. & a. gray virgin’s bower, leather-flower creeks, mountain ravines; spring delphinium carolinianum walter ssp. larkspur mountains, virescens (nutt.) r.e. brooks roadsides; spring [=delphinium penardii huth] myosurus minimus l. mouse tail streams, fields; early spring rhamnaceae ceanothus americanus l. new jersey tea mountains, streams; spring rosaceae crataegus crus-galli l. cock-spur haw, red raw rivers; spring crataegus viridis l. southern thorn streams; early spring geum canadense jacq. white avena mountain ravines; summer prunus americana marshall wild yellow or red plum rivers; spring prunus angustifolia marshall chickasaw plum roadside ditches, near rivers; spring rubus argutus link bramble rivers; spring rubus sp. bailey’s blackberry mountainsides; [=rubus baileyanus britton, misapplied] spring rubiaceae cephalanthus occidentalis l. button-bush streams, mountains; late spring diodia teres walter rough button-weed rivers; summer galium aparine l. cleavers mountains, streams; spring 30 oklahoma native plant record volume 14, december 2014 lottie opal baldock galium pilosum aiton hairy bedstraw mountains, streams; summer houstonia pusilla schoepf bluets creeks, pastures; [=houstonia minima beck] early spring stenaria nigricans (lam.) terrell narrow-leaved houstonia mountains; spring [=houstonia angustifolia michx.] rutaceae ptelea trifoliata l. tree-leaved hop-tree mountainsides; spring salicaceae populus deltoides w. bartram ex marshall cottonwood, necklace poplar creeks; pre-vernal salix nigra marshall black willow damp soils, streams; spring sapotaceae sideroxylon lanuginosum michx. chittim-wood, wooly buckthorn mountainsides; [=bumelia lanuginosa (michx.) pers.] summer smilacaceae [liliaceae] smilax bona-nox l. spiny-leaved greenbrier creeks; spring smilax herbacea l. carrion flower mountain ravines; summer smilax rotundifolia l. common greenbrier, creeks; spring horse-brier santalaceae comandra umbellata (l.) nutt. ssp. pallida bastard toad-flax dry sandy soils, (a. dc.) piehl near rivers; spring [=comandra pallida a. dc.] sapindaceae sapindus saponaria l. var drummondii wild china-tree, drummond’s creeks; spring (hook. & arn.) l.d. benson soapberry [=sapindus drummondii hook. & arn.] scrophulariaceae castilleja purpurea (nutt.) g. don var. indian paint brush mountains, lindheimeri (a. gray) shinners pastures; spring [=castilleja lindheimeri a. gray] castilleja sessiliflora pursh downy painted-cup mountainsides; summer collinsia violacea nutt. violet or narrow-leaved mountainsides; collinsia spring nuttallanthus canadensis (l.) d.a. sutton linaria mountains; spring [=linaria canadensis (l.) chaz.] oklahoma native plant record 31 volume 14, december 2014 lottie opal baldock penstemon cobaea nutt. beard-tongue prairies; late spring penstemon tubaeflorus nutt. funnel-shaped beard-tongue creeks; summer penstemon sp. sharp-leaved beard-tongue hillsides; summer [=penstemon acuminatus douglas ex lindl., misapplied] veronica agrestis l. field speedwell fields, pastures; spring veronica peregrina l. neckweed purslane, l creeks; early spring speedwell solanaceae chamaesaracha sp. hairy chamaesaracha roadsides; summer [=chamaesaracha sordida (dunal) a. gray, misapplied] datura stramonium l. jimson weed roadsides; summer physalis cinerascens (dunal) hitchc. ground-cherry creeks; spring [=physalis viscosa l., misapplied] physalis longifolia nutt.var. longifolia smooth ground-cherry creeks; spring physalis longifolia nutt. var. subglabrata smooth ground-cherry sandy soils, near {mack. & bush) cronq. rivers; summer [=physalis subglabrata mack. & bush] physalis mollis nutt. velvety ground-cherry roadsides; summer quincula lobata (torr.) raf. purple-flowered ground-cherry roadsides, prairies; [=physalis lobata torr.] spring, early summer solanum carolinense l. horse nettle prairies; common; spring, summer solanum elaeagnifolium cav. horse nettle pastures, roadsides; common; spring, summer solanum rostratum dunal buffalo bur abundant; summer, fall solanum sp. nightshade dry sandy soils; [=solanum nigrum l., misapplied] summer tamaricaceae tamarix sp. tamarish damp sandy soils; [=tamarix gallica l., misapplied] summer typhaceae typha latifolia l. broad-leaved cat-tail ponds; summer ulmaceae [urticaceae] celtis laevigata willd. southern hackberry creeks; spring celtis occidentalis l. rough-leaved hackberry mountains, streams; early spring ulmus americana l. white, american, or water elm creeks; early spring 32 oklahoma native plant record volume 14, december 2014 lottie opal baldock ulmus rubra muhl. slippery or red elm mountain ravines; [=ulmus fulva michx.] spring umbelliferae chaerophyllum tatinturieri hook. teinturier’s chervil roadsides; common; spring daucus pusillus michx. american carrot fields, pastures; spring lomatium foeniculaceum (nutt.) j.m. coult. carrot-leaved parsley rivers; summer & rose ssp. daucifolium (torr. & a. gray) w.l. theobald [=lomatium daucifolium (torr. & a. gray) j.m. coult. & rose] ptilimnium nuttallii (dc.) britton nuttall’s mock bishop-weed low places near mountains; summer sanicula canadensis l. short-styled snake-root mountainsides; summer spermolepis echinata (nutt. ex dc.) a. heller bristly-fruited spermolepis mountainsides; spring spermolepis inermis (nutt. ex dc.) mathias spreading spermolepis rivers; spring & constance [=spermolepis patens (nutt. ex dc.) b.l. rob.] valerianaceae valerianella radiata (l.) dufr. beaked corn salad creeks; spring verbenaceae glandularia bipinnatifida (nutt.) nutt. small-flowered verbena creeks, pastures; [=verbena bipinnatifida nutt.] early spring, summer glandularia canadensis (l.) nutt. large-flowered verbena sandy soils, near [=verbena canadensis (l.) britton] rivers; spring glandularia pumila (rydb.) umber dwarf verbena roadsides; summer [=verbena pumila rydb.] phyla cuneifolia (torr.) greene wedge-leaved fog-fruit hillsides; summer [=lippia cuneifolia (torr.) steud.] phyla lanceolata (michx.) greene fog-fruit creeks; summer [=lippia lanceolata michx.] phyla nodiflora (l.) greene spatulate-leaved fog-fruit streams; summer [=lippia nodiflora (l.) michx.] verbena bracteata cav. ex lag. & rodr. large-bracted verbena prairies; summer [=verbena bracteosa michx.] verbena stricta vent. hoary vervain mountainsides; summer violaceae viola bicolor pursh pansy or heart’s ease streams, pastures; [=viola rafinesquei greene] early spring oklahoma native plant record 33 volume 14, december 2014 lottie opal baldock viola sororia willd. violet damp sandy soils; [=viola papilionacea pursh] early spring vitaceae ampelopsis cordata michx. simple-leaved cissus rivers; spring [=cissus ampelopsis pers.] cissus trifoliata (l.) l. rock-grape mountains; summer [=cissus incisa des moulins, misapplied] parthenocissus quinquefolia (l.) planch. virginia creeper, creeks; summer five-leaved ivy vitis cinerea (engelm.) engelm. ex millard ashy or downy grape streams; spring vitis vulpina l. frost-grape, sweet scented creeks, rivers; [incl. vitis cordifolia michx.] grape spring zygophyllaceae kallstroemia parviflora j.b.s. norton greater caltrop rivers; summer [=kallstroemia maxima (l.) hook. & arn., misapplied] tribulus terrestris l. caltrop roadsides; common; summer 34 oklahoma native plant record volume 14, december 2014 lottie opal baldock appendix b tabular list of the families, kiowa county, ok [this table includes taxa as they were in the original thesis.] divisions, orders, families, etc. genera species varieties pteridophyta filicales polypodiaceae 4 5 marsileaceae 1 1 spermatophyta gymnospermae coniferales pinaceae 1 1 angiospermae monocotoledoneae pandales typhaceae 1 1 graminales gramineae 31 58 2 cyperaceae 6 11 xyridales commelinaceae 2 6 liliales juncaceae 1 6 liliaceae 7 11 amaryllidaceae 1 1 iridaceae 1 3 dicotyledoneae salicales salicaceae 2 2 juglandales juglandaceae 2 3 fagales fagaceae 1 6 urticales urticaceae 4 5 1 santalales santalaceae 1 1 polygonales polygonaceae 3 12 chenopodiales oklahoma native plant record 35 volume 14, december 2014 lottie opal baldock chenopodiaceae 4 5 1 amaranthaceae 3 5 phytolaccaceae 1 1 nyctaginaceae 1 4 illecebraceae 1 1 aizoaceae 1 1 caryophyllales caryophyllaceae 3 4 portulacaceae 1 1 ranunculales ranunculaceae 4 5 menispermaceae 1 1 papavervales papaveraceae 1 2 fumariaceae 2 2 cruciferae 8 16 capparidaceae 2 2 rosales crassulaceae 1 1 saxifragaceae 1 1 rosaceae 4 7 leguminosae 25 42 20 geraniales linaceae 1 3 oxalidaceae 1 3 geraniaceae 1 1 zygophyllaceae 1 2 rutaceae 1 1 polygalaceae 1 1 euphorbiaceae 5 16 1 sapindales anacardiaceae 2 4 sapindaceae 1 1 rhamnales rhamnaceae 1 1 vitaceae 3 6 malvales malvaceae 2 3 tamaricales tamaricaceae 1 1 violales violaceae 1 2 loasaceae 1 2 opuntiales 36 oklahoma native plant record volume 14, december 2014 lottie opal baldock cactaceae 2 2 myrtales lythraceae 1 1 onagraceae 4 17 1 umbellales umbelliferae 6 7 cornaceae 1 2 primulales primulaceae 2 2 ebenales sapotaceae 1 1 ebenaceae 1 1 gentianales oleaceae 1 2 gentianaceae 2 3 apocynaceae 2 2 asclepiadaceae 4 11 polemoniales convolvulaceae 3 6 polemoniaceae 2 2 hydrophyllaceae 1 2 boraginaceae 5 6 1 verbenaceae 2 8 labiatae 6 11 1 solanaceae 4 12 scrophulariaceae 5 9 martyniaceae 1 1 acanthaceae 1 2 phrymaceae 1 1 plantaginales plantaginaceae 1 4 rubiales rubiaceae 4 6 caprifoliaceae 4 2 valerianaceae 1 1 campanulales curcurbitaceae 1 1 campanulaceae 1 2 lobeliaceae 1 2 compositae 42 86 2 oklahoma native plant record 37 volume 14, december 2014 addenda [nomenclature has been updated according to the plants database (http://plants.usda.gov/plants).] the following plants were counted in the tabular list but are not given in the list of species: artemisia filifolia desmodium obtusum draba reptans [draba caroliniana] eleocharis obtusa gaillarida suavis [gaillardia trinervata] juncus tenuis physalis virginiana rudbeckia hirta scutellaria parvula symphyotrichum oblongifolium [aster oblongifollius] vicia minutiflora [vicia micrantha] xanthisma texanum the following plants listed in the stevens’ collection were not found by the author: artemisia ludoviciana spp. mexicana [artemisia mexicana] beta vulgaris bouteloua rigidiseta [bouteloua texana] carex gravida distichlis spicata dyssodiopsis tagetoides [dyssodia tagetoides] eleocharis rostellata muhlenbergia arenicola palafoxia sphacelata samolus ebracteatus lottie opal baldock flora of kiowa county, oklahoma, m.s. thesis by ms. lottie opal baldock oklahoma native plant record, volume 16, number 1, december 2016 4 oklahoma native plant record volume 16, december 2016 constance e. taylor https://doi.org/10.22488/okstate.17.100120 pollination ecology of sabatia cam pestr is nutt. (gentianaceae) unpublished report university of oklahoma biological station lake texoma 1972 constance e. taylor professor emeritus department of biological sciences southeastern oklahoma state university durant, ok 74701-0609 keywords: phenolog y, sabatia campestris, autog amy, allog amy abstract flower timing studies in june and july (1972) on populations of sabatia campestris nutt. show this plant to be allogamous (out crossing) under natural field conditions. however, when environmental factors reduce populations of solitary bees or when flower populations are particularly extensive and dense, the uncollected pollen causes retention of anthers into the period of style opening and stigma presentation. then autogamy (self-pollination) occurs. pollinators observed were solitary bees in the genera calliopsis, dialictus, and infrequently augochlorella. pollen viability is generally excellent. a chromosomal count of n=12 indicated the presence of aneuploid races in this plant species. the lengthening of petals from anthesis to wilting and calyx from bud to fruit production indicates flower size cannot be used as a taxonomic character to separate species. introduction sabatia campestris nutt. (gentianaceae), prairie rose gentian, is a common prairie annual found from illinois south through eastern texas and east to mississippi, with its greatest development in the prairie regions of the south central portion of the united states. the type specimen was collected from oklahoma (taylor and taylor 1994) and its occurrence cited as “in the open prairies of arkansas and red river, common, flowering in june and july” (nuttall 1836). the winged calyx that encloses the fruit is a unique species character. perry (1971), studying cross fertilization between species, found s. campestris to be reproductively isolated from other species. description of the flower flower timing studies in june and july (1972) on populations of s. campestris show this plant to be allogamous (out crossing) under natural field conditions. however, when environmental factors reduce populations of solitary bees or when flower populations are particularly extensive and dense, the uncollected pollen causes retention of anthers into the period of style opening and stigma presentation. then autogamy (self-pollination) occurs. the flowers are borne in a cymose inflorescence, the first flower terminal on a branch and subsequent flowers at the ends of opposite branches from below the first flower. the calyx is composed of five green oklahoma native plant record 5 volume 16, december 2016 constance e. taylor sepals fused and winged at their lower edges, the lobes being 2–6 times longer than the calyx tube. the calyx continues to grow during flowering and during fruit maturation. the winged calyx tube lengthens to 8 mm, and the calyx lobes grow to 25 mm long. the corolla is composed of five petals fused at the base, rose to pink or rarely white. petal lobes, like the calyx lobes, grow during flowering, reaching 23 mm long and 13 mm wide. the base of each petal lobe fades to white and has a rectangular yellow mark about the size and shape of the anther. this mark increases in length and intensity of yellow color during flowering. there are five anthers and one pistil with two branches, first green, then turning yellow with maturity. fruit capsules and seeds are numerous. methods the floral aspect of reproduction and pollinator behavior studies were done under field conditions during the height of blooming. two plots located at the northeast edge of durant, bryan county, oklahoma, approximately 1.21 and 4.86 hectares [3 and 12 acres] in size, were intensively observed. two plots in marshall county, oklahoma, one adjacent to the university of oklahoma biological station (u.o.b.s.) on lake texoma and another along a roadside 4.82 km [3 mi] north of willis were also observed. during the height of blooming in early june, flowers were marked by crewel yarns in the bud stage and checked three times daily, at a minimum, and during one night to determine floral presentation and movement of flower parts. climatic conditions were also noted. studies of pollinator presence were made by halfhour monitoring over several days, and behavior of bees were observed and recorded with a 35 mm single lens reflex camera. autogamy (self-pollination) was confirmed by the use of pollinator exclusion bags. no seed set was obtained when styles were excised. results observations the following description of floral presentation is for a sunny summer day in the presence of pollinators—solitary bees. s. campestris is protandric (the anthers functioning first) with the opening bud displaying straight anthers; the green pistil with tightly twisted styles lies in a horizontal position when the flower is fully open. by 8:30 a.m. (cdst), the anthers will have recurved at the tip, splitting open to release pollen from the anther chamber (figure 1). the timing of anther dehiscence is delayed by rain and cloudy weather, recurvation of anther tips and pollen presentation being delayed until as late as 11:30 a.m. under these conditions. the flower closes in the late evening, so no opening to floral parts is apparent. closure is not as tight as in the bud. the first night is the only night full closing occurs. on the second day, the anthers gradually curl further back and eventually tilt into a horizontal position. anther presentation of pollen is usually for two days but may be extended for a third day. on the morning of the third day, the anthers fall to the center of the bowl shaped flower. flowers presenting anthers for a third afternoon usually have anthers fall off the filaments in the evening. anther presentation is greatly modified by pollinator absence, being considerably prolonged for another one or two days. presentation of the stigmatic surface typically begins with the falling of the anthers. while still in a horizontal position, the stigmatic surfaces begin to turn yellow. staining with lactophenol aniline blue indicated receptivity is correlated with the appearance of the yellow color. the twined style branches begin untwisting gradually. over 24 hours is required for the style to 6 oklahoma native plant record volume 16, december 2016 constance e. taylor become erect and the stigmatic branches to completely untwist. stigmatic presentation is from the third day after opening (figure 2) until the flower wilts on the 6th or 7th day. some blossoms showed stigmatic wilt prior to petal wilt. the sepals are retained until the fruit dehisces some months later. figure 1 late day 2 showing curled dehiscence of anthers and unreceptive pistil with tightly twisted styles figure 2 day 4 or 5 showing erect style with receptive untwisted style branches, no anthers, and mimic anther lines at base of petals pollination pollinators visiting s. campestris were solitary bees in the genera calliopsis (andrenidae), dialictus (halictidae), and augochlorella (halictidae). all species of solitary bees were shorter than the anther length. visits were for pollen collection. day and time of visitation and plot visited are summarized in table 1. the flowers produced no nectar and seemed to hold little attraction for most other insects. occasional visits by various crab spiders and insects from leaf hoppers to butterflies were noted, but they were scattered and infrequent. in the durant plots, the size of the three principal pollinators diminished. the largest-sized species visited before june 10; the middle-sized species visited in midjune; and toward the last few days of june and first two weeks of july, the smallest bee species was the pollinator. the ecological interactions of bee and flower are pronounced. when flowers are open in the absence of pollinators, the anthers remain in an upright position for extended periods of time, and the total length of blooming is increased. flowers in paper pollinator exclusion bags and screen wire cages had, by the 5th and 6th days, accumulated piles of pollen heaped on the anther and scattered in the bowl of the blossom. the presence of this accumulation of dehisced pollen excluded wind pollination as an effective pollination agent. the presence of anthers remaining in an upright position delayed stigma presentation for several days, as late as six or seven days after opening. blooming time per flower was dramatically extended, one flower remaining unwilted for 14 days with the stigma still yellow and upright. collection of pollen was done in the same manner by all species of bees. after landing on the petal platform, they climbed up the anther, usually from the interior portion of the blossom. with the use of mouth parts, the pollen was collected and transferred by the front legs to the corbicula. after collecting for some minutes, the bee then packed the pollen into oklahoma native plant record 7 volume 16, december 2016 constance e. taylor table 1 times of bee visitation observed for plots pollinator visitation location date (1972) time (cdst) calliopsis sp. before june 10 12:30–3:30 durant dialictus sp. (black) june early morning u.o.b.s. dialictus sp. (black) late june all afternoon durant dialictus sp. (metallic rust) late june–early july all afternoon durant augochlorella sp. mid-june noon u.o.b.s. and n. of willis a firmer mass and was observed flying off for about one foot from the petal and then returning to the same blossom for further pollen. anthers were usually worked in a counter-clockwise manner, taken one after another until pollen from all five were collected. a blossom was worked sometimes two or three more times around. the longest timed collection of pollen from one flower by one bee was seven minutes. bees seemed to be unable to discriminate between flowers presenting anthers and those presenting stigmatic surfaces until landing on the flower. a bee quickly realized the absence of collectable pollen and would fly immediately to another flower. no bee was observed on a flower without anthers for more than 15 s, unless the petal platform was used when packing pollen. as the bee climbed on the style branches in the same manner it did the stamens, the pollen of the corbicula brushed the stigmatic surface causing pollination. quadrat sampling confirmed the hypotheses that pollen reward for bee visitation occurred in less than 40 percent of the blossoms open at any one time. later in the bloom season, sampling showed flowers presenting pollen fell to 27 percent of the total open blossoms. pollen viability counts were made on material stained with either lactophenol analine blue or snow’s stain. pollen viability was generally less than 1% non-viable pollen. highest count of non-viable pollen was 41.7%. lewis et al (1962) and perry (1971) reported chromosome numbers of n=13. collections of buds were made at three different times from the 1.21 ha [3 acre] durant plot. all three counts were n=12. chromosomal numbers for the genus as reported by perry (1971) are n=13, 14, 16, 17, 18, 19, 32, and 38. discussion perry (1971) reports the sequence of events in anthesis of sabatia flowers for all species. his data for s. campestris are bud (day 1), petals expanding and anthers recurving (day 2), anthers dehiscent (day 3), pollen shed (days 7–8), and stigma uncoiling and receptivity (days 7–8). i assume floral timing is based on greenhouse plants, as he states his pollination and fruit set studies are done under these conditions. his data for floral timing is the same as the data i recorded under pollinator exclusion bags, indicating that this species is generally crosspollinated. however, in the event of environmental influences which would hinder pollinator visits, the plants will set seed by self-pollination. 8 oklahoma native plant record volume 16, december 2016 constance e. taylor in the smaller plots, pollen collection was efficient throughout the blooming season. the 4.86 ha [12 acre] plot, densely populated by s. campestris, always had occasional blossoms from which pollen had not been collected. during observations, over a dozen flowers were observed with direct mechanical self-fertilization occurring by direct contact of the stigma surface with the anther. the only flowers not setting fruit were those which wilted during dry periods. all flowers in pollinator exclusion bags set fruit. in young plants with few blossoms, synchronization of flowers on the same plant was observed, all blossoms either presenting anthers or all presenting stigmas, but not presenting both on the same plant. older plants with eight or more flowers usually lost this synchronization. each s. campestris blossom has a small eye or star located at the center of the flower. each petal has a single yellow or green-yellow line notched at the apex, presented against a white background. the yellow line and white area enlarge and lines assume a bright yellow—at least to the human eye—as the stigmatic surfaces are presented. after anther shed, the yellow streaks and large yellow divergent style branches together provide a facsimile to the yellow flicker pattern of the presented anthers. the evolution of this pattern is often considered to be due to long and close bee pollinator association, with patterns on blossoms considered as nectar guides. this does not seem to be the case in s. campestris. the petals are pink, a color not generally associated with bee flowers. various shadings were common in all plots, ranging from white, light pink, pink, to dark pink. ultraviolet reflection did not occur on blossoms tested. the floral whorls, both calyx and corolla, grow after anthesis. this extremely unusual growth pattern is responsible for the enlargement of the eye. the largest flower on any plant will be that blossom which has been open longest. wilber (1955) noted that on some plants the central blossom was largest. neither wilbur (1955) nor perry (1971) mention the continued growth of the blossom while in bloom. the calyx also continues growth after the petals wither. length and width of the calyx and corolla have been used as taxonomic character to separate species. the phenomenon of continued growth also occurs in at least two other species. examination of fresh flowers of a population of s. angularis (l.) pursh, collected in june on the east side of the glover river at arkansas crossing, mccurtain county, oklahoma, showed the same size differentiation. the lengthening of petals from anthesis to wilting and calyx from bud to fruit production indicates flower size cannot be used as a taxonomic character to separate species. differences of individual blossoms on a single plant varied from 5 to 9 mm as measured from petal tip to petal tip tangent to the fruit. likewise, examination of specimens collected from a population of s. arenicola green, collected in april from a littoral area behind the dunes 12.87 km [8 mi] west of sabine pass, texas, showed size differences. measurement of the petal lobes showed they increased between 2 and 3 mm from early blooming flowers to old blossoms beginning to wilt. flowers in fruit had longer calyx lobes than those in flower. continued growth of blossoms is suspected for other species from descriptions and discussions in the monograph (1955) of the genus. acknowledgements thanks are due to dr. j. r. estes for direction in cytological work and to dr. r. w. thorp who identified the bees. special recognition to dick carson, stone post creations (rosecarson@juno.com), for the photographs of sabatia campestris. mailto:rosecarson@juno.com oklahoma native plant record 9 volume 16, december 2016 constance e. taylor literature cited lewis, w.h., h.l. stripling, and r.g. ross. 1962. chromosome numbers for some angiosperms of the southern united states and mexico. rhodora 64:147–161. nuttall, thomas. 1836. transactions of the philadelphia society. n.s. 5:197–198. perry, j.d. 1971. biosystematic studies in the north american genus sabatia (gentianaceae). rhodora 73:309–369. taylor, r. john and constance e.s. taylor. 1994. an annotated list of the ferns, fern allies, gymnosperms and flowering plants of oklahoma. 3rd edition. self-published. wilber, r.l. 1955. a revision of the north american genus sabatia (gentianaceae). rhodora 57:2–104. pollination ecology of sabatia campestris nutt. (gentianaceae) by dr. constance e. taylor journal of the oklahoma native plant society, volume 4, number 1, december 2004 oklahoma native plant record 57 volume 4, number 1, december 2004 editorial why do species’ names change? patricia a. folley the reason why scientific names change is because research is constantly correcting errors and scholarship is constantly untangling the related misconceptions. until the advent of the internet new names and name changes were approved by the international botanical congresses that met at ten-year intervals. between intervals, proposed new names were published by recognized publications like rhodora or sida. in 1994 john t. kartesz of the biota of north american program published a two-volume second edition of a synonymized checklist of the vascular flora of the united states, canada, and greenland, which became the established reference for names of north american plants on the date of its publication. this work made the flora of north america project practical by setting a base population against which the specialists could establish the limits of their work. with this resource there are two transforming innovations that are currently bringing about more rapid name changes in north american flora. first, the advent of the internet has vastly increased the speed of communication of scientific literature. results of research are published on the web within days of their discovery, and search engines make them accessible immediately. the u.s. department of agriculture has long-maintained a database for plant names for use by its agents and agencies. when that database became available on-line, with the inclusion of the kartesz checklist, any person with an internet connection could find out the current status of a plant name within a few minutes. the usda plants database http://plants.usda.gov/plants then became the publisher for all additions and corrections to the kartesz work, and changes are now posted daily. new names and combinations are also still published in print, including a detailed description of the plants involved. the impact on scholarship can be seen as the difference between the old “10 years or so” and the current “24 hours or so”. the second transformer is the flora of north america project (fna) which was begun in 1982 at the missouri botanical gardens. since the publication of vol. 1 in 1993 the flora project has driven both scholarship and research into the details of floristics in america. the list of contributors includes plant systematists and taxonomists still living today. conceived as a database project from the beginning, it both feeds and is fed by the internet. standards for the fna work have always compelled workers to research global archives. information based on past assumptions required verification, and the verification process yielded unexpected results. many contributors found themselves revising a lifetime of their own research before it could be accepted into the fna. verifying the work of contributors who have passed on is being continued by their successors. the majority of these efforts are being made by scholars and scientists who, while publicly funded for their teaching or research work, are not otherwise supported, and thus are volunteering their time and knowledge. as users of botanical information, we are often challenged to know what “today’s name” for a plant may be. but the outcome of the fna project, coupled with the unparalleled access to the literature provided by the internet, has made all of us better scholars with more reliable sources of information on the plants themselves. in time, the fna project will also become a printed reality, and the rate of change will slow. however, it will never cease as long as the real plants out in the real world continue to evolve. folley, p.a. https://doi.org/10.22488/okstate.17.100034 http://plants.usda.gov/plants� journal of the oklahoma native plantsociety, volume 3, number 1, december 2003 oklahoma native plant record volume 3, number 1, december 2003 68 three birds orchid and crane-fly orchid in oklahoma dr. lawrence magrath corresponding member, american orchid society conservation committee former chair, southwestern region orchid growers association curator, usao (ocla) herbarium chickasha, ok 73018-5358 the three birds orchid or nodding pogonia, triphora trianthophora (sw.) rydb., is one of the most beautiful jewels of the fall orchid collection in the eastern 1/3 of the country. it occurs from vermont and ontario, south to florida, west to texas, and north to michigan. pridgeon and urbatsch (1977) cite one collection from west feliciana parish in louisiana. in kansas it was long listed as part of the orchid flora, based on a report by popenoe. however, no one had seen a living specimen until rufus thompson, an algae specialist at the university of kansas, discovered them in baldwin woods south of the city of lawrence (douglas county) in late august 1971. after his report several botanists from the university of kansas also found it in baldwin woods and later in several other counties. still later, when working in the herbarium at the smithsonian institution in washington, d.c., i found a specimen collected by popenoe in topeka in shawnee county in 1876. in oklahoma it has been known to occur in cleveland county in central oklahoma since 1947. a collection from leflore county in the southeastern part of the state was made in 1967. since 1972 it has been found in choctaw, caddo, adair counties, and most recently in canadian county by dr. paul buck in 1993. there magrath, l. k. https://doi.org/10.22488/okstate.17.100023 have been several other unconfirmed reports in oklahoma. in arkansas it has been found in several counties in the ozark mountains, which cross the border into oklahoma. one of the reasons that so few people have reported seeing this delightful little orchid is probably that it blooms in late august and early september. that is a time when few orchid or wildflower enthusiasts are out in the woods because of the ticks, mosquitoes, and miserable hot weather. nevertheless, it is a flower well worth the trouble endured to find it. it grows in rich mixed deciduous woodlands or deciduous-pine woodlands in the deep humus or leaf mold of moist shaded areas. this species is often associated with other fall flowering orchids such as corallorhiza odontorhiza and tipularia discolor. it may occur as a single stem, a few scattered stems, or as large colonies up to 3 feet in diameter with hundreds or occasionally thousands of stems, such as are found at the battiest site in northern mccurtain county, oklahoma. the 7-30 cm (3-12 inches) plants produce from one to six (rarely seven) flowers about the size of a nickel at the tip of succulent green stems. typically the plants are about 1018 cm (4-7 inches) tall. the flowers open white with a delicate patch formed by three crests or lamellae of emerald green oklahoma native plant record 69 volume 3, number 1, december 2003 magrath, l.k. in the center of the lip. as the flowers age, they become flushed with pink or lavender. when observed under magnification the flowers appear to be sculpted out of transparent or translucent crystal. they are truly a delight to behold. the plants have underground stolons bearing fleshy tuberoids (medley 2002). the tuberoids rarely ever penetrate into the soil, but rather appear to be confined to the layer of decaying organic matter. any attempts to cultivate this plant should take this into account. oklahoma native plant society (onps) and southwestern region orchid growers association (swroga) conservation committees would appreciate knowing of additional locations for this orchid. it is probably more common than previously believed. however, proof is in the finding! the crane-fly orchid, tipularia discolor (prush) nutt., is one of the more interesting and elusive native orchids. its name is derived from the latin tippula “water-spider” + discolor “variegated, of different colors.” it is one of the late summer orchids found in arkansas, southeastern oklahoma, eastern texas, and louisiana. the genus contains three recognized species: tipularia josephi in the himalayan mountains, t. japonica in japan and t. discolor in the united states. it ranges from florida west to eastern texas through oklahoma, arkansas, missouri, southern illinois, and indiana east to pennsylvania, new jersey, and massahusetts, as well as along the atlantic coast. it may be the most common orchid in arkansas (slaughter 1993). in oklahoma it was first collected in 1968 two miles south of honobia by steve stephens from the university of kansas. the collection consisted of one flowering plant (magrath 1973). since then it has been found throughout the southeastern part of oklahoma in colonies often numbering in the thousands. according to luer (1975) “the plants are characterized by their series of undergournd tubers which are actually corms connected by slender rhizomes.” a new corm is produced each year. each new mature corm produces a solitary ovoid, overwintering leaf which disappears in may or june. the inflorescence is produced in august. the scape is slender and is terminated by a raceme of small, dull flowers. the sepals and petals are free but one petal partially overlaps the dorsal sepal. the lip is threelobed and has a spur at the base. homoya (1993) describes the inflorescence as “giving an impression of a swarm of flying gnats, mosquitoes, or small crane-flies.” to find a large colony of these plants in full bloom in a dimly lit woods and to watch the flowers dance with every little bit of breeze is a treat. then they truly seem like insects in flight. homoya (1993) notes that, “the flowers of tipularia are unique among north american orchids in that they are not bilaterally symmetrical. instead, the sepals and petals are positioned so that the flower is lopsided, with an unlike number of petals and sepals to either side of the column. moreover, the flowers are angled to one side of the main stem, some to the right, others to the left.” homoya (1993) further notes that tipularia commonly sets seed capsules. occasionally, isolated solitary plants may not be pollinated, but wherever there is a population, each plant normally will have between 80 to 100% capsule set. tipuolaria like aplectrum (adamand-eve, putty root) has a series of corms connected by a slender rhizome. both produce an over-wintering single leaf, although aplectrum is usually 2-5 times oklahoma native plant record volume 3, number 1, december 2003 magrath, l. k. 70 larger and accordion pleated, both are typically purplish on the underside of the leaf. aplectrum is, however, a late spring to early summer flowering plant, while tipulari is a late summer flowering plant. in oklahoma its late flowering time overlaps with golden plume, platanthera ciliaris, and three-bird orchid, triphora trianthophora. tipularia tends to grow in decaying leaf litter in relatively well drained areas often over a rocky substrate, and seems to prefer drier locations than does aplectrum. homoya (1993) notes that tipularia “is clearly advancing its range” in indiana. i feel that the same is true in oklahoma. the orchid can be cultivated in shaded areas where decaying leaf litter that is relatively moist, but well drained. basically the same type of habitat in which triphora trianthophora and malaxis unifolia (green adder’s mouth) would grow. i have also successfully grown it in terrarium culture. since it produces large numbers of seed capsules, it would seem to be a good candidate for growing in flask from seed and it is to be hoped that at some time in the near future it will be available in the form of nursery propagated plants, as opposed to collected plants. while it is hoped that this native will soon begin to come into cultivation and that the triphora trianthophora will be found in more locations, as always, we recommend that when in natural settings leave only footprints, being careful not to damage young seedlings, and take only memories and photos. references correll, d.s. 1950. native orchids of north american north of mexico. waltham, ma: chronica botanica company. homoya, m.a. 1993. orchids of indiana. indiana university press: indiana academy of science. luer, c.a. 1975. the native orchids of the united states and canada excluding florida. bronx, ny: new york botannical garden. magrath, l.k. 1973. the native orchids of the prairies and plains region of north america [dissertation]. lawrence, ks: university of kansas. medley, m. 2002. triphora in flora of north america. vol.26: magnoliophyta: liliidae: liliales and orchidales. flora of north america editoral committee. new york and oxford: oxford univ. press. pridgeon, a.m. & l.e. urbatsch. 1977. contributions to the flora of louisiana. 2. distribution and identification of orchidaceae. castanca 42:293-304. slaughter, carl r. 1993. wild orchids of arkansas. isbn: 0-9638497-0-0. published by the author. yatskievych, g. 1999. steyermark’s flora of missouri, vol 1, rev. ed. missouri dept. of conservation in cooperation with the missouri botanical garden press. st. louis, mo. oklahoma native plant record 71 volume 3, number 1, december 2003 magrath, l.k. three birds orchid, triphora trianthophora (sw.) rydb. photos courtesy charles lewallen. 72 oklahoma native plant record volume 3, number 1, december 2003 magrath, l. k. crane-fly orchid, tipularia discolor (prush) nutt. photos curtesy of charles lewallen. oklahoma native plant record, volume 14, number 1, december 2014 50 oklahoma native plant record volume 14, december 2014 katherine e. keil and karen r. hickman https://doi.org/10.22488/okstate.17.100105 mapping distribution in oklahoma and raising awareness: purple loosestrife (lythrum salicaria), multiflora rose (rosa multiflora), and japanese honeysuckle (lonicera japonica) katherine e. keil karen r. hickman oklahoma state university oklahoma state university stillwater, oklahoma 74074 stillwater, oklahoma 74074 katie.keil@okstate.edu karen.hickman@okstate.edu keywords: invasive plants, management, population, fact sheet abstract this paper includes updated oklahoma distribution maps and informational fact sheets for purple loosestrife, multiflora rose, and japanese honeysuckle to promote awareness of invasive plant issues. the current information on the internet contains discrepancies concerning the county-level distribution data of these three invasive plants in oklahoma. to gain a more accurate dataset, the authors created a survey and sent it to oklahoma state university extension educators, master gardeners, oklahoma association of conservation districts, and other knowledgeable, credible parties across oklahoma. once survey data were compiled, 3 distribution maps were created and 6 unique fact sheets were produced with the updated information. from the 22 survey responses, 7 new county records were documented and mapped. two new sightings were documented for purple loosestrife in canadian county and rogers county; 4 new sightings were documented for multiflora rose in the counties of atoka, johnston, payne, and pushmataha; and 1 new sighting was documented for japanese honeysuckle in garfield county. the findings in this research detail the need for updated distribution maps and increased education to prevent the spread of problem species and provide the public with methods of eradication. introduction according to the united states national arboretum, an invasive plant is a species that “has the ability to thrive and spread aggressively outside its natural range” (the united states national arboretum 2008). these invasive plants have a competitive edge on their native counterparts because the insects, diseases, and animals that naturally keep their numbers in check do not typically exist in the new environment. this situation is known as an “enemy release”, which allows the populations of an invasive species to increase to high levels, suppressing growth of native vegetation and altering their composition and structure (keane and crawley 2002). invasive plant species adversely affect the habitats they invade economically, environmentally, and/or ecologically by disrupting natural ecosystem processes (the united states national arboretum 2008). purple loosestrife (lythrum salicaria), multiflora rose (rosa multiflora), and japanese honeysuckle (lonicera japonica) are 3 problematic invasive species in oklahoma. they were chosen for this study because each represents a difference in abundance and growth form. according to the oklahoma vascular plants database (ovpd), purple loosestrife is the least abundant of the three and is known to occur in only 4 oklahoma counties. oklahoma native plant record 51 volume 14, december 2014 katherine e. keil and karen r. hickman japanese honeysuckle, however, has spread extensively across oklahoma and is listed in 45 counties. finally, multiflora rose is documented in 39 counties (oklahoma vascular plants database 2014). the three plants also differ in growth form: purple loosestrife is an herb, multiflora rose is a shrub, and japanese honeysuckle is a vine. our objective was to accumulate information about invasive species that demonstrate the wide variation in abundance, growth form, and distribution of invasive plant species in oklahoma. figure 1 purple loosestrife flower. courtesy of samuel roberts noble foundation, ardmore, oklahoma. purple loosestrife is an erect, perennial, wetland herb that is popular among gardeners because of its magenta-colored spikes of flowers (fig. 1) that bloom from june to september. purple loosestrife was brought to america from europe for ornamental and medicinal purposes in the 19th century but was also unintentionally imported in ship ballast water (bravo 2009a). in addition to anthropogenic vectors, the high reproductive ability of purple loosestrife has contributed to its success as an invasive species in the united states. a single flowering individual can produce approximately 2.5 million seeds a year. these seeds are dispersed by animals, humans, and other vectors that carry the propagules significant distances. for example, waterfowl can carry the seeds along waterways, or seeds can attach to boat bottoms and be transported downstream. purple loosestrife also spreads asexually by stem and root fragments that resprout when they settle in a new location (new hampshire department of environmental services 2010). purple loosestrife has spread extensively throughout the united states and is now documented in every contiguous state, excluding florida (blossey 2002). as the pennsylvania department of conservation and natural resources states, the rapid growth rate and resulting dense stands of purple loosestrife allow it to outcompete native vegetation, some of which may be rare or endangered species (e.g., federally endangered orchids). this causes significant ecological harm by reducing the biodiversity of an area and creating monocultures. these dense stands of purple loosestrife can reduce the native species’ habitats and available food sources. purple loosestrife can also inhibit stream flow, changing the hydrology of wetlands (pennsylvania department of conservation and natural resources, n.d.). economic impacts of purple loosestrife include reduced land values for properties infested by the plant and impediment of boating and other recreational activities. in addition, purple loosestrife invades irrigation systems and adversely impacts agricultural productivity (washington state department of ecology, n.d.). however, because it is valued as a garden plant, purple loosestrife typically stirs little concern. for example, this project unveiled that purple loosestrife has been planted in demonstration and teaching gardens in oklahoma, whose purpose is to educate the public about plants that grow well in the area (penn state extension 2014). promoting invasive species growth in gardens is potentially harmful because these plants can become garden escapees, alter valuable ecosystem functions, and threaten local vegetation (the united states national arboretum 2008). 52 oklahoma native plant record volume 14, december 2014 katherine e. keil and karen r. hickman figure 2 multiflora rose flower. courtesy of samuel roberts noble foundation, ardmore, oklahoma. multiflora rose is a multi-stemmed, sprawling shrub that can grow more than 4.5 m tall with long, arching stems that produce recurved thorns. in june and july, multiflora rose begins to bloom, displaying large clusters of fragrant white flowers (fig. 2) (the university of maine 2001). multiflora rose was first introduced to the united states from japan in 1866 as a rootstock for ornamental roses and was distributed approximately 70 years later by the u. s. soil conservation service to control soil erosion. since then, it has been intentionally spread to serve as wildlife habitat improvement, fences for livestock, vehicle crash barriers along roadways, and protection from the glare of oncoming vehicle headlights (swearingen et al. 2010). even without human aid, multiflora rose is prolific and can successfully spread by its own means. each adult plant can produce approximately 1 million seeds annually that are distributed by birds and other wildlife that eat the fruits, known as hips. multiflora rose can also spread asexually. as stems grow taller they begin to arch, and when they come into contact with the ground they form roots (forest invasive plants resource center 2005). although it is a practical shrub, multiflora rose remains highly invasive and its spread should be avoided if possible. it grows aggressively, creating dense, impenetrable thickets. these blankets of multiflora rose suppress native vegetation and lead to a loss in biological diversity by prohibiting nest construction by birds, altering habitat structure, and inhibiting forest tree regeneration (the university of maine 2001). figure 3 japanese honeysuckle flower. courtesy of samuel roberts noble foundation, ardmore, oklahoma. japanese honeysuckle is a woody vine with fragrant white flowers that yellow with age. the flowers consist of 5 fused petals, occur in pairs on axillary peduncles (fig. 3), and bloom from april to july. in autumn, mature japanese honeysuckle plants bear small black fruits containing two to three seeds each (forest invasive plants resource center 2005). japanese honeysuckle was introduced to the us from asia in the 1800s and first became naturalized in the northeast. since its arrival, it has been intentionally spread throughout the country because it is valued as a fragrant ornamental. japanese honeysuckle has also been anthropogenically distributed to offset erosion and serve as wildlife forage and cover (schierenbeck 2004). wildlife is of further assistance to distribution by consuming japanese honeysuckle fruits and dispersing seeds long distances (forest invasive plants resource center 2005). similar to the aforementioned plants, japanese honeysuckle is a successful invader oklahoma native plant record 53 volume 14, december 2014 katherine e. keil and karen r. hickman without external assistance. japanese honeysuckle’s lack of natural competitors, ability to grow rapidly, adaptability to varying habitats, and prolonged growth period allow it to spread successfully. it also has vegetative runners that root when they make contact with the soil (forest invasive plants resource center 2005). japanese honeysuckle impacts both individual plants and plant communities. due to its climbing nature, japanese honeysuckle winds around the stems and trunks of native woody plants, restricting the water flow to the plant and ultimately killing them. japanese honeysuckle also affects the herbaceous and understory plant communities by forming a dense blanket of growth that blocks sunlight and suppresses native plants growth, altering forest structure (bravo 2009b). distribution information for purple loosestrife, multiflora rose, and japanese honeysuckle was compiled from three sources: the oklahoma vascular plants database (ovpd), the oklahoma invasive plant council (okipc), and the united states department of agriculture plants database (usda). the ovpd is an online data source consisting of label data from specimens stored in oklahoma herbaria. these data are queryable and are used to produce county-level distribution maps for oklahoma plants. since an accessioned voucher is necessary for inclusion in their database (oklahoma vascular plants database 2014), the ovpd does not accept any public observation data in the form of online submissions, photos, or other reports. the usda plants database is a clearinghouse derived from scientific literature, herbarium specimen, and confirmed observations. the public is able to contribute by providing verifiable plant distribution information including locality, date collected, collector’s name, and at least one form of documentation of the invasive plant (usda, nrcs 2014). the okipc compiles data from the ovpd and observations provided by the oklahoma biological survey to generate their county distribution maps. okipc records of invasive plants are not of exact physical locations but consist of occurrences within oklahoma counties. their ultimate goal is “facilitating education and management for protection of our economic and natural resources” (oklahoma invasive plant council 2014). the county distribution maps for each plant species, however, differ between organizations, revealing discrepancies among the data sources. these differences can create confusion for oklahomans concerned about the spread of invasive plants throughout the state. as a contribution to remedying this discrepancy, this study’s objective is to update distribution data for purple loosestrife, multiflora rose, and japanese honeysuckle and promote awareness of invasive plant impacts in 6 new fact sheets. methods we created a survey in order to more accurately reflect the distribution and density of purple loosestrife, multiflora rose, and japanese honeysuckle in the 77 counties of oklahoma. the survey included 11 questions about the presence, specific location, and density of these three species and was distributed to oklahoma state university extension educators, master gardeners, oklahoma association of conservation districts, and other experienced parties across oklahoma. those who received the survey were encouraged to forward it to their colleagues and include it in listservs, so we estimate 250-350 surveys were distributed in total. survey responses were recorded in an excel® workbookfor summary and analysis. although data were collected on the density of specific invasive plant occurrences, the resources to accurately map abundance were lacking, and thus any further conclusions on 54 oklahoma native plant record volume 14, december 2014 katherine e. keil and karen r. hickman abundance in oklahoma counties are excluded from this report. these new distribution data were then compared with data from ovpd, okipc, and usda. three new maps were created to integrate the survey data with the preexisting data to enhance the accuracy of distribution maps for purple loosestrife, multiflora rose, and japanese honeysuckle. these maps were used in the production of two different formats of new fact sheets for each species of invasive plant. the first fact sheet follows the oklahoma state university extension services format and provides a general description of the species’ characteristics and life histories, invasive traits and impacts, and recommended management options (appendix a). the second fact sheet is unique, specific to oklahoma, and formatted to the style of an old western wanted poster. the authors designed this second fact sheet in a reader-friendly manner to attract attention and be memorable. this fact sheet includes a description of the plant and its habitat, its “crime”, “hometown”, classification in oklahoma, and the number of counties in which it is found (appendix b). results and discussion the results of the data collection were limited to the 22 responses the survey respondents provided, and thus some location data on the invasive plants are less detailed than desired. a comparison of the pre-existing data from the ovpd, okipc, and usda to survey data illuminates several discrepancies. comparisons among these data sources and analysis of newly acquired data are discussed for each plant below. purple loosestrife of the 22 surveys returned, three respondents provided data for purple loosestrife, noting that it exists in canadian, cleveland, and rogers counties. in canadian county, purple loosestrife was sighted in the city of edmond on morgan road, 0.25 miles north of edmond road. in cleveland county, purple loosestrife was located in a demonstration and teaching garden in the cleveland county fairgrounds. in rogers county, purple loosestrife was sighted around the edges of a homeowner’s pond on the western border of the county. the distribution maps of purple loosestrife (fig. 4) include the ovpd occurrences in four counties and the okipc in 3 counties. the usda plants database shows purple loosestrife as occurring in oklahoma, but does not provide county level occurrence data. this project’s data resulted in a six county distribution map with new occurrence records for rogers and canadian counties. multiflora rose a total of 8 surveys were returned for multiflora rose, noting invasive occurrences in atoka, bryan, choctaw, comanche, johnston, mccurtain, oklahoma, okmulgee, osage, payne, pittsburg, and pushmataha counties. a single respondent provided the county sightings in atoka, bryan, choctaw, mccurtain, pittsburg, and pushmataha, describing all 6 counties as having scattered occurrences. in comanche county, multiflora rose was sighted in the wichita mountains wildlife refuge, with the occurrence being approximately 53 m² in size. in johnston county, multiflora rose was sighted in the south-central part of the county, but no other information was provided. in oklahoma county, multiflora rose was sighted on tinker air force base with the invasion being described as light to moderate in density. there were several sightings in pastures and fence lines throughout okmulgee county. in osage county, multiflora rose was sighted in both the southeast and far northeast portions of the county with a sparse occurrence on the ground and in fence lines. in payne county, oklahoma native plant record 55 volume 14, december 2014 katherine e. keil and karen r. hickman multiflora rose was sighted in a forested area outside of the stillwater city limits and was noted as a light occurrence. the ovpd contains records for multiflora rose for 39 counties, the okipc for 35 counties, and the usda plants database for seven counties. combined with this project’s data, multiflora rose has a 43 county distribution with new occurrence records for atoka, johnston, payne, and pushmataha counties (fig. 5). it is evident from the number of records of multiflora rose that this invasive plant is widespread in oklahoma, and eradication efforts will be significantly more difficult than for purple loosestrife. thus, efforts should be focused primarily on limiting the spread of populations to the western tier counties of oklahoma or other uninhabited regions of the state. japanese honeysuckle nine surveys were returned on japanese honeysuckle, providing information for atoka, bryan, carter, choctaw, comanche, garfield, mccurtain, oklahoma, okmulgee, payne, pushmataha, and tulsa counties. the sightings of atoka, bryan, choctaw, mccurtain, and pushmataha counties were all recorded by one respondent, who described occurrences at “numerous locations” including the antlers industrial park property, which was described as a very dense infestation. in carter county, japanese honeysuckle was reported as a dense occurrence on a homeowner’s property east of ardmore. in comanche county, japanese honeysuckle was sighted in the wichita mountains wildlife refuge, encompassing approximately 4 m². no details were provided regarding the japanese honeysuckle occurrences in garfield county. in oklahoma county, japanese honeysuckle was sighted on tinker air force base and was described as moderately to highly dense. in okmulgee county, japanese honeysuckle was sighted across the county, with no specific location or density information mentioned by the respondent. in payne county, japanese honeysuckle was sighted in a forested area outside of stillwater city limits and was moderately dense. in tulsa county, japanese honeysuckle was sighted in mohawk park surrounding the tulsa zoo. for japanese honeysuckle, the ovpd records this species in 45 counties, the okipc in 43 counties, and the usda in 7 counties. in combination with this project’s data, japanese honeysuckle has a 46 county distribution with a new occurrence record for garfield county (fig. 6). japanese honeysuckle is not present in the northwestern portion of oklahoma, which is likely due to colder temperatures and lower precipitation that limits japanese honeysuckle growth (forest invasive plants resource center 2005). however, the rest of the state is widely inhabited by this species and thus efforts should be focused on preventing further spread of this highly invasive plant in these areas. distribution maps the differences in county distributions among the ovpd, okipc, and usda, (see figs. 4-6) can be attributed mostly to each organization’s differing data sources. the usda plants database distribution maps had the lowest documented occurrences of these species, while the ovpd had the most occurrences recorded. it is important to note that not all invasive occurrences are equally significant. for example, the record of purple loosestrife in the cleveland county demonstration gardens may hold less threat of spreading beyond the residential site; whereas, the invasion on the pond’s edge in rogers county is more problematic due to its increased likelihood of spreading through the watershed. purple loosestrife is readily transported through waterways, establishing communities downstream or in this example, potentially spreading around the 56 oklahoma native plant record volume 14, december 2014 katherine e. keil and karen r. hickman figure 4 the distribution of purple loosestrife in oklahoma between sources oklahoma native plant record 57 volume 14, december 2014 katherine e. keil and karen r. hickman figure 5 the distribution of multiflora rose in oklahoma between sources 58 oklahoma native plant record volume 14, december 2014 katherine e. keil and karen r. hickman figure 6 the distribution of japanese honeysuckle in oklahoma between sources oklahoma native plant record 59 volume 14, december 2014 katherine e. keil and karen r. hickman rogers county pond. once established, these purple loosestrife stands may outcompete the native vegetation and alter the pond’s wetland structure and function (new hampshire department of environmental services 2010). in addition, these larger communities of purple loosestrife have increased odds of spreading to other sites due to the greater number of offspring they produce. in the cleveland county gardens, however, the extent of the purple loosestrife population may be maintained, and there is reduced opportunity to spread through the waterways. the difference in each organization’s criteria for adding an invasive into their distribution maps may explain why the ornamental planting in the cleveland county gardens is not listed by any of the organizations. the authors, however, have included the cleveland county record in this report to show that residential ornamental plantings, albeit less pervasive than others, can still spread outside the garden’s borders and cause ecological impact. although it may not qualify as a record by the organizations listed in this report, it can be argued that no invasive plant should be viewed as acceptable and remain undocumented. overall, accurate distribution maps must be produced to inform the public and land managers where invasive populations exist in order to limit the spread of invasive populations to uninhabited areas of oklahoma. accurate fact sheets must also be available to guide conservationists to the best method(s) for their eradication. conclusion the objective of this study was to emphasize the need for more research in invasive plant distributions while increasing the number of known occurrences for purple loosestrife, multiflora rose, and japanese honeysuckle using survey responses. based on survey results, 7 new county-level occurrences were documented for these 3 invasive plants. the differences among existing data sources in this report currently provide conflicting impressions of invasive plant distributions. these discrepancies can potentially impede management and eradication efforts and become increasingly problematic without the availability of accurate data. it is evident that one person or research project alone cannot efficiently take on the burden of mapping invasive plants. invasive plant species play a role in many aspects of life — from the environment to the economy — and must be considered as a group. programs such as early detection and distribution mapping system (eddmaps®), a phone application and website, enable the citizen scientist aspect of research. this program allows a user to upload photographs, gps coordinates, and population data of a plant that they believe is invasive. these data are then submitted to an expert to confirm species identification. once the identification is verified, the occurrence is added to a statewide distribution map that is viewable online (eddmaps 2014). maps that contain more detailed occurrence records enhance our ability to provide information to the public about the threats these invasive species have on our natural resources. acknowledgments we would like to thank tracey miller for helping distribute the survey. we also appreciate the respondents, all of whom provided significant assistance in this project: chad webb, greg highfill, will cubbage, pearl pearson, dennis martin, scott johnson, loren sizelove, doug maxey, paul koenig, tracey miller, ray ridlen, tom smith, mindy mcnair, jay ross, danny cook, john hasse, john krupovage, todd fagin, mike porter, jim johnson, and jeanetta cooper. 60 oklahoma native plant record volume 14, december 2014 katherine e. keil and karen r. hickman literature cited blossey, b. 2002. purple loosestrife. ecology and management of invasive plants program. http://www.invasiveplants.net/plants/p urpleloosestrife.htm. accessed 20 april 2014. bravo, m.a. 2009a. purple loosestrife. plant conservation alliance. national park service. http://www.nps.gov/plants/alien/fact/ loja1.html. accessed 15 april 2014. bravo, m.a. 2009b. japanese honeysuckle. plant conservation alliance. national park service. http://www.nps.gov/plants/alien/fact/ loja1.htm. accessed 15 april 2014. eddmaps. 2014. early detection & distribution mapping system. the university of georgia – center for invasive species and ecosystem health. available online at http://www.eddmaps.org/; last accessed december 30, 2014. forest invasive plants resource center. 2005. usda forest service. na.fs.fed.us/spfo/invasiveplants/index.a sp. accessed 15 april 2014. keane, r.m.and m.j. crawley. 2002. exotic plant invasions and the enemy release hypothesis. trends in ecology and evolution 17:164-170. maryland department of natural resources. n.d. facts about purple loosestrife. http://dnr.maryland.gov/wildlife/plants _wildlife/purpleloosestrife/index.asp. accessed 19 april 2014. new hampshire department of environmental services. 2010. purple loosestrife: an exotic menace. http://des.nh.gov/organization/commi ssioner/pip/factsheets/bb/documents/ bb-45.pdf. accessed 10 april 2014. oklahoma invasive plant council. n.d. oklahoma’s problem species. http://ok-invasive-plantcouncil.org/species.html, accessed 20 march 2014. oklahoma vascular plants database. n.d. oklahoma biological survey. university of oklahoma, norman. http://www.oklahomaplantdatabase.org. accessed 3 february 2014. penn state extension. 2014. demonstration gardens. http://extension.psu.edu/plants/mastergardener/counties/allegheny/demonstrat ion-gardens. accessed 19 november 2014. pennsylvania department of conservation and natural resources. n.d. invasive plants in pennsylvania purple loosestrife. http://www.dcnr.state.pa.us/cs/groups /public/documents/document/dcnr_0 10234.pdf. accessed 19 november 2014. shierenbeck, k. 2004. japanese honeysuckle (lonicera japonica) as an invasive species: history, ecology, and context. critical reviews in plant sciences 23:391-400. swearingen, j., b. slattery, k. reshetiloff, and s. zwicker. 2010. plant invaders of mid-atlantic natural areas. 4th ed. the national park service, u.s. fish and wildlife service. pg. 69-70. the united states national arboretum. 2008. invasive plants. http://www.usna.usda.gov/gardens/in vasives.html. accessed 25 april 2014. the university of maine. 2001. maine invasive plants. http://umaine.edu/publications/2509e. accessed 1 may 2014. usda, nrcs. 2014. the plants database. http://plants.usda.gov. accessed 26 july 2014. washington state department of ecology. n.d. non-native invasive freshwater plants. http://www.ecy.wa.gov/programs/wq/ plants/weeds/aqua009.html. accessed 1 may 2014. oklahoma native plant record 61 volume 14, december 2014 katherine e. keil and karen r. hickman appendix a fact sheet: purple loosestrife common name: purple loosestrife scientific name: lythrum salicaria country of origin: europe and asia history of introduction: it was brought to north america and canada in the early 1800s from eurasia for ornamental and medicinal purposes. it was also imported accidentally as a contaminant on ship ballasts or as seeds on raw wool and sheep aboard. when the us expanded their road and canal systems, purple loosestrife expanded with these developments and now inhabits every contiguous state in the nation except florida. how it invades: purple loosestrife spreads by seeds, which an adult plant produces about 2.5 million a year. purple loosestrife is also able to spread by re-sprouting from roots and fragments. it is easily transported by animals, waterways, boats, cars, and many other vectors. species description: purple loosestrife is an erect perennial herb that stands typically 3-10 feet tall. it has showy magenta colored flower spikes consisting of 5-7 petals that bloom from july to september. the flower has a yellow-white center that contains nectar and is useful for bee-forage. purple loosestrife has tough stems, which can number as many as 50. its leaves are lance-shaped and heartshaped or rounded at the base with pubescent surfaces. population level traits promoting invasion: purple loosestrife is able to invade native communities successfully because it is able to adapt quickly, produces a large amount of offspring, thrives in a wide variety of wet habitats and conditions, has no natural predators, and spreads rapidly. community and ecosystem level effects of invasion: purple loosestrife is problematic because it outcompetes native vegetation creating monocultures, changes water flow that can cause sediment buildups, alters the nitrogen cycle and the water’s chemistry, grows in irrigation systems which blocks the flow of water, alters wetland structure and thus function, and forms dense stands which reduces native animals habitat and food sources. management: purple loosestrife can be managed through mechanical, chemical, and biological methods. if a small community exists, physically remove the plants and (if possible) burn them. for larger communities, spray with a glyphosate herbicide and/or use the beetle galerucella spp. that feeds on the purple loosestrife. ideal time for removal is in june-september due to plant’s noticeability and lack of seeds. references: 1.) http://des.nh.gov/organization/commissioner/pip/factsheets/bb/documents/bb-45.pdf 2.) http://www.nps.gov/plants/alien/fact/lysa1.htm 3.) http://www.invasiveplants.net/plants/purpleloosestrife.htm 4.) http://dnr.maryland.gov/wildlife/plants_wildlife/purpleloosestrife/index.asp 5.) http://plants.usda.gov/plantguide/pdf/pg_lysa2.pdf 62 oklahoma native plant record volume 14, december 2014 katherine e. keil and karen r. hickman fact sheet: multiflora rose common name: multiflora rose scientific name: rosa multiflora country of origin: japan, korea, and eastern china history of introduction: multiflora rose was introduced to the united states from japan in 1866 as a rootstock for grafted ornamental cultivars. in the 1930s, it was further distributed by the u.s. soil conservation service to control erosion. it also has been promoted as effective habitat for animals, crash barriers and headlight reduction for roadways, and fencing for livestock. multiflora rose has since spread significantly and now encompasses 30 states, including the d.c. area. how it invades: multiflora rose most commonly establishes from fruits that fall close to the original plant, which lead to dense thickets. however, animals that eat the plant can disperse seeds longer distances. a single adult plant can produce 1 million seeds annually. plants can also establish roots where their canes touch the ground. species description: multiflora rose is a perennial thorny shrub that can grow to upwards of 15 feet tall. it has clusters of white or tinted pink flowers consisting of 5 petals that appear in may or june. it is multi-stemmed with long, flexible stems containing re-curved thorns and large, alternate leaves. multiflora rose can sometimes be a climbing vine. population level traits promoting invasion: multiflora rose is able to invade native communities successfully because it has a tolerance for diverse soil conditions, grows aggressively, and produces a lot of offspring. it also has a long-lived seed bank that remains viable for 10-20 years that allows it to invade communities even after it is believed to be eradicated. community and ecosystem level effects of invasion: it forms dense, impenetrable thickets that outcompetes native vegetation for resources, including light. management: mechanical, chemical, and biological methods can be implemented to manage multiflora rose. cutting and hand-pulling can remove plants, but one must ensure that all roots are removed in order to be successful. frequent cuttings of 3-6 times a growing season may be necessary. glyphosate herbicides can be sprayed on the foliage or applied to stumps and is ideally used during the dormant season to minimize effect on native plants. rose-rosette disease, a virus, is transported by mites and has fatal effects on multiflora rose. it can kill plants in two years, but must be used with caution so that it does not also wipe out native plants. goats and other grazers can also aid in the control of multiflora rose. fire regimes can prevent plant establishment as well. references: 1.) http://www.nyis.info/index.php?action=invasive_detail&id=33 2.) http://www.nps.gov/plants/alien/fact/romu1.htm 3.) http://mdc.mo.gov/your-property/problem-plants-and-animals/invasive-plants/multiflora-rose-control oklahoma native plant record 63 volume 14, december 2014 katherine e. keil and karen r. hickman fact sheet: japanese honeysuckle common name: japanese honeysuckle scientific name: lonicera japonica country of origin: japan and korea history of introduction: japanese honeysuckle was introduced to long island, new york from japan in 1806 for ornamental and ground cover purposes. it was slow to spread, but once it escaped new york it took over the majority of the united states by the early 1900s. it has since been used for erosion control and wildlife forage and cover. how it invades: japanese honeysuckle invades ecosystems through a series of long runners that develop roots and underground rhizomes. their seeds can also be transported by birds and other wildlife that consume the berries. species description: japanese honeysuckle is a perennial woody vine that often remains evergreen. its white flowers contain 5 petals and bloom from april to october, turning yellow with age. these flowers occur in pairs at leaf junctures and are highly fragrant. japanese honeysuckle is notorious for twisting around objects, specifically stems and trunks. small, black fruits form in august. population level traits promoting invasion: japanese honeysuckle is able to invade native communities successfully because it has few natural enemies in north america, is tolerant to a wide range of environmental conditions, spreads by sending out vegetative runners that can root in a plethora of environments, forms dense thickets, and has a high growth rate. it also has a large seed bank, which can remain viable in the soil for long periods of time. community and ecosystem level effects of invasion: japanese honeysuckle inflicts damage on forest communities because it twines around stems and trunks, establishing dense blankets that block out light, inhibit water flow in native plants, and ultimately smother them. it can prevent growth of native vegetation and decreases the biological diversity of the area. because japanese honeysuckle largely remains evergreen, it remains physiologically active while other vegetation is dormant, allowing it to outcompete native plants. management: prevention is ideal, but both mechanical and chemical management options exist if japanese honeysuckle becomes established. for small communities, hand-pulling at the base of the plant to uproot it and cutting the vines can be successful if monitored regularly to ensure no new seedlings have established. mowing in both july and september can be beneficial for larger patches. glyphosate herbicides can be applied in autumn when the plant has healthy, green leaves, but should be carefully applied according to labels. burning can eliminate the ground cover, but since japanese honeysuckle contains underground rhizomes, prescribed burns remain a temporary solution. finally, animals such as goats have been successful in eating this invasive plant and preventing further spread of it. a combination of the practices listed above will be most effective. references: 1.) http://www.nps.gov/plants/alien/fact/loja1.htm 2.)error! hyperlink reference not valid. http://mdc.mo.gov/your-property/problem-plants-andanimals/invasive-plants/japanese-honeysuckle-control 3.) http://www.in.gov/dnr/files/japanese_honeysuckle.pdf 4.) http://plants.ifas.ufl.edu/node/239 64 oklahoma native plant record volume 14, december 2014 katherine e. keil and karen r. hickman appendix b wanted poster fact sheet: purple loosestrife lythrum salicaria – alias: purple loosestrife description: 3-10 feet tall perennial herb magenta colored flower spikes consisting of 5-7 petals from july to september, often has multiple tough stems, leaves are lance-shaped and heart-shaped or rounded at the base sightings: approximately 6 counties in oklahoma hometown: europe crime: suppression of native vegetation leading to loss of biological diversity, alteration of n cycle and flow of water, disruption of natural wetland function, elimination of food sources for animals instructions for “capture”: if small community, manually remove plant and (if possible) burn it. for larger communities, spray with a glyphosate herbicide or use the beetle galerucella spp. ideal time for removal is in june-august due to plant’s noticeability victims: unsuspecting wetlands or waterways habitat: wetland conditions (i.e., marshes, river banks, ditches, pond edges, roadsides, reservoirs, and wet meadows) citations: http://www.nps.gov/plants/alien/fact/pdf/lysa1.pdf http://www.dnr.state.mn.us/invasives/aquaticplants/purpleloosestrife/control.html http://www.sleloinvasives.org/about-invasives/target-species/purple-loosestrife/ oklahoma classification: declared aquatic nuisance species arrested under  29 o.s. section 6‐601 as of 2014 how you can help: prevent it, recognize it, report it, and remove it oklahoma native plant record 65 volume 14, december 2014 katherine e. keil and karen r. hickman wanted poster fact sheet: multiflora rose 66 oklahoma native plant record volume 14, december 2014 katherine e. keil and karen r. hickman wanted poster fact sheet: japanese honeysuckle lonicera japonica– alias: japanese honeysuckle description: perennial woody vine that twines around objects, flowers are fragrant with 5 white petals that occur in pairs and bloom from april to october, petals turn yellow with age, black fruits form in august, leaves are oblong or oval sightings: approximately 46 counties in oklahoma hometown: eastern asia crime: suppression of native vegetation by forming dense blankets, alteration of forest structure, encircling of trees and stems which cuts off water flow to plant instructions for “capture”: for small communities, hand-pulling the entire plant and mowing can be effective. for larger communities, applying a glyphosate herbicide when green leaves are present is recommended. victims: unsuspecting native herbs and shrubs habitat: open natural communities, but can thrive in a wide range of environmental conditions (i.e., successional fields, old home sites, forests) citations: http://plants.ifas.ufl.edu/parks/japanese_honeysuckle.html http://www.nps.gov/plants/alien/fact/loja1.htm http://www.cnseed.org/japanese-honeysuckle-seeds-lonicera-japonica-seeds.html oklahoma classification: no current legal status mugshot 2014 how you can help: prevent it, recognize it, report it, and remove it mapping distribution in oklahoma and raising awareness: purple loosestrife (lythrum salicaria), multiflora rose (rosa multiflora), and japanese honeysuckle (lonicera japonica) by ms. katherine e. keil and dr. karen r. hickman journal of the oklahoma native plantsociety, volume 3, number 1, december 2003 oklahoma native plant record volume 3, number 1, december 2003 4 black mesa flora study james k. mcpherson, ph.d. department of botany oklahoma state university 22 february 1993 summary of season’s work the following constitutes a report on field, laboratory, and library work done in 1992 on the flora of the state parks-the nature conservancy preserve property at black mesa. this property is north of the town of kenton; r1e, t6n, sections 28-33 (portions), and r1e, t5n, s6 (portion), cimarron county, oklahoma. i spent 14 full days collecting plants on the preserve, each time camping at the state park a few miles away the nights before and after, so very little travel time was used on collecting days. collecting dates in the 1992 growing season were 2-3 march, 6-7 april, 30 april-1 may, 14-16 may, 26 june, 2-3 september, and 21-22 september. during each trip an effort was made to visit and collect in as many different types of sites as possible. collections of 199 species were made. these were handled in the conventional way, with duplicate specimens being made. one set is deposited in the oklahoma state university herbarium, and the other in the bebb herbarium at the university of oklahoma. interpretation of findings flora. the families compositae, leguminosae, and gramineae are represented by the largest numbers of species. however, 47 other families are present. members of the gramineae mcpherson, j.k. https://doi.org/10.22488/okstate.17.100018 (grass) family clearly dominate most of the landscape. the pinaceae (in the inclusive sense) is the other dominant family, due to the numerous members of the genus juniperus in some areas. two species that are endemic were collected. the shrub glossopetalon planitierum (=forsellesia p.), celastraceae, which is known only from a few adjacent counties in the texas panhandle, one nearby county in new mexico and the black mesa area of cimarron county, ok. the type locality is “near the top of black mesa, cimarron co.” it is possible that the type locality is now on the preserve, though it probably is not possible to know with certainty. the other endemic collected was the perennial herb astragalus puniceus, leguminosae. it is known only from the mesa de maya area (las animas county, colorado; union county, new mexico; and cimarron county, oklahoma) and deaf smith county, texas. both species are fairly common locally, but can be considered rare in a general sense. four other species are worth mentioning in this context. i did not collect them, but know about them from the literature (rogers, 1953; harrington 1964; waterfall 1969; mcgregor et al. 1977; mcgregor et al. 1986, correll and johnston 1970). sarcostemma lobata, asclepiadaceae, is apparently known only from black mesa. it is likely that this species will be found on the preserve, and oklahoma native plant record 5 volume 3, number 1, december 2003 mcpherson, j.k. seems to be a legitimate rare species. lesquerella calcicola, cruciferae, palafoxia macrolepis, compositae, and swertia coloradensis, gentianaceae, are all endemic in southeastern colorado, but are at higher elevations and/or on soil types that are not found in oklahoma, so probably are not on the preserve. finally, pericome glandulosa, compositae, was collected and is described by rogers (1953) as being an endemic, but has been reduced to varietal status by harrington. thus it is now pericome caudata var. glandulosa. the reduction appears legitimate. the type locality for it is also black mesa. in my opinion, var. glandulosa is only a local variant of a widespread species. it occurs on sandstone hills which are common in the region and there does not seem to be any substantial distinct feature about it. concern about it is probably not justified. i collected 199 species. rogers’ (1953) list contains 578 species and 11 varieties, a total of 589 taxa. there are some caveats to be mentioned about the comparison of numbers, however. first, rogers collected from a much larger area. second, he included types of sites that are not on the preserve (elevations up to 6850 ft., cimarron river bed and floodplain, sand dunes, and a salt-pan). finally, some of his species seem questionable in view of present knowledge. the following is a list of species i collected that rogers (1953) did not. identifications will be rechecked. selaginellaceae selaginella underwoodii [1] polypodiacae cheilanthes lanosa asplenium serpentrionale[1] gramineae bromus unioloides eragrostis trichodes var. trichodes [1] cyperaceae scirpus validus (s. lacustris in waterfall 1969) lemnaceae lemna minor liliaceae allium canadense var. fraseri salicaceae salix interior forma wheeleri s. nigra (possibly rogers’ “salix species”) moraceae morus alba[1] chenopodiaceae suckleya suckleyana ranunculaceae clematis hirsutissima var. scottii[1] cruciferae arabis fendleri saxifragaceae ribes odoratum [1] leguminosae petalostemon tenuifolium linaceae linum rigidum var. rigidum vitaceae parthenocissus quinquefolia (ident. should be checked) vitis vulpina onagraceae oenothera triloba asclepiadaceae asclepias arenaria[1] sarcostemma crispum[1] boraginaceae cryptantha minima labiatae salvia azurea var. grandiflora rubiaceae galium texense compositae ambrosia linearis[1] (tentative) aster fendleri a. leucelene hymenoxys acaulis kuhnia chlorolepis solidago mollis oklahoma native plant record volume 3, number 1, december 2003 mcpherson, j.k. 6 most of these species are permanent resident, “climax” types. they probably would not have immigrated into the area since rogers made his collections in the late 1940’s. the most likely explanation is that rogers simply missed seeing them. vegetation. this is not a formal study of the vegetation or plant communities of the preserve, but i made observations on these attributes of the site on which i can report. two vegetation types, in the conventional sense of barbour and billings, 1988, are present on the preserve. these are juniper-pinyon woodland, which is on the steeper slopes of the mesa and rock outcrops, and shortgrass prairie, on level to gently sloping sites with deeper soil. within this general picture are some smaller-scale patterns. the most obvious is the presence of cooper’s arroyo, a stream with rare-intermittent flow. it does have a pool that contains water most of the time, and its bed provides conditions that support typical moist-soil plant species such as salix spp., tamarix gallica, and carex gravida. this can be termed a riparian community. two variants of shortgrass prairie are present. on the berthoud loam and portions of the travessilla stony loam (usda, 1960) in the low-lying parts of the preserve is a prairie with many weeds, especially erioneuron pilosum, bothriochloa sacchariodes, and ambrosia psilostachya. there is also a substantial amount of the cactus opuntia imbricata which here is associated with disturbance. this portion of the preserve was the most accessible to cattle when the land was ranched, and was where most of the water was provided. it appears that overgrazing is the main cause of the abundance of weedy species and partial loss of the dominants, buchloe dactyloides and bouteloua gracilis. on the apache stony clay loam (usda, 1960), which is found only on the basalt rock forming the top of the mesa, is a slightly different version of shortgrass prairie. the dominant grasses, buchloe dactyloides and bouteloua gracilis, are the same, but they are more dominant and there are fewer weeds. more of the native forbs such as castilleja sessiliflora, oenothera lavendulaeflora, and several compositae are present. in my judgment, the difference is caused by a history of less disturbance, and by the soil’s higher clay content. the contrast between the two variants of short grass prairie will probably diminish with time and the cessation of grazing, but differences due to the contrasting soils are likely to remain. the mesa-top community probably will have a higher diversity of climax species. on the sides of the mesa the soils are mapped as rough stony land and the higher parts of the travessilla stony loam (usda, 1960). this is where the juniperpinyon woodland is found. juniperus monosperma is the strong dominant here, with only a few pinus edulis trees, despite the traditional name of the vegetation type. there are differing communities within this area, but they are not as clearly separated as is the case with the prairie communities. the most noteworthy group of species here, after j. monosperma, is the shrubs. on the drier, open slopes are rhus aromatica, cercocarpus montanus, brickellia brachyphylla, and b. californica. also, opuntia imbricata is here, appearing less weedy than it does in the prairies. in one area near the east end of the preserve the endemic glossopetalon planitierrum is a component of the shrub flora. all are fairly widely spaced so that walking among them is easy. oklahoma native plant record 7 volume 3, number 1, december 2003 mcpherson, j.k. in the canyons where more moisture accumulates and there is some shelter from the wind is a denser shrub community. near the bottoms of the deeper canyons it is dense indeed, becoming impenetrable in places. most of the species just listed are present, and they are joined by prunus americana, p. virginiana, rubus deliciosus, ptelea trifoliata, and celtis reticulata. here also is juniperus scopulorum, a rocky mountain species, which is quite uncommon and is very close to the extreme edge of its range. throughout the juniper-pinyon vegetation is an array of grasses, mostly of different species from the prairie. very common are poa fendleriana and eragrostis cilianensis. in pockets of deep soil, often only a meter or two across, are andropogon gerardii, sorghastrum nutans, and schizachyrium scoparium. these are dominants of the tallgrass prairie 150 and more miles east, but grow well here in small, favorable sites. the juniper-pinyon woodlands are the least disturbed communities on the preserve. the only other local community that should be noted is the very weedy one that develops in and around the usuallydry, man-made “tanks” or stock-watering ponds. these ponds contain water so seldom that its main effect is to drown any climax species that invade the bed. the original construction work left a massive scar, and trampling by cattle has perpetuated the disturbance. species commonly found in and around the ponds include proboscidea louisianica, xanthium strumarium, cenchrus pauciflorus, and suckleya suckleyana. if left alone, without cattle trampling, the dams and margins of these ponds will slowly revert to shortgrass prairie. the beds will be weedy as long as the dams occasionally retain water. references cited barbour, m.c. and w.d. billings (eds.). 1988. north american terrestrial vegetation. cambridge, ma: cambridge univ. press. correll, d.s. and m.c. johnston. 1970. manual of the vascular plants of texas. renner, tx: texas research foundation. harrington, h.d. 1964. manual of the plants of colorado. athens, oh: sage books/swallow press. mcgregor, r.l. et al. 1977. atlas of the flora of the great plains. ames, ia: iowa state university press. mcgregor, r.l. et al. 1986. flora of the great plains. lawrence, ks: university press of kansas. rogers, c.m. 1953. the vegetation of the mesa de maya region of colorado, new mexico, and oklahoma. lloydia:257-290. united states department of agriculture. soil conservation service; soil survey: cimarron county, oklahoma. u.s. government printing office. 1960. waterfall, u.t. 1969. keys to the flora of oklahoma. 4th ed. stillwater, ok: [published by the author] oklahoma native plant record volume 3, number 1, december 2003 mcpherson, j.k. 8 black mesa flora study year two supplement james k. mcpherson 20 january 1994 introduction this is a supplement to my report on the same subject of last year. it is assumed that the present readers have that report and can refer to it. this paper is organized the same way and is in the same sequence as last year’s. summary of 1993 work i spent seven full days collecting, using the same plans & format as in 1992. the dates were; 25-26 april, 9-10 may, 31 may, and 6-7 october. collections of 30 species new for this project were made, bringing the total to date to 229. they were handled and distributed as before. interpretation of findings the count of families has risen to 53 from 50, because of collection of single members of the selaginellaceae, sapindaceae, and polemoniaceae. two species should be mentioned. (1) the parthenocissus at the mesa may be p. vitacea, the “western” species. it is known from a few places in the state, but on most herbarium specimens it cannot be distinguished from p. quinquefolia so it is hard to know how common it is. waterfall did not realize p. vitacea was in okla. (or did not accept it), so most people have assumed that it was all p. quinquefolia. it will be next season before i will know which we have at the mesa. (2) there is an ambrosia there that keys to a. linearis, which is “apparently restricted to a few localities in the open high plains of eastern colorado; rarely collected.” there are no specimens in ou’s or our herbaria, so ron tyrl and i sent it off to university of colorado for identification. we haven’t heard back from them yet. it looks very much like a. psilostachya, which is abundant that area; this may be why it is overlooked. my 1993 estimate of 250-260 species being present on the preserve still seems reasonable. since 229 have been collected, about 20-30 remain to be found. oklahoma native plant record 9 volume 3, number 1, december 2003 mcpherson, j.k. itors otes this paper is published with the courteous agreement of the nature conservancy for whom it was prepared. the approximate ps location of lack mesa state park is between latitudes 36.833 and 36.861 and longitudes 102.862 and 102.900. the elevation of the mesa ranges from 960 ft (1512 m) to 973 ft (1516 m). it is now contained within lack mesa state park which contains approximately 3 9 acres of land. the original species list has been updated as follows [1] on july 1, 199 , ten days before his death, jim mcpherson generated plant labels for 15 additional specimens he had collected on june 7 at lack mesa on his way to california. with the generous assistance of iris mcpherson, his wife, they are included in the flora and the taxa summary table below. amilies 55 enera 172 species 2 infraspecific taxa 1 xotic species 16 olley’s “additions to lack mesa lora study”, which follows mcpherson’s flora in this volume, includes areas of lack mesa state park not included in his study and lists only species that are not included here. [2] the international code of otanical nomenclature “conserved” several traditional family names when they standardized the family nomenclature. mcpherson used some of these traditional names in the lack mesa report, but since they are falling into disuse standardized names are provided here. current species’ names have also been provided. name changes are updates only. no specimens were reexamined for this publication. kartesz, j.r. (199 ). a synonymized checklist of the vascular flora of the united states, canada, and reenland. portland, or timber press. voss, . ., .m. urdet, w. . chaloner, v. demoulin, p. iepko, j. mcneill, r.d. meikle, d. . nicolson, r.c. rollins, p.c. silva, & w. reuter, 1983. international code of botanical nomenclature, adopted by the thirteenth international otanical congress, sydney, august 1981). [3] introduced species are indicated in this list. correll & johnston. 1970. manual of the vascular plants of texas. renner, t texas research oundation. taylor, r.j. & c. .s. taylor. 1991. an annotated list of the ferns, fern allies, gymnosperms and flowering plants of oklahoma. published by the authors at southeastern oklahoma state university . tyrl, r.j., susan arber, paul uck, wayne lisens, james stes, patricia olley, awrence magrath, constance taylor, and rahmona thompson. the flora of oklahoma. the lora of oklahoma ditorial oard. orthcoming. usda-nrcs 2003. the p ants database. (http plants.usda.gov plants.) oklahoma native plant record volume 3, number 1, december 2003 10 species by family of the black mesa reserve, cimarron county james k. mcpherson, 1992 (93) = species added in 1993 (94) = species added in 1994[1]) standardized name [2] pteridaceae dryopteridaceae aspleniaceae poaceae division/class/family selaginellaceae selaginella underwoodii (93) polypodiaceae cheilanthes eatoni cheilanthes feei (93) cheilanthes lanosa notholena standleyi pellaea atropurpurea var. purpurea (93) woodsia oregan (94) asplenium septentrionale (94) pinaceae juniperus monosperma juniperus scopulorum pinus edulis gramineae agropyron smithii var. smithii andropogon gerardii aristida longiseta aristida purpurea aristida wrightii bothriochloa saccharoides bouteloua curtipendula bouteloua eriopoda bouteloua gracilis bouteloua hirsuta var. hirsutea bromus anomalus var. lanatipes bromus tectorum bromus unioloides buchloe dactyloides cenchrus carolinianus chloris verticillata echinochloa cruzgalli elymus virginicus var. jejunus elymus canadensis (94) eragrostis cilianensis mcpherson, j.k. common family name spikemoss family spikemoss true fern family eaton's lip fern slender lip fern hairy lip fern star cloak-fern cliff-brake oregon woodsia forked spleenwort pine family one-seed juniper rocky mtn. juniper pinyon pine grass family western wheatgrass[3] big bluestem fendler three-awn purple three-awn wright three-awn silver bluestem side-oats grama black grama blue grama hairy grama nodding brome cheat[3] rescue grass[3] buffalo grass sandbur windmill grass barnyard grass[3] virginia wildrye canadian wild rye stinkgrass[3] 11 mcpherson, j.k. elymus elymoides eragrostis trichodes var. var. trichodes (94) erioneuron pilosum hilaria jamesii hordeum pusillum lycurus phleoides muhlenbergia torreyi oryzopsis hymenoides oryzopsis micrantha panicum capillare var. capillare panicum hallii (93) panicum obtusum poa fendleriana schedonnardus paniculatus schizachyrium scoparium setaria leucopila sitanion hystrix sorghastrum nutans sporobolus cryptandrus stipa comata stipa scribneri vulpia octoflora cyperaceae carex gravida cyperus schweinitzii (93) scirpus americanus var. polphyllus scirpus validus scirpus tabernaemontanus salix exigua commelinaceae commelina erecta var. angustifolia (94) tradescantia occidentalis lemnaceae lemna minor liliaceae allium canadense var. fraseri yucca glauca salicaceae populus deltoids salix amygdaloides salix interior forma wheeleri salix nigra sand love grass fluffgrass[3] galleta little barley wolftail ring muhly indian ricegrass little-seed ricegrass common witchgrass hall panic grass vine-mesquite muttongrass tumblegrass little bluestem plains bristlegrass squirreltail indian grass sand dropseed thread-and-needle scribner needlegrass six-weeks fescue sedge family sedge umbrella sedge bulrush bulrush spiderwort family erect dayflower western spiderwort duckweed family duckweed lily family wild onion plains yucca willow family cottonwood peach-leaf willow sandbar willow black willow oklahoma native plant record volume 3, number 1, december 2003 oklahoma native plant record volume 3, number 1, december 2003 mcpherson, j.k. 12 ulmaceae elm family celtis reticulate hackberry santalaceae sandalwood family commandra pallida bastard toad-flax commandra umbellata ssp. pallida urticaceae nettle family parietaria pennsylvanica pennsylvania pellitory polygonaceae buckwheat family eriogonum jamesii james wild buckwheat eriogonum lachnogynum wild buckwheat polygonum lapathifolium pale smartweed polygonum ramosissimum knotweed rumex crispus curly dock chenopodiaceae goosefoot family ceratoides lanata winterfat chenopodium album (93) lamb's quarters chenopodium incanum (93) goosefoot kochia scoparia kochia salsola kali var. tenuifolia russian thistle salsola kali var. tragus suckleyla suckleana poison suckleya amaranthaceae pigweed family amaranthus retroflexus rough pigweed[3] nyctaginaccae four-o'clock family mirabilis carletonii (93) carleton’s four-o'clock mirabilis linearis var. subhispida narrowleaf four-o'clock portulacaceae purslane family portulaca retusa purslane portulaca oleracea ssp. oleracea ranunculanceae buttercup family clematis hirsutissima var. scottii(93) virgin's bower delphinium virescens var. penardi prairie larkspur delphinium carolinianum var. virscens (93) ranunculus sceleratus cursed crowfoot fumariaceae fumitory family corydalis aurea golden corydalis capparidaceae caper family polanisia dodecandra clammy-weed oklahoma native plant record 13 volume 3, number 1, december 2003 mcpherson, j.k. cruciferae mustard family brassicaceae arabis fendleri rock cress descurania pinnata tansy mustard var. intermedia erysimum capitatum wallflower lepidium densiflorum peppergrass[3] lesquerella ovalifolia bladderpod saxifragaceae saxifrage family grossulariaceae ribes cereum western red currant ribes odoratum buffalo currant ribes aureum var. villosum (93) rosaceae rose family cercocarpus montanus var. argenteus mountain mahogany physocarpus monogynus (93) mountain ninebark prunus americana var. americana wild plum prunus virginiana choke cherry rubus deliciosus boulder raspberry leguminosae pea family fabaceae amorpha canescens lead plant forma canescens (94) astragalus crassicarpus ground-plum var. paysoni (93) astragalus gracilis slender milk-vetch astragalus lotiflorus lotus milk-vetch astragalus missouriensis missouri milk-vetch astragalus mollissimus wooly locoweed astragalus puniceus trinidad milk-vetch dalea aurea golden prairie-clover dalea candida white prairie-clover var. oligophylla dalea enneandra nine-anther prairie-clover dalea formosa (93) feather plume dalea jamesii james dalea glycyrrhiza lepidota (93) wild licorice[3] hoffmannseggia drepanocarpa (93) sicklepod rush-pea caesalpinia drepanocarpa hoffmannseggia jamesii james rush-pea caesalpinia jamesii krameria lanceolata ratany krameriaceae melilotus officinalis yellow sweet clover[3] mimosa borealis pink mimosa petalostemum tenuifolia slimleaf prairie-clover dalea tenuifolia psoralea argophylla (93) silver-leaf scurf pea pediomelum argophylla psoralea tenuiflorum scurf pea psoralidium tenuiflorum vicia americana american vetch oklahoma native plant record volume 3, number 1, december 2003 mcpherson, j.k. 14 linaceae flax family linum lewisii blue flax linum rigidum var. rigidum stiff flax zygophyllaceae caltrop family tribulus terrestris goat head[3] rutaceae citrus family ptelea trifoliata wafer-ash polygalaceae milkwort family polygala alba milkwort euphorbiaceae spurge family argythamnia humilis wild mercury argythamnia mercurialina wild mercury croton texensis texas croton euphorbia fendleri fendler spurge chamaesyce fendleri euphorbia lata hoary spurge chamaesyce lata euphorbia dentata toothed spurge forma cuphosperma euphorbia marginata snow-on-the-mountain tragia ramosa noseburn anacardiaceae sumac family rhus aromatica var. pilosissima lemon sumac toxicodendron radicans poison ivy celastraceae staff-tree family crossosomataceae glossopetalon planitierum grease-bush sapindaceae soap-berry family sapindus drummondii (93) soap-berry sapindus saponaria var. drummondii vitaceae grape family parthenocissus quinquefolia virginia creeper vitis vulpina fox grape vitis riparia malvaceae mallow family sphaeralcea angustifolia globe mallow sphaeralcea coccinea scarlet globe mallow tamaricaceae tamarisk family tamarix gallica salt cedar[3] violaceae violet family hybanthus verticillatus green violet oklahoma native plant record 15 volume 3, number 1, december 2003 mcpherson, j.k. loasaceae stick-leaf family mentzelia decapetala blazing star cactaceae cactus family echinocereus viridiflorus green-flowered hedgehog mammillaria vivipara (93) pincushion cactus escobaria vivipara var. vivipara opuntia imbricata cholla opuntia phaeacantha var. major prickly pear opuntia trichophora (93) prickly pear opuntia polyacantha var. trichophora onagraceae evening primrose family gaura coccinea var. coccinea scarlet butterfly flower oenothera serrulata evening primrose calyophus serrulatus oenothera albicaulis (93) evening primrose oenothera lavendulaefolia evening primrose calyophus lavandulifolius oenothera triloba stemless evening primrose umbelliferae parsley family apiaceae cymopteris acaulis (93) (no common name) cymopteris montanus (no common name) asclepiadaceae milkweed family asclepias arenaria (94) sand milkweed asclepias asperula low milkweed var. decumbens asclepias macrotis (94) longhood milkweed asclepias pumila threadleaf milkweed asclepias uncialis (93) dwarf milkweed sarcostemma crispum (94) convolvulaceae morning glory family convolvulus incanus field bindweed[3] convolvulus arvensis evolvulus nuttallianus nuttall evolvulus ipomoea leptophylla (94) bush morning-glory polemoniaceae phlox family gilia laxiflora (93) gilia ipomopis laxiflora boraginaceae borage family cryptantha jamesii popcorn flower cryptantha cineria var. jamesii cryptantha minima small popcorn flower cryptantha thyrsiflora popcorn flower oklahoma native plant record volume 3, number 1, december 2003 mcpherson, j.k. 16 lappula redowskii stickseed lappula occidentalis var. occidentalis var. occidentalis lithospermum incisum cutleaf puccoon onosmodium molle var. occidentale false gromwell verbenaceae vervain family verbena canadensis rose vervain glandularia canadensis verbena bracteata prostrate vervain labiatae mint family lamiaceae monarda pectinata spotted beebalm salvia azurea var. grandiflora pitcher sage solanaceae nightshade family chamaesaracha conioides false nightshade physalis virginiana virginia ground cherry var. sonorae (94) physalis lobata ground cherry quincula lobata solanum elaeagnifolium silverleaf nightshade solanum rostratum (93) buffalo bur scrophulariaceae figwort family castilleja sessiliflora downy indianpaintbrush penstemon albidus white beardtongue penstemon ambiguous (94) veronica anagallis-aquatica water speedwell[3] martyniaceae unicorn-plant family proboscidea louisianica devil's claw plantaginaceae plantain family plantago purshii var. purshii wooly plantain plantago purshii var. spinulosa (93) wooly plantain rubiaceae madder family galium texense texas bedstraw cucurbitacaeae cucumber family cucurbita foetidissima buffalo gourd compositae sunflower family asteraceae agoseris cuspidate false dandelion nothocalais cuspidata ambrosia sp. (93) ragweed ambrosia psilostachya western ragweed artemisia filifolia sandsage artemisia glauca silky wormwood artemisia dracunculus oklahoma native plant record 17 volume 3, number 1, december 2003 mcpherson, j.k. ssp. glauca artemisia ludoviciana louisiana sagewort aster ericoides heather aster aster fendleri fendler's aster aster leucelene white aster chaetoppa ericoides aster oblongifolius aromatic aster berlandiera lyrata green eyes brickellia brachyphylla (no common name) brickellia californica (no common name) chrysopsis villosa var. villosa golden aster heterotheca villosa var. villosa chrysothamnus nauseosus rabbit brush cirsium undulatum wavy-leaf thistle conyza canadensis var. canadensis horseweed dyssodia papposa fetid marigold engelmannia pinnatifida engelmann's daisy erigeron divergens var. cinereus fleabane erigeron colomexicanus evax prolifera rabbit-tobacco gaillardia pinnatifida blanket flower grindelia squarrosa var. nuda curly-top gumweed gutierrezia sarothrae snakeweed haplopappus spinulosus cutleaf ironplant machaeranthera pinnatifida helianthus annuus annual sunflower hymenopappus flavescens yellow plainsman hymenopappus tenuifolius white plainsman hymenoxys acaulis stemless bitterweed tetraneuris acaulis hymenoxys scaposa var. linearis bitterweed tetraneuris scaposa kuhnia chlorolepis false boneset brickellia eupatorioides var. chlorolepis liatris punctata var. punctata dotted gayfeather lygodesmia juncea (94) skeleton plant lygodesmia pauciflora skeletonweed stephanomeria pauciflora machaeranthera tanacetifolia (93) tansy aster melampodium leucanthemum black-foot daisy pericome caudate (no common name) ratibida columnifera mexican hat ratibida tagetes (94) prairie coneflower senecio douglasii var. longilobus shrub groundsel senecio flaccidus senecio plattensis prairie ragwort senecio tridenticulatus ragwort solidago mollis soft goldenrod solidago petiolaris (93) downy goldenrod thelesperma megapotamicum greenthread townsendia exscapa easter daisy oklahoma native plant record volume 3, number 1, december 2003 mcpherson, j.k. 18 tragopogon major (93) goatbeard[3] tragopogon dubius verbesina encelioides golden crownbeard xanthium strumarium cocklebur zinnia grandiflora wild zinnia moraceae morus alba (94) white mulberry[3] oklahoma native plant record, volume 13, number 1, december 2013 1 oklahoma n ative plant record journal of the oklahoma native plant society p. o. box 14274 tulsa, okla homa 74159-1274 volume 13, december 2013 issn 1536-7738 http://ojs.library.okstate.edu/osu/ managing editor: sheila strawn production editor: paula shryock electronic production editor: sandy graue technical advisors: amy buthod, kristi rice the purpose of onps is to encourage the study, protection, propagation, appreciation, and use of the native plants of oklahoma. membership in onps is open to any person who supports the aims of the society. onps offers individual, student, family, and life memberships. 2013 officers and board members president: adam ryburn vice-president: joe roberts secretary: sandy graue treasurer: mary korthase membership coordinator: tina julich historian: vacant past president: lynn michael board members: brooke bonner pearl garrison elaine lynch bruce smith janette steets jay walker chapter chairs: central: joe roberts cross timbers: mark fishbein mycology: steve marek northeast: alicia nelson southwest: carrie reed color oklahoma chair: pearl garrison conservation chair: chadwick cox gaillardia editor: chadwick cox website manager: adam ryburn http://www.oknativeplants.org award chairs: harriet barclay: rahmona thompson anne long: gloria caddell service: sue amstutz photography contest chair: lynn michael librarian: karen haworth mailings chair: karen haworth publicity chair: alicia nelson cover photo: aesclepias speciosa by leslie cole, onps photo contest winner . articles (c) the authors journal compilation (c) oklahoma native plant society except where otherwise noted, this work is licensed under a creative commons attributionnoncommercial-sharealike4.0 international license, https://creativecommons.org/licenses/ by-nc-sa/4.0/, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly attributed, not used for commercial purposes, and, if transformed, the resulting work is redistributed under the same or similar license to this one. https://doi.org/10.22488/okstate.17.100094 http://ojs.library.okstate.edu/osu/ http://www.oknativeplants.org/ oklahoma native plant record volume 13, december 2013 2 oklahoma native plant record volume 13 table of contents foreword ................................................................................................................................................. 3 ecology and taxonomy of water canyon, canadian county, oklahoma, m.s. thesis .............. 4 dr. constance e. taylor a checklist of the vascular flora of the mary k. oxley nature center, tulsa county, oklahoma ................................................................................................................... 29 ms. amy k. buthod smoke-induced germination in phacelia strictaflora ........................................................................ 48 dr. stanley a. rice and dr. sonya l. ross critic’s choice essay: a calvacade of oklahoma botanists in oklahoma − contributors to our knowledge of the flora of oklahoma ........................................................ 55 dr. ronald j. tyrl and ms. paula shryock editorial policies and procedures ................................................................................................... 101 five year index to oklahoma native plant record ........................................... inside back cover oklahoma native plant record, journal of the oklahoma native plant society, volume 13, december 2013 title page table of contents foreword oklahoma native plant record, volume 11, number 1, december 2011 oklahoma native plant record volume 11, december 2011 parkhurst, m. j., et al. https://doi.org/10.22488/okstate.17.100083 33 spatial genetic structure of the tallgrass prairie grass dich an t h elium oligosanth es (scribner’s panicum) molly j. parkhurst1 1oklahoma state university andrew doust1 department of botany margarita mauro-herrera1 301 physical sciences jeffrey m. byrnes2 stillwater, ok 74078 janette a. steets1 janette.steets@okstate.edu 2oklahoma state university boone pickens school of geology 105 noble research center stillwater, ok 74078 keywords: m ixed-mating system, population differentiation, population g enetic structure abstract the spatial genetic structure within plant populations and genetic differentiation among populations can vary in strength due to the forces of natural selection, gene flow and genetic drift. in this study, we investigate the level of genetic structure and differentiation present in oklahoma populations of dichanthelium oligosanthes (schult.) gould (scribner’s panicum), a c3 grass native to the united states and a frequent member of the tallgrass prairie. to examine fine-scale spatial genetic structure of d. oligosanthes, we collected leaves from 48 spatially separated plants in a population in stillwater, ok. to examine genetic differentiation among adjacent populations, we sampled leaf tissue from eight individuals at each of three populations in stillwater, ok. dna was extracted from these samples and inter-simple sequence repeats (issr) markers were amplified. within a single population of d. oligosanthes, we found a weak and non-significant negative relationship between genetic similarity and geographical distance. in contrast, we found evidence for moderate and significant genetic differentiation among populations. introduction genetic variation is the sum total of all genetically based variation within and among species and represents an important component of biodiversity. maintaining genetic variation within and among native plant populations is a central goal of conservation biology, as genetic variation provides the raw material for plants to evolve in response to environmental change and contributes to population fitness (leimu et al. 2006, reed and frankham 2003, wagner et al. 2011). given the importance of genetic variation for the maintenance and evolution of plant populations, it is critical to understand the levels of genetic diversity within species and determine how this variation is organized spatially, both within and between populations. the way in which genetic variation is organized within plant populations (spatial genetic structure) is affected by many factors, including selection pressures within a population, mating system (relative production of selfed to outcrossed individuals), and whether gene flow is restricted (loiselle et al. 1995, loveless and hamrick 1984, miyazaki and isagi 2000, perry and knowles 1991). plants are sessile organisms and gene flow can only occur through pollen and seed movement. restricted dispersal of seed or pollen can oklahoma native plant record volume 11, december 2011 parkhurst, m. j., et al. 34 occur due to the dispersal mechanism utilized by the plant, the presence of physical barriers, and when rates of inbreeding are high. under these conditions, population genetic structure is predicted to develop, with a clustering of genetically related individuals among plants within a population and a high level of genetic differentiation among geographically separated populations (epperson and li 1997, hamrick and nason 1996). dichanthelium oligosanthes is a short, c3 perennial grass native to the united states that is commonly found in open prairies, meadows, and disturbed areas (kansas state university libraries 2011). although it is not a dominant species of the tallgrass prairie, it is a highly consistent member of this community (adams and wallace 1985). the mating system and ecology of d. oligosanthes are likely to affect population genetic structure. first, d. oligosanthes has a mixed mating system resulting from the production of two types of flowers within a single individual: closed, self-fertilizing cleistogamous flowers and open, potentially outcrossing chasmogamous flowers. the chasmogamous flowers form on a terminal panicle in may – june, are open-pollinated for a short time, then close and self-fertilize in the absence of pollination (bell and quinn 1985, freckmann and lelong 2003). the cleistogamous flowers extend from within the sheath and appear from june – november (bell and quinn 1985, freckmann and lelong 2003). grasses in the genus dichanthelium tend to reproduce proportionately more through cleistogamy than through chasmogamy (bell and quinn 1985, bell and quinn 1987). this high rate of self-fertilization should lead to significant spatial genetic structure within populations and genetic differentiation among populations. second, as an element of the tallgrass prairies, d. oligosanthes has a relatively short stature of less than 45 cm. in tallgrass prairies, d. oligosanthes is imbedded within a matrix of the dominant grass species (andropogon gerardii, panicum virgatum, sorgastrum nutans, and schizachyrium scoparium). these dominant grasses can reach heights of up to 2 m (usda, nrcs 2011) and thus may serve as a physical barrier to d. oligosanthes pollen and seed dispersal, further contributing to spatial genetic structure within populations. in contrast to the first two factors, which would tend to increase fine-scale spatial genetic structure within d. oligosanthes populations, one aspect of d. oligosanthes fruit dispersal could lead to reduced genetic structure. in particular, the fruits of d. oligosanthes can disperse great distances when the inflorescence breaks off of the plant, leading to a “tumbleweed” dispersal mechanism (campbell et al. 1983). we examined spatial genetic structure and population differentiation of d. oligosanthes in stillwater, ok. through our study we aimed to answer the following questions: (1) how is genetic diversity distributed spatially within a population of d. oligosanthes? (2) are populations of d. oligosanthes genetically differentiated? to address these questions, we examined genetic diversity in inter-simple sequence repeats (issr )markers in three d. oligosanthes populations. within a single population of d. oligosanthes, we expected genetically related individuals to be aggregated spatially, and thus we expected to find a negative correlation between genetic similarity and geographical distance. among populations of d. oligosanthes, we expected to find a significant degree of genetic differentiation. materials and methods how is genetic diversity distributed spatially within a population of d. olig osanthes? we collected leaf samples from 48 representatives of d. oligosanthes from the southern portion of the oklahoma state university (osu) cross country field south in stillwater, ok (ccs population: 36°08’12.0”n, 97°04’36.6”w; figure 1). thirty-eight of these samples were located across three approximately parallel transects, oklahoma native plant record volume 11, december 2011 parkhurst, m. j., et al. 35 each approximately 36 m in length and five to ten m apart. material from an additional ten plants was collected at increasing distances from these transects. locations of all plants were recorded using a trimble 2008 geoxh handheld gps with an external zephyr antenna and were differentially corrected relative to a stationary base station to increase the accuracy of the gps data. the locations of each of the 48 plant samples were mapped on an aerial photograph of the site (see figure 1). we extracted dna from each individual following the procedure of junghans and metzlaff (1990). for each sample, genomic dna concentrations were quantified using nanodrop spectrophotometry and 1% agarose gel electrophoresis. we tested 12 issr markers (zietkiewicz et al. 1994) from the ubc primer set #9 (university of british columbia nucleic acid-protein service unit) and successfully amplified five of these via polymerase chain reaction (pcr) (table 1). we ran 10 µl pcrs using 1 μl of diluted genomic dna (from a 1:30 dilution corresponding to 10 to 20 ng of genomic dna), dntps at 100 μm each, issr primer at 0.5 µm, 1x green gotaq® flexi buffer (promega, madison, wi), mgcl2 at 1.5 mm, and 0.6 units of promega go taq polymerase (promega, madison, wi). amplifications were performed in an eppendorf mastercycler pro thermal cycler using the following touchdown conditions: single initial denaturation step at 95˚ c for two minutes; followed by 32 cycles each with three steps: a denaturation step at 94˚ c for 40 seconds, followed by an annealing touchdown step (starting at 56˚ c or 58˚ c, depending on the primer, for two cycles; then 54˚ c or 56˚ c for two cycles to reach 52˚ c or 54˚ c for 28 cycles) for 40 seconds, and an extension step at 72˚ c for 50 seconds; and a final extension at 72˚ c for eight minutes. pcr products were resolved electrophoretically on 1% agarose gels run at 150 v in tbe buffer, visualized by staining with ethidium bromide, and photographed under uv illumination. fragment sizes were estimated using hyper ladder ii (bioline, tauton, ma). issr bands were scored as present or absent for each plant sample. bands were scored and compared by two different people to reduce subjectivity in the scoring procedure. twelve individuals showed poor amplification of some of the issr markers, resulting in missing data in the dataset. as the statistical analyses described below do not allow for missing data, we eliminated these individuals from further analysis. thus, the analysis of genetic structure in the ccs population used a total of 36 individuals. geographical distances between plants were estimated using euclidean distances. genetic similarities between plants were estimated using a matrix of dice genetic similarity coefficients created with the past program (hammer et al. 2001). the dice coefficient, which weighs positive matches between plant samples and ignores negative matches, was used because issrs are dominant markers, and therefore only the presence of a pcr product is meaningful. a mantel test was performed to determine whether there was a relationship between geographical distance and genetic similarity with both the past program and the zt program (bonnet and van der peer 2009), using 10,000 permutations of the data. finally, to examine hierarchal clustering in the samples, we generated an unweighted pair group method with arithmetic mean (upgma) tree of the genetic distance data using the past program (hammer et al. 2001). a bootstrap analysis using 1000 replications was performed to determine the support for the hierarchical clustering. are populations of d. olig osanthes genetically differentiated? to determine whether nearby populations of d. oligosanthes in stillwater, ok are genetically differentiated from one another, we examined the diversity of six issr markers (see table 1) in three populations (figure 2): osu cross country field south (ccs; oklahoma native plant record volume 11, december 2011 parkhurst, m. j., et al. 36 36°08’12.0”n, 97°04’36.6”w), osu cross country field north (ccn; 36°08’20.8”n, 97°04’39.9"w), and lakeview road west (lw; 36°08’42.6”n, 97°05’38.9”w). populations ccs and ccn were separated from one another by 0.3 km, ccn and lw by 1.6 km and ccs and lw by 1.8 km. within each of the three populations, we collected leaf material from eight randomly chosen individuals. we extracted dna, amplified issr markers via pcr, and visualized pcr products via gel electrophoresis as described above. we used the program hickory version 1.1 (holsinger and lewis 2003, holsinger et al. 2002) to estimate genetic differentiation among populations of d. oligosanthes. hickory allows for estimation of heterozygosity within populations and genetic differentiation among populations using dominant markers without assuming hardy-weinberg equilibrium (holsinger and lewis 2003, holsinger et al. 2002). this program uses bayesian methods to estimate the average heterozygosity within subpopulations (hs; an analog of the expected heterozygosity) and test for genetic differentiation among populations through the unbiased estimate θii, which is analogous to weir and cockerham’s (1984) fst (holsinger and lewis 2003). θii measures the amount of genetic differentiation among contemporaneous populations (holsinger and lewis 2003); values close to 0 suggest little genetic differentiation between populations (i.e., complete panmixia), whereas values close to one suggest genetic isolation between populations. the d. oligosanthes issr data were fitted to four models: (1) full (uses non-informative priors for f ), (2) f = 0 (assumes no inbreeding; f is analogous to the inbreeding coefficient fis), (3) θ ii = 0 (assumes no population differentiation), and (4) f-free (decouples the estimation of θ ii from the estimation of f ). in each model run, the default parameters were used (burn-in = 5,000; number of samples = 25,000; thinning = 5). the models were compared to one another based on the deviance information criterion (dic) of spiegelhalter et al. (2002) implemented in hickory. the full model best fit our dataset as it provided the lowest dic value; thus, we present the results of the full model in the results section. results how is genetic diversity distributed spatially within a population of d. olig osa nthes? the five issr primers produced a total of 21 loci that could be reliably scored in the ccs population, of which 13 were polymorphic (table 2). the mantel test revealed a non-significant negative relationship between genetic similarity and geographic distances, with a negative correlation between geographical distance and similarity (r = -0.042, p = 0.755). the upgma analysis revealed weak hierarchical clustering among the plant samples, consistent with the results of the mantel test (data not shown). thus, we find no evidence for significant spatial genetic structure in the ccs population of d. oligosanthes. oklahoma native plant record volume 11, december 2011 parkhurst, m. j., et al. 37 figure 1 map of the oklahoma state university cross country field south (ccs) dichanthelium oligosanthes population in stillwater, ok. yellow points indicate the location of d. oligosanthes plants sampled; base image is an aerial photograph acquired by the usda farm service agency. oklahoma native plant record volume 11, december 2011 parkhurst, m. j., et al. 38 figure 2 map of dichanthelium oligosanthes populations in stillwater, ok; base image from google earth. ccs = oklahoma state university (osu) cross country field south, ccn = osu cross country field north, lw = lakeview road west. table 1 issr primers used for dna amplification from ubc primer set #9 (university of british columbia nucleic acid-protein service unit). the right-most column indicates whether the given primer was used in the study of genetic diversity in the oklahoma state university cross country field south population of d. oligosanthes (a) and/or the study of genetic differentiation among three stillwater, ok populations of d. oligosanthes (b). primer sequence (5' 3') study using primer ubc – 808 aga gag aga gag aga gc b ubc – 809 aga gag aga gag aga gg a and b ubc – 810 gag aga gag aga gag at a and b ubc – 816 cac aca cac aca cac at a and b ubc – 817 cac aca cac aca cac aa a and b ubc – 818 cac aca cac aca cac ag a and b oklahoma native plant record volume 11, december 2011 parkhurst, m. j., et al. 39 table 2 numbers of loci reliably scored, numbers of polymorphic loci, and percentage of polymorphic loci for each of the five issr primers used in the study of genetic diversity in the oklahoma state university cross country field south (ccs) population of d. oligosanthes. primer number of loci reliably scored1 number of reliably scored polymorphic loci percentage polymorphic loci2 ubc – 809 6 3 50.0 ubc – 810 4 2 50.0 ubc – 816 5 2 40.0 ubc – 817 3 3 100.0 ubc – 818 3 3 100.0 total 21 13 61.9 1 for all primers, additional bands were present but could not be reliably scored (i.e., band was weak or too close to adjacent band). the number of loci reliably scored does not include these additional unscored bands. 2 given that bands that could not be reliably scored were excluded from this calculation, this is an approximate percentage of polymorphic loci. table 3 number of loci reliably scored, number of polymorphic loci, and percentage of polymorphic loci for each of the six issr primers used in the study of genetic differentiation among three d. oligosanthes populations in stillwater, ok. primer number of loci reliably scored1 number of reliably scored polymorphic loci percentage polymorphic loci2 ubc 808 13 4 30.8 ubc 809 6 3 50.0 ubc 810 7 5 71.4 ubc 816 8 5 62.5 ubc 817 4 4 100.0 ubc 818 3 3 100.0 total 41 24 58.5 oklahoma native plant record volume 11, december 2011 parkhurst, m. j., et al. 40 are populations of d. olig osanthes genetically differentiated? the six issr primers produced a total of 41 loci that could be reliably scored, of which 24 were polymorphic across the three populations (table 3). in the ccs and lw populations 51.2% of the 41 issr loci were polymorphic and in the ccn population 53.7% were polymorphic. the average heterozygosities (hs) within each of the three populations were similar (mean ± s.d.; ccs: 0.36 ± 0.022, ccn: 0.35 ± 0.023, lw: 0.37 ± 0.020), indicating that genetic diversity does not differ drastically among the populations. on average, hs was 0.36 ± 0.016 across the three populations. in the bayesian analysis of population genetic differentiation using the full model, θii (analogous to weir and cockerham’s (1984) fst) was estimated to be 0.134 (s.d. = 0.0428), indicating a moderate proportion of differentiation among populations. discussion given the central role of genetic diversity in the evolutionary process, it is critical to understand how genetic diversity is distributed within and among populations. in this study, we found no evidence for fine-scale spatial genetic structure within a single population of d. oligosanthes. however, we found significant levels of genetic differentiation among three d. oligosanthes populations. below we expand on these findings and discuss potential factors contributing to the patterns of genetic structure and differentiation observed in d. oligosanthes. in the ccs population of d. oligosanthes, we found a negative but non-significant relationship between genetic similarity and geographic separation of plants. the lack of significance is contrary to our expectations, although the trend fits our hypothesis that plants in close spatial proximity to one another were more genetically similar than plants that were more spatially separated. such findings can be contrasted with those of other plant species, such as the selfcompatible annual herb polygonum thunbergii. in this species, konuma and terauchi (2001) found a significant negative correlation between genetic similarity and geographic distance (r = -0.64). the difference in finescale spatial genetic structure of these plant species may be due to differences in seed dispersal. grasses in the genus dichanthelium may disperse fruits over greater distances as the inflorescence breaks off of the plant, leading to a “tumbleweed” dispersal of the fruits (campbell et al. 1983). in contrast, p. thunbergii shows restricted seed dispersal, with seeds being dispersed in close proximity to the maternal plant (konuma and terauchi 2001). an additional factor that may contribute to the non-significant relationship found between genetic similarity and geographic distance in the ccs population of d. oligosanthes is low germination and recruitment of selfed (i.e., cleistogamous) individuals. however, in a related dichanthelium species, d. clandestinum, bell and quinn (1985) found that cleistogamous seeds germinated and emerged, both in the greenhouse and in the field, at a higher rate than chasmogamous seeds. future work is needed in d. oligosanthes to determine whether differences in cleistogamous and chasmogamous germination and recruitment may contribute to the observed patterns in spatial genetic structure. we found significant levels of genetic differentiation among three d. oligosanthes populations. this pattern of genetic differentiation could be due to the cleistogamous mating system of d. oligosanthes. grasses in the genus dichanthelium tend to reproduce proportionately more through cleistogamy than through chasmogamy (bell and quinn 1985, bell and quinn 1987). with elevated levels of inbreeding, genetic differentiation between populations is expected. the differentiation we observed amongst three neighboring populations of d. oligosanthes indicates that it is necessary to oklahoma native plant record volume 11, december 2011 parkhurst, m. j., et al. 41 conserve multiple populations of this species to maintain genetic diversity. acknowledgements funding for this study was provided by national science foundation through oklahoma louis stokes alliance for minority participation and oklahoma state university. special thanks to theresa henley for the aerial photograph used as the base for figure 1. literature cited adams, d. e. and l. l. wallace. 1985. nutrient and biomass allocation in five grass species in the oklahoma tallgrass prairie. american midland naturalist 113:170-181. bell, t. j. and j. a. quinn. 1985. relative importance of chasmogamously and cleistogamously derived seeds of dichanthelium clandestinum (l.) gould. botanical gazette 146:252-258. bell, t. j. and j. a. quinn. 1987. effects of soil moisture and light intensity on the chasmogamous and cleistogamous components of reproductive effort of dichanthelium clandestinum populations. canadian journal of botany 65:2243-2249. bonnet, e. and y. van der peer. 2009. zt: a software tool for simple and partial mantel tests. (http://www.psb.ugent.be/ ~erbon/mantel/). campbell, c. s., j. a. quinn, g. p. cheplick, and t. j. bell. 1983. cleistogamy in grasses. annual review of ecology and systematics 14:411-441. epperson, b. k. and t. li. 1997. gene dispersal and spatial genetic structure. evolution 51:672-681. freckmann, r. w. and m. g. lelong. 2003. dichanthelium. m. e. barkworth, k. m. capels, s. long, and m. b. piep (eds.). flora of north america north of mexico, vol. 25. new york. hammer, o., d. a. t. harper, and p. d. ryan. 2001. past: paleontological statistics software package for education and data analysis. paleontologia electronica 4:9. (http://palaeo-electronica.org/2001 _1/past/issue1_01.htm). hamrick, j. l. and j. d. nason. 1996. consequences of dispersal in plants. in: o. e. rhodes, r. k. chesser, and m. h. smith (eds.). population dynamics in ecological space and time. the university of chicago press, chicago, illinois. holsinger, k. e. and p. o. lewis. 2003. hickory: a package for analysis of population genetic data, version 1.1. department of ecology and evolutionary biology, university of connecticut storrs, connecticut. (http://darwin.eeb.uconn.edu/ hickory/hickory.html). holsinger, k. e., p. o. lewis, and d. k. dey. 2002. a bayesian approach to inferring population structure from dominant markers. molecular ecology 11:1157-1164. junghans, h. and m. metzlaff. 1990. a simple and rapid method for the preparation of total plant dna. biotechniques 8:176. kansas state university libraries. 2011. scribner dichanthelium. (m. haddock, editor) retrieved june 29, 2011, from kansas wildflowers and grasses: www.kswildflowers.org. konuma, a. and r. terauchi. 2001. population genetic structure of the selfcompatible annual herb; polygonum thunbergii (polygonaceae) detected by multilocus dna fingerprinting. american midland naturalist 146:22-127. leimu, r., p. mutikainen, j. koricheva, and m. fischer. 2006. how general are positive relationships between plant population size, fitness, and genetic variation. journal of ecology 94:942-952. loiselle, b. a., v. l. sork, j. nason, c. graham. 1995. spatial genetic structure of a tropical understory shrub, psychotria officinalis (rubiaceae). american journal of botany 82:1420-1425. loveless, m. d. and j. l. hamrick. 1984. ecological determinants of genetic oklahoma native plant record volume 11, december 2011 parkhurst, m. j., et al. 42 structure of plant populations. annual review of ecology and systematics 15:65-95. miyazaki, y. and y. isagi. 2000. pollen flow and the intrapopulation genetic structure of heloniopsis orientalis on the forest floor as determined using microsatellite markers. theoretical applied genetics 101:718-723. perry, d. j. and p. knowles. 1991. spatial genetic structure within three sugar maple (acer saccharum marsh.) stands. heredity 66:137-142. reed, d. h. and r. frankham. 2003. correlation between fitness and genetic diversity. conservation biology 17:230-237. spiegelhalter, d. j., n. g. best, b. r. carlin, and a. van der linde. 2002. bayesian measures of model complexity and fit. journal of the royal statistical society, b, methodological 64:583-616. usda, nrcs. 2011. the plants database (http://plants.usda.gov, 23 june 2011). national plant data center, baton rouge, la. wagner, v., w. durka, and i. hensen. 2011. increased genetic differentiation but no reduced genetic diversity in peripheral vs. central populations of a steppe grass. american journal of botany 98:1173-1179. weir, b. s. and c. c. cockerham. 1984. estimating f-statistics for the analysis of population structure. evolution 38:13581370. zietkiewicz, e., a. rafalski, and d. labuda. 1994. genome fingerprinting by simple sequence repeat (ssr)-anchored polymerase chain reaction amplification. genomics 20:176–183. spatial genetic structure of the tallgrass prairie grass dichanthelium oligosanthes (scribner’s panicum) by ms. molly j. parkhurst, dr. andrew doust, dr. margarita mauro-herrera,dr. janette a. steets, and dr. jeffrey m. byrnes journal of the oklahoma native plant society, volume 2, number 1, december 2002 4 oklahoma native plant record volume 2, number 1, december 2002 vascular plants of the wichita mountains wildlife refuge (1977) paul buck department of life sciences the university of tulsa tulsa, oklahoma introduction this list is an effort to update a popular list of the vascular plants of the wichitamountains wildlife refuge prepared by the author approximately ten years ago. the earlier work was compiled from lists published by eskew (1938), osborn and allen(1949) and from studies of several regional herbaria. the changes over the previous list reflect new species found during field activities in the refuge through the years,additions reported in the literature and modifications of scientific names as published in floras, keys, and manuals. common names are always a problem,frequently changing several times within relatively small geographic areas. in an effort to avoid this difficulty the common names have been taken from a guide to plant names, (rechenthin, 1954) which was prepared for use by the regional soil conservation service. the very few not listed in this publication were obtained from fernald (1950)and the updated version of britton and brown (gleason 1952). the asterisks (*) indicate species that are questionable as present inhabitants of the refuge for a couple of reasons: (1) many were reported in the area by earlier workers but recent literature indicates their ranges do not extend into the wichita mountains system. it might be inferred these were misidentified,and in several cases where the specimens were located, this proved to be the case. buck, p. https://doi.org/10.22488/okstate.17.100011 (2) during the 1930’s thousands of seedlings of non-native woody species were introduced and planted (example: mahonia aquifolium, barberry; spiraea arguta, spirea; hibiscus syriacus, rose of sharon; lagerstroemia indica, crape myrtle, etc.). most of these were subsequently moved to other locations within the southwest. probably none of those left in the refuge survived the competition of the native flora. the identity of the maple within the refuge has been a subject of debate for many years. opinions have been divided between the bigtooth maple (acer grandidentatum) and sugar maple (a. saccharum). the writer felt, when the earlier list was prepared, the wichita maple was the latter and deleted the bigtooth maple from the list. dent(1969) made a study of the sugar maples of oklahoma and concluded the caddo canyon and wichita mountains groups are both acer saccharum. references cited dent, thomas curtis 1969 relationships of two isolated groups of sugar maple in central oklahoma to eastern and western species. unpublished phd dissertation, university ofoklahoma, norman. eskew, cletis t. 1938 the flowering plants of the wichita mountains wildlife refuge, oklahoma. amer. mid. nat. 20:695-703. fernald, m.l. 1950 gray’s manual of botany. american book company, new york. oklahoma native plant record 5 volume 2, number 1, december 2002 buck, p. gleason, henry a. 1952 the new britton and brown illustrated flora of the northeastern united states and adjacent canada. the new york botanical garden, new york. osborn, b. and p. f. allan 1949 vegetation of an abandoned prairie-dog town in tall-grass prairie. ecology 30:322-332. rechenthin, c.a. 1954 a guide to plant names in texas, oklahoma, louisiana, arkansas. united states department of agriculture, soil conservation service. species list equisetum laevigatum equisetaceae kansas horsetail selaginella peruviana selaginellaceae sheldon selaginella selaginella rupestris rock selaginella isoetes butleri isoetaceae butler quillwort isoetes melanopoda blackfoot quillwort asplenium trichomanes polypodiaceae spleenwort cheilanthes eatoni eaton lipfern cheilanthes fendleri* fendler lipfern cheilanthes lanosa hairy lipfern cheilanthes lindheimeri lindheimer lipfern cheilanthes vestita lipfern cheilanthes wootoni* wooton lipfern dryopteris marginalis standley’s lipfern notholaena standleyi standley’s lipfern pellaea atropurpurea purple cliffbrake pellaea ternifolia wright’s cliffbrake woodsia obtuse woodsia pilularia americana marsileaceae american pillwort marsilea mucronata pepper wort juniperus virginiana pinaceae eastern red cedar pinus echinata* shortleaf pine pinus edulis* nut pine pinus ponderosa* western yellow pine pinus taeda* loblolly pine thuja orientalis* cupressaceae oriental arbor-vitae typha angustifolia typhaceae narrow leaf cattail typha domingensis cattail typha latifolia cattail zosteraceae potamogeton amplifolius largeleaf pondweed potamogeton diversifolius waterthread pondweed potamogeton nodosus longleaf pondweed potamogeton pusillus baby pondweed najadaceae najas guadalupensis southern naiad alismataceae echinodorus cordifolius erect burhead sagittaria calycina longlobe arrowhead sagittarria latifolia arrowhead agropyron repens* poaceae couch grass agrostis ellottiana 6 oklahoma native plant record volume 2, number 1, december 2002 buck, p. elliott bentgrass agrostis hymenalis winter bentgrass alopecurus caroliniana carolina foxtail andropogon barbinodis cane bluestem andropogon gerardi big bluestam andropogon saccharoides silver bluestem andropogon scoparius little bluestem aristida dichotoma var. dichotoma churchmouse threeawn aristida dichotoma var. curtissii curtis threeawn aristida longiseta red threeawn aristida longespica slimspike threeawn aristida oligantha oldfield threeawn aristida purpurea purple threeawn aristida wrightii wright threeawn bouteloua curtipendula sideoats grama bouteloua gracilis blue grama bouteloua hirsute hairy grama bouteloua rigidiseta texas grama bromus japonicus japanese brome bromus arvensis* field brome bromus purgans canada brome bromus tectorum cheatgrass bromus uniloides rescuegraas buchloe dactyloides buffalo grass cenchrus pauciflorus mat sandbur chloris verticillata tumble windmillgrass chloris virgata showy chloris cynodon dactylon bermuda grass dactylis glomerata orchard grass digitaria sanguinalis hairy crabgrass diplachne fascicularis bearded spranglegrass echinochloa crusgalli barnyard grass elusine indica goosegrass elymus canadensis canada wildrye elymus virginicus virginia wildrye eragrostis capillaris lacegrass eragrostis curtipedicellata gummy lovegrass eragrostis hypnoides teal lovegrass eragrostis intermedia plains lovegrass eragrostis cilianensis stinkgrass eragrostis oxylepsis var. oxylepis red lovegrass eragrostis oxylepsis var. beyrichii wichita lovegrass eragrostis pectinacea carolina lovegrass eragrostis pilosa india lovegrass eragrostis reptans creeping lovegrass eragrostis sessilispica tumble lovegrass eragrostis spectabilis purple lovegrass eragrostis trichodes sand lovegrass eriochloa contracta prairie cutgrass festuca octoflora sixweeks fescue festuca pratensis suiter fescue festuca versuta texas fescue hordeum pusillum little barley leersia oryzoides rice grass leptochloa dubia oklahoma native plant record 7 volume 2, number 1, december 2002 buck, p. green sprangletop leptoloma cognatum fall witchgrass limnodea arkansana ozark grass manisuris cylindrica carolina jointtail melica nitens threeflower melic muhlenbergia capillaris hairawn muhly muhlenbergia frondosa wirestem muhly muhlenbergia racemosa green muhly panicum agrostoides redtop panicum panicum anceps beaked panicum panicum capillare witchgrass panicum depauperatum poor panic grass panicum dichotomiflorum fall panicum panicum lanuginosum wooly panicum panicum linearifolium slimleaf panicum panicum malacophyllum shortleaf panicum panicum obtusum vine-mesquite panicum obligosanthes var. scribnerianum scribner panicum panicum perlongum elongate panic grass panicum philadelphicum philadelphia witchgrass panicum virgatum switchgrass paspalum pubiflorum var. publiflorum hairyseed paspalum paspalum publiflorum var. glabrum smoothseed paspalum paspalum setaceum fringeleaf paspalum phalaris caroliniana carolina canary grass poa annua annual bluegrass poa arachnifera texas bluegrass poa compressa canada bluegrass schedonnardus paniculatus tumblegrass setaria geniculata knotroot bristlegrass setaria viridis green bristlegrass sorgastrum nutans indiangrass sorghum halapense johnson grass spartina pectinata var. suttiei prairie cordgrass sphenopholis obtusata prairie wedgescale sporobolus airoides alkali sacaton sporobolus asper tall dropseed sporobolus cryptandrus sand dropseed sporobolus pyramidatus whorled dropseed sporobolus vaginiflorus var. vaginiflorus poverty dropseed sporobolus vaginiflorus var. neglectus puffsheath dropseed trichachne californica arizona cottontop tridens albescens white tridens tridens flavus purpletop tridens muticus var. muticus slim tridens tridens muticus var. elongatus rough tridens tridens pilosus hairy tridens tridens strictus longspike tridens tripsacum dactyloides eastern gamagrass uniola latifolia broadleaf uniola bulbostylis capillaris cyperaceae hairsedge carex amphibola narrowleaf sedge carex annectens yellowfruit sedge 8 oklahoma native plant record volume 2, number 1, december 2002 buck, p. carex austrina southern sedge carex blanda woodland sedge carex festucacea fescue sedge carex frankii frank sedge carex gravida var. lunellina lunnell sedge carex joori hummock sedge carex microrhyncha littlesnout sedge carex praegracilis slender sedge carex stricta var. elongata emory sedge carex vulpinoidea fox sedge cyperus acuminatus taperleaf flatsedge cyperus aristatus bearded flatsedge cyperus erythrorhizos redroot flatsedge cyperus esculentus chufa cyperus filiculmis slender flatsedge cyperus odoratus fragrant flatsedge cyperus ovularis globe flatsedge cyperus schweinitzii schweinitz flatsedge cyperus setigerus bristled spike flatsedge cyperus strigosus false nut sedge cyperus virens green flatsedge eleocharis compressa flatstem spikesedge eleocharis engelmanii englemann spikesedge eleocharis macrostachya longspike spikesedge eleocharis montevidensis sand spikesedge eleocharis obtusa blunt spikesedge eleocharis parvula dwarf spikesedge eleocharis quadrangulata squarestem spikesedge eleocharis tenuis slender spikesedge fimbristylis spadicea plains fimbry fimbristylis vahlii vahl fimbry fuirena simplex western umbrella sedge hemicarpha micrantha hemicarpha scirpus acutus hardstem bulrush scirpus americanus american bulrush scirpus atrovirens green bulrush scirpus lacustris softstem bulrush scirpus lineatus rusty bulrush scirpus paludosis alkali bulrush scirpus pauciflora fewflower nutrush arisaema dracontium araceae dragonroot or jack-in-the-pulpit lemna minor lemnaceae duckweed commelina erecta commelinaceae var. angustifolia narrowleaf dayflower commelina virginica virginia dayflower tradescantia hirsutiflora hairyflower spiderwort tradescantia occidentalis prairie spiderwort tradescantia ohiensis smoothstalk spiderwort tradescantia tharpii tharp spiderwort heteranthera limosa pontederiaceae blue mudplantain juncos acuminatus juncaceae knotleaf rush oklahoma native plant record 9 volume 2, number 1, december 2002 buck, p. juncus balticus baltic rush juncus interior inland rush juncus marginatus grassleaf rush juncus tenuis var. tenuis poverty rush juncus tenuis var. dudleyi dudley rush juncus torreyitorrey rush allium canadense liliaceae canada garlic allium drummondii drummond onion allium stellatum prairie onion androstephium coeruleum blue funnellily camassia scilloides atlantic camass erythronium americanum fawn lily nothoscordum bivalve yellow false garlic smilax bona-nox saw greenbrier smilax herbacea carrionflower smilax rotundifolia greenbrier smilax tamnoides bristly greenbrier yucca glauca small soapweed agave lata amaryllidaceae agave zephyranthes brazosensis evening start or rain lily nemastylis geminiflora iridaceae prairie iris sisyrinchium angustifolium blueeye grass spiranthes cernua orchidaceae nodding ladiestresses populus deltoids salicaceae cottonwood salix amygdaloides peachleaf willow salix caroliniana ward's willow salix interior sandbar willow salix matsudana* willow salix nigra black willow carya cordiformis juglandaceae bitternut hickory carya illinoensis pecan juglans nigra black walnut juglans rupestris little walnut quercus falcate fagaceae southern red oak quercus macrocarpa bur oak quercus marilandica blackjack oak quercus mohriana mohrs oak quercus muhlenbergii chinquapin oak quercus rubus var. borealis red oak quercus shumardii var. shumardii shumard’s oak quercus shumardii var. schneckii schneck's oak quercus stellata post oak quercus velutina black oak quercus virginiana live oak celtis laevigata ulmaceae sugar hackberry celtis occidentalis hackberry celtis reticulata netleaf hackberry 10 oklahoma native plant record volume 2, number 1, december 2002 buck, p. ulmus americanus american elm ulmus pumila* chinese elm ulmus rubra slippery elm maclura pomifera moraceae osage orange morus microphylla texas mulberry morus rubra red mulberry morus tatarica* russian mulberry boehmeria cylindrica urticaceae smallspike falsenettle parietaria pensylvanica pennsylvania pellitory comandra pallida santalaceae western comandra loranthaceae phoradendron serotinum mistletoe aristolochiaceae aristolochia tomentosa wooly dutchmanpipe eriogonum annum polygonaceae annual wild buckwheat eriogonum longifolium longleaf wild buckwheat polygonum aviculare prostrate knotweed polygonum bicorne longstype smartweed polygonum coccinium bigroot smartweed polygonum convolvulus dullseed cornbind polygonum hydropiper marshpepper smartweed polygonum hydropiperoides var. hydropiperoides swamp smartweed polygonum hydropiperoides var. opelousanum opelousas swamp smartweed polygonum lapathifolium curlytop smartweed polygonum pensylvanicum pennsylvania smartweed polygonum punctatum dotted smartweed polygonum ramosissimum bushy knotweed polygonum scandens hedge cornbind polygonum tenue pleatleaf knotweed rumex altissimus pale dock rumex crispus curly dock chenopodium album chenopodiaceae lambsquarters chenopodium hybridum var. gigantospermum bigseed goosefoot chenopodium leptophyllum slimleaf goosefoot chenopodium standleyanum standley goosefoot salsola kali var. tenuifolia russian thistle monolepis nutalliana nuttall monolepis alternanthera repens amaranthaeceae mat chaff flower amaranthus graecizans tumbleweed amaranthus hybridus slim amaranth amaranthus palmeri palmer amaranth amaranthus retroflexus redroot amaranth amaranthus tamarascinus waterhemp brayulinea densa* cottonflower froelichia floridana florida snakecotton floelichia gracilis slender snakecotton gossypianthus lanuginosus woolly cottonflower gossypianthus tenuiflorus lanceleaf cottonflower oklahoma native plant record 11 volume 2, number 1, december 2002 buck, p. mirabilis albida nyctaginaceae pale umbrella-wort mirabilis inearis narrowleaf umbrella-wort mirabilis nyctaginea heartleaf umbrella-wort phytolacca americana phytolaccaceae pokeberry mollugo verticillata aizoaceae green carpetweed claytonia virginica portulacaceae virginia spring beauty portulaca oleracea purslane portulaca parvla shaggy purslane portulaca umbraticola wingpod purslane talinum calycinum rockpink talinum parvifolorum prairie fameflower talinum rugospermum* roughseed fameflower arenaria patula caryophyllaceae pitcher’s sandwort arenaria stricta var. texana texas sandwort cerastium brachypodum shortstalk chickweed sagina decumbens trailing pearlwort silene antirrhina sleepy catchfly paronychia jamesii illecebraceae james nailwort paronychia virginica virginia nailwort ceratophyllum demersum ceratophyllaceae coontail ranunculaceae anemone caroliniana carolina anemone anemone decapetala tenpetal anemone clematis pitcheri pitcher clematis delphinium virescens plains larkspur berberidaceae mahonia aquifolium* barberry cocculus carolinus menispermaceae carolina snailseed menispermum canadense moonseed argemone polyanthemos papaveraceae prickle poppy corydalis aurea fumariaceae var. occidentalis goldon corydalis cleome serrulata capparidaceae bee spider flower cleomella angustifolia narrowleaf rhombopod polanisia dodecandra roughseed clammywort cruciferae arabis missouriensis green rockcress capsella bursa–pastoris shepherdspurse descurainia pinnata pinnate tansymustard dithyrea wislizenii wislizen's spectacle pod draba brachycarpa shortpod draba draba cuneifolia wedgeleaf draba draba reptans carolina draba erysimum capitatum plains erysimum lepidium oblongum veiny pepperwort lepidium virginicum virginia pepperwort lesquerella auriculata earleaf bladderpod 12 oklahoma native plant record volume 2, number 1, december 2002 buck, p. lesquerella gordonii gordon bladderpod lesquerella gracilis lax bladderpod lesquerella ovalifolia var. alba roundleaf bladderpod nasturtium officinale watercress rorippa islandica bog marshcress rorippa sessilifolia stalkless yellowcress sibara virginica virginia rockcress sedum nuttallianum crassulaceae yellow stonecrop sedum pulchellum texas stonecrop ribes odoratum saxifragaceae clove current argimonia parviflora rosaceae mayflower grovebur amelanchier arborea service berry crataegus crus-galli cockspur hawthorn crataegus engelmannii engelmann hawthorn crataegus reverchonii reverchon hawthorn geum canadense white avens geum virginiana* bristly avens potentilla arguta cinquefoil prunus americana american plum prunus angustifolia chickasaw plum prunus armeniaca* apricot prunus persica peach prunus gracilis oklahoma plum prunus mexicana mexican plum prunus virginiana chokecherry rosa foliolosa* leafy rose rosa multiflora* multiflora rose rubus aboriginum northern dewberry rubus idaeus red raspberry rubus louisanus louisiana blackberry rubus occidentalis blackcap raspberry rubus trivialis lowbush blackberry spiraea arguta* spirea spirea cantoniensis* spirea spirea vanhouttei* spirea acacia angustissima leguminosae prairie acacia amorpha canescens leadplant amorpha fruticosa indigobush amorpha amorpha microphylla* smallleaf amorpha amorpha nana* fragrant false indigo apios americana groundnut astragalus crassicarpus groundplum milkvetch astragalus lindheimeri lindheimer milkvetch astragalus plattensis platte milkvetch baptisia australis var. minor blue wild indigo baptisia leucophaea plains wild indigo baptisia sphaerocarpa golden wild indigo cassia fasciculate showy partridgepea cassia marilandica wild senna cercis canadensis redbud clitoria mariana atlantic pigeonwings oklahoma native plant record 13 volume 2, number 1, december 2002 buck, p. colutea arborescens* bladder senna crotalaria sagittalis arrow crotalaria dalea aurea siltop aurea dalea enneandra bigtop dalea dalea purpurea purple prairie-clover desmanthus illinoensis illinois bundleflower desmodium ciliare littleleaf tickclover desmodium glutinosum sticky tickclover desmodium nudiflorum barestem tickclover desmodium paniculatum panicled tickclover desmodium sessilisovium sessile tickclover galactia volubilis downy milkpea gleditsia triacanthos honeylocust glycyrrhiza lepidota american licorice gymnocladus dioica kentucky coffee tree hoffmanseggia densiflora indian rushpea indigofera miniata var. leptosepala coast indigo krameria secundiflora trailing ratany lespedeza capitata roundhead lespedeza lespedeza intermedia intermediate lespedeza lespedeza procumbens trailing lespedeza lespedeza virginica slender lespedeza lotus americanus deervetch melilotus alba white sweet clover melilotus officianalis yellow sweet clover neptunea lutea yellow neptunea oxytropis lamberti stemless loco petalostemum candidum white prairie clover petalostemum multiflorum roundtop prairie clover petalostemum purpureum purple prairie clover petalostemum tenuifolium slimleaf prairie clover prosopis juliflora mesquite psoralea cuspidata tallbread scurfpea psoralea esculenta indian breadroot psoralea lineariflolia slimleaf scurfpea psoralea tenuiflora wild alfalfa robinia pseudoacacia black locust shrankia uncinata catclaw sensitive brier strophostyles helvola trailing wildbean strophostyles leiosperma slickseed wildbean stylosanthes riparia endbeak pencilflower trifolium reflexum buffalo clover vicia ludoviciana louisiana vetch vicia sparsifolia stiffleaf vetch geraniaceae geranium carolinianum carolina geranium erodium cicutarium storksbill oxalidaceae oxalis stricta oxalis oxalis violacea violet woodsorrel linaceae linum hudsonioides tufted flax linum rigidum stiffstem flax linum sulcatum grooved flax zygophyllaceae tribulus terrestris 14 oklahoma native plant record volume 2, number 1, december 2002 buck, p. puncturevine rutaceae ptelea trifoliate wafer ash simaroubaceae ailanthus altissima* tree-of-heaven polygalaceae polygala alba white milkwort polygala verticillata whorled milkwort euphorbiaceae acalypha gracilens slender copperleaf acalypha ostryaefolia hophornbeam copperleaf croton capitatus woolly croton croton glandulosus tropic croton croton lindheimerianus threeseed croton croton monanthogynus oneseed croton croton texensis texas croton euphorbia commutata tinted euphorbia euphorbia corollata flowering spurge euphorbia dentata toothed euphorbia euphorbia glyptosperma ridgeseed euphorbia euphorbia hexagona six-angle euphorbia euphorbia marginata snow-on-the-mountain euphorbia missourica missouri euphorbia euphorbia nutans spotted euphorbia euphorbia obtusata roughpor euphorbia euphorbia prostrata prostrate euphorbia euphorbia spathulata warty euphorbia euphorbia supina milk purslane phyllanthus caroliniensis carolina leafflower phyllanthus polygonoides knotweed leafflower stillingia sylvatica queen's delight tragia ramose branching noseburn callitriche heterophylla callitrichaceae larger waterstar wort anacardiaceae rhus aromatica skunkbush rhus copallina flameleaf sumac rhus glabra smooth sumac rhus toxicodendron poison ivy celastraceae celastrus scandens american bittersweet euonymus atropurpureus eastern wahoo euonymus europaeus* european spindle-tree acer saccharinum* aceraceae silver maple acer saccharum sugar maple hipposcastanaceae aesculus glabra ohio buckeye sapindaceae sapindus drummondii* western soapberry rhamnaceae ceanothus herbaceus var. pubescens fuzzy ceanothus rhamnus frangula* alder buckthorn ampelopsis cordata vitaceae heartleaf ampelopsis cissus incisa ivy treebine parthenocissus quinquefolia virginia creeper oklahoma native plant record 15 volume 2, number 1, december 2002 buck, p. vitis aestivalis summer grape vitis cinerea sweet grape vitis palmata cat grape vitis rotundifolia muscadine grape vitis rupestris sand grape vitis vulpine riverbank grape malvaceae callirhoe involucrata low poppymallow callirhoe leiocarpa tall poppymallow hibiscus syriacus* rose of sharon sphaeralcea coccinea scarlet mallow guttiferae hypericum drummondii nits and lice hypericum mutilum dwarf st. john's wort tamaricaceae tamarix hispida* kashgar tamarisk tamarix odessana* tamarisk tamarix pentandra* tamarisk lechea tenuifolia cistaceae narrowleaf pinwheel violaceae hybanthus verticillatus whorled nodviolet viola missouriensis missouri violet viola papilionacea butterfly violet viola rafinesquii johnny-jump-up loasaceae mentzelia oligosperma chickthief mentzelia stricta sandlily cactaceae echinocereus baileyi wichita pin cushion echinocereus reichenbachii lace echinocereus opuntia compressa prickly pear lythraceae ammannia auriculata earleaf ammannia ammannia coccinea purple ammannia lagerstroemia indica* crape myrtle lythrum alatum winged lythrum gaura biennis var. pitcheri onagraceae pitcher gaura gaura coccinea scarlet gaura gaura parviflora smallflower gaura gaura sinuata wavyleaf gaura guara suffulta roadside gaura gaura tripetala var. triangulata three petal gaura jussiaea decurrens wingleaf water primrose jussiaea peploides var. glabrescens smooth water primrose ludwigia alternifolia bushy boxseed ludwigia palustris marsh boxseed oenothera biennis evening primrose oenothera heterophylla varileaf evening primrose oenothera jamesii trumpet evening primrose oenothera laciniata var. laciniata cutleaf evening primrose oenothera laciniata var. grandiflora largeflowered cutleaf evening primrose oenothera linifolia threadleaf sundrops 16 oklahoma native plant record volume 2, number 1, december 2002 buck, p. oenothera macrocarpa var.macrocarpa ozark sundrops oenothera macrocarpa var. incana gray sundrops oenothera missouriensis missouri evening primrose oenothera serrulata halfshrub sundrops oenothera speciosa showy sundrops oenothera triloba stemless evening primrose stenosiphon virgatus false gaura haloragaceae myriophyllum brasiliensis parrot feather myriophyllum exalbescens parrot feather umbelliferae ammoselinum butleri butler sandparsley ammoselinum popei plains sandparlsey chaerophyllum tainturieri var. dasycarpum hairyfruit chervil cicuta maculata spotted water hemlock daucus carota* wild carrot daucus pusillus southwestern carrot eryngium diffusum bushy eryngo eryngium leavenworthii leavenworth eryngo limnosciadum pinnatum arkansas dogshade lomatium daucifolium carrotleaf lomatium polytaenia nuttallii prairie parsley ptilimnium nuttallii nuttall mockbishop sanicula canadensis canada sanic1e spermolepis divaricata forked scaleseed spermolepis echinata bristly scaleseed cornaceae cornus drummondii roughleaf dogwood cornus florida* flowering dogwood primulaceae androsace occidentalis western rockjasmine dodecatheon meadia shooting star samolus parviflorus thinleaf brookweed sapotaceae bumelia lanuginosa woollybucket bumelia ebenaceae diospyros virginiana persimmon oleaceae forsythia suspensa var. fortunei* golden bells fraxinus americana white ash fraxinus pennsylvanica var. pennsylvanica red ash fraxinus pennsylvanica var. subintegerrima green ash fraxinus velutina* arizona ash jasminum nudiflorum* jasmine ligustrum amurense* privet ligustrum vulgare* privet sabatia angularis gentianaceae squarestem rosegentium sabatia campestris prairie rosegentian amsonia ciliata var. texana apocynaceae texas slimpod amsonia tabernaemontana willow slimpod apocynum cannabinium var. glaberrimum hemp dogbane oklahoma native plant record 17 volume 2, number 1, december 2002 buck, p. asclepias asperula asclepiadaceae var. decumbens trailing milkweed asclepias latifolia broadleaf milkweed asclepias pumila plains milkweed asclepias stenophylla slimleaf milkweed asclepias sullivantii sullivant milkweed asclepias tuberosa butterfly milkweed asclepias verticillata whorled milkweed asclepias viridiflora green acerates asclepias viridis green antelope horn convolvulus arvensis convolvulaceae small bindweed convolvulus incanus gray bindweed cuscuta coryli hazel dodder cuscuta gronovii gronovius dodder cuscuta indecora showy dodder cuscuta cuspidata cusp dodder evolvulus nuttallianus silver evolvulus ipomoea shumardiana narrowleaf morning glory gilia rubra polemoniaceae texasplume phlox pilosa downy phlox hydrophyllaceae nama hispidum rough nama phacelia congesta spike phacelia phacelia hirsuta hairy phacelia boraginaceae lithospermum arvense corn gromwell lithospermum incisum narrowleaf gromwell myosotis macrosperma southern forget-me-not myosotis verna forget-me-not onosmodium molle marbleseed verbenaceae phyla cuneifolia wedgeleaf fogfruit phyla incisa sawtooth fogfruit verbena bipinnatifida dakota verbena verbena bracteata bigbract verbena verbena canadensis rose verbena verbena pumila pink verbena verbena stricta woolly verbena verbena urticifolia white verbena vitex agnus-castus* chaste tree labiatae hedeoma drummondii drummond hedeoma hedeoma hispida rough hedeoma lamium amplexicaule henbit lycopus americanus american bugleweed monarda citriodora lemon beebalm monarda clinopodioides basil beebalm monarda fistulosa shrubby beebalm monarda punctata spotted beebalm nepeta cataria catnip salvia azurea var. grandiflora azure sage salvia reflexa lanceleaf sage scutellaria drummondii 18 oklahoma native plant record volume 2, number 1, december 2002 buck, p. drummond skullcap scutellaria resinosa resindot skullcap scutellaria wrightii wright skullcap teucrium canadense american germander teucrium laciniatum cutleaf germander trichostema brachiatum fluxweed solanaceae physalis lobata purple groundcherry physalis virginiana var. sonorae virginia groundcherry physalis viscosa var. cinerascens beach groundcherry physalis viscosa var. mollis field groundcherry solanum carolinense carolina horsenettle solanum citrullifolium* watermelon leaved solanum solanum elaegnifolium silverleaf nightshade solanum rostratum buffalobur solanum torreyi torrey horsenettle scrophulariaceae bacopa rotundifolia disc waterhyssop castilleja citrina citron paintbrush castilleja sessiliflora downy paintbrush collinsia violacea violet collinsia gratiola virginiana virginia hedgehyssop leucospora multifida narrowleaf conobea linaria canadensis var. texana oldfield toadflax lindernia anagallidea c1asping false-pimpernel lindernia dubia yellowseed false-pimpernel penstemon albidus white penstemon penstemon cobea cobaea penstemon penstemon oklahomensis oklahoma penstemon penstemon tubaeflorus tube panstemon scrophularia lanceolata narrowleaf figwort veronica arvensis speedwell veronica peregrina purslane speedwell verbascum thapsus flannel mullein catalpa speciosa bignoniaceae catalpa chilopsis linearis* desert willow martyniaceae proboscidea louisianica unicorn plant lentibulariaceae utricularia biflora bladderwort utricularia vulgaris bladderwort acanthaceae justicia americana justicia ruellia humilis low ruellia ruellia pedunculata stalked ruellia phrymaceae phryma leptostachya lopseed plantaginaceae plantago aristata bottlebrush plantain plantago elongata elongate plantain plantago purshii var. purshii slender plantain plantago purshii var. spinulosa bristlebract plantain plantago rhodosperma redseed plantain plantago virginica paleseed plantain plantago wrightiana wright plantain oklahoma native plant record 19 volume 2, number 1, december 2002 buck, p. rubiaceae cephalanthus occidentalis buttonbush diodia teres var. setifera rough buttonweed galium aparine catchweed bedstraw galium pilosum hairy bedstraw galium texense texas bedstraw galium triflorum fragrant bedstraw galium virgatum southwest bedstraw hedyotis crassifolia tiny bluet hedyotis nigricans narrowleaf bluet hedyotis purpurea var. longifolia purple bluet lonicera japonica* caprifoliaceae honeysuckle lonicera tatarica* tatarian honeysuckle symphoricarpos orbiculatus coralberry viburnum prunifolium var. ferrugineum rusty blackhaw valerianella radiata valerianaceae var. radiata beaked cornsalad valerianella radiata var. parviflora narrowcell cornsalad cucurbita foetidissima cucurbitaceae buffalo gourd cyclanthera dissectra cutleaf cyclanthera ibervillea lindheimeri lindheimer globeberry melothria pendula drooping me1onette campanulaceae specularia biflora small venus-looking glass specularia leptocarpa slimpod venus-looking glass specularia perfoliata clasping venus-looking glass lobeliaceae lobelia appendiculata earflower lobelia lobelia cardinalis cardinal flower lobelia spicata palespike lobelia compositae achillea lanulosa western yarrow achillea millefolium yarrow agoseris cuspidate wavyleaf agoseris ambrosia artemisiifolia ragweed ambrosia psilostachys western ragweed ambrosia trifida giant ragweed antennaria plantaginifolia plantain-leaf pussy toes aphanostephus pilosus hairy dozedaisy aphanostephus ramosissimus plains dozedaisy aphanostephus skirrhobasis arkansas dozedaisy artemisia carruthii carruth sagewort artemisia filifolia figleaf sagewort artemisia glauca false terragon sagewort artemisia ludoviciana var. ludoviciana louisiana sagewort artemisia ludoviciana var. mexicana american sagewort artemisia serrata* sawtooth sagewort aster communtatus* cluster aster aster ericoides heath aster aster fendleri fendler aster aster oblongifolius aromatic aster 20 oklahoma native plant record volume 2, number 1, december 2002 buck, p. aster patens skydrop aster aster subulatus awl shaped aster baccharis salicina willow groundsel-tree berlandiera texana texas greeneyes bidens cernua nodding beggarticks bidens frondosa devil’s beggarticks centaurea americana basketflower chaetopappa asteroides least daisy chrysopsis pilosa soft goldaster chrysopsis villosa var. villosa hairy goldaster chrysopsis villosa var. canescens hairy goldaster chrysopsis villosa var. stenophylla narrowleaf goldaster cirsium ochrocentrum yellowspine thistle cirsium undulatum wavyleaf thistle conyza canadensis horseweed fleabane conyza ramoisissima conyza coreopsis grandiflora var. longipes bigflower coreopsis coreopsis lanceolata var. villosa lanceleaf coreopsis coreopsis tinctoria plains coreopsis dyssodia tagetioides marigold dogweed echinacea angustifolia black sampson echinacea pallida pale purple coneflower eclipta alba yerba de tajo englemannia pinnatifida engelmann daisy erigeron bellidiastrum western fleabane erigeron divergens spreading fleabane erigeron strigosus prairie fleabane eupatorium serotinum late eupatorium evax multicaulis manystem evax evax prolifera bighead evax gaillardia pulchella rosering gaillardia gaillardia suavis sweet gaillardia gnaphalium chilense cottonbatting cudweed gnaphalium purpureum purple cudweed gnaphalium wrightii wright cudweed grindelia squarrosa var. squarrosa curlycup gumweed grindelia squarrosa var. nuda rayless gumweed gutierrezia dracunculoides broom weed gutierrezia sarothrae* broom snakeweed haplopappus ciliatus wax goldenweed haplopappus spinulosis cutleaf goldonweed helenium amarum var. amarum bitter sneezeweed helenium amarum var. badium basin sneezeweed helenium autumnale sneezeweed helenium flexuosum flexuous sneezeweed helenium microcephalum smallhead sneezeweed helianthus annuus annual sunflower helianthus hirsutus hairy sunflower helianthus laetiflorus stiff sunflower helianthus maximilani maximilian sunflower helianthus petiolaris prairie sunflower hieracium longipilum longbeard hawkseed hymenopappus scabiosaeus flattop woollywhite oklahoma native plant record 21 volume 2, number 1, december 2002 buck, p. hymenopappus tenuifolius chalkhill woollywhite hymenoxys linearifolia fineleaf actinea hymenoxys scaposa var. linearis plains actinea iva ciliata seacoast sumpweed krigia dandelion tuber dwarf dandelion krigia oppositifolia weedy dwarf dandelion kuhnia eupatorioides var. corymbulosa plains kuhnia lactuca canadensis canada lettuce lactuca pulchella* chickory lettuce lactuca scariola prickly lettuce liatris aspera rough gayfeather liatris punctata dotted gayfeather liatris scariosa tall gayfeathcr palafoxia texana texas palafoxia pluchea camphorata camphor pluchea pyrrhopappus carolinianus carolina false-dandelion pyrrhopappus scaposus tuber false-dandelion ratisida columnifera upright prairie coneflower rudbeckia hirta blackeyed susan santolina chamaecyparissus* lavender cotton senecio plattensis prairie groundsel senecio riddellii riddell groundsel silphium asteriscus sand rosinweed silphium laciniatum compass plant solidago bootii* boot goldenrod solidago gigantea var. leiophylla november goldenrod solidago missouriensis var. fasciculate missouri goldenrod solidago mollis ashy goldenrod solidago petiolaris downy goldenrod solidago radula rough goldenrod taraxacum officinale dandelion thelesperma ambiguum colorado greenthread thelesperma filifolium plains greenthread thelesperma megapotamicum rayless greenthread townsendia excapa stemless townsendia tragopogon major yellow goat's beard verbescina virginica white crownbeard vernonia baldwinii baldwin ironweed vernonia missurica missouri ironweed xanthisma texanum texas sleepydaisyxanthium strumarium cocklebur editor’s note: this paper, written in 1977, was originally distributed by the wichita mountains wildlife refuge as an informational handout but never published. scientific names that were originally underlined have been italicized and species epithets derived from a person’s name are not capitalized as they were in the original work to abide with current taxonomic practice. the author emphasizess that there is a great need to update this twenty-five year old floristic list and, together with wmwr, encourages new research on this site. oklahoma native plant record, volume 13, number 1, december 2013 oklahoma native plant record volume 13, december 2013 stanley a. rice and sonya l. ross https://doi.org/10.22488/okstate.17.100098 48 smoke-induced germination in ph acelia str ictaflor a stanley a. rice department of biological sciences southeastern oklahoma state university durant, ok 74701-0609 srice@se.edu sonya l. ross department of biological sciences southeastern oklahoma state university durant, ok 74701-0609 sross76@student.se.edu key words: butenolides, cross-timbers, fire, karrikens, reg eneration abstract seedlings of phacelia strictiflora a.l. de jussieu (hydrophyllaceae) germinate profusely after major fires in the cross timbers forest of oklahoma and rarely at other times and places. seed germination was greatly enhanced under laboratory conditions by exposure to a water suspension of smoke chemicals. this is the first report of smoke enhancement of germination in a native oklahoma plant species. many plant species grow abundantly after disturbances including fires, but smoke enhancement of germination allows p. strictiflora to grow abundantly after fires and only rarely after other kinds of disturbance. introduction seeds of some plant species rarely germinate except after fires. these species grow in ecological communities that depend upon a fire cycle for regeneration. examples include the chaparral (keeley and fotheringham 1998a; keeley et al. 2012), the shrublands of western australia (dixon et al. 1995; thomas et al. 2007; turner et al. 2009), and the south african fynbos (delange and boucher 1990). germination after fire may confer a great advantage on seedlings because the adult plants often form a dense cover against which the seedlings would be unable to successfully compete for light, water, and nutrients. post-fire germination also allows seedlings to benefit from a flush of mineral nutrients provided by the ashes. moreover, in firecycle communities, seeds may require exposure to one or more chemical components of smoke in order to germinate. in some cases, these chemicals may be oxidizing gases such as no2 (keeley and fotheringham 1998b), while in other cases they may be a group of butenolides known as karrikins, which are growth regulators produced by the combustion of cellulose (flematti et al. 2004; chiwocha et al. 2009; dixon et al. 2009). in many cases, profuse germination after fires may result mainly from the sudden and greater abundance of light, water, and nutrients at the ground level, rather than from exposure to smoke chemicals. many forms of disturbance other than fire also provide flushes of these resources. in this paper, we report that seed germination of phacelia strictiflora a. l. de jussieu (hydrophyllaceae) is very strongly enhanced by smoke and rarely occurs without smoke stimulation. we also present evidence that post-fire germination of p. strictiflora seeds is not simply due to availability of a flush of resources or to stratification. this is the first report of mailto:srice@se.edu mailto:sross76@student.se.edu oklahoma native plant record 49 volume 13, december 2013 smoke enhancement of germination in a native oklahoma plant species. phacelia strictiflora is a native oklahoma plant species that is a close relative of the chaparral p. grandiflora, in which smoke strongly enhances germination (keeley and fotheringam 1998a). p. strictiflora grows in poor soils in the south central united states. in oklahoma, p. strictiflora is normally a rare spring annual in the cross timbers forest, which is noted for its poor soils. however, after major summer fires, p. strictiflora may grow and bloom profusely, forming nearly a monoculture in some areas, in the following spring (figures 1, 2). the strong association between fire and germination led us to hypothesize that smoke chemicals greatly enhance the germination of p. strictiflora seeds. materials and methods we collected seeds from mature, dry phacelia strictiflora plants in a postfire area of the blue river wildlife conservation area maintained by the oklahoma department of wildlife conservation, near the junction of state highway 7 and the blue river in bryan county (34º 21.50’ n, 96º 35.41’ w). a wildfire destroyed most of the adult trees in 2011. we collected seeds of mature, senescent plants in may 2012 from ten different 25 m2 areas within the burned forest. we stored the dry seeds in plastic bags in the laboratory, at first at room temperature and later in a refrigerator, but we did not moisten the seeds except in the stratification treatment (see below). we used a dissecting microscope to carefully select healthy seeds (plump and free of discoloration) to use in the following experiments. figure 1 phacelia strictiflora at the blue river wildlife conservation area the april following the 2011 fire. stanley a. rice and sonya l. ross oklahoma native plant record volume 13, december 2013 stanley a. rice and sonya l. ross 50 figure 2 abundant post-fire growth of phacelia stricitflora in the cross timbers forest at the blue river wildlife conservation area in april 2012. oklahoma native plant record 51 volume 13, december 2013 stanley a. rice and sonya l. ross we first explored the likely range of conditions that may induce germination of p. strictiflora seeds. we exposed seeds to a broad range of conditions, including: heat and smoke from a grill in which we burned dried branches of q. stellata; ground ashes; physical scarification with sandpaper; a 3 mg/ml solution of miracle-gro® complete fertilizer; a 5% solution of colgin brand liquid smoke®; sulfuric acid (diluted to ph = 3); and ammonium hydroxide (diluted to ph = 10). we used 30 petri plates, each with 20 seeds, for a total of 600 seeds. we kept all petri plates on racks underneath fluorescent lights in a temperaturecontrolled laboratory for a month. for the two main experiments, we produced an aqueous suspension of smoke molecules in the following manner. we placed 150 ml (5.07 oz) of water in the reservoir of a commercial hookah pipe and burned post oak wood in the bowl. we drew smoke through the water using a nasal aspirator for three hours, during which time we replenished the oak wood as it burned. the result was a water suspension that was visibly amber and smelled like smoke. we diluted some of this suspension to halfstrength, and some of it to quarter-strength. we kept the three dilutions in refrigerated test tubes sealed with parafilm to discourage the diffusion of volatile smoke chemicals and performed the experiments during storage of the water suspension. in the first of these experiments, we compared stratified with unstratified seeds. we stratified some seeds by keeping them for a month in a refrigerator in moist paper towels inside of open plastic bags. for each of the following conditions, we then prepared four petri plates, each with 25 stratified seeds kept under the following conditions: • distilled water (control) • quarter-strength smoke solution • half-strength smoke solution • full strength smoke solution we did the same with petri dishes of unstratified seeds. the result was a sample of 400 stratified seeds (in sixteen plates) and 400 unstratified seeds (in sixteen plates), for a total sample size of 800 seeds. we kept all petri plates on racks underneath fluorescent lights in a temperature-controlled laboratory for a month. as needed, we added distilled water to the plates, but no additional smoke solution. in the second of these experiments, we used only unstratified seeds that had been stored in a refrigerator. we placed two control plates, two quarter-strength plates, two half-strength plates, and two full strength plates, each with 25 seeds, under the fluorescent lights (n = 200) and in darkness (a drawer adjacent to the lights; n = 200), for a total sample size of 400 seeds. temperature under the lights was slightly higher (26º c, 78.8º f) than temperature in darkness (22º c, 71.6º f). we analyzed the germination results using a chi-square analysis. we could not use a parametric test because in many of the petri dishes no seeds germinated. the preponderance of zeroes made parametric analysis invalid. results none of the 600 seeds germinated in the initial studies. from this we concluded that seed dormancy in phacelia strictiflora prevented germination in response to a flush of nutrients or a change in ph. we also concluded that dormancy of p. strictiflora seeds could not be easily broken by heat or physical abrasion. in the experiment that compared stratified with unstratified seeds, no control seeds germinated, and only one seed germinated when exposed to full-strength smoke suspension. the greatest germination occurred in half-strength and quarterstrength smoke suspension (table 1). this pattern was significant for both stratified oklahoma native plant record volume 13, december 2013 stanley a. rice and sonya l. ross 52 (p = 0.001) and unstratified (p = 0.001) seeds, analyzed separately. not only did stratified seeds display lower total germination than unstratified seeds, but they germinated more slowly. unstratified seeds began to germinate after 7 days, while stratified seeds did not begin to germinate until 21 days. contrasting with the preliminary experiments, the experiment that compared seeds exposed to light and darkness resulted in 11 of the 50 dark control seeds germinating. however, no seeds germinated in full-strength smoke suspension, and the greatest germination still occurred in halfstrength and quarter-strength smoke suspensions, as in experiment 1 (p = 0.001) (table 2). overall germination was greater in darkness than in light (p = 0.001) ___________________________________________________________________________ table 1 number and percentage of stratified and unstratified phacelia strictiflora seeds that germinated. each treatment contained 100 seeds (n = 800). unstratified seeds cold-stratified seeds full-strength smoke suspension 0 (0%) 1 (1%) half-strength smoke suspension 54 (54%) 16 (16%) quarter-strength smoke suspension 37 (37%) 5 (5%) control 0 (0%) 0 (0%) ___________________________________________________________________________ table 2 number and percentage of phacelia strictiflora seeds that germinated in the light and the dark. each treatment contained 50 seeds (n = 400). light dark full-strength smoke suspension 0 (0%) 0 (0%) half-strength smoke suspension 13 (26%) 21 (42%) quarter-strength smoke suspension 13 (26%) 27 (54%) control 0 (0%) 11 (22%) ___________________________________________________________________________ discussion and conclusions very few control seeds of phacelia strictiflora germinated. we cannot explain why 11 control seeds germinated in darkness in the final experiment, while no control seeds germinated in previous experiments. one possibility is that smoke chemicals diffused through the air from plates with smoke suspensions into some control plates, since all the plates were in the same drawer. but we have observed that a few phacelia seedlings germinate in the field in years without fire. even the post-fire herbaceous plants of the chaparral have a low level of seed germination under control conditions; a few control seeds germinated in seven of twelve chaparral species (including p. grandiflora) investigated by keeley and fotheringam (1998a). we have nevertheless demonstrated a strong smoke enhancement of p. strictiflora seed oklahoma native plant record 53 volume 13, december 2013 stanley a. rice and sonya l. ross germination, similar to that of post-fire species in fire-dependent ecological communities. halfor quarter-strength smoke suspension greatly enhanced seed germination. the failure of seeds to germinate upon exposure to the fullstrength suspension agrees with the conclusions of drewes et al. (1995) that high concentrations of smoke chemicals can inhibit germination. the possibility that the seeds require exposure to light in order to respond to smoke (todorović et al. 2005) was not supported in this species by our experiments. greater germination in the dark in the final experiment may have resulted from the slightly lower temperature. growth chambers with full temperature control were not available for this research. we demonstrated that cold stratification was unnecessary as well as insufficient for germination. in southern oklahoma, many annual species germinate and grow during the mild winters, and the same may be true of p. strictiflora. in fact, cold stratification appeared to mildly inhibit germination. in the tallgrass prairie of north america, recurring fires destroy woody vegetation and promote the re-growth of perennial grasses and forbs. however, smoke enhances germination in only about one-third of prairie species (jefferson et al. 2008). this may be due to the fact that most post-fire re-growth in the tallgrass prairie comes from the re-sprouting of perennials rather than the germination of seeds. seed germination in prairies often occurs after soil disturbance by animals. the cross timbers forest is an ecotonal community between the eastern deciduous forest and the tallgrass prairie. however, even though its common name is “prairie phacelia,” p. strictiflora appears to be more common in deciduous forest than in prairie habitats. butenolides such as karrikins may stimulate germination and promote seedling vigor even in species of plants, including agricultural plants, that do not require them (stevens et al. 2007; ghehebriot et al. 2008; lindon and menges 2008; nelson et al. 2009; hong and kang 2011). unlike these species, however, smoke enhancement of p. strictiflora germination appears to be strong enough to effectively limit its growth to post-fire conditions. in this way, p. strictiflora more closely resembles the post-fire plant species of fire-dependent ecological communities than it does the other plant species of the deciduous forest. a seed bank, produced by postfire growth after previous fires, is the only likely explanation of the massive postfire growth of p. strictiflora in 2012. too few individuals grow without fire to permit such profuse germination without a persistent seed bank. the seed bank remains largely, but not completely, dormant until a fire occurs. further, we have observed massive blooms of p. strictiflora only in cross timbers forests growing on granite substrate, but no such blooms in forests growing on limestone. the thinner soils that develop over granite may have, over time, favored the growth of p. strictiflora, allowing it to build up a seed bank in those soils. investigation of the phacelia seed bank is an opportunity for future research. smoke enhancement of seed germination has not been reported for other native oklahoma plant species. we have observed another species, selenia aurea (brassicaceae), growing abundantly after the same fire that promoted the mass flowering of p. strictiflora, but the possibility that s. aurea benefits from or requires smoke for germination has not been investigated. references chiwocha, s., k. dixon, g.r. flematti, e.l. ghisalberti, d.j. merritt, d.c. nelson, and j. stevens. 2009. karrikins: a new family of plant growth regulators in smoke. plant science 177 (4):252–256. oklahoma native plant record volume 13, december 2013 stanley a. rice and sonya l. ross 54 delange, j.h. and c. boucher. 1990. autecological studies on audouinia capitata (bruniaceae). i. plant-derived smoke as a seed germination cue. south african journal of botany 56 (6):700–703. dixon, k.w., s. roche, and j.s. pate. 1995. the promotive effect of smoke derived from burnt native vegetation on seed germination of western australian plants. oecologia 101 (2):185–192. dixon, k., d. merritt, g. flematti, and e. ghisalberti. 2009. karrikinolide: a phytoreactive compound derived from smoke with applications in horticulture, ecological restoration, and agriculture. acta horticulturae 813:155–170. drewes, f.e., m.t. smith, and j. van staden. 1995. the effect of a plantderived smoke extract on the germination of light-sensitive lettuce seed. plant growth regulation 16:205–209. flematti, gavin r. et al. 2004. a compound from smoke that promotes seed germination. science 305:977. ghehebriot, h.m. et al. 2008. smoke-water and a smoke-isolated butenolide improve germination and seedling vigour of eragrostis tef (zucc.) trotter under high temperature and low osmotic potential. journal of agronomy and crop science 194:270–277. hong, e. and h. kang. 2011. effect of smoke and aspirin stimuli on the germination and growth of alfalfa and broccoli. electronic journal of environmental, agricultural, and food chemistry 10 (2):1918–1926. jefferson, l.v., m. pennacchio, k. havens, b. forsberg, d. sollenberger, and j. ault. 2008. ex situ germination responses of midestern usa prairie species to plant-derived smoke. american midland naturalist 159 (1):251–256. keeley, j.e. and c.j. fotheringham. 1998a. smoke-induced seed germination in california chaparral. ecology 79 (7):2320– 2336. keeley, j.e. and c.j. fotheringham. 1998b. mechanism of smoke-induced seed germination in a post-fire chaparral annual. journal of ecology 86 (1):27–36. keeley, j.e., c.j. fotheringham, and p.w. rundel. 2012. postfire chaparral regeneration under mediterranean and non-mediterranean climates. madroño 59 (3):109–127. lindon, h.l. and e. menges. 2008. effects of smoke on seed germination of twenty species of fire-prone habitats in florida. castanea 73 (2):106–110. nelson, d.c., j.a. riseborough, g.r. flematti, j. stevens, e.l. ghisalberti, k.w. dixon, and s.m. smith. 2009. karrikins discovered in smoke trigger arabidopsis seed germination by a mechanism requiring gibberellic acid synthesis and light. plant physiology 149:863–873. stevens, j.c., d.j. merritt, g.r. flematti, e.l. ghisalberti, and k.w. dixon. 2007. seed germination of agricultural weeds is promoted by the butenolide 3-methyl2h-furo[2,3-c]pyran-2-one under laboratory and field conditions. plant soil 298:113–124. thomas, p.b., e.c. morris, and t.d. auld. 2007. response surfaces for the combined effects of heat shock and smoke on germination of 16 species forming soil seed banks in south-east australia. austral ecology 32:605–616. todorović, s., z. giba, s. żivković, d. grubišić, and r. konjević. 2005. stimulation of empress tree seed germination by liquid smoke. plant growth regulation 47 (2-3):141–148. turner, s.r., d.j. merritt, m.s. renton, and k.w. dixon. 2009. seed moisture content affects afterripening and smoke responsiveness in three sympatric australian native species from fireprone environments. austral ecology 34:866–877. smoke-induced germination in phacelia strictaflora by dr. stanley a. rice and dr. sonya l. ross oklahoma native plant record, volume 16, number 1, december 2016 64 oklahoma native plant record volume 16, december 2016 laura e. jardine, adam k. ryburn, and anthony j. stancampiano https://doi.org/10.22488/okstate.17.100123 effects of fire severity on habitat recovery in a mixed grass prairie ecosystem laura e. jardine adam k. ryburn anthony j. stancampiano department of biology oklahoma city university oklahoma city, ok 73106 ajstancampiano@okcu.edu key words: wichita m ountains, disturbance interaction, herbivory, competition abstract we assessed the recovery and current status of three mixed grass prairie sites 5 yr post burn in the wichita mountains wildlife refuge, indiahoma, oklahoma. these sites represent three burn histories: moderate burn, severe burn, and unburned. we used a modified point-intercept method to sample 38 habitat variables at 280 points along three transects at each site. these data were subjected to principal components analysis to assess trends in habitat structure among the sites. the first three components explained 66.6% of the variation in the dataset. component i represents a gradient from short forbs, lichen covered rocks, and minimal disturbance to areas of tall grasses and ungulate disturbance. component ii represents a gradient from tall forbs and water disturbance to areas with woody shrubs, short herbaceous litter, and graminoid and moss ground cover. component iii represents a gradient from areas with mid-level forbs, fecal matter and herbaceous litter ground cover to areas with tall grasses and bare ground. projections of the burn treatment sites onto principal components i–iii indicate that the moderate and unburned sites cluster closely on component i but are distinct along components ii and iii. we interpret our results as supporting a relationship between high severity fire and more complete nutrient cycling from accumulated litter, leading initially post fire to dense grass cover followed by increasing forb cover. this increase in forage density potentially alters the grazing patterns of large herbivores, which inflicts higher levels of disturbance. conversely, the unburned and moderate burn sites had a greater diversity of herbaceous species at lower coverage densities, perhaps resulting from reestablshiment from surviving shoots and seeds. introduction prairie ecosystems are maintained primarily through disturbance, herbivory, and competition. fire is the principal disturbance type and can be manipulated and controlled by humans, or it can have a completely uncontrolled influence on the landscape. historically, fire has been perceived in a negative context as having a detrimental effect on livestock, timber, and other human-desired resources, and has subsequently been suppressed (bland et al. 1973; archer 1989; allen and palmer 2011). this attitude has softened somewhat in recent years, and fire is commonly used as a range management tool in an attempt to maximize forage quality, remove nonpalatable tissues, and to control encroachment of woody species (archer 1994; raynor et al. 2015; collins 2016). controlled burns are typically undertaken oklahoma native plant record 65 volume 16, december 2016 laura e. jardine, adam k. ryburn, and anthony j. stancampiano when winds are low and humidity is high. perimeters are established, and, if conducted properly, specific areas are evenly burned in terms of both areal extent and fire severity (gibson and hulbert 1987; rideout-hanzak et al. 2011; gill et al. 2013; winter 2013). these types of fires allow researchers to conduct before and after studies regarding a variety of ecological effects (collins and calabrese 2012; winter et al. 2013; larson 2014). studies such as these produce valuable information due, in part, to the ability of researchers to replicate them. however, there are limits imposed on the various treatments involved by the fact that they must be controlled. this includes variables such as areal extent, fire intensity, burned patch shapes, and nonrandom site selection. wildfires, on the other hand, whether human caused or natural, more closely represent the environmental pressures under which communities have evolved. all human controls are lost, and fires take their natural course as determined by climatic conditions (e.g. drought), wind direction, wind speed, fuel volume and quality, time since last burn, and topography (gibson and hulbert 1987). for example, spring fires generally tend to increase above ground biomass production by a few dominant grass species. this results in low species richness and diversity of forbs as competition for light increases (gibson and hulbert 1987; collins and calabrese 2011; winter et al. 2013). lowland areas support increased grass biomass and lower species diversity than upland prairie. these lower areas tend to have more available nutrients and soil moisture. upland areas tend to have lower quality soils and therefore less dense vegetation. this combination of biotic (fuel volume and quality) and abiotic (elevation and moisture) factors, in addition to other physical factors such as wind speed and direction, determine fire characteristics. because studies following these natural events are initiated after the fact and as such cannot be replicated, sampling cannot be entirely randomized. additionally, there are no pre-established controls available for before and after comparison (wiens and parker 1995). in this study, we compared the recovery of plant communities, assessed by sampling horizontal and vertical habitat structure, subjected to different burn treatments five years after a wildfire (ferguson fire) in the special use area (sua) of the wichita mountains wildlife refuge (wmwr) in indiahoma, oklahoma. the objective of this paper is to describe the broad gradients of variation in the physical structure of these mixed grass prairie communities. methods and materials the wichita mountains wildlife refuge is located in comanche county, oklahoma (figure 1). it covers 23,885 ha of the central great plains ecoregion (woods et al. 2005). the sua covers 14,136 ha on roughly the northern 2/3 of the refuge. it consists of low, rounded granite mountains permeated by mixed grass prairie. mesophytic forests border streams and xeric forests consisting mostly of blackjack oak (quercus marilandica münchh.), post oak (q. stellata wangenh.), and eastern red cedar (juniperus virginiana l.) and occur on lower granite hills. the ferguson fire started on 1 september 2011, approximately 900 m east of the wmwr visitor center. southerly winds rapidly pushed the fire northward into the sua where the landscape was subjected to an incinerating burn that resulted in no remaining living vegetation. as the fire moved northward, it completely jumped small pockets of the landscape leaving them unburned. after burning through the refuge and exiting the north boundary, a northerly wind shift occurred pushing the fire southwest and back onto the refuge. this wind-shifted leg of the fire was less intense than the initial blaze due to light precipitation and light winds resulting in a moderately burned 66 oklahoma native plant record volume 16, december 2016 laura e. jardine, adam k. ryburn, and anthony j. stancampiano fi gu re 1 l oc at io n of th e w ic hi ta m ou nt ai ns w ild lif e r ef ug e, c om an ch e c ou nt y, o kl ah om a (in se t) a nd th re e st ud y si te s oklahoma native plant record 67 volume 16, december 2016 laura e. jardine, adam k. ryburn, and anthony j. stancampiano fi gu re 2 f ire s ev er ity in th e w ic hi ta m ou nt ai ns w ild lif e r ef ug e, c om an ch e c ou nt y, o kl ah om a 68 oklahoma native plant record volume 16, december 2016 laura e. jardine, adam k. ryburn, and anthony j. stancampiano landscape where not all vegetation was destroyed. a total of 11,270 ha was burned on the wmwr. the ferguson fire followed the hottest summer on record in oklahoma since 1895 and moderate to extreme drought conditions in comanche county since may 2010 (noaa 2010). we established three survey sites on 7 may 2016, one for each burn treatment (severe, moderate, and unburned), in mixed grass prairie of the sua. each site consisted of three 90 m transect arms extending from a center node (c-node) with one arm oriented in a north-south direction (0 to 180°). the other two arms extended from the c-node to the southeast (135°) and southwest (225°) for a linear total of 270 m/site. the maximum distance between sites was 2,351 m (unburned to moderate burn), and the minimum distance was 935 m (moderate to severe burn). the distance from the unburned to the severe burn site was 1,605 m (see figure 1). site elevations were within a 6.7 m range with the severe burn at 589.5 m, moderate burn at 582.8 m, and the unburned site at 585.5 m. we sampled the physical structure at each site from 5–11 june 2016. we assigned fire impact as unburned, moderate burn, and severe burn as determined by stambaugh et al. (2015) (figure 2). these classifications were derived through a combination of remote sensing, ground truthing, and modelling. unburned indicates that the area after the fire was indistinguishable from pre-fire conditions. the moderate burn class represents a mixture of effects on the dominant vegetation with some patches of above ground cover completely removed while others show little or no change and low mortality of the dominant vegetation. high severity burn indicates complete consumption of the canopy (stambaugh et al. 2015). we used a modified point-intercept method to sample 38 habitat variables along each transect at each site. these variables included measures of ground disturbance, ground cover, and vertical cover (table 1). to determine ground disturbance and cover, we passed a 3 mm x 1 m rod vertically through the vegetation and onto the substrate at 0.5 m horizontal intervals along each transect. we recorded the ground disturbance and cover type at the point of contact. ground disturbance type was determined by obvious alteration of ground cover, if any. at the same time, we determined vertical structure in decimeter intervals (1–10) by recording the interval at which any vertical cover contacted the rod. we sampled a total of 270 points at each site (figure 3). we used these data in a principalcomponents analysis (pca) to assess patterns in habitat structure 5 yr post fire. pca is an unconstrained ordination method that is useful for visualizing broad patterns of covariation in a multivariate data set (anderson and willis 2003). all calculations were performed using nt-sys (rohlf 1998). we mean-centered the raw data and calculated correlations among the variables. we then projected the standardized data onto eigenvectors projected from the correlation matrix. oklahoma native plant record 69 volume 16, december 2016 laura e. jardine, adam k. ryburn, and anthony j. stancampiano table 1 categories and description of variable codes used in point-intercept sampling of three burn treatments in the wmwr. vertical cover (vc) is measured in decimeter categories. category variable no. variable code habitat variable description disturbance (dist) 1 2 3 4 n un w h none ungulate water human ground cover (gc) 5 6 7 8 9 10 11 12 13 14 15 16 cg cf l m ac lh gr co bo wa bg fm crown graminoid crown forb lichen moss algae/cyanobacteria litter herbaceous gravel <7.5cm cobble >7.5-25cm boulder >25cm water bare ground fecal matter vertical cover (vc) 17-20 lhv herbaceous litter vertical hits 21-28 fcg graminoid foliage cover vertical hits 29-34 fcf forb foliage cover vertical hits 35-38 fcs shrub foliage cover vertical hits 70 oklahoma native plant record volume 16, december 2016 laura e. jardine, adam k. ryburn, and anthony j. stancampiano figure 3 author laura jardine sampling vertical structure at intense burn site in wichita mountains wildlife refuge we surveyed the flora at each site by recording the presence of each species encountered (table 2). plant species identification followed the flora of oklahoma: keys and description (tyrl et al. 2015). results principal components analysis of 38 habitat variables produced three axes that accounted for 66.6% of the variation. principal component i (pc i) explained 32.8%, pc ii 19.5%, and pciii 14.3% of the variation. component i represents a gradient from short forbs, lichen covered cobble and boulders, and low disturbance to areas of tall grasses and ungulate disturbance (table 3). component ii represents a gradient from tall forbs and water disturbance to areas with woody shrubs, herbaceous litter near the surface, and graminoid and moss ground cover. component iii represents a gradient from areas with mid-level forbs, fecal matter, and herbaceous litter ground cover to areas with tall grasses and bare ground cover. projections of the burn treatment sites onto pc i, pc ii, and pc iii indicate that the moderate and unburned sites cluster closely on pc i but are distinct along pc ii and pc iii (figure 4). the severe burn has the highest positive loadings along pc i and is intermediate with respect to pc ii (see figure 4). the three transects for unburned and moderate burn sites cluster tightly within sites along pc iii, but the two sites themselves are separated. the transects in the severe burn are widely separated along pc iii. the plant species composition of the three sites is as follows: unburned – 40 species of 20 families; moderate burn – 40 species of 23 families; and severe burn – 28 species of 13 families (see table 2). discussion there have been few studies that inventory the flora of the wichita mountains (eskew 1938; osborn and allan 1949; buck 1977; collins and barber 1986; carter et al. 2008). other studies associate mixed grass prairie floristic components of the wmwr with specific mammal assemblages (osborn and allan 1949; stancampiano and caire 1995). stancampiano and schnell (2004) assessed small mammal distributions across nearby fort sill using, among others, vertical structure of vegetation. it appears that no studies have been published of the vertical structure or cover types on the wmwr prior to this study. floristic composition across all sites is consistent with unpublished seasonal checklists and published floras of the area (buck 1977; oklahoma native plant record 71 volume 16, december 2016 laura e. jardine, adam k. ryburn, and anthony j. stancampiano table 2 plant community composition of three burn treatments in the wichita mountains wildlife refuge species common name family moderate burn severe burn no burn allium canadense canada garlic amaryllidaceae x x x daucus carota wild carrot apiaceae x x ptilimnium nuttallii nuttall's mockbishopweed apiaceae x asclepias viridis green antelope horn apocynaceae x x x yucca glauca small soapweed asparagaceae x achillea millefolium yarrow asteraceae x x x ambrosia psilostachya western ragweed asteraceae x x x artemisia ludoviciana louisiana sagewort asteraceae x x chaetopappa asteroides least daisy asteraceae x x x cirsium undulatum wavyleaf thistle asteraceae x x coreopsis lanceolata lanceleaf coreopsis asteraceae x x echinacea angustifolia black sampson asteraceae x x gaillardia pulchella indian blanket asteraceae x x x helenium amarum bitter sneezeweed asteraceae x x thelesperma filifolium plains greenthread asteraceae x vernonia baldwinii baldwin ironweed asteraceae x x lepidium virginicum virginia pepperrwort brassicaceae x x x paysonia auriculata earleaf bladderpod brassicaceae x echinocereus reichenbachii lace hedgehog cactus cactaceae x opuntia humifusa var. humifusa prickly pear cactaceae x x triodanis perfoliata ssp. biflora small venus looking-glass campanulaceae x symphoricarpos orbiculatus buckberry caprifoliaceae x x x valerianella radiata cornsalad caprifoliaceae x tradescantia ohiensis smoothstalk spiderwort commelinaceae x x x cuscuta cuspidata cusp dodder convolvulaceae x sedum nuttallii yellow stonecrop crassulaceae x x juniperus virginiana eastern red cedar cupressaceae x carex sp. sedge cyperaceae x x eleocharis montevidensis sand spikesedge cyperaceae x x amorpha canescens leadplant fabaceae x x baptisia australis blue wild indigo fabaceae x x x lespedeza virginica slender lespedeza fabaceae x 72 oklahoma native plant record volume 16, december 2016 laura e. jardine, adam k. ryburn, and anthony j. stancampiano table 2 (continued) species common name family moderate burn severe burn no burn mimosa nuttallii catclaw sensitive brier fabaceae x x quercus marilandica blackjack oak fagaceae x quercus stellata post oak fagaceae x geranium carolinianum carolina geranium geraniaceae x x juncus sp. rush juncaceae x callirhoe involucrata low poppymallow, winecup malvaceae x oenothera glaucifolia false guara onagraceae x x x oenothera suffrutescens scarlet beeblossom onagraceae x x x castilleja purpurea var. citrina citron paintbrush orobanchaceae x oxalis stricta sheep sorrel oxalidaceae x nuttallanthus texanus texas toadflax plantaginaceae x x plantago aristata bottlebrush plantain plantaginaceae x x plantago virginica paleseed plantain plantaginaceae x alopecurus carolinianus carolina foxtail poaceae x bromus japonicus japanese brome poaceae x bromus tectorum cheatgrass poaceae x x x dichanthelium oligosanthes scribner's panicum poaceae x x elymus repens quackgrass poaceae x hordeum pusillum little barley poaceae x x mnesithea cylindrica carolina jointtail grass poaceae x panicum virgatum switch grass poaceae x x schizachyrium scoparium little bluestem poaceae x x geum canadense white avens rosaceae x prunus angustifolia chickasaw plum (sand plum) rosaceae x x stenaria nigricans var. nigricans narrowleaf bluet rubiaceae x selaginella peruviana sheldon selaginella selaginellacea x solanum carolinense carolina groundcherry solanaceae x x glandularia canadensis rose verbena verbenaceae x x oklahoma native plant record 73 volume 16, december 2016 laura e. jardine, adam k. ryburn, and anthony j. stancampiano table 3 summary of principal components analysis of 38 habitat variables for nine burn treatment sites variable pci pcii pciii vc-lhv 1 0.3276 0.5352 0.4793 vc-fcg 1 -0.7487 -0.4238 0.0202 vc-fcf 1 -0.8446 0.1512 -0.0202 vc-lhv 2 -0.0607 0.3614 0.3851 vc-fcg 2 -0.5746 -0.3075 0.1936 vc-fcf 2 -0.2223 -0.0764 -0.5285 vc-lhv 3 -0.2355 0.3395 -0.3065 vc-fcg 3 0.9065 0.3400 -0.1269 vc-fcf 3 0.0143 -0.0071 -0.7783 vc-lhv 4 -0.4187 0.4370 0.0380 vc-fcg 4 0.8107 0.3034 0.2569 vc-fcf 4 0.5395 -0.5924 -0.1209 vc-fcs 4 -0.3047 0.7298 -0.1569 vc-fcg 5 0.9105 0.0613 0.1999 vc-fcf 5 0.2135 -0.5646 -0.1058 vc-fcs 5 -0.3047 0.7298 -0.1569 vc-fcg 6 0.4988 0.6896 0.4904 vc-fcf 6 0.6083 -0.3815 -0.2211 vc-fcs 6 -0.3047 0.7298 -0.1569 vc-fcg 7 0.8268 0.1727 0.5080 vc-fcs 7 -0.3047 0.7298 -0.1569 vc-fcg 9 0.5711 0.3764 0.6516 74 oklahoma native plant record volume 16, december 2016 laura e. jardine, adam k. ryburn, and anthony j. stancampiano table 3 (continued) variable pci pcii pciii dist-n -0.9067 0.2242 0.0895 dist-un 0.9394 -0.1228 -0.0914 dist-w -0.6050 -0.5713 0.4695 dist-h 0.6083 -0.3815 -0.2211 gc-cg 0.1550 0.8001 0.0796 gc-cf -0.5041 -0.1689 0.0406 gc-l -0.8289 -0.0900 0.3338 gc-m -0.3086 0.8362 -0.1531 gc-ac -0.2352 0.3120 -0.3838 gc-lh 0.5002 -0.2972 -0.7984 gc-gr -0.2206 0.2877 -0.4824 gc-co -0.8596 0.0177 0.2127 gc-bo -0.8100 -0.2332 0.4047 gc-wa -0.6050 -0.5713 0.4695 gc-bg 0.3922 -0.1612 0.7967 gc-fm 0.2043 -0.0715 -0.5141 % total variance 32.77 19.54 14.28 cumulative % 32.77 52.31 66.58 oklahoma native plant record 75 volume 16, december 2016 laura e. jardine, adam k. ryburn, and anthony j. stancampiano figure 4 projections of 3 study plots (nc=north to central node; sec=southeast to central node; swc=southwest to central node) based on 38 variables onto principal components i, ii, and iii in the special use area of the wichita mountains wildlife area nc sec swc nc sec swc nc sec swc -0.6 -0.4 -0.2 0 0.2 0.4 0.6 0.8 1 -0.6 -0.4 -0.2 0 0.2 0.4 0.6 0.8 1 1.2 pc ii moderate burn unburned severe burn nc sec swc nc sec swc nc sec swc -0.6 -0.4 -0.2 0 0.2 0.4 0.6 0.8 -0.6 -0.4 -0.2 0 0.2 0.4 0.6 0.8 1 1.2 pc ii i pc i moderate burn unburned severe burn 76 oklahoma native plant record volume 16, december 2016 laura e. jardine, adam k. ryburn, and anthony j. stancampiano carter et al. 2008). the unburned and moderate burn sites had higher spring species composition and cover of forbs when compared to the severe burn site which had higher cover of grasses. the gradients produced by the pca are consistent with other prairie fire studies with regard to plant species richness and the physical structure of the plant community (gibson and hulbert 1987; collins and calabrese 2012; winter et al. 2013). our study involved three study sites located within 2.5 km of each other, which reflects similar abiotic and biotic conditions. many studies measure differences in post fire prairie communities based on frequency of fires (gibson and hulbert 1987; collins and calabrese 2012; winter et al. 2013). using controlled fires in the tallgrass konza prairie, gibson and hulbert (1987) concluded that time since the last fire was the greatest determinant of prairie species composition. they also found that cover of grasses decreased over time, while cover of forbs and woody species increased. as in most controlled burns, fire severity was not taken into account. their study took place prior to the reintroduction of bison to the konza prairie; therefore, there was no effect on vegetation from grazing. we made the assumption, a priori, that grazing by large herbivores (bison, elk, and longhorn cattle) was equal across all three burn treatments, post fire. our analysis infers that large herbivores do indeed prefer the severe burn site forage at this point in recovery. we did, however, observe these large herbivores at all three sites. as indicated in studies of tallgrass ecosystems (fuhlendorf and engle 2004; allred et al. 2011), it is possible that fire and grazing interact in landscapes to increase heterogeneity, as fire concentrates grazing activity to certain burned patches thereby reducing grazing in others. our study supports the findings of many others in that fire severity also affects the recovery of vegetation, including not only composition but also its vertical and horizontal structure (gibson and hulbert 1987; collins and calabrese 2012; winter et al. 2013). this follows the pattern of allogenic change due to fire fostering an increased probability of autogenic change (e.g., grazing) and its subsequent effects across the landscape. acknowledgements we would like to thank the wichita mountains wildlife refuge and dan mcdonald for access to the special use area and for logistical support. this study was supported, in part, by a 2016 cairs ocu undergraduate research grant and the beta beta beta research scholarship fund. we thank the dean’s office in the petree college of arts and science and the department of biology at ocu for use of a field vehicle and financial support for field equipment and travel. we thank matt white for creating the site location map (figure 2). finally, thanks to two reviewers for insightful comments and constructive suggestions. literature cited allen, m.s. and m.w. palmer. 2011. fire history of a prairie/forest boundary: more than 250 years of frequent fire in a north american tallgrass prairie. journal of vegetation science 22:436–444. allred, b.w., s.d. fuhlendorf, d.m. engle, and r.d. elmore. 2011. ungulate preferences for burned patches reveal strength of fire-grazing interaction. ecology and evolution 1:132–144. anderson, m.j. and t.j. willis. 2003. canonical analysis of principal coordinates: a useful method of constrained ordination for ecology. ecology 84:511–525. archer, s. 1989. have southern texas savannas been converted to woodlands in recent history? american naturalist 134:545–561. oklahoma native plant record 77 volume 16, december 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winter, s.l., k.r. hickman, c.l. goad, s.d. fuhlendorf, and m.s. gregory. 2013. seasonal fires, bison grazing, and the tallgrass prairie forb arnoglossum plantagineum raf. natural areas journal 33:327–338. 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 photographs) (map scale 1:1,250,000). reston (va): u.s. geological survey. https://www.ncdc.noaa.gov/sotc/drou ght/201005. https://www.ncdc.noaa.gov/sotc/drought/201005 https://www.ncdc.noaa.gov/sotc/drought/201005 effects of fire severity on habitat recovery in a mixed grass prairie ecosystem by ms. laura e. jardine, dr. adam k. ryburn, and dr. anthony j. stancampiano 2019 oklahoma native plant record 1 oklahoma native plant r ecord journal of the okla hom a native plant society p. o. box 14274 tulsa, oklahoma 74159-1274 volume 19, december 2019 issn 1536-7738 http://ojs.library.okstate.edu/osu/ editor: gloria caddell production editor: paula shryock electronic production editor: sandy graue manuscript editor: chad king technical advisor: erica corbett the purpose of onps is to encourage the study, protection, propagation, appreciation, and use of the native plants of oklahoma. membership in onps is open to any person who supports the aims of the society. onps offers individual, student, family, and life memberships. officers and board members president: bill farris vice-president: donna horton secretary: connie murray treasurer: mary korthase past president: bruce smith directors at large: kathy doss jim elder ray luth joe roberts janet thomas rahmona thompson chapter chairs: central: patrick bell cross timbers: elaine lynch northeast: teresa blue mycology: nancy hamill committee chairs: historian: fran stallings publicity/merchandise: barbara klein betty kemm award: sue amstutz awards: constance murray membership database: sandy graue membership database ed.: tina julich mailings/printings: sandy graue gaillardia editor: lynn michael color oklahoma: alicia nelson webmaster: adam ryburn web editors: joe roberts, sandy graue http://www.oknativeplants.org articles (c) the authors journal compilation (c) oklahoma native plant society except where otherwise noted, this work is licensed under a creative commons attribution noncommercial-sharealike4.0 international license, https://creativecommons.org/licenses/ by-nc-sa/4.0/, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly attributed, not used for commercial purposes, and, if transformed, the resulting work is redistributed under the same or similar license to this one. https://doi.org/10.22488/okstate.20.100000 http://ojs.library.okstate.edu/osu/ http://www.oknativeplants.org/ https://webmail.suddenlink.net/do/redirect?url=https%253a%252f%252fcreativecommons.org%252flicenses%252f&hmac=c55f81a2d88a183f74295dc18f7bbb4a https://webmail.suddenlink.net/do/redirect?url=https%253a%252f%252fdoi.org%252f10.22488%252fokstate.19.100000&hmac=a73ec5e971d5bca0ed3e9607a4f15f73 2 oklahoma native plant record volume 19, december 2019 oklahoma native plant record volum e 19 table of contents foreword .................................................................................................................................................... 3 historical land cover along the deep fork river: an analysis of vegetation composition and distribution of the deep fork national wildlife refuge, okmulgee county, oklahoma, circa 1897 ........................................................................................... 4 bruce hoagland, rick thomas, and daryn hardwick a floristic inventory of the john w. nichols scout ranch, canadian county, oklahoma ................................................................................................................................................. 17 abby crosswhite and adam k. ryburn a walk through the mcloud high school oak-hickory forest with a checklist of the woody plants. ............................................................................................................ 30 bruce a. smith sexual reproduction of kudzu (pueraria montana [lour.] merr.) in oklahoma ............................. 52 eric b. duell and karen r. hickman critic’s choice essay: seeking a special plant ......................................................................................... 58 paul buck† editorial policies and procedures ......................................................................................................... 60 five year index to oklahoma native plant record ............................................... inside back cover † indicates an author who is deceased cover photo: dalea purpurea vent. (purple prairie clover) by john cleal for the 2008 onps photo contest five year index to oklahom a n ative plant r ecord volume 13 4 ecology and taxonomy of water canyon, canadian county, oklahoma, m. s. thesis, constance a. taylor 29 a checklist of the vascular flora of the mary k. oxley nature center, tulsa county, oklahoma, amy k. buthod 48 smoke-induced germination in phacelia strictaflora, stanley a. rice and sonya l. ross 55 critic’s choice essay: a calvacade of oklahoma botanists in oklahoma – contributors to our knowledge of the flora of oklahoma, ronald j. tyrl and paula a. shryock volume 14 4 flora of kiowa county, oklahoma, m. s. thesis, lottie opal baldock 38 gardens of yesteryear, sadie cole gordon 43 oklahoma deciduous trees differ in chilling enhancement of budburst, stanley a. rice and sonya l. ross 50 mapping distribution in oklahoma and raising awareness: purple loosestrife (lythrum salicaria), multiflora rose (rosa multiflora), and japanese honeysuckle (lonicera japonica), katherine e. keil and karen r. hickman 67 non-twining milkweed vines of oklahoma: an overview of matelea biflora and matelea cynanchoides (apocynaceae), angela mcdonnell 80 critic’s choice essay: pollination ecology of our native prairie plants, gloria m. caddell volume 15 4 preface to first flowering dates for central oklahoma, wayne elisens 6 first flowering dates for central oklahoma, ben osborn 19 forest structure and fire history at lake arcadia, oklahoma county, oklahoma (1820–2014), chad king 31 interplanting floral resource plants with vegetable plants enhances beneficial arthropod abundance in a home garden, chrisdon b. bonner, eric j. rebek, janet c. cole, brian a. kahn, and janette a. steets 49 contributions to the flora of cimarron county and the black mesa area, amy k. buthod and bruce w. hoagland 78 antifungal activity in extracts of plants from southwestern oklahoma against aspergillus flavus, tahzeeba frisby and cameron university students 96 kudzu, pueraria montana (lour.) merr. abundance and distribution in oklahoma, marli claytor and karen r. hickman 105 critic’s choice essay: mistletoe, phoradendron serotinum (raf.) johnston, paul buck volume 16 4 pollination ecology of sabatia campestris nutt. (gentianaceae), constance e. taylor 10 the structure of the gynostegium, breeding system, and pollination ecology of spider milkweed, asclepias viridis walt. (apocynaceae), m. s. thesis, shang-wen liaw 45 a floristic inventory of the university of oklahoma’s kessler atmospheric and ecological field station, mcclain county, oklahoma, amy k. buthod and bruce w. hoagland 64 effects of fire severity on habitat recovery in a mixed grass prairie ecosystem, laura e. jardine, adam k. ryburn, and anthony j. stancampiano 78 critic’s choice essay: a conversation with a small beetle, paul buck volume 17 4 a study of the flowering plants of tulsa county, oklahoma, exclusive of the grasses, sedges, and rushes, m.s. thesis, maxine b. clark† 37 laboratory studies of allelopathic effects of juniperus virginiana l. on five species of native plants, erica a. corbett and andrea lashley 53 vascular flora of e. c. hafer park, edmond, oklahoma, gloria m. caddell, katie christoffel, carmen esqueda, and alonna smith 69 first record of chorioactis geaster from oklahoma, clark l. ovrebo and sheila brandon 72 critic’s choice essay: allelopathy, paul buck† oklahoma native plant society p.o. box 14274 tulsa, oklahoma 74159-1274 _________________________________________________________________________ in this issue of oklahoma native plant record volume 18, december 2018: _________________________________________________________________________ 4 characteristics of a bottomland hardwood forest at arcadia lake, edmond, oklahoma, with special emphasis of green ash (fraxinus pennsylvanica marshall) chad b. king and joseph a. buck 19 presence of pueraria montana (lour.) merr. var. lobata (willd.) maesen & s.m. almeida ex sanjappa & predeep (kudzu vine) in tulsa county, oklahoma isaac walker and paulina harron 24 comparative transpiration studies on the invasive eastern redcedar (juniperus virginiana l.) and adjacent woody trees adjoa r. ahedor, bethany spitz, michael cowan, j’nae miller, and margaret kamara 38 new record of myriopteris lindheimeri (hook.) j. sm. in kiowa county, oklahoma bruce a. smith 45 anther number, anther apical appendages, and pollination biology of calyptocarpus vialis (heliantheae: asteraceae) james r. estes 52 critic’s choice essay: myrmecochory paul buck five year index to okla homa native plant r ecord – inside back cover oklahoma native plant record, volume 15, number 1, december 2015 oklahoma native plant record 31 volume 15, december 2015 chrisdon b. bonner, et al. https://doi.org/10.22488/okstate.17.100113 interplanting floral resource plants with vegetable plants enhances beneficial arthropod abundance in a home garden chrisdon b. bonner1 eric j. rebek2 janet c. cole3 brian a. kahn3 janette a. steets1 janette.steets@okstate.edu 1oklahoma state university department of botany 301 physical sciences stillwater, ok 74078 2oklahoma state university department of entomology and plant pathology 127 noble research center stillwater, ok 74078 3oklahoma state university department of horticulture and landscape architecture 358 agricultural hall stillwater, ok 74078 keywords: cowpea, herbivory, native ornamentals, natural enemies, pollinators, polyculture, tomato abstract we examined whether interplanting vegetable and ornamental flowering plants reduces herbivory and enhances photosynthetic rate, plant growth, natural enemy abundance, and pollinator visitation relative to monoculture plantings. we found no evidence of physiological or growth costs due to growth in polyculture. herbivore damage to plants did not differ with planting regime. natural enemies occurred in greater abundance in polycultures compared to monocultures. pollinator diversity was enhanced in some polyculture plots. we suggest that interplanting vegetable and flowering ornamental plants at small spatial scales may improve plant health and reproduction through natural pest control and a diversified pollinator pool. introduction habitat manipulation strategies regulate pest populations in managed landscapes by enhancing the abundance of arthropod predators and parasitoids (natural enemies) by provisioning additional plant-based resources (i.e., nectar, pollen, alternative prey, or shelter) (rebek et al. 2005, 2006; fiedler et al. 2008). these same strategies may also have beneficial effects for pollinator abundance and diversity due to an increased abundance of flowering plants in the managed landscape (tuell et al. 2008). a common habitat manipulation strategy that often benefits natural enemies and pollinators in managed landscapes is the use of polycultures, the cultivation of multiple plant species together. relative to most monoculture plantings, polycultures offer beneficial arthropods (i.e., natural enemies and pollinators) greater floral resources (i.e., nectar and pollen rewards) throughout the growing season, alternative prey, and increased habitat structure and availability of nesting sites (andow and risch 1985; andow 1991; landis et al. 2000, 2005; hooks and johnson 2003). polycultures may also provide improved microhabitats for plants and arthropods, such as increased shade and protection from wind, relative to most monocultures (andow 1991; landis et al. 2000). in addition to enhancing beneficial mailto:janette.steets@okstate.edu mailto:janette.steets@okstate.edu 32 oklahoma native plant record volume 15, december 2015 chrisdon b. bonner, et al. arthropod abundance, polycultures often support lower pest arthropod abundance than monocultures (kloen and altieri 1990; nicholls and altieri 2004; ponti et al. 2007; isaacs et al. 2009). at small spatial scales, such as those of home gardens, polycultures are a particularly attractive alternative to cultivation techniques that require heavy pesticide applications to control pest arthropods. home gardeners commonly use pesticides to control pest arthropods (sadof et al. 2004); in the united states, 16% of all insecticides applied annually are used in residential gardens and lawns (u. s. epa 2011). widespread residential pesticide use poses significant threats to human health and the environment by increasing the incidence of pesticide poisonings (pimentel et al. 1992; u. s. epa 2009), reducing stream and ground water quality (cohen 2010), and killing non-target organisms (e.g., insect pollinators, aquatic fauna) (johansen and mayer 1990; pimentel et al. 1992; relyea 2009). effective alternatives to residential pesticide applications are needed to improve safety, minimize effects on nontarget organisms, and reduce environmental contamination. to date, most studies of polyculture techniques have examined the role of plantbased resources for natural enemy ecology and in regulating natural enemy populations (fiedler et al. 2008). what remains less well studied is whether planting polyculture gardens of vegetable and flowering ornamental plants has other beneficial effects for garden crops. we hypothesized that plants grown in polycultures will have higher rates of pollinator visitation as well as higher abundance of natural enemies relative to monoculture plantings. with an increase in natural enemy abundance (landis et al. 2000), we hypothesized that plants grown in polycultures will experience reduced rates of herbivory compared to monoculture plantings. herbivore damage is known to adversely affect photosynthesis and plant growth (crawley 1997; zangerl et al. 2002). thus, if growing plants in polycultures reduces herbivory, then we hypothesized that plants in polycultures will have higher rates of photosynthesis and growth relative to monoculture plantings. accordingly, the first objective of this study was to examine whether polycultures of vegetable and flowering ornamental plants reduce herbivory relative to monocultures. our second objective was to examine whether polycultures of vegetable and ornamental plants enhance photosynthetic rate and growth relative to monocultures. our third objective was to examine whether polycultures enhance pollinator visitation and pollinator diversity relative to monoculture plantings. our fourth objective was to examine whether polycultures enhance natural enemy abundance relative to monocultures. materials and methods garden design we conducted this study at the botanic garden at oklahoma state university (stillwater, ok; 36°07'08.6" n, 97°06'04.5" w) from april 23, 2009, to september 1, 2009. seven plant species were included in the study. four native, commonly cultivated ornamental species were largeflower tickseed (coreopsis grandiflora hogg ex sweet ‘early sunrise’), purple coneflower (echinacea purpurea (l.) moench), blanketflower (gaillardia x grandiflora van houtte ‘arizona sun’), and goldenrod (solidago sp. ‘wichita mountains’). three commonly cultivated vegetable species were cilantro (coriandrum sativum l.), tomato (solanum lycopersicum l. ‘mountain fresh plus’), and cowpea (vigna unguiculata (l.) walp ‘early scarlet’). we chose vegetable species that would be typical of an oklahoma or southern u.s. home garden (hillock and simons 2002). while tomato and cowpea are self-fertile, visitation by insects, primarily bees, improves oklahoma native plant record 33 volume 15, december 2015 chrisdon b. bonner, et al. reproductive success (free 1993). all flowering plants included in our study provide nectar and pollen to beneficial arthropods, and the chosen species overlap in blooming period, ensuring a continuous supply of floral resources. we used a randomized complete block design consisting of four blocks of nine experimental plots each. plots within each block were randomly assigned to one of nine planting treatments. each plot measured 1 m x 2 m and was separated from other plots by a 1 m mulched border. all plots and borders were kept free of weeds by hand pulling. all plots were composed of native soil (norge loam, finesilty, mixed, thermic udic paleustolls) and were provided supplemental water by drip irrigation. plants were not fertilized, as adequate plant mineral nutrients were available from fertilization of previous trials. the nine planting treatments included monocultures of each of the plant species (seven plots) and two different polycultures to add more generality to our results. one polyculture consisted of largeflower tickseed, goldenrod, cilantro, and tomato (one plot; ‘polyculture one’), and the other polyculture consisted of purple coneflower, blanketflower, goldenrod, and cowpea (one plot; ‘polyculture two’). monocultures of the four ornamental species were planted with 18 plants/plot on april 23–24, 2009, using established nursery stock. tomato monocultures were planted on april 25, 2009, using established nursery stock and included two plants/plot; plants were centered in each plot and spaced 60 cm apart within the row. we seeded the monocultures of cilantro on april 25, 2009, at a density of 240 seeds per 1 m row, with six rows per plot. monocultures of cowpea were seeded on may 22, 2009, at a density of 20 seeds per 1 m row, with two rows per plot. we later thinned cowpea to 10 plants per 1 m row where stands permitted. within polyculture one plots, we planted six largeflower tickseed, three goldenrod, two tomatoes, and seeded one row of cilantro at a density of 240 seeds per 1 m row. we planted goldenrod and largeflower tickseed on april 23–24, 2009. we planted tomatoes and seeded cilantro on april 25, 2009. within polyculture two plots, we planted three purple coneflower, three blanketflower, three goldenrod, and seeded two rows of cowpea at a density of 20 seeds per 1 m row. we later thinned cowpeas to 10 plants per row where stands permitted. we planted cowpeas on may 22, 2009, and purple coneflower, blanketflower, and goldenrod on april 23–24, 2009. the plant species were sown at densities recommended by the oklahoma cooperative extension service. different species were not planted on the same date because 1) a planting date of april 25 was too early for cowpea, which was directseeded and requires warm soils for proper germination; and 2) ornamental plants (purple coneflower, blanketflower, and goldenrod) were planted at later dates as a result of plant availability. we did not observe any shading of later-planted species by those planted earlier. cilantro and goldenrod did not establish in monoculture or polyculture. in addition, several plots of the other plant species did not establish well. thus, our analyses included four plots of largeflower tickseed, three plots of purple coneflower, four plots of blanketflower, three plots of tomato, two plots of cowpea, two plots of polyculture one, and three plots of polyculture two. herbivory to determine whether planting regime influenced rates of herbivory, we quantified leaf damage on two plants of each species per plot twice during the growing season (june and july). for each plant, we estimated herbivore damage on one standard module per plant (turcotte et al. 2014) by counting the total number of leaves and the number of leaves with 34 oklahoma native plant record volume 15, december 2015 chrisdon b. bonner, et al. herbivore damage on the module (i.e., branch, rosette). the plant module varied for each species based on plant morphology; we counted all of the leaves on purple coneflower, only the leaves of the basal rosette on blanketflower, the leaves of one stem on largeflower tickseed, and the leaves of one lower branch on tomato and cowpea. for these same plants, we then recorded the number of leaves on the module damaged by herbivory. we calculated percent of damaged leaves for each plant as the total number of damaged leaves divided by the total number of leaves per module. we quantified herbivore damage on plants rather than inventorying herbivores, as herbivore damage represents a more comprehensive temporal perspective on herbivory in these plots; however, common herbivores in these crops included aphids (family aphididae), tomato hornworms (manduca quinquemaculata), flea beetles (family chrysomelidae, tribe alticini), squash bugs (anasa tristis), cucumber beetles (acalymma sp. and diabrotica sp.), and spider mites (family tetranychidae). plant height and photosynthetic measurements to determine whether planting regimes (i.e., monoculture versus polyculture) affected traits related to plant health, we quantified height to the nearest centimeter and measured light-saturated photosynthetic rate using an infrared gas analyzer (li-6400, li-cor, inc.; lincoln, ne). photosynthetic rate was quantified for one newly expanded leaf from each of two plants per species per plot. we recorded these measurements twice during the growing season (july and august). each month, all measurements were taken within a three-day period between 09:00– 13:00 cdst on sunny days. we standardized leaf chamber conditions with a temperature of 30c, photosynthetically active radiation (par) at 1500 μmol m-2s-1, and co2 concentration of 400 ppm. because calculations of photosynthetic rate are based in part on leaf surface area, we collected leaves that did not fill the entire leaf chamber and later determined leaf surface area using image analysis software (imagej, national institutes of health freeware; bethesda, md). pollinator abundance and composition throughout the summer, we observed insect visitation to flowers within our experimental plots during 15 min observation periods. observations were limited to sunny days when the wind was calm. throughout the summer, observation times varied throughout the day (between 07:00–17:00 cdst) to capture a wider diversity of insect visitors. on a given day, we rotated observations among experimental plots (kearns and inouye 1993). during each 15 min observation period, we recorded insect visitation at the flower or inflorescence level, recording visits to all open flowers or inflorescences on several plants within each experimental plot. observations of tomato and cowpea were conducted at the flower level; whereas, observations of all other plant species were conducted at the inflorescence level. within a plot, we observed as many flowers or inflorescences on as many plants as was possible at one time, including simultaneous observations of several plant species in polycultures. we recorded floral visitors from four insect orders: beetles (coleoptera); wasps, honey bees, bumble bees, and small-bodied bees (hymenoptera); true flies (diptera); and butterflies (lepidoptera). we observed monoculture plots for a total of 7.75 h over the course of the experiment. total duration of observations varied among species in monocultures (tomato, 1.25 h; cowpea, 1 h; purple coneflower, 1.5 h; largeflower tickseed, 1.5 h; blanketflower, 2.5 h). we observed polyculture plots for a total of 9 h over the oklahoma native plant record 35 volume 15, december 2015 chrisdon b. bonner, et al. course of the experiment. as with monocultures, the total duration of observations varied among species in polycultures (tomato, 1.25 h; cowpea, 1.75 h; purple coneflower, 1.5 h; largeflower tickseed, 2.5 h; blanketflower, 3 h). natural enemy abundance and composition we sampled natural enemies using 7.5 x 13 cm yellow sticky cards (hoback et al. 1999) every two weeks throughout the growing season, for a total of seven sample dates over the course of the experiment. around mid-morning on selected days, two sticky cards per plot were placed 1 m above ground level on stakes and left for 48 h. we used a compound stereomicroscope to identify and sort specimens from the sticky cards into twelve groups of arthropods: spiders (order araneae), rove beetles (family staphylinidae), lady beetles (family coccinellidae), hover flies (family syrphidae), tachinid flies (family tachinidae), minute pirate bugs (family anthocoridae), nabid bugs (family nabidae), other predators, parasitic wasps, other wasps, bees, and other pollinators. arthropods were sorted and identified to family level and/or functional group (e.g., parasitic wasps) for comparison among plots. we defined total natural enemy abundance as the sum of individuals of all arthropod classes found on the sticky cards, excluding bees and o