( aax JOURNAL 2 OF THE KENTUCKY. ACADEMY OF SCIENCE Official Publication of the Academy Volume 66 Number 2 Fall 2005 The Kentucky Academy of Science Founded 8 May 1914 | GOVERNING BOARD 2005 EXECUTIVE COMMITTEE 2005 President: Bruce Mattingly, Morehead State University/b.mattin@morehead-st.edu President Elect: Miriam Steinitz-Kannan, Northern Kentucky University/kannan@nku.edu Vice President: Nigel Cooper, University of Louisville Medical Center/nigelcooper@louisville.edu Past President: Robert W. Kingsolver, Bellarmine University /kingsolver@bellarmine.edu Secretary: Kenneth Carstens, Murray State University/kenneth.carstens@murraystate.edu Treasurer: Kenneth Crawford, Western Kentucky University/kenneth.crawford@wku.edu Executive Director (ex officio): Linda Hightower, KAS Office, Science Outreach Center, _ University of Kentucky/kasexecutivedirector@yahoo.com Editor, JOURNAL (ex officio): John W. Thieret. Northern Kentucky University/thieretj@nku.edu DIVISION AND AT-LARGE REPRESENTATIVES Biological Sciences (2005): Thomas C. Rambo, Northern Kentucky University/rambot@nku.edu Biological Sciences (2007): Philip Lienesch, Western Kentucky University/philip.lienesch@wku.edu Physical Sciences (2006): Mark Blankenbuehler, Morehead State University/m.blanken@morehead-st.edu | Physical Sciences (2008): Scott Nutter, Northern Kentucky University/nutters@nku.edu Social & Behavioral Sciences (2006): David Hogan, Northern Kentucky University/hogan@nku.edu Social & Behavioral Sciences (2008): David Olson, Morehead State University/d.olson@morehead-st. edu At-Large (2005): Ralph Thompson, Berea College/ralph.thompson@berea.edu | At-Large (2007): Christopher Lorentz, Thomas More College/chris.lorentz@thomasmore.edu Program Coordinator (ex officio): Robert Creek, Eastern Kentucky University/rcreek@chpl.net Director, Kentucky Junior Academy of Science (ex officio): Elizabeth Sutton, Campbellsville University/ eksutton@campbellsville.edu Editor, NEWSLETTER (ex officio): Susan Templeton, Kentucky State University/stempleton@gwmail. kysu.edu Editor, KAS Webpage (ex officio): Claire Rinehart, Western Kentucky University/claire.rinehart@wku.edu AAAS/NAAS Representative (ex officio): Guenter Schuster, Eastern Kentucky University/guenter.schuster@eku. edu COMMITTEE ON PUBLICATIONS Editor and Chair: John W. Thieret, Northern Kentucky University/thieretj@exchange.nku.edu Abstract Editor: | Robert J. Barney, Kentucky State University/rbarney@gwmail.kysu.edu Index Editor: Varley E. Wiedeman/varleyw@mac.com 7 Editorial Board: | Ralph Thompson, Berea College/ralph_thompson@berea.edu Susan Templeton, Kentucky State University/stempleton@gwmail.kysu.edu Claire Rinehart, Western Kentucky University/claire.rinehart@wku.edu | All manuscripts and correspondence concerning manuscripts should be addressed to the Editor. © The JOURNAL is indexed in BIOSIS, Cambridge Scientific Abstracts, Selected Water Resource Abstracts, ‘State Academies of Science Abstracts, and Zoological Record. Membership in the Academy is open to interested persons upon nomination, payment of dues, and dechion! Asslidation forms for membership may be obtained from the Secretary. The JOURNAL is sent free to all members in good standing. Annual dues are $25.00 for Active Members; $15.00 for Student Members; $35.00 for Family; $350.00 for Life Mem- bers. Subscription rates for nonmembers are: $50.00 domestic; $60.00 foreign. Back issues are $30.00 per volume. The JOURNAL is issued semiannually in spring and fall. Two numbers comprise a volume. Correspondence concerning memberships or subscriptions should be addressed to the Executive Secretary. This paper meets the requirements of ANSI/NISO Z39.48-992 (Permanence of Paper). Volume 66 of the Journal of the Kentucky Academy of Science is dedicated to the memory of John W. Thieret, Editor (1996-2005) Journal of the Kentucky Academy of Science 66(2) JOHN W. THIERET (1926-2005) JOURNAL OF THE KENTUCKY ACADEMY OF SCIENCE ISSN 1098-7096 Continuation of Transactions of the Kentucky Academy of Science Volume 66 Fall 2005 Number 2 ]. Ky. Acad. Sci. 66(2):73-81. 2005. John W. Thieret (1926-2005) Ralph L. Thompson Berea College Herbarium, Berea College, Berea, Kentucky 40404-2121 ABSTRACT John W. Thieret (1926-2005), an internationally recognized American plant taxonomist, is remembered as a consummate field botanist, exemplary teacher, acclaimed author, superb editor, fine herbarium director, inspiring mentor to students and colleagues, and noble friend. One of the most renowned American plant taxonomists of the 20th century has died. It is with great sadness and a profound sense of loss that I write this tribute about the life and career of John W. Thieret, Professor Emeritus of Biological Sciences at Northern Kentucky University, retired Director of the Northern Kentucky University Herbarium, and Editor of the Journal of the Kentucky Academy of Science (JKAS). John suffered a brain aneu- rysm at his home in Alexandria, Kentucky, on 6 December 2005, while editing a manuscript for the JKAS. He never regained conscious- ness and died on 7 December at the age of 79. John Thieret was a gifted botanist, excellent educator, wise scholar, and gentleman. He dearly loved his family, and after family, his great passion was plants. Although John was focused on botany his entire life, he had other diverse interests including classical music, par- ticularly opera. He also enjoyed sharing his knowledge far beyond botany and the natural sciences through verbal discourse about di- verse topics in history, literature, art, religion, and the social sciences. John William Thieret was born on 1 August 1926, in Chicago, Illinois, the only child of Hans and Lorena Thieret. Growing up, he was 73 interested in plants and became an avid stu- dent of botany during his school days at Hyde Park High School. At Hyde Park, John met his future wife, Mildred Wolf, fittingly in a botany class. After working briefly in Chicago, John moved to Logan, Utah, to study at Utah State University. Three years later, Mildred also moved to Logan and attended Utah State. They were married on 13 March 1950, by one of their professors, after completing their Evo- lution final exams. Both earned their B.S. de- grees in 1950: John’s in Botany and Mildred’s in Bacteriology. They remained at Utah State University for graduate work and in 1951, John earned his M.S. in Botany with focus on barley genetics and Mildred completed her M.S. in Bacteriology. They returned to Chi- cago, where John attended the University of Chicago to work on his doctorate under Theo- dor K. Just, Chief Curator of the Chicago Field Museum of Natural History. John re- ceived his Ph.D. in Botany in 1953. The title of John’s dissertation was “Gross Morphology of the Seeds of the Scrophulariaceae and Clas- sification of the Family.” Later in 1953, John became Assistant Cu- rator of Economic Botany at the Chicago Field Museum and then Curator of Economic Botany from 1954 to 1961. While at the Field Journal of the Kentucky Academy of Science 66(2) In Memoriam eas Museum, he made collecting trips to Cuba, Mexico, the northern Great Plains of the Unit- ed States, and the Northwest Territories of Canada. John authored 26 publications during that time and his special interests in the Po- aceae and Scrophulariaceae were evident in his published works. John published five new nomenclatural combinations while at the Field Museum. He also wrote three articles on the flora and vegetation of the Canadian North- west Territories. John left the Field Museum to become As- sociate Professor and later Professor of Biol- ogy at the University of Southwestern Louisi- ana, Lafayette (USL), from 1961 to 1973. At USL, he sponsored undergraduate research projects and directed six M.S. theses and one Ph.D. dissertation. During this period, John conducted most of his personal research in the southeastern United States, with an emphasis on the Louisiana flora. At USL, he authored or co-authored 47 articles. John named four plant species new to science that he discov- ered in Louisiana: Cyperus brevifolioides Thieret & Delahoussaye; Cyperus louisianen- sis Thieret; Isoetes louisianensis Thieret; and Limnophila Xludoviciana Thieret. From his work at USL, John published 11 nomencla- tural combinations. He also published his first two generic flora treatments of the southeast- ern United States, five more articles from the Canadian Northwest Territories, and several North American plant records. John made his final academic career move in 1973, when he joined the faculty at North- ern Kentucky University (NKU) in Highland Heights, as Professor and Chair of the De- partment of Biological Sciences. The oppor- tunity to teach at NKU, the presence of the Lloyd Library in Cincinnati, and relocating his family to live in a cooler climate, were espe- cially appealing to him. John served as Chair until 1980 and continued as Professor until re- tiring in 1992, with the title Professor Emer- itus of Biological Sciences. John loved teaching and was an exemplary teacher in the classroom, laboratory, and field. His classes were challenging, enjoyable, infor- mative, popular, and inspired many students toward their full potential. He always was available to students and colleagues for dis- cussions about botany, academics, or just about life. John was a strong believer in pro- viding students with actual plant specimens, either fresh or dried, for a “hands-on” ap- proach, and he went to great effort to accom- plish that goal. In the field, John’s passion for botany was especially contagious and he al- ways felt rejuvenated after a field trip with stu- dents. At NKU, John taught 15 different courses for the Department of Biological Sciences in- cluding Agrostology, Aquatic Vascular Plants, Dendrology, Field Botany, General Botany, General Biology Lab, Horticultural Plants, Li- brary Resources in Biology, Plants and People, Plants in Winter, Plant Taxonomy, Spring Flo- ra of Kentucky, Summer Flora of Kentucky, Trees of Kentucky, and Woody Plants. John generously shared his expertise and enthusi- asm for botany, especially plant taxonomy, with students, colleagues, and the general public for 32 years. His exuberance, magnetic personality, and unparalleled depth and breadth of knowledge were inspirations to ev- eryone his activities touched. For many years, John’s summer activities were focused on teaching at various biological field stations. He served as Visiting Lecturer in Botany at the Itasca Biological Station, Uni- versity of Minnesota; the Ollahoms Biological Station, University of Oklahoma; the Michigan Biological Station, University of Michigan; and the Franz Theodore Stone Laboratory, Ohio State University. John enjoyed the flora of cooler climates, and biological stations provid- ed Mildred and his active children (Robert, Nancy, Richard, Jeffrey, and Jennifer) new en- vironments to experience. He also conducted major field travels in the southeastern, south- western, and northwestern United States, the Great Basin of Nevada and Utah, and the Ca- nadian Arctic, Newfoundland, and Ontario. Scientists are often judged by the number of articles by them or about them. If this is any measure of a person, John stands taller than a coastal redwood. During his profession- al career, John authored at least 157 refereed journal articles and book articles. Many of his articles appeared in Sida, Contributions to Botany. He also published in Bartonia, Ca- nadian Field-Naturalist, Castanea, Economic Botany, Journal of the Arnold Arboretum, Rhodora, Taxon, and the Transactions/Journal of the Kentucky Academy of Science. In ad- dition to books, journal articles, and 19 no- 76 Journal of the Kentucky Academy of Science 66(2) menclatural combinations, John wrote 136 book reviews, 65 articles for Encyclopaedia Britannica, 46 articles for Encyclopedia Amer- icana, and many popular science articles. During his tenure at NKU, John was the author or co-author of five books. His books were entitled: Louisiana Ferns and Fern AIl- lies; Aquatic and Wetland Plants of Kentucky; Trees: A Quick Reference Guide to Trees of North America; Assessment and Management of Plant Invasions; and National Audubon So- ciety Field Guide to North American Wild- flowers: Eastern Region. His books continue to serve many audiences, from professional botanists to amateur plant enthusiasts. John authored or co-authored with other botanists, 58 refereed journal articles at NKU. He frequently collaborated with younger NKU biology colleagues in research activities and publications. John also generously shared his research interests with current and former undergraduate and graduate students. He did not restrict his research activities to colleagues and students. Indeed, John was a general mentor for botanists from other institutions to pursue research activities in Kentucky and elsewhere. His collaborative research with Kentucky botanists ranged from the 150-year chronology of Amur honeysuckle (Lonicera maackii) and people, to the history of the medlar (Mespilus germanica), to the invasive spread of Coincya (Coincya monensis) in the eastern United States, to “coffee” from the Kentucky coffee- tree, to new Kentucky plant distribution rec- ords from two of his favorite weedy haunts, Silver Grove Railroad Yard in Campbell Coun- ty and Latonia Railroad Yard in Kenton Coun- In the 1980s, John initiated an effort with other Kentucky botanists to produce a manual of the Kentucky flora. Several published con- tributions to that effort resulted, but at a rate too slow to produce a complete state flora. In the mid-1990s, inspired by John’s continual encouragement and editorial assistance, Ron- ald L. Jones of Eastern Kentucky University became the author of the Kentucky flora, with other botanists contributing selected group treatments. As Editorial Associate, John worked closely with Ronald over the next de- cade by editing many drafts of the manuscript and contributing the treatment of the grasses. In 2005, Jones’ comprehensive book, Plant Life of Kentucky, was published. One of John Thieret’s crowning achieve- ments was establishing the Northern Ken- tucky University Herbarium (KNK) in 1973. As the first Director of the KNK Herbarium, he built the herbarium from his personal col- lections, additions by colleagues and students, and through an active specimen exchange with various national and international herbaria. Currently, KNK has over 35,000 mounted specimens with strong representation from Kentucky, the Southeast, and the Midwest. John’s career specimen accession numbers were over 62,000. John accurately identified all plants deposited into the herbarium and meticulously mounted his own plant speci- mens. The KNK collection has the highest species diversity among Kentucky herbaria collections. Because of his efforts, the KNK herbarium is also the best-curated herbarium in Kentucky. John donated his reference li- brary (over 600 books) to the herbarium. John served several significant editorships and advisory roles during his professional ca- reer. He was a Member of the Editorial Board of Economic Botany from 1959-1965, Book Editor of Economic Botany from 1959-1984, Editor from 1986-1990, and Associate Editor from 1992-2005. He was a founding Member of the Editorial Board for the Vascular Flora of the Southeastern United States project from 1981-2005. John served as the Associate Editor of Sida, Contributions to Botany, from 1971-2005, and contributed to its excellence and prestige through dedicated work. Barney L. Lipscomb, current Editor of Sida, Botanical Research Institute of Texas, notes, “John was a towering lighthouse to the editors of Sida. His steadfast editorial ‘light’ was a never end- ing source of guidance and navigation in keep- ing Sida on a positive course.” During 1983-2005, one of John’s most sig- nificant roles was as a Member of the Editorial Committee of the monumental multi-volume project, Flora of North America North of Mex- ico. John edited the first 10 published volumes and prepared 25 family and generic treat- ments. He also was an Advisor in Botany for Encyclopaedia Britannica from 1959-2005 and a Member of the Advisory Committee at Lloyd Library in Cincinnati from 1992-2005. John loved editing and his ability as an ed- In Memoriam Wat itor was extraordinary. He was a perfectionist and a superlative editor second to none, but his efforts always brought out the best in au- thors. Those who submitted manuscripts for John’s editorial scrutiny often found them re- turned with a profusion of red ink on the printed text. When his recommended changes were made, the greatly improved manuscripts always told the story better. Many Kentuckians may best remember John for his devoted service to the Kentucky Academy of Science (KAS). He served as Ab- stract Editor of Transactions of the Kentucky Academy of Science (TKAS) from 1981-1995. John then assumed the position of Editor of TKAS in 1996. He had the title changed from Transactions to Journal of the Kentucky Acad- emy of Science (JKAS) in 1998 to better reflect its mission and content. Under John’s editor- ship, all JKAS manuscripts were peer-re- viewed to meet the highest standards. Since John worked without assistance, he was en- tirely responsible for all organizational aspects of each JKAS issue. The stature of the JKAS as a multi-disciplinary journal of state and re- gional scientific literature was greatly en- hanced because of his dedicated efforts. John received many awards and honors dur- ing his career. One of his most esteemed hon- ors was having a mint in the Lamiaceae that he discovered in Louisiana named for him. In 1964, botanist Lloyd H. Shinners named this new species Scutellaria thieretii Shinners. John received the 1984 Distinguished Ken- tucky College/University Scientist Award from the Kentucky Academy of Science for his sig- nificant academic research and teaching con- tributions to the Commonwealth. Most re- cently, John was presented the 2005 Outstand- ing Academy Service Award from the Ken- tucky Academy of Science for his outstanding editorial contributions to the JKAS. To commemorate his contributions to the Northern Kentucky University Herbarium, it was officially renamed the John W. Thieret Herbarium by the Northern Kentucky Uni- versity Board of Regents on 22 March 2006. In 1994, The John W. Thieret Research Award was established by John as an annual award to the NKU student who accomplished the most significant research. The Thieret family would like to continue this student hon- or, and have asked that donations in his mem- ory be sent to the Northern Kentucky Uni- versity Foundation, designated to that award. ohn is survived by his devoted wife of 55 years, Mildred Thieret, his five children, Rob- ert, Nancy, and Jeffrey in Minnesota, Richard in China, and Jennifer in Highland Heights, seven grandchildren, and five great-grandchil- dren. John Thieret was one of the patriarchs in North American plant taxonomy, and one of the last great field naturalists. His death marks the ending of a botanical era, but his legacy continues through the work of many former students and colleagues. John was an inspiring and stabilizing mentor who enriched the lives of those who knew him. He will be greatly missed by all. ACKNOWLEDGMENTS I am especially grateful to Mildred Thieret for graciously sharing her memories about John and providing photographs of her hus- band. I also acknowledge contributions from Ronald L. Jones, Eastern Kentucky University, David M. Brandenburg, Dawes Arboretum, James O. Luken, Coastal Carolina University, Debra K. Pearce, Northern Kentucky Univer- sity, and Barney L. Lipscomb, Botanical Re- search Institute of Texas. PUBLICATIONS Baird, J. R., and J. W. Thieret. 1985. Notes on Themeda quadivalvis (Poaceae) in Louisiana. Iselya 2:129-137. Baird, J. R., and J. W. Thieret. 1988. The bur gherkin (Cucumis anguria var. anguria, Cucurbitaeae). Econ. Bot. 42:447-451. Baird, J. R., and J. W. Thieret. 1989. The medlar (Mes- pilus germanica, Rosaceae) from antiquity to obscurity. Econ. Bot. 43:328-372. Baird, J. R., and J. W. Thieret. 1993. Spartina. Pages 1296-1297 in J. C. Hickman (ed). The Jepson manual: higher plants of California, University of California Press, Berkeley and Los Angeles. Beal, E. O., and J. W. Thieret. 1986. Aquatic and wetland plants of Kentucky. Kentucky State Nature Preserves Commission, Scientific and Technical Series, Number 5, Frankfort. Brandenburg, D. M., W. H. Blackwell, and J. W. Thieret. 1991. Revision of the genus Cinna (Poaceae). Sida 14: 581-596. Brandenburg, D. M., J. E. Estes, S. B. Russell, and J. W. Thieret. 1991. One-nerved paleas in Cinna arundinacea L. (Poaceae). Trans. Kentucky Acad. Sci. 52:94—-96. Brandenburg, D. M., J. E. Estes, and J. W. Thieret. 1991. 78 Journal of the Kentucky Academy of Science 66(2) Hard grass (Sclerochloa dura, Poaceae) in the United States. Sida 14:369-376. Brandenburg, D. M., and J. W. Thieret. 1996. Sclerochloa dura (Poaceae) in Kentucky. J. Kentucky Acad. Sci. 57: 47-48. Brandenburg, D. M., and J. W. Thieret. 2000. Cinna and Limnodea (Poaceae): not congeneric. Sida 19:195-200. Brandenburg, D. M., and J. W. Thieret. 2003. Epipactis helleborine (Orchidaceae) in Kentucky, with overview of literature on biology of the species. J. Kentucky Acad. Sci. 64:55-74. Buddell, G. F., UL, and J. W. Thieret. 1985. Notes on Er- igenia bulbosa (Apiaceae). Bartonia 51:69-76. Buddell, G. F., II, and J. W. Thieret. 1997. Saururaceae. Pages 37-38 in Flora of North America, Vol. 3. Oxford Univ. Press, New York and Oxford. Buddell, G.F., H, and J. W. Thieret. 2004. Lobed leaves in Salix exigua, sandbar willow (Salicaceae), in Ken- tucky. J. Kentucky Acad. Sci. 65:51. Calaway, M. L., and J. W. Thieret. 1985. Amphibromus scabrivalvis (Gramineae) in Louisiana. Sida 11:207— 214. Clark, H. L., and J. W. Thieret. 1968. The duckweeds of Minnesota. Michigan Bot. 7:67—76. Clark, R. C., R. L. Jones, T. J. Weckman, R. L. Thompson, J. W. Thieret, Kentucky State Nature Preserves Com- mission, and K. Feeman. 2005. State records and other noteworthy collections for Kentucky. Sida 21:1909- 1916. Cranfill, R., and J. W. Thieret. 1981. Thirty additions to the vascular flora of Kentucky. Sida 9:55-58. Delahoussaye, A. J., and J. W. Thieret. 1967. Cyperus sub- genus Kyllinga (Cyperaceae) in the continental United States. Sida 3:128-136. Eckenwalder, J. E., and J. W. Thieret. 1993. Keys to gym- nosperm families. Pages 345-346 in Flora of North America, Vol. 2. Oxford Univ. Press, New York and Ox- ford. Evers, R. A., and J. W. Thieret. 1957. New plant records: Illinois and Indiana. Rhodora 59:181. Hall, D. W,, and J. W. Thieret. 2003. Chrysopogon. Pages 633-636 in Flora of North America, Vol. 25. Oxford Univ. Press, New York and Oxford. Hartman, R. L., J. W. Thieret, and R. K. Rabeler. 2005. Paronychia. Pages 30-43 in Flora of North America, Vol. 5. Oxford Univ. Press, New York and Oxford. Hils, M. H., J. W. Thieret, and J. D. Morefield. 2003. Sarcobatus. Pages 387-389 in Flora of North America, Vol. 4. Oxford Univ. Press, New York and Oxford. Kartesz, J. T., P. Allen, and J. W. Thieret. 1997. Epilobium brachycarpum (Onagraceae) in Kentucky. Trans. Ken- tucky Acad. Sci. 58:99. Kartesz, J. T., and J. W. Thieret. 1991. Common names for vascular plants: guidelines for use and application. Sida 14:421—434. Landry, G., and J. W. Thieret. 1973. Isoetes louisianensis (Isoetaceae), a new species from Louisiana. Sida 5:129— 130; Luken, J. O., and J. W. Thieret. 1987a. Linum grandiflo- rum (Linaceae), Papaver dubium (Papaveraceae), and Salvia pratensis (Labiatae): additions to the Kentucky flora. Trans. Kentucky Acad. Sci. 48:26. Luken, J. O., and J. W. Thieret. 1987b. Sumac-directed patch succession on northern Kentucky roadside em- bankments. Trans. Kentucky Acad. Sci. 48:51-54. Luken, J. O., and J. W. Thieret. 1988. A life-form spec- trum for Ohio. Trans. Kentucky Acad. Sci. 49:38-39. Luken, J. O., and J. W. Thieret. 1995. Amur honeysuckle (Lonicera maackii; Caprifoliaceae): its ascent, decline, and fall. Sida 16:479-503. Luken, J. O., and J. W. Thieret. 1996. Amur honeysuckle, its fall from grace. BioScience 46:18-24. Luken, J. O., and J. W. Thieret. 1997. Amur honeysuckle, its fall from grace. Arnoldia 57:2-12. Luken, J. O., and J. W. Thieret. 2001. Floristic compari- sons of mud flats and shorelines at Cave Run Lake, Kentucky. Castanea 66:336-351. Luken, J. O., and J. W. Thieret (eds). 1997. Assessment and management of plant invasions. Springer-Verlag Environmental Management Series, New York, NY. Luken, J. O., J. W. Thieret, and J. R. Kartesz. 1993. Er- ucastrum gallicum (Brassicaceae): invasion and spread in North America. Sida 15:569-582. Medley, M. E., H. Bryan, J. MacGregor, and J. W. Thieret. 1985. Achyranthes japonica (Miq.) Nakai (Amarantha- ceae) in Kentucky and West Virginia: new to North America. Sida 11:92—95. Medley, M. E., R. Cranfill, and J. W. Thieret. 1983. Vas- cular flora of Kentucky: additions and other noteworthy collections. Sida 10:114—122. Medley, M. E., and J. W. Thieret. 1991. Ulmus parvifolia (Ulmaceae) naturalized in Kentucky. Sida 14:610-613. Mohlenbrock, R. H., and J. W. Thieret. 1987. Trees: a quick reference guide to trees of North America. Col- lier Books, Macmillan Publishing Company, New York, NY. Naczi, R. F. C., and J. W. Thieret. 1996a. Addition to the flora of Bradford County, Pennsylvania. Bartonia 59:81— 85. Naczi, R. F. C., and J. W. Thieret. 1996b. Invasion and spread of Coincya monensis (Brassicaceae) in North America. Sida 17:43-53. Naczi, R. F. C., and J. W. Thieret. 1996c. The gold-cone tamarack (Larix larcina forma lutea, Pinaceae) in Penn- sylvania. Bartonia 59:123-124. Naczi, R. F. C., and J. W. Thieret. 2000. Additions to the flora of Potter County, Pennsylvania. Bartonia 60:117— 120. Nienaber, M. A., and J. W. Thieret. 2003. Phytolaccaceae. Pages 3-11 in Flora of North America, Vol. 4. Oxford Univ. Press, New York and Oxford. Pearce, D. K., and J. W. Thieret. 1991. Japanese-quince (Chaenomeles speciosa, Rosaceae), a dual-use shrub. Econ. Bot. 45:285—288. Pearce, D. K., and J. W. Thieret. 1993. Persimmon (Di- ospyros virginiana, Ebenaceae) and mayapple (Podo- In Memoriam 79 phyllum peltatum), Berberidaceae): proximate analysis of their fruits. Trans. Kentucky Acad. Sci. 54:30-31. Rabeler, R. K., and J. W. Thieret. 1988. Comments on the Caryophyllaceae of the southeastern United States. Sida 13:149-156. Rabeler, R. K., and J. W. Thieret. 1997. Sagina (Cary- ophyllaceae) range extensions in Canada: S. japonica new to Newfoundland, S. procumbens, new to the Northwest Territories. Canad. Field-Naturalist 111: 309-310. Reece, W. D., and J. W. Thieret. 1966. Botanical study of the Five Islands of Louisiana. Castanea 31:251—277. Smith, C. E, Jr, and J. W. Thieret. 1959a. An English obituary account of Thomas Nuttall. Bartonia 29:10. Smith, C. E., Jr, and J. W. Thieret. 1959b. Thomas Nuttall (1786-1859): an evaluation and bibliography. Leafl. West. Bot. 9:33—-42. Spaeth, J. P., and J. W. Thieret. 2004. Notes on “coffee” from the Kentucky coffeetree (Gymnocladus dioicus, Fabaceae). Sida 21:345-356. Spjut, R. W,, and J. W. Thieret. 1989. Confusion between multiple and aggregate fruits. Bot. Rev. 55:53-72. Thieret, J. W. 1953. Gross morphology of the seeds of the Scrophulariaceae and classification of the family. Ph.D. Dissertation. The University of Chicago. Thieret, J. W. 1953. Dipsacus laciniatus in Illinois. Rho- dora 55:268. Thieret, J. W. 1954. The tribes and genera of Central American Scrophulariaceae. Ceiba 4:164—184. Thieret, J. W. 1955a. The seeds of Veronica and allied genera. Lloydia 18:37—45. Thieret, J. W. 1955b. The status of Berendtia A. Gray. Ceiba 4:304—305. Thieret, J. W. 1956a. Bryophytes as economic plants. Econ. Bot. 10:75-91 Thieret, J. W. 1956b. Nardoo. Amer. Fern Jour. 46:108— 109. Thieret, J. W. 1956c. Stenardrium Nees versus Gerardia L. Taxon 5:58-59. Thieret, J. W. 1957. Plants new to Illinois and to the Chi- cago region. Rhodora 59:289. Thieret, J. W. 1958a. Agalinis Rafinesque versus Chytra Gaertn. Taxon 7:142-143. Thieret, J. W. 1958b. Castilleja Mutis ex L. versus Bartsia L. Taxon 7:83-84. Thieret, J. W. 1958c. Economic botany of the cycads. Econ. Bot. 12:3-41. Thieret, J. W. 1959a. Grassland vegetation near Fort Proy- idence, Northwest Territories. Canad. Field-Naturalist TSA6I—L67. Thieret, J. W. 1959b. Scrophulariaceae. Pages 3-10 in J. Angely (ed). Catalogo e estatastica dos géneros botani- cos fanerogamicos, Vol. 49. Thieret, J. W. 1960a. Calamovilfa longifolia and its variety magna. Amer. Midl. Nat. 63:169-176. Thieret, J. W. 1960b. The formaldehyde method of col- lecting plant specimens. Turtox News 38:114—-115. Thieret, J. W. 1961a. A collection of plants from the Horn Plateau, District of Mackenzie, Northwest Territories. Canad. Field-Naturalist 75:77-83. Thieret, J. W. 1961b. New plant records for southwestern District of Mackenzie. Canad. Field-Naturalist 75:111— IDeA Thieret, J. W. 1961c. The Scrophulariaceae-Buchnerae of Central America. Ceiba 8:92-101. Thieret, J. W. 1961d. The specific epithet of the pecan. Rhodora 63:296. Thieret, J. W. 1962a. Exceptional height for Rhododen- dron lapponicum. Canad. Field-Naturalist 76:123. Thieret, J. W. 1962b. New plant records from District of Mackenzie, Northwest Territories. Canad. Field-Natu- ralist 76:206—208. Thieret, J. W. 1963a. Additions to the flora of Louisiana. Castanea 28:169—170. Thieret, J. W. 1963b. Additions to the flora of the North- west Territories. Canad. Field-Naturalist 77:126. Thieret, J. W. 1963c. Botanical survey along the Yellow- knife Highway, Northwest Territories, Canada. I: Cat- alogue of the flora. Sida 1:117-170. Thieret, J. W. 1963d. Life-forms in the plains flora of southern Mackenzie, Northwest Territories. Rhodora 65: 1492157, Thieret, J. W. 1963e. The correct name for the water- melon. Taxon 12:37. Thieret, J. W. 1964a. Botanical survey along the Yellow- knife Highway, Northwest Territories, Canada. II: Veg- etation. Sida 1:187—239. Thieret, J. W. 1964b. Eriogonum annuum (Polygonaceae): biennial in Nebraska. Sida 1:382. Thieret, J. W. 1964c. Fatoua villosa (Moraceae) in Loui- siana: new to North America. Sida 1:248. Thieret, J. W. 1964d. Lysimachia japonica (Primulaceae) and Clinopodium gracile (Labiatae) in Louisiana: new to the United States. Sida 1:294—295. Thieret, J. W. 1964e. More additions to the Louisiana flo- ra. Sida 1:294—295. Thieret, J. W. 1966a. Additions to the Louisiana flora. Sida 2:264—265. Thieret, J. W. 1966b. Habit variation in Myrica pensyl- vanica and M. cerifera. Castanea 31:183-184. Thieret, J. W. 1966c. Seeds of some United States Phy- tolaccaceae and Aizoaceae. Sida 2:352-—360. Thieret, J. W. 1966d. Synopsis of the genus Calamovilfa (Gramineae). Castanea 31:145-152. Thieret, J. W. 1967a. Life-forms in the flora of Minnesota. J. Minnesota Acad. Sci. 34:251-277. Thieret, J. W. 1967b. Neogaerrhinum kelloggii (Greene) Thieret, comb. nov. (Scrophulariaceae). Sida 3:187. Thieret, J. W. 1967c. Supraspecific classification in the Scrophulariaceae: a review. Sida 3:87—106. Thieret, J. W. 1967d. Thirty additions to the Louisiana flora. Sida 3:123-127. Thieret, J. W. 1968. Additions to the vascular flora of Lou- isiana. Proc. Louisiana Acad. Sci. 31:91-97. Thieret, J. W. 1969a. Baptisia lactea (Rafinesque) Thieret, comb. nov. (Leguminosae). Sida 3:446. 80 Journal of the Kentucky Academy of Science 66(2) Thieret, J. W. 1969b. Notes on Epifagus. Castanea 34: 397-402. Thieret, J. W. 1969c. Rumex obovatus and Rumex para- guayensis (Polygonaceae) in Louisiana: new to North America. Sida 3:445-446. Thieret, J. W. 1969d. Sagittaria guayanensis (Alismata- ceae) in Louisiana: new to the United States. Sida 3: 445. Thieret, J. W. 1969e. Trifolium vesiculosum (Legumino- sae) in Mississippi and Louisiana: new to North Amer- ica. Sida 3:446—447. Thieret, J. W. 1969f. Twenty-five species of vascular plants new to Louisiana. Proc. Louisiana Acad. Sci. 32:78-82. Thieret, J. W. 1970a. Bacopa repens (Scrophulariaceae) in the conterminous United States. Castanea 35:132-136. Thieret, J. W. 1970b. Orobanchaceae. Pages 331-337 in C. L. Lundell (ed). Flora of Texas, Vol. 2. Thieret, J. W. 1970c. Scrophulariaceae—Figwort family. Pages 316-321 in E. A. Menninger (ed). Flowering vines of the world. Thieret, J. W. 1971a. Additions to the Louisiana flora. Cas- tanea 36:219-222. Thieret, J. W. 1971b. Eriocaulon cinereum R.Br. in Lou- isiana. Southwest. Nat. 15:391. Thieret, J. W. 1971c. Observations on some aquatic plants in northwestern Minnesota. Michigan Bot. 10:117—124. Thieret, J. W. 1971d. Physalis lagascae (Solanaceae) in Louisiana: new to the conterminous United States. Sida ATT. Thieret, J. W. 197le. Quadrat study of a bottomland forest in St. Martin Parish, Louisiana. Castanea 36:174—181. Thieret, J. W. 1971f. The genera of Orobanchaceae in the southeastern United States. J. Arnold Arbor. 52:404— 434. Thieret, J. W. 1972a. Aquatic and marsh plants of Loui- siana: a checklist. J. Louisiana Soc. Hort. Res. 13:1-45. Thieret, J. W. 1972b. Checklist of the vascular flora of Louisiana: Part 1. Fern and fern allies, gymnosperms, and monocotyledons. Lafayette Natural History Muse- um Techn. Bull. 2. Thieret, J. W. 1972c. Rotala indica (Lythraceae) in Loui- siana. Sida 5:45. Thieret, J. W. 1972d. Synopsis of Hemichaena, including Berendtiella (Scrophulariaceae). Fieldiana Bot. 34:89-— 99. Thieret, J. W. 1972e. The Phrymaceae in the southeastern United States. J. Arnold Arbor. 53:226-233. Thieret, J. W. 1972f. Zeuxine strateumatica in Louisiana. Amer. Orch. Soc. Bull. 41:413. Thieret, J. W. 1973. Sex and the angiosperms. Sida 5:59-— 60. Thieret, J. W. 1974. Allium ampeloprasum (Liliaceae) and Trifolium vesiculosum (Leguminosae) in Oklahoma. Sida 5:286-287. Thieret, J. W. 1975a. Hemigraphis reptans (Acanthaceae), a greenhouse weed in Louisiana. Sida 6:115. Thieret, J. W. 1975b. The Mayacaceae in the southeastern United States. J. Arnold Arbor. 56:248—255. Thieret, J. W. 1976a. Floral biology of Proboscidea louis- ianica (Martyniaceae). Rhodora 78:169-179. Thieret, J. W. 1976b. Vascular plants new to Ohio. Cas- tanea 41:181—183. Thieret, J. W. 1977a. Cyperus louisianensis (Cyperaceae), a new species from southern Louisiana. Proc. Louisiana Acad. Sci. 40:23-26. Thieret, J. W. 1977b. Juvenile leaves in Oklahoma Mar- silea (Marsileaceae). Sida 7:218-219. Thieret, J. W. 1977c. Life-forms in the Michigan flora. Michigan Bot. 16:27-33 Thieret, J. W. 1977d. The Martyniaceae in the southeast- ern United States. J. Arnold Arbor. 58:25-39. Thieret, J. W. 1979. Hyptis mutabilis (Labiatae) in south- eastern United States. Sida 8:202—209. Thieret, J. W. 1980. Louisiana ferns and fern allies. La- fayette Natural History Museum. Published in conjunc- tion with The University of Southwestern Louisiana, Lafayette, LA. Thieret, J. W. 1982. The Sparganiaceae in the southeast- ern United States. J. Arnold Arbor. 63:341-355. Thieret, J. W. 1988. The Juncaginaceae in the southeast- ern United States. J. Arnold Arbor. 69:1-23. Thieret, J. W. 1989. Picea abies (Pinaceae) naturalized in southeastern Minnesota. Sida 13:505. Thieret, J. W. 1993a. Calocedrus. Page 412 in Flora of North America, Vol. 2. Oxford Univ. Press, New York and Oxford. Thieret, J. W. 1993b. Pinaceae. Pages 352-354 in Flora of North America, Vol. 2. Oxford Univ. Press, New York and Oxford. Thieret, J. W. 1993c. Psilotaceae. Pages 16-17 in Flora of North America, Vol. 2. Oxford Univ. Press, New York and Oxford. Thieret, J. W. 2003a. Arthraxon. Pages 677 in M. E. Bark- worth, K. M. Capels, S. Long, and M. B. Piep (eds). Flora of North America, Vol. 25. Oxford Univ. Press, New York and Oxford. Thieret, J. W. 2003b. Calamovilfa. Pages 140-144 in M. E. Barkworth, K. M. Capels, S. Long, and M. B. Piep (eds). Flora of North America, Vol. 25. Oxford Univ. Press, New York and Oxford. Thieret, J. W. 2003c. Coix. Pages 703-704 in M. E. Bark- worth, K. M. Capels, S. Long, and M. B. Piep (eds). Flora of North America, Vol. 25. Oxford Univ. Press, New York and Oxford. Thieret, J. W. 2003d. Eremochloa. Pages 688-690 in M. E. Barkworth, K. M. Capels, S. Long, and M. B. Piep (eds). Flora of North America, Vol. 25. Oxford Univ. Press, New York and Oxford. Thieret, J. W. 2003e. Hackelochloa. Pages 691-693 in M. E. Barkworth, K. M. Capels, S. Long, and M. B. Piep (eds). Flora of North America, Vol. 25. Oxford Univ. Press, New York and Oxford. Thieret, J. W. 2003f. Microstegium. Pages 623-624 in M. E. Barkworth, K. M. Capels, S. Long, and M. B. Piep (eds). Flora of North America, Vol. 25. Oxford Univ. Press, New York and Oxford. In Memoriam Sl Thieret, J. W. 2003g. Monoanthochloé. Pages 29-30 in M. E. Barkworth, K. M. Capels, S. Long, and M. B. Piep (eds). Flora of North America, Vol. 25. Oxford Univ. Press, New York and Oxford. Thieret, J. W. 2005. Agrostemma. Pages 214-215 in Flora of North America, Vol. 5. Oxford Univ. Press, New York and Oxford. Thieret, J. W., and C. M. Allen. 1974. Setaria pallide-fusca (Gramineae) in Louisiana. Castanea 39:290-291. Thieret, J. W., and J. R. Baird. 1985. Thalaspi alliaceum (Cruciferae) in Kentucky and Indiana. Trans. Kentucky Acad. Sci. 46:143-145. Thieret, J. W., and D. M. Brandenburg. 1986. Scaevola (Goodeniaceae) in southeastern United States. Sida 11: 445-453. Thieret, J. W., and R. A. Evers. 1957. Notes on Illinois grasses. Rhodora 59:123-124. Thieret, J. W., and S. F. Glassman. 1958. Grasses new to Illinois and the Chicago Region. Rhodora 60:264. Thieret, J. W., R. L. Hartman, and R. K. Rabeler. 2005. Herniaria. Pages 43-45 in Flora of North America, Vol. 5. Oxford Univ. Press, New York and Oxford. Thieret, J. W., and J. T. Kartesz. 1997. Lardizabalaceae. In: Pages 293-294 in Flora of North America, Vol. 3. Oxford Univ. Press, New York and Oxford. Thieret, J. W., and B. L. Lipscomb. 1985. Scaevola sericea Vahl var. taccada (Gaertn.) Thieret and Lipscomb. Sida EOS. Thieret, J. W., and J. O. Luken. 1996. The Typhaceae in the southeastern United States. Harvard Pap. Bot. 8: O16: Thieret, J. W., and R. S. Maples. 1979. Thelypteris inter- rupta (Polypodiaceae) new to Louisiana. Iselya 1:55. Thieret, J. W., W. A. Niering, and N. C. Olmstead. 2001. National Audubon Society Field Guide to North Amer- ican wildflowers: eastern region, revised ed. National Audubon Society, Alfred A. Knopf, Inc., New York, NY. Thieret, J. W., and R. K. Rabeler. 2005a. Corrigiola. Pages 48—49 in Flora of North America, Vol. 5. Oxford Univ. Press, New York and Oxford. Thieret, J. W., and R. K. Rabeler. 2005b. Polycarpaea. Pages 23-25 in Flora of North America, Vol. 5. Oxford Univ. Press, New York and Oxford. Thieret, J. W., and R. K. Rabeler. 2005c. Polycarpon. Pag- es 25-26 in Flora of North America, Vol. 5. Oxford Univ. Press, New York and Oxford. Thieret, J. W., and R. K. Rabeler. 2005d. Saponaria. Pages 157-158 in Flora of North America, Vol. 5. Oxford Univ. Press, New York and Oxford. Thieret, J. W., and R. K. Rabeler. 2005e. Scleranthus. Pag- es 149-151 in Flora of North America, Vol. 5. Oxford Univ. Press, New York and Oxford. Thieret, J. W., and R. K. Rabeler. 2005f. Vaccaria. Page 156 in Flora of North America, Vol. 5. Oxford Univ. Press, New York and Oxford. Thieret, J. W., and R. L. Thompson. 1984. Cleome orni- thopodioides (Capparaceae): adventive and spreading in North America. Bartonia 50:25-26. Thieret, J. W., and S. B. Young. 1988. The Kerguelen- cabbage, Pringlea antiscorbutica (Brassicaceae). Econ. Bot. 42:288-291. Thompson, R. L., and J. W. Thieret. 1986. Alopecurus arundinaceus (Poaceae) established in Kentucky. Trans. Kentucky Acad. Sci. 47:138. Vincent, M. A., and J. W. Thieret. 1987. Thymelaea pas- serina (Thymelaeaceae) in Ohio. Sida 12:51-54. Wessel, M. V., and J. W. Thieret. 2000. Agrimonia (Ro- saceae) in Kentucky with notes on the genus. J. Ken- tucky Acad. Sci. 61:146-162. Woodward, R. W., and J. W. Thieret. 1953. A genetic study of complementary genes for purple lemma, palea, and pericarp in barley (Hordeum vulgare L.). J. Agron- omy 45:162-185. J. Ky. Acad. Sci. 66(2):82-88. 2005. Cepaea nemoralis (Gastropoda, Helicidae): The Invited Invader Maggie Whitson Department of Biological Sciences, Northern Kentucky University, Highland Heights, Kentucky 41099 ABSTRACT Marauding snails may not immediately come to mind when considering invasive species, but many non- native snails have successfully colonized the U.S. The wood snail, Cepaea nemoralis (L.), is one of the most striking of these introductions, due in part to its attractive shell coloration. This is one of the few snails that people have purposely introduced into their gardens. Several populations are now established in Kentucky, including a newly discovered population in Kenton County. The bright, striped or solid, yellow, pink, and brown shells of this species have long caught the eyes of natural historians and biologists. Populations of these snails are classic model systems for ecological genetics studies. While introduced populations of wood snails seem to have had only minor impact as agricultural pests, they may have the potential to competitively exclude some native species of snails. INTRODUCTION Though snails may not be the speediest of beasts, several species have launched success- ful invasions of the U.S. (Cowie and Robinson 2001; Dundee 1974; Mead 1971). Perhaps the most glamorous of these invaders is the wood snail, Cepaea nemoralis (L.). While many gas- tropods rely on stealth and interstate shipping to fuel their spread, wood snails have another weapon in their arsenals: charm. Their color- ful shells (see Figure 1) are nearly irresistible to small children, nostalgic malacologists, and many an evolutionary biologist. TAXONOMY The wood snail belongs to the family Heli- cidae, which includes the bulk of the Euro- pean edible snails. It is a Linnaean species de- scribed in 1758 and was originally Helix ne- moralis, until Held established the genus Ce- paea in 1837 (Abbott 1989). Currently, four species are included in the genus, of which C. nemoralis is the type. The specific epithet means ‘of the woods’ or ‘inhabiting woods/ groves’ (Pilsbry 1939; Reeve 1863; Rimmer 1907). Cepaea hortensis, the white-lipped grove snail, is considered the sister species of C. ne- moralis (Jones et al. 1977). Historically, tax- onomists often treated C. hortensis as a variant of C. nemoralis (Step 1901). The two species are primarily differentiated by lip color of the shells, which seems a minor feature in light of the fact that C. nemoralis is the most variably colored species in its genus and perhaps even among European land snails. However, Rim- 82 mer (1907) argued in support of recognizing C. hortensis, having observed several mixed populations and noting that of the many snails seen paired on tree trunks, he saw no “mat- rimonial alliances between these two forms.” Current taxonomists also take this view, and the occasional hybrids produced by these spe- cies are sterile (Jones et al. 1977). Both taxa occur in the U.S., and though wood snails are known to be introduced, there is disagreement on whether C. hortensis is native or was also introduced from Europe (Burch 1962; Dun- dee 1974; Jones et al. 1977; Mead 1971). LIFE HISTORY AND NATIVE RANGE Native to central and western Europe, wood snails are widespread in disturbed habitats, from woodlands to fields and yards, but are also found on chalk cliffs and even coastal dunes (Reeve 1863). They are known by a va- riety of common names, the English ones in- cluding banded grove snail, banded wood snail, brown-lipped snail, and girdled snail (Abbott 1989; Reed 1964; Step 1901; Turton 1857). This species has been widely intro- duced and now has a nearly worldwide distri- bution (Abbott 1989). Wood snails are obligately outcrossing her- maphrodites, with both individuals exchanging sperm during mating, and both individuals able to lay eggs afterward (Stine 1989). Like other members of the Helicidae, Cepaea snails have a bizarre courtship behavior in which the courting pair stabs each other with sharp, cal- careous structures, aptly named darts, before mating (Abbott 1989; Pilsbry 1939). Wood Cepaea nemoralis (Gastropoda. Helicidae)—Whitson 83 Figure 1. Shells of Cepaea nemoralis (L.), the wood snail, showing solid pink (left), solid yellow (top), and striped morphs. Note the brown lip characteristic of this species. These European snails have been introduced throughout the northeastern U.S. and occur in at least three counties of Kentucky. snails often mate multiple times prior to egg laying and can store sperm for up to 15 months (Murray 1964). It is not unusual for one clutch of eggs to include offspring from two different fathers (Murray 1964). Eggs are buried in moist soil, hatching after about 3 weeks (Abbott 1989). The snails reach matu- rity in 4 years and may live as long as 5-9 years (Abbott 1989; Jones et al. 1977). Like those of most land snails, wood snail shells are dextral (spiraling to the right), though rare sinistral individuals are sometimes seen (Rimmer 1907; Turton 1857). Mature in- dividuals of C. nemoralis reach 2—2.5 cm in diameter and have five whorls to the shell (Pilsbry 1939). When the snail reaches full size and ceases to grow, a reflexed lip forms around the aperture of the shell. The dark brown col- oration of this lip differentiates C. nemoralis from the similar C. hortensis. Shell color varies from yellow, to pinkish, brown, or oc- casionally even white (Step 1901; Turton 1857). Shells are also generally augmented with 1-5 dark brown bands, though unstriped shells are seen as well (Figure 2). Multiple col- or variants are commonly found within the same population. Wood snails, feeding primarily at night, eat a variety of plants, though they often prefer dead plant material to living, and may even forage on dead organisms such as worms or other snails (Thompson 1996; Turton 1857). Among. living plant materials, they prefer broad-leaved plants over the tougher grasses 84 Journal of the Kentucky Academy of Science 66(2) Figure 2. Typical five-striped wood snail morph. There is such great variation in shell striping for this species that a coding system has been developed to catalogue it (Howe 1898; Jones et al. 1977). Stripes are numbered from top to bottom. This shell would be coded 12345. A shell with only the third stripe present (Figure 3) would be 00300. When partial or total fusion of bands is taken into account, there are hundreds of possible variants. which are common in their habitats, and avoid species with high concentrations of secondary compounds or physical defenses against her- bivory such as hairs (Grime et al. 1968; Thompson 1996). Oddly enough, they are said to particularly favor the leaves of stinging net- tles (Step 1901). They are adaptable in the lab or under cultivation and happily eat lettuce, carrots, fruit, pure cellulose filter paper, and even (occasionally) mutton (Grime 1968; Judd 1953; Murray 1964; Sowerby 1825; Thompson 1996). Though these snails lack operculums which would allow them to close their shells, they have a relatively high drought tolerance due to their ability to aestivate. After feeding, they generally crawl up onto the plant or a nearby shrub or stone wall, stick themselves down with a dab of slime, and remain inactive until the next moist evening. During dry spells, these organisms can remain dormant for long periods of time until conditions improve. (Aes- tivation in many species of snails can last for months, and even for years in some [Abbott 1989].) For the winter months, the snails bury into the soil and remain dormant until spring (Lovell 1884). In Great Britain, song thrushes can be a major predator of adult wood snails, crushing their shells on stones to get at the soft snail within. Other birds, including chickens, will sometimes eat wood snails (Howe 1898). Sev- eral snail predators are invertebrate organ- isms, including certain beetles, glowworm lar- vae (related to fireflies), and even predatory snails (Jones et al. 1977; Woodward 1913). Small mammals such as shrews, moles, and hedgehogs also enjoy these slow-moving mor- sels (Dees 1970; Reed 1964; Woodward 1913). SPREAD IN THE USS. Since the arrival of Europeans, many spe- cies of molluscs have been both purposely and accidentally introduced into the U.S. In recent years, the number one pathway for the intro- duction of new land snail species seems to be via infested horticultural materials (Cowie and Robinson 2001). Eggs and small individuals such as juveniles can be difficult to see when intermixed with soil, mulch, or other plant ma- terial (Cowie and Robinson 2001). Many snail species can also self-fertilize or store sperm for up to a year after mating, so one over- looked adult may be all it takes to pioneer an invasion (Cowie and Robinson 2001; Thomp- son 1996). Wood snails specifically have also been found by the USDA stuck to vehicles and military cargo (Dundee 1974). The helicid snails, which include C. nemor- alis, have a somewhat more colorful history of introduction, as many of these were purposely established in new habitats (Dees 1970; Mead 1971). Helicids, such as Helix pomatia, the French escargot, are often prized as choice co- mestibles. Many helicid introductions can be traced back to the kitchen gardens of Euro- pean immigrants desiring a renewable source of snails (Mead 1971). As a small and unusually colorful species, C. nemoralis has the distinction of being more often introduced for ornament than for food. The earliest U.S. introduction of this species was made by malacologist William Binney in 1857 (Pilsbry 1939). Binney collected snails in Sheffield, England, returned to the U.S. and then released them in his Burlington, NJ, gar- den, where they proceeded to flourish (Binney and Bland 1869). The U.S. populations of C. nemoralis orig- inate from multiple sources, however. A Lex- ington, VA, population was attributed either to an introduction of Italian snails in packing ma- Cepaea nemoralis (Gastropoda. Helicidae)—Whitson 85 ay) p fie ‘= ese | Be B. Figure 3. A. The wood snail has successfully colonized most of the northeastern U.S. and has also been found in some western states, including California and Texas (Ab- bott 1950; Burch 1962; Dundee 1974; Reed 1964). B. In Kentucky, populations of the wood snail have been found in Fayette, Jefferson, and Kenton Counties. (Maps cour- tesy of the online National Atlas of the United States [2005 ].) terials or of British snails in imported ivy (Bar- ber 1918; Howe 1898). After the turn of the century, imported shrubs from the Nether- lands and Ireland were probably the source of other snail populations discovered in Virginia and Massachusetts, respectively (Reed 1964). By 1974, populations of wood snails had been documented in at least 15 states and through- out the northeastern U.S. (Figure 3A). KENTUCKY POPULATIONS Documentation of land snail diversity and distributions in Kentucky is scanty, but scat- tered populations of C. nemoralis are reported from the state (Figure 3B). Reed (1964) cited a specimen found in Ohio River drift at Lou- isville, and at least three Lexington collections have been recorded (Branson and Batch 1969; FNMH 2005). Specimens have also been not- ed from Cincinnati, Ohio (Reed 1964; FMNH 2005), so it is not surprising that a large pop- ulation of C. nemoralis was recently found in northern Kentucky near Ft. Mitchell, Kenton Figure 4. Living wood snail from Kenton County, Ken- tucky. These snails have four pairs of tentacles, with the eyes located at the tips of the retractable upper pair. A shorter pair of sensory tentacles bracket the snail’s mouth. County (pers. obs.; see Figure 4). Surveys of snail fauna from Mammoth Cave National Park in Edmonson County and the Doe Run Creek Area of Meade County did not list C. nemoralis among the species found, perhaps because these are less disturbed areas than those noted above (Hubricht 1968; Kaplan and Minckley 1960). ECONOMIC IMPORTANCE While considered one of the European ed- ible snails, its small size, and the belief that species with striped shells are inferior in flavor have limited the popularity of C. nemoralis among gastronomes (Lovell 1884). None the less, the relative hardiness of this species com- pared to larger species of escargot, the ease of culture, and the nearly worldwide availability has kept them on the lists of species with po- tential for cultivation (Dees 1970; Thompson 1996). Though wood snails eat a variety of plant materials, their apparent preference for dead material has limited their impact as agricul- tural pests (Dees 1970; Thompson 1996). Oc- casional note has been made of the fact that even in areas with many snails, they appear to do little damage to the flora (Abbott 1950; Brooke 1897; Judd 1953). However, in high enough densities, they have the potential to damage landscaping or crops. For example, one Virginia population ranged from 50-100 86 Journal of the Kentucky Academy of Science 66(2) snails per square meter, with an estimated to- tal of 2500-5000 individuals (Stine 1989). Wood snail shells are carried by some shell dealers, but generally tropical landshells and marine shells are more popular with collec- tors. At least one U.S. population may have been introduced to serve as a shell source (Cowie and Robinson 2001). Wood snails also make easy to care for, if unusual, pets. However, most land snails are considered potential pest species by the USDA, and there are restrictions even on state-to-state transport of living snails (Dees 1970; Thompson 1996), dashing the hopes of those in the exotic pet industry hoping to spark a nationwide snail craze. ECOLOGICAL AND GENETIC STUDIES The variety of shell colors seen among banded wood snails has long fascinated natu- ralists, and many papers catalogue diversity within populations (Brooke 1897; Howe 1898; Johnson 1928; Judd 1953). The genetics of most color variations have been determined via crossing studies (Cain et al. 1968). At least five shell color loci are linked into a “super- gene” (Jones et al. 1977). These control the shell’s base color and four banding features: presence or absence, intensity of band and lip color, whether bands are continuous or dotted, and their spread (Jones et al. 1977). Four oth- er unlinked loci also affect banding, with the number of bands controlled by two, one con- trolling darkening along the length of the bands, and one determining whether bands are black or orange (Cain et al. 1968; Jones et al. 1977). Epistasis between some loci also plays a role (Jones et al. 1977). Considering that there are no fewer than six alleles for base color of the shell, and that banding is affected by at least eight loci and 18 alleles (Jones et al. 1977), it is not surprising that early workers enumerated hundreds of shell varieties (Howe 1898). Researchers have wondered how such high levels of variation are maintained. With long distance gene flow often limited by the slow spread of individuals, and many populations founded by small numbers of snails, one would expect to commonly see fixation of shell morphs through loss of alleles. However, fixed populations are rare. For example, a survey of 1000 French populations revealed only two that were monomorphic for shell coloration (Murray 1964). In a similar survey of 3000 British populations, fewer than 20 were mono- morphic (Jones et al. 1977). Two factors are thought to play a crucial role in maintaining this diversity. Because these snails are her- maphrodites, mating is possible between any two individuals, increasing the potential allele combinations available to offspring (Murray 1964). Also, wood snails generally mate at least twice prior to laying eggs, and can store sperm from multiple matings, effectively increasing the population size (Murray 1964). Thus, even small populations of snails may harbor more genetic diversity than would be seen in other types of organisms. Founder effects do have an impact on di- versity, though, especially in U.S. populations, most of which arose from introductions of small numbers of individuals (Brussard 1975). A study scoring shell polymorphisms and nine isozyme loci showed that the major differenc- es between U.S. populations seemed to be based on which part of Europe the snails had been introduced from, rather than the envi- ronment they were currently in (Brussard 1975). Later isozyme studies have also sup- ported the founder effect as having a major impact on the genetic variation within U‘S. populations (Selander and Foltz 1981). Climate also has great influence on the di- versity of shell colors. Wood snails, commonly found in cool temperate climates, are sensitive to overheating (Armold 1969; Jones et al. 1977). One study of populations on sand dunes found a disproportionate number of brown and pink shelled individuals dying from heat shock (Jones et al. 1977). Climatic selec- tion is thought to play a major role in large- scale patterns of shell color, with pale shells being selected for in hotter climates (Jones et al. 1977). Indeed, there is a cline for shell col- or across Europe, and in the hottest parts of their European range, yellow shelled wood snails are the most common type (Jones et al. 1977). Additionally, observation of shells dug from archaeological sites in England shows that, historically, brown shells were more com- mon during periods with colder climates (Jones et al. 1977). However, interpretation of the interplay between climate and color is complicated by the fact that small scale envi- ronmental conditions may also have an effect Cepaea nemoralis (Gastropoda. Helicidae)—Whitson 87 (Arnold 1968, 1969; Jones et al. 1977). For example, even in regions with generally warm climates, brown shells may be favored in cer- tain cool, shady microhabitats, because brown individuals absorb heat faster than pale indi- viduals and can thus become active more quickly (Jones et al. 1977). Early workers observed that wood snails had a tendency to “mimic” their backgrounds, with the shell colors that best blended into the background being the most common (Howe 1898). Later workers have shown that visual selection by predators can produce this effect (Currey et al. 1964; Davison 2002; Jones et al. 1977). In Great Britain, song thrushes are ef- ficient snail predators, crushing the shells on stones to get to the snail. Birds see in color, and in areas where song thrushes are com- mon, shells which contrast with their back- grounds are preferentially eaten (Currey et al. 1964; Jones et al. 1977). Other predators such as mammals and glowworms have also been shown to prefer certain shell morphs over oth- ers (Jones et al. 1977). With such a wealth of information on the genetic control of shell color, diversity of nat- ural populations, and factors influencing shell morphs, wood snails have become wonderful model systems for study of evolutionary mech- anisms and ecological genetics (Davison 2002). These organisms have the added ad- vantage of being common and easy to work with in both the field and the lab. ECOLOGICAL IMPACT Some authors have expressed concern about the potential impact of non-native snails upon populations of our native species (Cowie and Robinson 2001; Mead 1971). Several years ago, wood snails were introduced to the Stone Lab area of Gibraltar Island (Ohio) via land- scaping activities, and Dr. Michael Hoggarth of Otterbein College has since noted an ap- parent decrease in the numbers of native snails seen there (pers. comm.). This is an is- sue that calls for further study. CONCLUSION While recent surveys of C. nemoralis pop- ulations in the U.S. are limited, it is obvious that this species has become widely estab- lished. Several populations from Kentucky have been noted, and further searching would undoubtedly uncover more. Though the spe- cies is apparently not a major agricultural pest, the potential impact of these very successful aliens on our native snail populations should be of concern. However, now that the wood snail has come to stay, its potential for use in the classroom or for ecological genetics stud- ies is an opportunity not to be overlooked. ACKNOWLEDGMENTS Thanks to Dr. Michael Hoggarth of Otter- bein College in Westerville, Ohio for critiqu- ing this paper; to Merritt Gillilland of Michi- gan State University for advice on finding some of the early literature on Cepaea intro- ductions in the U.S.; to Dr. Debra Pearce, Northern Kentucky University, for aid; and to Dr. John Thieret and the staff of the Lloyd Library in Cincinnati for providing access to a variety of delightful and very vintage malaco- logical works. LITERATURE CITED Abbott, R. T. 1950. Snail invaders. Nat. Hist. 59(2):80—85. Abbott, R. T. 1989. Compendium of landshells. American Malacologists, Melbourne, FL. Arnold, R. 1968. Climatic selection in Cepaea nemoralis (L.) in the Pyrenees. Philosoph. Trans. Roy. Soc. Lon- don, Ser. B, Biol. Sci. 253:549-593. Arnold, R. 1969. The effects of selection by climate on the land-snail Cepaea nemoralis. Evolution 23:370-378. Barber, M. D. 1918. Helix nemoralis in Knoxville, Tenn. The Nautilus 31:107. Binney, W. G., and T. Bland. 1869. Land and fresh water shells of North America, Vol. 1. Smithsonian Misc. Coll. 194. Branson, B. A., and D. L. Batch. 1969. Notes on exotic mollusks in Kentucky. The Nautilus §2:102-106. Brooke, J. M. (Mrs.) 1897. The colony of Helix nemoralis at Lexington, Va. The Nautilus 10:142-143. Brussard, P. F. 1975. Geographic variation in North Amer- ican colonies of Cepaea nemoralis. Evolution 29:402- 410. Burch, J. B. 1962. How to know the eastern land snails. W. C. Brown Co., Dubuque, IA. Cain, A. J., P. M. Sheppard, F. R. S. King, and J. M. B. King. 1968. Studies on Cepaea: the genetics of some morphs and varieties of Cepaea nemoralis (L.). Philo- soph. Trans. Roy. Soc. London, Ser. B, Biol. Sci. 253: 383-396. Cowie, R. H., and D. G. Robinson. 2001. Pathways of introduction of nonindigenous land and_ freshwater snails and slugs. California Department of Food and Agriculture, Plant Pest Diagnostic Center, 2001 Inva- sive Mollusc Conference, Sacramento, CA. Accessed at 88 Journal of the Kentucky Academy of Science 66(2) http://www.cdfa.ca.gov/phpps/ppd/Entomology/Snails/ pathwayspub.htm on 26 Oct 2005. Currey, J. D., R. W. Arnold, and M. A. Carter. 1964. Fur- ther examples of variation of populations of Cepaea ne- moralis with habitat. Evolution 18:111—117. Davison, A. 2002. Land snails as a model to understand the role of history and selection in the origins of bio- diversity. Population Ecology 44:129-136. Dees, L. T. 1970. Edible land snails in the United States. United States Department of the Interior, U.S. Fish and Wildlife Service, Resource Publ. 91. Washington, D.C. Dundee, D. S. 1974. Catalog of introduced molluscs of Eastern North America (North of Mexico). Sterkiana 5D 137, [FNMH] Florida Museum of Natural History. 2005. In- vertebrate Zoology Database. Accessed at http://www. flmnh.ufl.edu/scripts/dbs/malacol_pub.asp on 27 Oct 2005. Grime, J. P., S. F. MacPherson-Stewart, and R. S. Dear- man. 1968. An investigation of leaf palatability using the snail Cepaea nemoralis L. Ecol. 56:405-420. Howe, J. L. 1898. Variation in the shell of Helix nemoralis in the Lexington, Va., colony. Am. Naturalist 32:913— 923. Hubricht, L. 1968. The land snails of Mammoth Cave National Park, Kentucky. The Nautilus 82:24—28. Johnson, C. W. 1928. Further notes on the colony of Helix nemoralis in Massachusetts. The Nautilus 41:47—49. Jones, J. S., B. H. Leith, and P. Rawlings. 1977. Polymor- phism in Cepaea: a problem with too many solutions? Annual Rev. Ecol. Syst. 8:109-143. Judd, W. W. 1953. A colony of the land snail Cepaea ne- moralis (L.) (Helicidae) in the vicinity of London, On- tario. Canad. Field-Naturalist 67:87—-89. Kaplan, M. F., and W. L. Minckley. 1960. Land snails from the Doe Run Creek area, Meade County, Kentucky. The Nautilus 74:62-65. Lovell, M. S. 1884. The edible mollusca of Great Britain and Ireland, 2nd ed. L. Reeve and Co., London. Mead, A. R. 1971. Helicid land mollusks introduced into North America. The Biologist 53:104-111. Murray, J. 1964. Multiple mating and effective population size in Cepaea nemoralis. Evolution 18:283-291. National Atlas of the United States. Accessed at http:// nationalatlas.gov on 31 Oct 2005. Pilsbry, H. A. 1939. Land mollusca of North America (north of Mexico), Vol. 1, part 1. Acad. Nat. Sci. Phil- adelphia. Monogr. 3. Reed, C. F. 1964. Cepaea nemoralis (Linn.) in eastern North America. Sterkiana 16:11-18. Reeve, L. 1863. The land and freshwater mollusks indig- enous to, or naturalized in, the British Isles. Reeve and Co., London. Rimmer, R. 1907. Shells of the British Isles. Land and Freshwater. John Grant, Edinburgh. Selander, R. K., and D. W. Foltz. 1981. Gametic disequi- librium between esterase loci in populations of Cepaea nemoralis in western New York. Evolution 35:190-192. Sowerby, J. D. C. 1825. Helix nemoralis, a carnivorous animal? Zool. J. 1:284-285. Step, E. 1901. Shell life. An introduction to the British mollusca. Frederick Warne & Co., London. Stine, O. C. 1989. Cepaea nemoralis from Lexington, Vir- ginia: the isolation and characterization of their mito- chondrial DNA, the implications for their origins and climactic selection. Malacologia 30:305-315. Thompson, R. 1996. Raising snails. Special Reference Briefs Series # SRB 96-05. Alternative Farming Systems Information Center, National Agricultural Library, Beltsville, MD. Accessed at http:/Avww.nal.usda.gov/ afsic/AFSIC_pubs/srb96-05.htm on 26 Oct 2005. Turton, W. 1857. Manual of the land and fresh-water shells of the British Islands. Longman, Brown, Green, Longmans, and Roberts, London. Woodward, B. B. 1913. The life of the Mollusca. Methuen & Co., London. J. Ky. Acad. Sci. 66(2):89-93. 2005. Location of Rediae of Proterometra macrostoma (Trematoda: Azygiidae) in the Snail Elimia semicarinata (Gastropoda: Pleuroceridae), and Daily Emergence of its Cercaria Ronald Rosen, Jonathan Fleming, Bojana Jovanovic, Aishe Sarshad, Emilie Throop, Fady Zaki, and Andy Ammons Department of Biology, Berea College, Berea, Kentucky 40404 ABSTRACT The objectives of this study were to describe the (1) location of Proterometra macrostoma rediae in the snail intermediate host, Elimia semicarinata, and (2) daily emergence of cercariae from individual snails over 3 weeks. Decalcification and histological sections of infected snails revealed rediae restricted to the bottom whorl within the mantle cavity in close association with the host gills. A continuous, low-level emergence of cercariae (0.25—0.833 larvae/snail/day) was observed over 21 days. Both moderately frequent (7-12 days) and infrequent (2-6 days) shedding of cercariae were observed (i.e., mean + SE = 7.3 + 0.4 days; range = 2— 12 days), but no consistent pattern was apparent. The average + SE number of rediae was significantly smaller in snails shedding only 2-6 days (11.9 + 2.9) than snails shedding between 7-12 days (35.2 + 8.5) at the termination of the experiment. INTRODUCTION The original work on morphology and gen- eral biology of the cercaria of Proterometra macrostoma was completed by Horsfall (1934) and Dickerman (1945). Horsfall (1934) noted the presence of larval forms of P. macrostoma in the snail body cavity. Dickerman (1945), based on his study of crushed snails, indicated that the size and number of emerging cercar- iae may cause the body cavity wall to rupture, thus introducing cercariae into the mantle cav- ity from where they can exit into the outside environment. However, Hyman (1967) and Voltzow (1994), in extensive reviews of pros- obranch anatomy, did not mention the pres- ence of a body cavity associated with these snails. Thus, these gross observations should be reassessed by histological techniques aimed at describing the correct location of the P macrostoma rediae and cercariae within their snail host. Lewis (1988) determined that an average of only 0.35 and 0.29 P macrostoma cercariae/ snail were released daily in the field and lab- oratory from sample populations of snails, re- spectively. This low production of P. macro- stoma cercariae was also documented in a 3- week study of daily emergence (Lewis et al. 1989). According to Lewis et al. (1989), this is 100-—1000-fold less than most other digeneans, 89 representing an adaptation for the creation of a few, large, conspicuous cercariae. However, no hypothesis was offered for this low cercarial output. Evaluation of cercarial emergence pat- terns from individual snails and a reexamina- tion of P. macrostoma redial stages may fur- ther our understanding of this phenomenon. The objectives of this study were (1) to de- termine the location of P. macrostoma rediae in the snail intermediate host and (2) to de- scribe the daily emergence of cercariae during 3 weeks from individual snails naturally in- fected with this worm. MATERIALS AND METHODS Snails of the species Elimia semicarinata were collected from North Elkhorn Creek in Scott County, Kentucky (lat 38°11’00” N, long 84°29'19" W), during summer 2004. They were then screened for patent infections (i.e., shedding cercariae) as described by Rosen et al. (2000). Thirty-six infected snails were si- multaneously fixed and decalcified in Cal-Ex II (Fisher) for routine paraffin sectioning to describe redial location in the snail host. Serial sections (5-10 ww) were stained with hematox- ylin and Gomori’s trichrome. Emergence of cercariae was assessed for 36 additional in- fected snails that were individually isolated, held at 20°C under a 12 hr light: 12 hr dark cycle, and checked at the end of each of these “"PUROII0 PUODES B JO Apoq O} SULIOYpr BLIROIGO QUO JO JOYONS [RIO = x *[RUS JO IOYM WOYWOG = A “ToyONsS [eIUSA = SA ‘BLIBOIGD aINYeU JO |e} = UI} ‘sa}so} = 9} ‘eIpot Jo yuswinse} = } ‘oeIpOI = I -Aoyons [R10 = SO -eLIeOIOD oINQeUL = OUT SAYIABO opurul = Ww ‘sqEs — 8 ‘euroi1s0 — 0 ‘omyerode [meus = B ‘suOnRIASIGGY ‘oWOIYOL} sLIOW0s) pue uydxoyeuray ‘11 OT !x 10.0 1.0-2.0 MGS/GS =10.0 2.0—5.0 MG/GS 10.0 2,.0-5.0 SG/GS 2 a) MG/PDA ~~ 2),0 5.0-10.0 MnCl,-4H,O, FeCl,-6H,O, ZnSO,-H,O, and CuSO,, 80 wg each. Stocks of each mineral nutrient (macronutrients, 100; micronutri- ents, 1000) were autoclaved separately, as were glucose solutions. A commercially-for- mulated herbicide was used as our paraquat (PQ) source (Gramoxone Extra, 2.5 Ibs of PQ per gallon, Syngenta Corporation, 2200 Con- cord Pike, P.O. Box 8353, Wilmington, DE 19803). Two commercially available neem (NM) preparations, both listing clarified hy- drophobic extract of neem oil as the active in- gredient (a.i.), were purchased locally. These were Green Light Tomato and Vegetable Spray, Ready-To-Use, containing 0.9% ai. (NMRTU; Green Light Company, San Anto- nio, TX 75418) and Garden Safe® Fungicide 3™ Concentrate containing 70% a.i. (NMC; Schultz Company, Bridgeton, MO 63044). Polystyrene Petri plates (100 < 15 mm) and polystyrene 24-well BD Falcon culture plates were purchased from Fisher. Fluorescent lighting tubes were purchased locally. The ger- micidal ultraviolet lamp (model UVG-54) em- ployed for visualization of SA fluorescence was purchased from UVP, Inc., 2066 W. 11th Street, Upland, CA 91786. Characteristics, Sources, and Maintenance of Microorganisms The names, in-text abbreviations, character- istics, and sources of the microorganisms em- ployed in this research are presented in Table 1. We isolated all fungi (except Colletotrichum graminicola, CG) from naturally-infected plant materials collected in the area of Lex- 120 ington, KY. Fungi were maintained by serial transfer on PDA, typically at 1-4 week inter- vals, although sometimes more frequently. Stock and experimental cultures were main- tained at room temperature, typically 20- 22°C. Botrytis cinerea (BC) and Pestalotia sp. (PESP) stock cultures were commonly main- tained in the dark, although PDA cultures of PESP and CG were sometimes maintained under standard fluorescent lights (12 hr light/ dark and 24 hr light, respectively) to promote spore germination. Spore suspensions of CG and PESP were prepared by agitating small plugs (ca. 3 X 10 mm) cut from sporulating cultures on PDA in sterile distilled water (SDW) with a vortex mixer. Spores of Moni- linia fructicola (MF) were produced by cul- turing the fungus on commercially-available unsulphured dried apricots, which were re- hydrated prior to sterilization by autoclaving. Spores of MF were transferred to a small vol- ume of SDW with a transfer loop, and the suspension was agitated with a vortex mixer. Because the presence or absence of light was otherwise found to have no effect on the out- comes of our experiments, inoculated plates were maintained under either condition, as space permitted. Two mixed cultures of bacteria that exhib- ited antifungal activity in preliminary co-cul- ture experiments with BC and PESP were designated as “brown” or “white” based on their appearance in PDA culture. The brown culture was isolated from rose leaves exhibit- ing symptoms of the black spot disease (caused by Diplocarpon rosae, DR) and was found to completely prevent DR spore ger- mination on PDA when present in inoculum drops. This brown culture begins to produce a visually detectable, brown diffusible pigment 3—4 days after inoculation to PDA or PDB cul- ture media. The white bacterial culture was obtained as a laboratory contaminant. It also appeared to produce one or more diffusible colorless antifungal substances on PDA. Bac- terial culture fluids (BCF) from prolonged (18 days) shake cultivation of the brown and white cultures in PDB (designated BRCF and WCF, respectively) were employed. This lengthy cul- tivation period appeared to have resulted in the death of bacteria, based on the absence of growth from aliquots of culture fluids plated onto PDA. Bacterial cultures were maintained Journal of the Kentucky Academy of Science 66(2) by serial transfer on PDA, usually at 14 week intervals. Assessment of SA Effects on Growth of Fungi on PDA Because SA is not stable at autoclave tem- peratures, culture media were amended with SA after autoclaving. Petri plates containing ~20 ml of PDA were amended with 200 pl of SDW or 200 wl of sterile aqueous stock solutions of SA applied to the solidified PDA and allowed to diffuse evenly throughout the agar for 3-4 days. The extent of SA diffusion in these plates was assessed by brief visuali- zation of SA fluorescence under a germicidal UV lamp. Plugs (typically 3-5 mm on a side) were cut from the margin of 3-5 day fungal cultures on PDA and placed in the center of Petri plates. The resultant fungal colonies were measured when the fastest-growing col- onies neared the edges of Petri plates, often after 2-3 days incubation at 20-22°C in the dark. Long and short diameters of the typically ovoid individual colonies were averaged for statistical analyses. Detection and Assessment of Synergistic Interactions of SA with Other Antifungal Materials Dose-response studies with individual ma- terials were conducted with each of the test fungi employed in these experiments (BC, CG, and PESP) to guide the selection of suit- able doses for interaction studies. For experi- ments conducted on PDA, SA and PQ were added after autoclaving. For experiments con- ducted in 24-well culture plates, aliquots of the liquid GS medium (0.4, 0.5, or 1.0 ml for mycelial growth assays, 0.4 ml for spore ger- mination assays) were added to wells and then amended with stock solutions of antifungal materials. In studies of the interactive effects of SA and PQ on BC growth in well-plate cul- ture, incubation continued until wells that supported significant growth of BC were filled with fungal mycelia (typically 15 days). Growth was assessed visually and evaluated as present/strong (+), absent (—), or weak (+). In studies of SA interactions with Cu or BCF in 24-well plates, fungal growth in well-plate assays was assessed by several techniques, usu- ally after 3-4 days for mycelial growth assays and after 1 day for spore germination assays. Synergistic Inhibition—Strobel and Porter The presence or absence of growth was first assessed visually, and absence of growth then verified by microscopic observation. Radial mycelial growth from agar plugs was assessed visually with a millimeter rule, by viewing plates from beneath by transmitted ceiling light, or sublit on a microscope stage upon which the rule had been placed. In some ex- periments (“NMRTU time-course studies”), addition of SA and inoculation with fungi were delayed relative to introduction of NMRTU. Mycelial growth of BC in the NMRTU time- course study presented in this paper was eval- uated relative to that of controls according to the following rating system: 0 = no growth; 1 = >90% inhibition (weak growth); 2 = 75% inhibition (weak-moderate growth); 3 = 50% inhibition (moderate growth); 4 = 10-25% in- hibition (moderate-strong growth); 5 = 90- 100% of maximal growth (strong growth). Germination of CG and PESP spores was as- sessed with a microscope at 100X total mag- nification. In experiments with CG spores, a minimum of 120 spores per well were evalu- ated. The nature of interactions between SA and these other antifungal materials was deter- mined according to the method of Kosman and Cohen (1996). For example, in Table 3, the observed means of various treatments in- volving doses of single agents (such as 2.0 mM SA or 4.0 or 8.0 mM PQ) are first converted into percentages of the observed mean values (OPCs) for controls (which received neither SA nor PQ). Then a simple additive model of interaction (AMI) is employed to calculate the values (predicted percent of control values, PPC) one would expect to see (in the absence of either antagonistic or synergistic interac- tions between test compounds). The additive model assumes the independent impact of each agent on a test organism (such as the fungus, BC), and the impact of two agents to- gether is calculated as the product of the im- pact of the first times the impact of the sec- ond. Thus, in Table 3, we find that 2.0 mM salicylate alone reduced BC growth to 80% of the water controls, whereas 4.0 mM PQ re- duced BC growth to 31% of the water con- trols. The PPC predicted by the AMI for the combination of 2.0 mM SA and 4.0 mM PQ is determined by multiplying 0.80 times 0.31, which yields a PPC of 0.25, or 25%. An ob- 12] served percent of control (OPC) value larger than 25% would indicate a lesser degree of inhibition than predicted by a simple additive model of interaction, and would constitute ev- idence of an antagonistic (ANT) interaction between the two test substances. In the pres- ent example, the OPC for the SA-PQ combi- nation mentioned above (12%) is lower than that predicted by the AMI (25%), and the in- teraction is thus judged to have been syner- gistic (a greater than predicted or expected degree of inhibition of BC by the SA-PQ com- bination was observed). RESULTS Dose-Dependent Inhibition of the Growth of Plant-Pathogenic Fungi by SA A summary of the sensitivities of BC and other fungi to SA added to PDA or GS is pre- sented in Table 1. SA concentrations compa- rable to those that may occur in infected plant tissues (10.0—-100.0 4M endogenous SA, data not shown) had no observable effect on BC mycelial growth in our experiments. Rather, SA concentrations required for complete in- hibition (IC,,,) of mycelial growth on PDA or GS media were typically =10.0 mM, whereas between 1.0 and 5.0 mM SA were required to inhibit mycelial growth on PDA by 50% (IC3,) for most fungi, and the IC;, of PYSP was 5.0-— 10.0 mM. Germinating CG spores were rela- tively more sensitive to SA than were spores of MF or PESP. The consistent nature of the dose-dependent inhibition of fungal growth in SA-amended PDA is shown by Table 2, which summarizes data from three consecutive ex- periments with BC. Synergistic Inhibition of BC by SA and the Pro-oxidant Herbicide, PQ SA and PQ were found to inhibit growth of BC on PDA in a synergistic manner (Table 3, Figure 1). Growth of BC on PDA amended with combinations of 2.0 mM SA and 4.0 or 8.0 mM PQ was substantially less than that predicted by an additive model of interaction, and these interactions were judged to be syn- ergistic in nature. (See Material and Methods section for details of the evaluation of inter- actions.) Synergistic inhibition of BC on PDA was also observed for other combinations of SA and PQ (5.0 mM SA with 4.0 or 8.0 mM PQ; data not presented). In a well-plate test 122 Table 2. Journal of the Kentucky Academy of Science 66(2) Dose-dependent inhibition of Botrytis cinerea by sodium salicylate (SA) in potato-dextrose agar. Data pre- sented are from three replicate experiments, and each value is the mean + SE colony diameter (mm) of four replicate Petri plates. Trial 0.0 mM SA 0.2 mM SA 0.5 mM SA 1 C( 007 65.9 + 0.8 60.5 + 1.4 2 66.5 = 0.7 63:3 = 0:6 Dio 220.5 5) 67 .0r 2) 66.0 + 1.1 62335222 7 of the interactive effects of SA and PQ on BC, the fungus grew to fill all six wells per treat- ment that had been amended individually with 0.0 or 16.0 mM PQ or 2.0 or 5.0 mM SA, but did not grow in wells amended with 16.0 mM PQ and 2.0 or 5.0 mM SA (data not shown). Synergism of SA with Cupric Chloride Results of two replicate trials of an experi- ment designed to evaluate the potential inter- active effects of SA and Cu on BC mycelial growth in well-plate culture are presented in Table 4. SA alone caused only a slight reduc- tion in mycelial growth of BC, whereas growth of BC decreased gradually with increasing concentrations of Cu alone, and complete in- hibition of BC growth was observed only with 10.0 mM Cu. In contrast, complete inhibition of BC growth occurred with 5.0 mM Cu (first experiment) or 2.0 mM Cu (second experi- ment) in the presence of 2.0 mM SA. Data regarding the synergistic inhibition of CG Table 3. celial growth in potato-dextrose agar amended with sodi- um salicylate (SA) and/or paraquat (PQ). Data presented are mean colony diameters (mm) + standard error for three replicate plates per treatment combination. PQ was mixed into partially cooled molten agar prior to pouring Synergistic inhibition of Botrytis cinerea my- into Petri plates (20 ml/plate), whereas SA was introduced as 0.2 ml of a concentrated stock (100) to the centers of plates and permitted to diffuse evenly throughout agar (as assessed by SA fluorescence under ultraviolet illumi- nation). Other abbreviations: OPC, observed percent of control; AMI, additive model of interaction; PPC, pre- dicted percent of control; SYN, synergistic interaction. PQ 0.0 mM PQ 4.0 mM PQ 8.0 mM SA 0.0 mm 83.3 204 222) 9 13672732 OPC 100 31 AN SA 2.0 mM 01.202 P10 S IG 42.= 01 AMI — (80)(31) (80)(17) DEC — 25 14 OPC 80 12 5 — SYN SYN 1.0 mM SA 2.0 mM SA 5.0 mM SA 10.0 mM SA D2 eS O04 2 lo 24 4 218s WAL Se, ere 50.0 + 0.4 AES 25.6) al 19D 23 56:4. 22156 36.8 + 1.8 27 ae 15.6 + 3.6 spore germination by SA and Cu are present- ed in Table 5. SA alone, at concentrations of 0.2 and 0.5 mM, did not affect the frequency of spore germination, whereas 0.75 mM SA inhibited mean germination by 61%, relative to controls. No germination of CG spores was observed in the presence of 1.0 or 2.0 mM SA (data not presented). In the absence of SA, Oe spore germination was progressively in- hibited by increasing Cu concentrations. Strong synergistic interactions were observed in combinations of 0.75 mM SA with Cu. For example, although percent spore germination relative to controls (OPC) was 94 and 39 for 0.2 mM Cu and 0.75 mM SA, respectively, this combination resulted in an OPC of 4% spore germination, much lower than the 37% ger- mination predicted by the additive model. In- terestingly, 0.2 and 0.5 mM SA effectively abolished the inhibition of spore germination by 0.2, 0.35, and 0.5 mM Cu, and SA thus appeared to act as an antagonist of Cu fungi- toxicity toward CG spores in these combina- tions. In a preliminary experiment with PESP, the combination of 2.0 mM SA with 0.5 mM Cu completely prevented spore germination, whereas these concentrations of SA or Cu alone appeared to have little or no effect on the frequency of spore germination (data not shown). In contrast to results obtained with CG, the presence of 0.2 or 0.56 mM SA did not alleviate the moderate inhibition of PESP germ tube elongation occasioned by 0.2 or 0.5 mM Cu (data not shown). Synergistic Inhibition of BC and PESP by SA in Combination with Culture Fluids of Antagonistic Bacteria The antifungal activities of BRCF and WCF toward BC and PESP were synergistically en- hanced by 2.0 mM SA (Table 6; Figure 2). BC growth was not prevented by exposure to SA Synergistic Inhibition—Strobel and Porter 1238 2 Mum Voy \ if Se o Figure 1. Synergistic inhibition of Botrytis cinerea (BC) mycelial growth in potato-dextrose agar PDA amended with sodium salicylate (SA) and/or paraquat (PQ). Pictured are all ee replicate plates from the SA 2.0 mM-PQ 4.0 mM (1/400) subset of a larger experiment whose data are presented in Table 3. The leftmost column of Petri plates (column 1) shows growth of BC in the absence of SA and PQ. Column 2 shows growth of BC in the presence of 2.0 mM SA. Column 3 shows growth of BC in the presence of 4.0 mM PQ. Column 4 shows growth of BC in the combined presence of 2.0 mM SA and 4.0 mM PQ. Table 4. Synergistic inhibition of Botrytis cinerea mycelial growth in a liquid glucose-salts medium amended with sodium salicylate (SA) and cupric chloride (Cu). Data are of two replicate trials in which each SA-Cu treatment combination was represented twice in each of three replicate 24-well culture plates (n = 6 for each treatment com- bination). Values presented are mean + standard error extent of radial mycelial growth (mm) from inoculum plugs, measured at 72 hours or 48 hours after BC inoculation (Trials I and II, respectively). Other abbreviations: OPC, observed percent of control; SYN, synergistic interaction. Cu 0.0 mM Cu 0.5 mM Cu 1.0 mM Cu 2.0 mM Cu 5.0 mM Cu 10.0 mM Trial I SA 0.0 mM Do 2 O02 45 20.1 3.7 + 0.1 2.0 + 0.0 1G 22 Oa1 0.0 + 0.0 OPC 100 fot) 70 38 34 0 SA 2.0 mM 46+ 0.1 3. + 0.1 2 EO O:7-22 Oa 0.0 + 0.0 O02 00 OPC 87 66 43 3 0 0) — SYN SYN SYN SYN — Trial I SA 0.0 mM 7 Vip mens 0 | So mena.) all 2.8 + 0.1 2.0 O10 | a Gemet § Re | O:0222.0:0 OPC 100 78 62 44 38 0 SA 2.0 mM 43+ 0.1 pAb fae 6 A | eS 2 O"1 0.0 + 0.1 0.0 + 0.0 0.0 + 0.0 OPC 96 60 25 0) 0) 0 — SYN SYN SYN SYN — 124 Journal of the Kentucky Academy of Science 66(2) Table 5. Synergistic inhibition of Colletotrichum graminicola spore germination in a liquid glucose-salts medium amended with sodium salicylate (SA) and cupric chloride (Cu). Germination was assessed microscopically based on the presence (germinated) or absence (ungerminated) of an appresorium. No germ tubes were observed in the absence of appresoria. All spores in one microscopic field at 100 total magnification (>120 spores per well) were evaluated. Values presented are mean * standard error for two replicate wells per treatment (n = 2). Other abbreviations: OPC, observed percent of control; ADD, ANT, and SYN denote additive, antagonistic, and synergistic interactions, respec- tively. No germination was observed in the presence of 1.0 or 2.0 mM SA, and these data are not presented in the body of the table. Cu 0.0 mM Cu 0.2 mM Cu 0.35 mM Cu 0.5 mM SA 0.0 mM GID okt 86.0 + 2.4 40.0 + 2:1 SiO) BaD) OPC 100 94 Th Al SA 0.2 mM 89.0 + 0.0 84.0 + 2.8 84.0 + 2.8 86.5 + 1.1 OPC 97 92 92 95 = ADD ANT ANT SA 0.5 mM 96.5 + 0.4 96.0 + 0.0 Ol: <2 18 89.5 + 0.4 OPC 105 105 100 98 — ANT ANT ANT SA 0.75 mM 36.0.2 7.1 4.0 + 0.0 1.0 + 0.0 0.0 + 0.0 OPC 39 4 ih 0 — SYN SYN SYN or BRCF alone. However, the combination of SA with 100 or 200 wl of BRCF prevented BC growth in 2/3 and 3/3 wells, respectively. Neither BRCF nor SA alone prevented PESP growth at the doses employed. However, the addition of SA prevented PESP growth in 2/3 wells that received 50 wl of BRCF, and in all three wells that received 100 or 200 wl of BRCF. Only a slight effect of SA on inhibition of BC by WCF was observed, whereas growth of PESP was uniformly prevented in all com- binations of SA and WCF. The Interactive Effects of SA and NM These effects varied with NM source [NM concentrate (NMC) or ready-to-use (NMRTU) formulations], SA and NM doses employed, timing of fungal inoculation following addition of NMRTU to GS, and species of test fungus. In trials with NMC, little or no inhibition of BC mycelial growth was observed with NMC alone or in combination with SA, although a transient delay in pigmentation was commonly observed in NMC-SA combinations (data not presented). In an experiment in which NMRTU was added to wells immediately pri- or to inoculation with BC, only a slight inhi- bition of BC by the highest dose of NMRTU was observed in the absence of SA (Table 7). Amendment with 2.0 mM SA only slightly en- hanced inhibition by NMRTU dilutions of 1/10X and 1/5 (data not shown). However, Table 6. Synergistic inhibition of mycelial growth of Botrytis cinerea (BC) and Pestalotia sp. (PESP) in a liquid glucose- salts medium amended with sodium salicylate (SA) and bacterial culture fluids (BCF). Single 24-well culture plates were filled with 0.5 ml GS per well to which were added 0, 50, 100, or 200 wl of BCF from the 18 d potato-dextrose broth shake cultures of an unidentified mixed white bacterial culture (WCF) or an unidentified mixed brown culture (BRCF). Fungal growth was assessed as present or absent for each of three replicate wells per treatment (one 24-well culture plate per BCF-fungus combination). WCE (lAvell) BRCF (wlAvell) 0 50 100 200 0 50 100 200 Incidence of BC mycelial growth SA 0.0 mM 3/3 3/3 3/3 2/3 3/3 3/3 3/3 3/3 SA 2.0 mM 3/3 2/3 3/3 WS oo 3/3 1/3 0/3 Incidence of PESP mycelial growth SA 0.0 mM ofS 3/3 By8} 2/3 o/o 3/3 3/3 3/3 SA 2.0 mM 3/3 0/3 0/3 0/3 3/3 1/3 0/3 0/3 Synergistic Inhibition—Strobel and Porter 125 Figure 2. Synergistic inhibition of Pestalotia sp. (PESP) in a liquid glucose-salts medium amended with sodium salicylate (SA) and/or fluids from the 18 day potato-dextrose broth culture of an unidentified mixed brown bacterial culture (BRCF). Columns 1, 3, and 5 contained no SA, whereas columns 2, 4, and 6 were amended with 2.0 mM SA (final concentration). Rows A, B, C, and D were amended with 0, 50, 100, 200 wl of BRCF, respectively. Plating of BRCF aliquots onto potato-dextrose agar revealed the absence of live bacteria (no growth). a strong synergism was observed in combina- both NMC and NMRTU products (when plat- tions of 4.0 mM SA with the higher NMRTU _ ed onto PDA), but only those in NMRTU ap- concentrations employed (1/20, 1/10, and peared to multiply after addition to GS. Be- 1/5X dilutions). Microscopic investigations re- cause a review of several preliminary experi- vealed that diverse bacteria were detectable in ments revealed that inhibition of fungi by Table 7. Synergistic inhibition of Botrytis cinerea mycelial growth by sodium salicylate (SA) and a ready-to-use com- mercial formulation of neem oil extract (NMRTU). NMRTU dilutions were achieved by adding 0-100 pl NMRTU to wells containing sufficient liquid glucose-salts medium to yield 0.5 ml total volumes. Final SA concentrations of 4.0 mM were achieved by adding 20 wl of a 100 mM SA stock per well. The extent of mycelial growth (mm) from inoculum plugs was measured after a 48 hour incubation period. Data are mean + standard error of six replicate wells (n = 6), for each SA 0.0 mM-NMRTU combination, and three replicate wells (n = 3) for each SA 4.0 mM-NMRTU combi- nation. Other abbreviations: OPC, observed percent of control. NMRTU 0 NMRTU 1/100X NMRTU 1/50x NMRTU 1/20x NMRTU 1/10x NMRTU 1/5x SA 0.0 mM A cae Os 42+ 0.1 3.625 0:1 oof = 01 3-2, Oell 3,0 02 OPC 100 98 88 86 86 70 SA 4.0 mM Oa 702 Bo e2 Qe ae 02 0.7 + 0.4 0.0 + 0.0 0.0 + 0.0 OPC (86) es 63 16 0 0 — SYN SYN SYN SYN SYN 126 Journal of the Kentucky Academy of Science 66(2) Table 8. Effect of timing of addition of sodium salicylate (SA) and a commercial ready-to-use formulation of a neem oil extract (NMRTU), relative to inoculation with Botrytis cinerea, on synergistic inhibition of mycelial growth. Data presented are growth incidence (GI, presence or absence) in three replicate wells per treatment combination (n = 3); and growth rate (GR) as mean ~ standard error for growth relative to controls, with a rating of 0 indicating no growth; 1,=10%; 2, 10-50%; 3, 51-75%; 4, 75-90%; or, 5, 91-100% growth relative to controls. No pretreatment indicates that NMRTU and SA were added to wells on the same day as BC inoculation, whereas 3-day pretreatment indicates that NMRTU and SA were introduced 3 days before BC inoculation. Other abbreviations: OPC, observed percent of control; ADD, ANT, and SYN denote additive, antagonistic, and synergistic interactions, respectively. 1/100 NMRTU dilutions 1/5X GI GR GI GR GI GR GI GR No Pretreatment SA 0.0 mM 3/3 5.0 + 0.0 3/3 5.0 + 0.0 3/3 4.0 + 0.0 wo To) BE Oat OPC 100 100 80 46 SA 2.0 mM 3/3 4.0 + 0.0 3/3 4.0 + 0.0 3/3 4.0 + 0.0 3/3 2.0 + 0.0 OPC 80 80 80 40 — ADD ANT ANT Three Day Pretreatment SA 0.0 mM 3/3 5.0 70:0 3/3 4.0 + 0.0 3/3 3.0 + 0.0 3/3 o.0) = 0:0 OPC 100 80 60 60 SA 2.0 mM ofS o0/z2 O20 3 0:3 =. 0 1/3 Osrs07 0/3 0.0 + 0.0 OPC 100 Onl On 0 — SYN SYN SYN combinations of NMRTU and SA was most consistent, frequent, and extensive when in- oculation with BC had occasionally been de- layed by one or more days after amendment of wells with NMRTU and SA, the effect of varied duration (0, 1, 2, or 3 days) of such delay was formally investigated. Data regard- ing the effects of a 0 or 3 d delay are pre- sented in Table 8. Plates that received NMRTU, SA, and BC on the same day exhib- ited mycelial growth in all wells of each treat- ment combination, and interactions of SA and NMRTU were found to be either additive or slightly antagonistic. In contrast, when BC in- oculation was delayed until 3 days after addi- tion of SA and NMRTU, synergistic inhibition of BC growth was observed in all SA-NMRTU treatment combinations. Turbidity associated with bacterial growth was much greater in plates that received NMRTU (with or without SA) 3 days prior to inoculation with BC than in wells that received NMRTU and SA im- mediately prior to BC inoculation. These dif- ferences in turbidity were observed both at the time of inoculation with BC and at 3 days post-inoculation, when BC growth was as- sessed. Interestingly, the addition of SA and NMRTU 1 day before BC inoculation resulted in only slight development of turbidity and no inhibition of BC, whereas turbidity and extent of BC inhibition were of intermediate degree when BC inoculation occurred 2 days after ad- dition of SA and NMRTU to wells (data not shown). The interactive effects of NMRTU and SA on mycelial growth of PESP were also examined. Again, little impact of NMRTU (alone or in combinations with SA) was ob- served when NMRTU and SA amendment oc- curred on the same day as BC inoculation (data not presented). In an experiment in which NMRTU and SA addition preceded BC inoculation by 2 days (Table 9), NMRTU alone reduced mean mycelial growth of PESP relative to controls in a dose-dependent man- ner, and the addition of SA resulted in a mod- est synergistic enhancement of fungal inhibi- tion with most of the NMRTU doses em- ployed. DISCUSSION Millimolar concentrations of SA were found to inhibit the growth and development of sev- en plant-pathogenic fungi in vitro, suggesting that sensitivity of fungal plant pathogens to mM SA concentrations may be a general phe- nomenon. Thus, SA concentrations equivalent Synergistic Inhibition—Strobel and Porter 127 Table 9. Synergistic inhibition of mycelial growth of Pestalotia sp. by sodium salicylate (SA) and a ready-to-use com- mercial formulation of a neem oil extract (NMRTU). NMRTU dilutions were achieved by adding 0-100 wl NMRTU to wells containing sufficient liquid glucose-salts medium (GS) to yield 0.5 ml total volumes. SA concentrations were achieved by adding 10 or 20 wl of a 100 mM stock to yield final concentrations of 2.0 mM or 4.0 mM. The extent of mycelial growth (mm) from inoculum plugs was measured after a 48 hour incubation period. Data are mean + standard error of four (SA 0.0 mM) or two (SA 2.0 mM) replicate wells per treatment. No PESP growth occurred in the presence of 4.0 mM SA (n = 2 per combination with NMRTU), and these data are omitted from the body of the table. Other abbreviations: OPC, observed percent of control; ADD and SYN denote additive and synergistic interac- tions, respectively. NMRTU 0 NMRTU 1/100X NMRTU 1/50 NMRTU 1/20 x NMRTU 1/10 NMRTU 1/5x SA 0.0 mM 6.0 + 0.2 4.0 + 0.0 3.0 = 0.1 2.8 = 0:1 L9 = 0.1 0.9 + 0.1 OPC 100 67 47 32 15 SA 2.0 mM 4.5 + 0.4 3:0° 0:0 20 = 0.0 15 + 0.4 1.0 + 0.4 0.0 + 0.0 OPC 75 50 3k 25 MZ 0 — ADD SYN SYN SYN SYN to those commonly used for the experimental induction of disease resistance in plants (2.0- 10.0 mM) are potentially directly antifungal as well. Sensitivity of plant-pathogenic fungi to SA is consistent with prior published reports of the inhibition of saprophytic fungi by SA (Cruess and Irish 1931). However, concentra- tions of SA equivalent to those reported to oc- cur in infected plant tissues (10.0-100.0 wM endogenous SA) did not inhibit the growth of BC in vitro. Thus, it seems unlikely that en- dogenous levels of SA in infected plant tissues (10-100 4M) would alone be capable of in- hibiting growth of pathogens in plants, as pro- posed by Ruffer et al. (1995). This does not rule out the possibility that SA may act in con- cert with other substances produced by plants to limit pathogen activity (see below). We observed multiple instances of in vitro synergism of SA with diverse other antifungal materials (PQ, Cu, BCF, and NMRTU). The mechanisms underlying these synergistic in- teractions remain to be determined. BC pro- duces a wide variety of enzymes that degrade reactive oxygen species (Steel and Nair 1993; Choi et al. 1997; Gil-ad et al. 2000) that might be inhibited by SA as are some related plant enzymes (Chen et al. 1993; Durner and Kles- sig 1995; Slaymaker et al. 2002). Another plau- sible mechanism for the observed effects of SA is the suppression by SA of mitochondrial generation of ATP (Norman et al. 2004), which supplies energy necessary for many as- pects of cellular life, including the production, operation, and maintenance of various detox- ification mechanisms. In the case of Cu, SA may also increase entry of the toxicant into fungal cells via chelation. However, this would not likely account for the observed antagonis- tic interactions of lower concentrations of SA with Cu seen with CG but not with PESP. In the case of NMRTU, matters seem more com- plex, and may involve multiple interactions of NMRTU active or inert ingredients, contami- nating bacteria, and SA. It is also plausible that acidification of the growth medium by anti- fungal materials or by fungi in response to same (not examined in the present studies) may result in increased uptake of salicylate, which is more lipid-soluble and more fungi- toxic in the protonated form (Cruess and Irish 1931). Whether SA may usefully synergize the antifungal activities of commercial synthetic fungicides also remains to be determined. We speculate that SA may be of practical utility as a multi-functional component (induc- er of local disease resistance, direct antifungal agent, modifier of the composition and size of microfloral populations on plant surfaces, and synergist of other antifungal materials) of new plant disease-control formulations that com- bine ingredients of synthetic chemical and/or biological origin. Such combinations may delay or prevent development of pathogen resis- tance to conventional fungicides, and may also enable reduced application rates for these fun- gicides (Ye et al. 1995). For example, fixed copper fungicides such as Bordeaux mixture are often applied at rates that deliver approx- imately 1-2 kg of elemental copper per acre per application, sometimes resulting in the ac- cumulation of copper to phytotoxic levels in 128 soils. Enhancement of the fungitoxicity of cu- pric ions by SA may permit large reductions in the total amount of Cu applied for plant disease management, thereby reducing or eliminating problems from Cu accumulation. Evaluation of these possibilities requires fur- ther testing of the effects of SA and other ma- terials on the outcomes of plant-pathogen in- teractions in controlled and field environ- ments. ACKNOWLEDGMENTS We gratefully acknowledge the financial support of two mini-grants from the Bluegrass Community and Technical College, 470 Coo- per Drive, Lexington, KY 40506-0235, without which our research would not have been pos- sible. We also give thanks to Dr. William Witt, Department of Agronomy, University of Ken- tucky, Lexington, KY 40506, for his gift of Gramoxone (PQ), and to Dr. Lisa Vaillancourt, Department of Plant Pathology, University of Kentucky, Lexington, Kentucky 40506, who provided us with a culture of Colletotrichum graminicola (CG). LITERATURE CITED Chen, Z., H. Silva, and D. Klessig. 1993. Active oxygen species in the induction of plant systemic acquired re- sistance by salicylic acid. Science 262:1883-1886. Choi, G. J., H. J. Lee, and K. Y. Cho. 1997. Involvement of catalase and superoxide dismutase in resistance of Botrytis cinerea to the dicarboximide fungicide vinclo- zolin. Pestic. Biochem. Phys. 59:1—10. Cruess, W. V., and J. H. Irish. 1931. Further observations on the relation of pH value to toxicity of preservatives to microorganisms. J. Bacteriol. 19:163—166. Durner, J., and D. F. Klessig. 1995. Inhibition of ascorbate peroxidase by salicylic acid and 2,6-dichloroisonicotinic acid, two inducers of plant defense responses. Proc. Nat. Acad. Sci. USA 92:11312-11316. Journal of the Kentucky Academy of Science 66(2) Gil-ad, N. L., N. Bar-Nun, T. Noy, and A. M. Mayer. 2000. Enzymes of Botrytis cinerea capable of breaking down hydrogen peroxide. Fed. Euro. Microbiol. Soc. Lett. 190:121—-126. Gil-ad, N. L., and A. M. Mayer. 1999. Evidence for rapid breakdown of hydrogen peroxide by Botrytis cinerea. Fed. Euro. Microbiol. Soc. Lett. 176:455—-461. Kosman, E., and Y. Cohen. 1996. Procedures for calcu- lating and differentiating synergism and antagonism in action of fungicide mixtures. Phytopathology 86:1263- ISAT Ees Norman, C., K. A. Howell, A. H. Millar, J. M. Whelan, and D. A. Day. 2004. Salicylic acid is an uncoupler and inhibitor of mitochondrial electron transport. Plant Phy- siol. 134:492-501. Ruffer, M., B. Steipe, and M. H. Zenk. 1995. Evidence against specific binding of salicylic acid to plant catalase. Fed. Euro. Biochem. Soc. Lett. 377:175-180. Shirasu, K., H. Nakajima, V. K. Rajasekhar, R. A. Dixon, and C. Lamb. 1997. Salicylic acid potentiates an ago- nist-dependent gain control that amplifies pathogen sig- nals in the activation of plant defense mechanisms. PI. Cell 9:261—270. Slaymaker, D. H., D. A. Navarre, D. Clark, O. del Pozo, G. B. Martin, and D. F. Klessig. 2002. The tobacco salicylic acid-binding protein 3 (SABP3) is the chloro- plast carbonic anhydrase, which exhibits antioxidant ac- tivity and plays a role in hypersensitive defense re- sponse. Proc. Nat. Acad. Sci. USA 99:11640-11645. Steel, C. C., and N. G. Nair. 1993. The physiological basis of resistance to the dicarboximide fungicide ipridione in Botrytis cinerea. Pest. Biochem. Phys. 47:60-68. Strobel, N. E., and L. A. Porter. 2005. Dose-dependent effects of salicylate on the plant-pathogenic fungus Bo- trytic cinerea. J. Kentucky Acad. Sci. 66:59. [Abstract] Xie, X., and Z. Chen. 1999. Salicylic acid induces rapid inhibition of mitochondrial electron transport and oxi- dative phosphorylation in tobacco cells. Pl. Phys. 120: 217225: Ye, X. S., N. E. Strobel, and J. A. Kuc. 1995. Induced systemic resistance (ISR): activation of natural defense mechanisms as part of integrated pest management (IPM). Pages 95-113 in R. Reuveni (ed). Novel ap- proaches to integrated pest management. Lewis Pub- lishers, Boca Raton. J. Ky. Acad. Sci. 66(2):129-136. 2005. Kentucky Baccalaureate Origins of Doctorate Recipients in the Biological Sciences, Chemistry, and Physics, 1978 through 2002 Fitzgerald B. Bramwell Department of Chemistry, University of Kentucky, Lexington, Kentucky 40506-0055 and Elinor L. Brown Department of Curriculum and Instruction, University of Kentucky, Lexington, Kentucky 40506-0017 ABSTRACT This paper presents the Kentucky baccalaureate origins of men, women, and under-represented group U.S. citizen doctoral recipients in the biological sciences, chemistry, and physics from 1978 through 2002. Coun- terintuitive results are reported for chemistry and physics. For these disciplines, institutions with an under- graduate focus and smaller science infrastructure have outperformed Kentucky’s research universities. Further, the results suggest that little or no attention has been paid in a systemic manner to any Kentucky under-represented minority group during this period. Conversely, there has been significant growth in the percentage and absolute numbers of U.S. women earning doctoral degrees in the biological sciences and in chemistry after receiving Kentucky baccalaureate degrees. INTRODUCTION Higher education graduate and undergrad- uate programs play a pivotal role in develop- ing the intellectual capital of the scientific and technical workforce. This paper presents the Kentucky baccalaureate origins of men, wom- en, and underrepresented group doctoral re- cipients in the biological sciences, chemistry, and physics from 1978 through 2002. It builds upon similar efforts in chemistry by Hall (1985) by using an expanded dataset that in- cludes underrepresented groups and the addi- tional disciplines of the biological sciences and physics. Results for the biological scienc- es, chemistry, and physics were targeted since these academic disciplines represent core competencies for any undergraduate program in the physical sciences. The importance of undergraduate training in the sciences in production of a technolog- ical infrastructure and as a prelude to graduate study is supported by all Carnegie research- extensive and research-intensive universities as well as by the worldwide scientific com- munity. For example, Dr. Frank Rhodes, for- mer president of Cornell University, stated, ‘while research and teaching both contribute to the strength and vitality of the U.S. research university, it is undergraduate teaching and learning, that is the central task. Undergrad- 1 uate education is fundamental to the existence of the university. . .’’ (Rhodes 1994). Kentucky’s transformation to a competitive, high-tech economy requires the development and enhancement of the scientific and tech- nological workforce. The critical nature of this effort is reflected in numerous state-supported and federally partnered initiatives such as the Experimental Program to Stimulate Competi- tive Research (NSF-EPSCoR Program 2005), Institutional Development Award (NIH-[DeA Program 2005), the Small Business Innova- tion Research/Small Business Technology Transfer (NSF-SBIR/STTR Program 2005). In recent years, comprehensive statewide efforts to improve higher education include signifi- cant budget reallocations through the Re- search Challenge Trust Fund initiative (STTR- EPSCoR Program 2005). The vast majority of these funds are directed toward improving the scientific and technological resources of Ken- tucky-based institutions of higher education. The basis for such efforts appears driven by research competitiveness within higher edu- cation (Teich 1996) and the need to update and enhance the Kentucky technical, scientif- ic, and educational infrastructure. The re- search or discovery undertakings at universi- ties and colleges can lead to patents and commercial development. Therefore, technol- e) 130 ogy transfer, intellectual property, and royalty income are among the important issues di- rectly linking the development of intellectual capital to the state’s economic growth and viability. METHODS The Kentucky baccalaureate origins of doc- torate recipients in the biological sciences, chemistry, and physics are derived from data provided by the Survey of Earned Doctorates (SED) and entered into WebCASPAR (Com- puter-Aided Science Policy Analysis and Re- search). The SED is conducted annually by the University of Chicago National Opinion Research Center for the Federal sponsors of the survey: the National Science Foundation (NSF), the National Institutes of Health (NIH), the U.S. Department of Education (USED), the National Endowment for the Hu- manities (NEH), the U.S. Department of Ag- riculture (USDA), and the National Aeronau- tic and Space Administration (NASA). The SED does not report professional degrees such as the D.D.S., D.V.M., J.D, M.D., and O.D. Information from this survey becomes part of the Doctorate Records File (DRF) and con- tains data on all earned doctorates granted by regionally accredited U.S. universities in all fields from 1920 to the present. WebCASPAR is a database system (NSF- WebCASPAR 2005) available on the World Wide Web and containing information about academic science and engineering resources. Information from academic surveys of the Di- vision of Science Resources Statistics (SRS) of the National Science Foundation and infor- mation from a variety of other sources, in- cluding but not limited to the SED, are in- cluded in the database. WebCASPAR currently can provide critical demographic data through access to most DRF doctoral rec- ords in the biological sciences, chemistry, and physics from 1966 through 2003. Data for the SED are collected directly from individual doctorate recipients. The questionnaire is distributed through the co- operation of graduate deans to those who are completing their doctorates. The data for a given year include all doctorates awarded in the 12-month period ending on 30 June. Over the past decade, the self-report response rates to the SED questionnaire varied from a high Journal of the Kentucky Academy of Science 66(2) of 94.7% in 1993 to a low of 91.2% in 2002. The response rate for a specific year may vary slightly over time as late responses are added. To minimize the effect of late responses, we used 2002 as the last year for our data survey. Since, ‘“‘differences in survey and item re- sponse rates from year to year can produce numerical fluctuations that are unrelated to true trends’’ (NSF Science Resources Statis- tics 2005), we used a 25-year time line (1978-— 2002) to minimize the effect of such fluctua- tions. Through this paradigm we addressed the stated concern that *‘increasing or decreasing numbers in a citizenship or racial/ethnic group . reflect to some degree any upward or downward change in both overall survey re- sponse and item response’’ (NSF Science Re- sources Statistics 2005). Until 1990, SED records with missing gen- der were imputed using the first name of the recipient only if the first name could be pos- itively identified as male or female. Records that could not be identified by first name were set to male. In 1990, follow-up procedures were implemented for cases still missing after imputation. Records for which gender still could not be identified in the follow-up were set to male. In 1997, it was decided to dis- continue this practice. Therefore, data with unknown gender occur in 1988 and in 1990 and later years. For each year beginning in 1990, however, the response rate is greater than 99.2% (Hill, S. T. 2005. Director, Doc- torate Data Project, NSF Division of Science Resources Statistics, pers. comm., |1 Feb.). As a result of new follow-up procedures implemented during the 1990 survey cycle, there was greater adjustment to the numbers for race/ethnicity and citizenship than in ear- lier years. Although these new procedures have improved the completeness of the data, they also created a break in trend data that should be taken into consideration when an- alyzing sensitive data such as race/ethnicity after 1989. The effect on the trend line for citizenship is extremely small, almost negli- gible. The effect on the trend line for race is minimal. To the extent that there has been an effect, blacks tend to be slightly overrepre- sented in institution-completed responses, so blacks would trend toward a larger proportion after the new procedures were implemented (Hill, S.T. 2005. Director, Doctorate Data Proj- Kentucky Baccalaureate Origins—Bramwell and Brown Table 1. KY Baccalaureate Institution for National Biology Doctorates Totals KY Rank — Rank/1434 University of Kentucky 161 i 120 University of Louisville 84 2 223 Centre College 54 3 324 Western Kentucky University 52 4 333 Murray State University 48 5 351 Thomas More College 4] 6 385 Eastern Kentucky University 36 7 42] Bellarmine College 28 8 488 Northern Kentucky University 25 fe) 528 Berea College 20 10 588 Georgetown College 20 10 588 Transylvania University 18 12 628 Morehead State University 17 13 649 Asbury College 14 14 716 Kentucky Wesleyan College 6 15 953 Kentucky State University 4 16 1043 Union College 3 17 1101 Brescia College 2 18 1161 Cumberland College 2 18 1161 Campbellsville University D 18 1161 Spalding University 1 21 1230 Total Academic Instiution 638 ect, NSF Division of Science Resources Sta- tistics, pers. comm. 11 Feb.). RESULTS Table 1 presents the Kentucky baccalaure- ate origins of doctoral recipients in the bio- logical sciences for men and women from 1978 through 2002. A total of 638 baccalau- reate degrees were awarded to individuals who received a doctorate in the biological sci- ences with 399 (63%) going to men and 239 (37%) going to women. Columns two, three, and four, respectively, present the total num- ber of doctorates awarded, the rank among Kentucky institutions of higher education, and the national rank out of 1434 institutions in the SED database. Columns five, six, and sev- en, respectively, present the number of doc- torates awarded to men, the rank among Ken- tucky institutions of higher education, and the national rank out of 1284 institutions in the SED database. Columns eight, nine, and ten, respectively, present the number of doctorates awarded to women, the rank among Kentucky institutions of higher education, and the na- tional rank out of 1288 institutions in the SED database. Data entries of ‘‘na’’ indicate infor- mation not available or not entered in the SED database. Table 2 presents the Kentucky baccalaure- ate origins of doctoral recipients in chemistry 131 Kentucky baccalaureate origins of U.S. citizen doctorate recipients in the biological sciences (1978-2002). Totals National Totals National Men KY Rank = Rank/1284 | Women KY Rank ~~ Rank/1288 56 i 132 34 2 214 29 3 243 23 4 301 13 5 428 8 11 596 2, 6 455 11 7 483 9 8 508 9 8 508 9 § 508 7 12 634 5 14 735 7 12 634 4 2 15 951 3 1 17 1064 3 na na na 2 0 20 1246 0 eo, 15 951 1 1 17 1064 0 1 17 1064 239 for men and women from 1978 through 2002. A total of 325 baccalaureate degrees were awarded to individuals who earned a doctorate in chemistry with 243 (75%) going to men and 82 (25%) going to women. Columns two, three, and four, respectively, present the total number of doctorates awarded, the rank among Kentucky institutions of higher edu- cation, and the national rank out of 1215 in- stitutions in the SED database. Columns five, six, and seven, respectively, present the num- ber of doctorates awarded to men, the rank among Kentucky institutions of higher edu- cation, and the national rank out of 1215 in- stitutions in the SED database. Columns eight, nine, and ten, respectively, present the number of doctorates awarded to women, the rank among Kentucky institutions of higher edu- cation, and the national rank out of 967 insti- tutions in the SED database. Data entries of ‘‘na’’ indicate information not available or not entered in the SED database. Table 3 presents the Kentucky baccalaure- ate origins of doctoral recipients in physics for men and women from 1978 through 2002. A total of 107 baccalaureate degrees were awarded to individuals who received a doc- torate in physics, with 98 (92%) going to men and 9 (8%) going to women. Columns two, three, and four, respectively, present the total number of doctorates awarded, the rank 132 Table 2. KY Baccalaureate Institution for National Chemistry Doctorates Totals KY Rank ~— Rank/1215 Western Kentucky University 48 i 185 University of Kentucky 46 2 191 University of Louisville 35 3 265 Murray State University 26 4 345 Centre College 25 5 356 Transylvania University 22 6 394 Berea College 18 7 449 Northern Kentucky University i 8 466 Bellarmine College 13 fe) 548 Eastern Kentucky University 13 9 548 Kentucky Wesleyan College 12 1] 567 Thomas More College 9 12 647 Morehead State University fe) 12 647 Cumberland College 9 12 647 Georgetown College 8 15 677 Asbury College 3 16 887 Union College 3 17 887 Campbellsville University 2 18 960 Brescia College 2 18 960 Kentucky State University 2 18 960 Pikeville College 2 18 960 Spalding University 1 22 1054 Total Academic Institution 825 among Kentucky institutions of higher edu- cation, and the national rank out of 955 insti- tutions in the SED database. Columns five, six, and seven, respectively, present the num- ber of doctorates awarded to men, the rank among Kentucky institutions of higher edu- cation, and the national rank out of 823 insti- tutions in the SED database. Columns eight, nine, and ten, respectively, present the number of doctorates awarded to women, the rank among Kentucky institutions of higher edu- Journal of the Kentucky Academy of Science 66(2) Kentucky baccalaureate origins of U.S. citizen doctorate recipients in chemistry (1978-2002). Totals National Totals National Men KY Rank Rank/1215 Women KY Rank Rank/967 37 1 185 11 2, 178 31 2 231 15 1 120 25 3 275 10 3 212 20 4 343 6 5 332 19 5 357 6 i) 332 15 6 419 ¢ 4 301 14 fi 446 4 8 442 13 8 472 4 8 442 11 10 515 2 11 627 12 9 49] 1 14 746 8 11 587 4 8 449. 4 15 741 5 ri 376 af 2 615 2 11 627 uf 12 615 Dw) ll 627 7 12 615 1 14 746 3 16 789 na na na 2 17 855 1 14 746 2 17 855 na na na 2, 17 855 na na na 2 17 855 na na na 2 17 855 na na na na na na 1 14 746 243 82 cation, and the national rank out of 474 insti- tutions in the SED database. Data entries of ‘*na’’ indicate information not available or not entered in the SED database. Table 4 presents the Kentucky baccalaure- ate origins of doctoral recipients in the bio- logical sciences from 1978 through 2002 for black men and black women. During that pe- riod, U.S. citizens in the biological sciences earned 81,951 doctorate degrees, of which 1538 (1.9%) went to black U.S. citizens. Ken- Table 3. Kentucky baccalaureate origins of U.S. citizen doctorate recipients in physics (1978-2002). KY Baccalaureate Institution for National Physics Doctorates Totals KY Rank ~— Rank/955 Thomas More College 20 1 176 Western Kentucky University ily 2 200 University of Kentucky 14 3 2.34 Eastern Kentucky University 13 4 252 Murray State University 11 5 285 University of Louisville fe) 6 324 Berea College 7 7 374 Centre College 6 8 399 Bellarmine College 3 fs) 542 Northern Kentucky University 2 10 590 Transylvania University 2 10 590 Cumberland College ih 12 672 Kentucky Wesleyan College 1 12 672 Union College 1 12 672 Georgetown College na 15 na Morehead State University na 15 na Asbury College na 15 na Total Academic Institution 107 Totals National Totals National Men KY Rank ~~ Rank/823 Women KY Rank Rank/474 1 1 3 227 yy) na na na 3 3 1 104 4 3 1 104 4 1 3 NT 6 1 3 227 i na na na 8 0 6 418 9 na na na na na na na na na na na na na na na na na na na na na na na na na na na 9 Kentucky Baccalaureate Origins—Bramwell and Brown Table 4, Kentucky baccalaureate origins of black U.S. citizen doctorate recipients in the biological sciences (1978-2002). KY Baccalaureate Institution for Biology Doctorates Awarded to Black Students Totals KY Rank University of Kentucky Berea College Kentucky State University Murray State University University of Louisville Total Academic Institution coe Ol cell ell cell ee ©) NWwWNNWwWNNW Fe tucky produced seven black baccalaureate re- cipients of doctoral degrees in the biological sciences. Table 5 presents the Kentucky baccalaure- ate origins of doctoral recipients in the bio- logical sciences from 1978 through 2002 for Hispanic men and Hispanic women. During that period U.S. citizens in the biological sci- ences earned 81,951 doctorate degrees, of which 1930 (2.4%) went to Hispanic U.S. cit- izens. Kentucky produced seven Hispanic baccalaureate recipients of doctoral degrees in the biological sciences. Table 6 presents the Kentucky baccalaure- ate origins of doctoral recipients in chemistry from 1978 through 2002 for black men and black women. During that period, U. S. citi- zens earned 30,864 doctorate degrees in chemistry, with 553 (1.8%) of that number go- ing to U.S. blacks. Kentucky produced four black baccalaureate recipients of doctoral de- grees in chemistry. Table 7 presents the Kentucky baccalaure- ate origins of doctoral recipients in chemistry from 1978 through 2002 for Hispanic men and Hispanic women. During that period U. S. cit- izens earned 30,864 doctorate degrees in chemistry, with 758 (2.5%) of that number go- Table 5. Kentucky baccalaureate origins of Hispanic U.S. citizen doctorate recipients in the biological sciences (1978-2002). KY Baccalaureate Institution for Biology Doctorates Awarded to Hispanic Students Totals KY Rank University of Kentucky 3 1 Centre College yy Murray State University 1 2, Thomas More College 1 ) University of Louisville 1 2 Total Academic Institution 7 133 Table 6. Kentucky baccalaureate origins of black U.S. citizen doctorate recipients in chemistry (1978-2002). KY Baccalaureate Institution for Chemistry Doctorates Awarded to Black Students Totals KY Rank Berea College 1 1 Centre College 1 1 University of Kentucky 1 1 Western Kentucky University 1 1 Total Academic Institution 4 ing to U.S. Hispanics. Kentucky produced three Hispanic baccalaureate recipients of doctoral degrees in chemistry. Within Kentucky, institutions of higher ed- ucation produced one black (from Centre Col- lege, 1979) and no Hispanic baccalaureate re- cipients who earned a doctorate in physics from 1978 through 2002. During that period U.S. citizens earned 15,938 doctorate degrees in physics. Of that number, black U. S. citi- zens earned 196 (1.2%) and Hispanic U.S. cit- izens earned 296 (1.9%). Table 8 presents the average number of doctorates earned annually by U. S. citizens matriculating from baccalaureate institutions over two time periods: 1978 through 1994 (17 years) and 1995 through 2002 (8 years). Col- umn one presents the discipline cohorts. Col- umn 2 presents the average number of doc- torates earned annually by individuals matriculating from baccalaureate institutions nationwide from 1978 through 1994 and from 1995 through 2002, as well as the percentage change between these averages. Column 3 presents similar data for individuals matricu- lating from Kentucky baccalaureate institutions. DISCUSSION Table 1 reports results for the production of doctorates whose baccalaureates are in the bi- ological sciences. The University of Kentucky Table 7. Kentucky baccalaureate origins of Hispanic U.S. citizen doctorate recipients in chemistry (1978— 2002). KY Baccalaureate Institution for Chemistry Doctorates Awarded to Hispanic Students’ Totals KY Rank Centre College 1 i Eastern Kentucky University il 1 Western Kentucky University 1 1 Total Academic Institution 8) 134 Journal of the Kentucky Academy of Science 66(2) Table 8. Comparison of national average annual doctorates annual doctorates (1978-1994) and (1995-2002). National Discipline (1978-1994) (1995-2002) % Change Biological Sciences Overall 3148 3555 12.93% Women 1109 1670 50.59% Black 43.6 99.5 128.21% Hispanic 54.2 126 132.47% Chemistry Overall 1276 1148 —10.03% Women 263 355 34.98% Black 17 34 100.00% Hispanic 28 35 25.00% Physics Overall 636 641 0.79% Women 48.4 80.1 65.50% Black 6.35 11 73.23% Hispanic 10.3 15.1 46.60% and the University of Louisville, as might be expected due to their size and available re- sources, produce the greatest number of bac- calaureates who earn doctorates. Kentucky in- stitutions appear to lag behind national benchmarks with the highest ranking (Univer- sity of Kentucky) in the biological sciences being 120 out of 1434 schools in the SED database. Although this ranking is above many institutions with far fewer resources and significantly smaller student populations, it ranks significantly below benchmark institu- tions of comparable mission and student pop- ulation. For example, it ranks significantly be- low the University of Georgia ranked at 58, the lowest of 20 benchmarks (UK Bench- marks 2005). Achieving parity with leading benchmark institutions appears to require implementation of comprehensive institutional approaches to strengthen STEM teaching and learning. At a minimum, such institutional approaches must necessarily focus on support for faculty, ap- propriate instrumentation, and improvement of access, retention, and graduation rates for all students. Further analysis of benchmark programs relative to those in Kentucky should yield greater insight into the substantial ob- served differences. Tables 2 and 3 report results that appear counterintuitive for the production of bacca- laureates in chemistry and physics. Western Kentucky University and Thomas More Col- lege, institutions primarily devoted to under- with Kentucky baccalaureate-granting institutions average Kentucky (1978-1994) (1995-2002) %Change 23.3 30.3 30.04% 7.88 13.1 66.24% 0.24 0.38 58.33% 0.18 0.5 177.78% 12.5 14.125 13.00% 2.88 4.13 43.40% 0.12 0.25 108.33% 0.18 0 —100.00% 4.47 3.88 —13.20% 0.29 0.50 72.41% 0 0.125 — 0 0 0% graduate education with relatively limited re- sources and relatively smaller enrollments, have assumed leadership in the production of intellectual capital, respectively, in chemistry and physics. In physics, Thomas More Col- lege and Western Kentucky University togeth- er produced 60% more baccalaureates that earned doctorates than the University of Ken- tucky and the University of Louisville combined. Given the robust nature of the science in- frastructure at the University of Kentucky and at the University of Louisville, (e.g., seminars, information technology resources, research opportunities for undergraduate students, and targeted NSF and NIH scientific infrastructure support), the relatively low numbers of bac- calaureates earning doctorates in chemistry and physics are surprising. Detailed analysis of undergraduate programs and institutional commitment offer promising areas for future study. Kentucky institutions appear to lag far behind national benchmarks, with the highest ranking in chemistry being 185 out of 1215 schools in the SED database. The highest ranking for a Kentucky research-extensive university is 191, significantly below the low- est of 20 benchmarks, the University of Iowa at 109. Kentucky institutions appear to lag far be- hind national benchmarks, with the highest ranking in physics being 176 out of 955 schools in the SED database. The highest ranking for a Kentucky research-extensive Kentucky Baccalaureate Origins—Bramwell and Brown university 1s 234, above the lowest ranked benchmark, Georgia at 305, but significantly below the next lowest benchmark, University of Iowa at 59. Tables 4 through 7 present results for the production of doctorates from underrepresent- ed minority group candidates whose baccalau- reates are in the biological sciences and chem- istry. No results were tabulated for physics since only one doctorate was earned over the 25 years of collected data. The results suggest that little or no attention was paid in a systemic manner to any Ken- tucky underrepresented minority group from 1978 through 2002. During that time, the per- centage and absolute numbers of U.S. under- represented minority group students receiving baccalaureate degrees and then earning doc- torates in the biological sciences, chemistry, and physics roughly doubled, but little change was reported for such individuals in Kentucky. Table 8 presents the annual averages of Kentucky baccalaureate origins of doctorate production from 1978 through 1994 (17 years) and from 1995 through 2002 (8 years). These time periods were chosen to reflect the impact of large-scale or systemic funding during the late 1980s and early 1990s by the NIH and the NSF. Of particular interest were the NSF- funded Statewide Systemic Initiative pro- grams, including the Rural Systemic Initiative Program (targeting pre-college students) and the Alliances for Minority Participation Pro- gram (targeting college undergraduates). These programs were introduced nationwide and targeted to address perceived deficiencies in the growth of the scientific and technolog- ical workforce. The effects of these programs should have been evident as early as 1995 on the production of doctorates in sciences (Hicks, A. J. 2004. Program Director, NSF- LS AMP Program, Division of Human Re- source Development, pers. comm., 11 Dec.). Thus, we chose to look at changes in doctorate degree production during these periods and compare them to national trends. Of the NSF Systemic programs that would directly affect doctorate production from baccalaureate insti- tutions, Kentucky universities participated only in the Rural Systemic Initiative, a pro- gram that focused on the needs of women and of the rural economically disadvantaged. 135 Our data with respect to women doctorates are consistent with those reported by Everett and DeLoach (1991) and more recently by Freeman et al.(2004). There has been signifi- cant growth in the percentage and absolute numbers of U.S. women receiving baccalau- reate degrees and then earning doctorates in the biological sciences and chemistry both na- tionwide and in Kentucky. The rate of growth, using results solely from 2002 and 1978, seems more rapid than Table 8 would suggest. If 2002 is used as a measure, then in the bi- ological sciences, 47.1% of U.S. doctorates (up from 25.1% in 1978) were awarded to women, while in Kentucky 51.4% (up from 20.8% in 1978) were awarded to women. In chemistry a similar trend increase was observed with 34.3% of U.S. doctorates (up from 12.6% in 1978) awarded to women, while in Kentucky 27.3% (up from 0% in 1978) were awarded to women. In physics, the overall number of doctorates awarded decreased nationwide, with women earning 4.8% of U.S. doctorates (down from 11.5% in 1978), while in Kentucky women earned 0% in 2002 and 0% in 1978. The results with respect to underrepresented minorities suggest that most numerical anal- yses would be questionable since so very few individuals who earned a baccalaureate in Kentucky subsequently earned a doctoral de- gree in the sciences. However, these results are consistent with (a) a highly successful NSF-sponsored and state-partnered Appalachian Rural Systemic Initiative program designed to increase the numbers of women and economically disad- vantaged Kentuckians entering the scientific and technological workforce; and (b) the lack of any systemic program (e.g., NSF—Louis Stokes Alliances for Minority Participation, NIH-Minority Access to Research Careers) designed to increase the numbers of underrep- resented minority Kentuckians entering the scientific and technological workforce. The baccalaureate origins of doctoral recip- ients in core competencies (the biological sci- ences, chemistry, and physics) in the physical sciences provide reasonable benchmarks to test the quality of science, technology, engi- neering, and mathematics (STEM) baccalau- reate programs. 136 SUMMARY These findings support the conclusion that the Kentucky higher education system failed to enhance the production of black and His- panic physical science majors who later earned doctorates. Further, these findings speak to a lost opportunity during the prom- ising equal opportunity era (ca. 1970 through 1995) to engage, more fully, underrepresented minority groups in the scientific and techno- logical workforce. Conversely, during this pe- riod significant progress was made to support and encourage female and economically dis- advantaged representation in the intellectual leadership of the physical sciences. These results suggest that constructive en- hancements to the Kentucky science and tech- nology policies should be made to address its deficiencies in order to realize the potential of the intellectual capital inherent in minority communities. The training and development of intellectual leadership in the physical sci- ences for women and the economically dis- advantaged has already yielded significant benefits. Similar efforts in the black and His- panic communities should help Kentucky re- alize the full extent of economic and quality of life advantages provided by an educated populace. ACKNOWLEDGMENTS Useful conversations with Dr. Luther Wil- liams, the William T. Kemper Director of Ed- ucation of the Missouri Botanical Garden, are acknowledged. Journal of the Kentucky Academy of Science 66(2) LITERATURE CITED Everett, K.G., and W.S. DeLoach. 1991. Chemistry doc- torates awarded to women in the United States. J. Chem. Education 68:545-—547. Freeman, R.B., E. Jin, and S. Chia-Yu. 2004. Where do new US-trained science-engineering PhDs come from? Page 23 in Working paper 10554, National Bureau of Economic Research, Cambridge, MA. Hall, A.E. 1985. Baccalaureate origins of doctorate recip- ients in chemistry: 1920-80. J. Chem. Education 62: 406-408. NIH-IDeA Program. 2005. Institutional development award program retrieved 9 May 2005 from http:// www.ncrr.nih.gov/resinfra/ri_idap.asp. NSF-EPSCoR Program. 2005. Experimental program To stimulate competitive research (EPSCoR) retrieved 9 May 2005 from __http://www.nsf.gov/ehr/epscor/ about.jsp. NSF-SBIR/STTR Program. 2005. Program description re- trieved 9 May 2005 from http://www.nsf.gov/eng/sbir/ program.jsp Summary. NSF Science Resources Statistics. 2005. Survey of doc- torate recipients retrieved 9 May 2005 from http:// www.nsf.gov/statistics/showsrvy.cfm?srvy-CatID = 2&srvy-Seri = 1. NSF-WebCASPAR. 2005. WebCASPAR database re- trieved 9 May 2005 from http://caspar.nsf.gov/. STTR-EPSCoR Program. 2005. Kentucky’s system of post-secondary education retrieved 9 May 2005 from http://kctcs.edu/legislative/legislativebooklet.pdf. Rhodes, F.H.T. 1994. Pages 180-181 in J.R. Barber, E.G. Cole, and S.R. Graubard (eds). The research university in time of discontent. Johns Hopkins Press, Baltimore, MD. Teich, A.H. (ed). 1996. Competitiveness in academic re- search. American Association for the Advancement of Science, Washington, DC. UK Benchmarks. 2005. Institutional research, planning and effectiveness retrieved 9 May 2005 from http:// www.uky.edu/OPIE/benchlist.shtml. J. Ky. Acad. Sci. 66(2):137-138. 2005. NOTES Host Occurrence of Eastern Mistletoe (Phoraden- dron leucarpum, Viscaceae) in Robertson County, Kentucky—A survey of eastern mistletoe (Phoradendron leucarpum)-infested host trees in Robertson County, Ken- tucky, was conducted from 27 Dec 2002 to 6 Jan 2003. All paved roads and passable gravel roads in the county were traveled by vehicle. Host trees were identified and tallied by species, and the number of clumps of mistletoe were recorded. Voucher specimens were obtained for each representative host tree species by a extendable 12 m fiberglass linesman pole and then mounted, labeled, and deposited in the Berea College Herbarium (BEREA). Four mistletoe studies have been published in Ken- tucky (1, 2, 3, 4). The only published report for mistletoe in Robertson County had been a sight record on Juglans nigra by Reed and Reed (1). Robertson County is located in northeastern Kentucky (Figure 1). With 259 km? of land area, it is the smallest Kentucky county. Mount Olivet, the county seat, is cen- trally located at latitude 38°31'55" N and_ longitude 84°02'14” W. Robertson County is the state’s least popu- lous county, having 2266 people; 287 people reside in Mount Olivet according to the 2000 census (5). The county is located entirely in the Hills of the Blue- grass ecoregion of the Interior Plateau Region of Ken- tucky (6). The hills consist of upland rolling topography with moderately steep slopes and broad ridges. Small ra- vines form seasonally intermittent streams leading to larg- er streams, e.g., North Fork Licking River and Licking River. Elevations in the county range from 170 m to 308 m. In Robertson County, Upper Ordovician limestone and shale bedrock belong to the Clays Ferry Formation, Lex- ington Limestone, Kope Formation, and Fairview For- mation (7). The Eden-Nicholson-Lowell soil association is found throughout the Robertson County upland terrain (8). Braun (9) classified forest vegetation as Western Me- sophytic Forest in this region of the eastern deciduous forest; Kiichler (10) placed northeastern Kentucky in the Quercus-Carya forest. I observed forest vegetation on up- land limestone and shale hills as Quercus alba-Carya spp. forest with scattered large stands of Juniperus virginiana and a riparian forest of Acer negundo-Platanus occiden- talis-Acer saccharinum in alluvial areas. A majority of the county is composed of pasturelands and agricultural crop- lands. The continental climate in northeastern Kentucky is characterized by cool to cold winters and warm, humid summers with precipitation well distributed throughout the year (11). Climate data, 1977-2000, are from the Fal- mouth weather station in Pendleton County, 32 km west of Robertson County (12). Mean annual precipitation is 111 cm with the lowest precipitation, 7.2 em, in October and the highest, 11.8 cm, in May and July. Mean annual temperature is 11.7°C with the mean lowest temperature, —1.6°C, in January and the highest temperature, 23.8°C, in July. The mean growing season is 175 days with a range of 132 days to 211 days. The median first fall frost is 15 October and the last spring frost is 24 April (12). Phoradendron leucarpum was found only on 45 trees from 7 host tree species in 6 families. Juglans nigra ac- counted for 23 host trees (51.1%). Other host trees were 10 Ulmus americana (22.2%), 6 Robinia pseudoacacia (13.4%), 3 Acer saccharinum (6.7%), and 1 each of Gle- ditsia triacanthos, Fraxinus americana, and Prunus sero- tina (6.6%). All host trees were situated east of Mount Olivet in the eastern half of the county (Figure 1). The sparse number of host trees in Robertson County was very significant when compared to other recent mistletoe stud- ies in central and south central Kentucky, i.e., Lexington- Blue Grass Army Depot in Madison County with 1837 host trees (2), Rockcastle County with 3502 trees (3), and Garrard County with 1740 (4). Low winter temperatures are the principal factor re- sponsible for the sparse number of mistletoe-infested trees in Robertson County, e.g., in January 1994 the low- est temperature was —34.4°C and in February 1996 it was —22.1°C. My observations of host trees in mistletoe sur- veys showed extensive mistletoe die-back to the endo- phytic root system when extremely low temperatures have occurred. Spooner (13) found that eastern mistletoe reaches its northernmost distribution range in southern Ohio. He produced evidence that the main reason for the northern limits of eastern mistletoe corresponded to the mean minimum January temperature of —4.5°C. My ex- tensive reconnaissance has shown that the incidence of mistletoe is sparse not only in Robertson County, but in all of the northern Kentucky counties within the Hills of the Bluegrass and Outer Bluegrass ecoregions. In several of these northern counties, I have observed eastern mis- tletoe only once or twice from one to three host tree spe- cies for the entire county, e.g., Bracken, Campbell, Lewis, Gallatin, Kenton, Owen, and Pendleton counties. Eastern mistletoe has not been observed in Grant County despite repeated searches. Other mistletoe studies have documented the effects of low temperatures on spread and mortality of mistletoes. Garman (14) reported that severe winters destroyed much eastern mistletoe in Kentucky, but mistletoe gradually re- appeared under more favorable temperatures in later years. Deam (15) observed the detrimental effects of low temperatures on eastern mistletoe and the restriction of the plant’s Indiana range to the southern part of the state. Lightle (16) documented that low temperatures were fac- tors in injury and mortality to three Phoradendron spp. in southern Arizona and New Mexico. Wagener (17) con- cluded that very low temperatures restricted two Phora- dendron spp. to their present distribution limits in Cali- fornia and explained their absence in other parts of the hosts’ range. The dioecious characteristic of mistletoe may be anoth- er important factor in the overall sparse numbers of mis- 137 138 Figure 1. Robertson County, in northeastern Kentucky. The forty-five mistletoe-infested tree locations are indi- cated with solid circles. tletoe-infested trees. The chance of mature fruits from a pistillate plant becoming dispersed by birds and estab- lished in new host trees with both sexes present in the near vicinity may be rare especially when there are so few mistletoe-infested trees in the county. Most the 45 host trees tended to be lightly infested; 19 trees had only a single clump, 7 trees had two clumps, and the remaining 19 trees had from 3 to 15 clumps. Thus, few trees had both staminate or pistillate plants in the same tree or near- by infested trees. Several host trees were isolated in up- land pasturelands or around residences. Other mistletoe- infested trees of the same species along upland roads or fencerows tended to show an aggregated or clustered pat- tern (Figure 1). LITERATURE CITED. (1) Reed, C.F., and P.G. Reed. 1951. Host distribution of mistletoe in Kentucky. Castanea Journal of the Kentucky Academy of Science 66(2) 16:7-15. (2) Thompson, R.L. 1992. Host occurrence of Phoradendron leucarpum in the Lexington-Blue Grass Army Depot, Blue Grass Facility, Madison County, Ken- tucky. Trans. Kentucky Acad. Sci. 53:170-171. (3) Thomp- son, R.L., and F.D. Noe Jr. 2003. American mistletoe (Phoradendron leucarpum, Viscaceae) in Rockcastle County, Kentucky. J. Kentucky Acad. Sci. 64:29-35. (4) Thompson, R.L., and D.B. Poindexter. 2005. Host speci- ficity of American mistletoe (Phoradendron leucarpum) in Garrard County, Kentucky. J. Kentucky Acad. Sci. 66:40- 43. (5) Wikipedia, the free encyclopedia. 2005. Robertson County, Kentucky. http:// en.wikipedia.org/wiki/Robert- son. County%2 C_Kentucky. Accessed 23 Mar 2005. (6) Woods, A.J., J.M. Omernik, W.H. Martin, G.J. Pond, W.M. Andrews, S.M. Call, J.A. Comstock, and D.D. Tay- lor. 2002. Ecoregions of Kentucky (color poster with map, descriptive text, summary tables, and photographs). U.S. Geological Survey, Reston, VA. (7) McDowell, R.C., G_]. Grabowski Jr., and S.L. Moore. Geologic map of Ken- tucky. U.S. Geological Survey, Washington, D.C. (8) Bai- ley, H.H., and J.H. Winsor. 1964. Kentucky soils. Univ. Kentucky Agric. Exper. Sta. Misc. 308. (9) Braun, E.L. 1950. Deciduous forests of eastern North America. Haf- ner Press, New York, NY. (10) Kiichler, A.W. 1964. Man- ual to accompany the map of potential natural vegetation of the conterminous United States. Am. Geogr. Soc. Spec. Bull. 36. (11) Trewartha, G.T., and L.H. Horn. 1980. An introduction to climate. 5th ed. McGraw-Hill Book Co., New York, NY. (12) Kentucky Climate Center. 2001. The Kentucky Climate Center at Western Kentucky University Station Climate Summaries—Falmouth, Pendleton Coun- ty station. http:// kyclim.wku.edu/cgi-bin/stations/152775. Accessed 29 Mar 2005. (13) Spooner, D.M. 1983. The northern range of eastern mistletoe, Phoradendron sero- tinum (Viscaceae), and its status in Ohio. Bull. Torrey Bot. Club 110:489-493. (14) Garman, H. 1913. Woody plants of Kentucky. Univ. Kentucky Agric. Sta. Bull. 169:3-62. (15) Deam, C.C. 1924. Shrubs of Indiana. Indiana De- partment of Conservation, Bloomington, IN. (16) Lightle, P.C., D. Wiens, and F.G. Hawksworth. 1964. Low-tem- perature injury to Phoradendron in Arizona and New Mexico. Southwest. Naturalist 8:204—209. (17) Wagener, W.W. 1957. The limitation of two leafy mistletoes of the genus Phoradendron by low temperatures. Ecology 38: 142-145.—Ralph L. Thompson, Herbarium, Depart- ment of Biology, Berea College, Berea, KY 40404. List of Recent Reviewers We gratefully acknowledge the contribution of time and expertise provided by the following individuals in reviewing manuscripts submitted for consideration by the Journal of the Kentucky Academy of Science. David M. Brandenburg James O. Luken Guenther Schuster Jerry H. Carpenter Landon McKinney Thomas Sproat Nigel G.F. Cooper Joseph I. Orban Ralph L. Thompson Merritt Gilliland III Stefan Paula Jerry W. Warner Susan K. Gregurick James S. Pringle Mary Kathryn Whitson Ronald L. Jones James O. Ryan Fred L. Wyatt Bobby Lee 139 J. Ky. Acad. Sci. 66(2):140-142. 2005. Press Release: Kentucky Academy of Sciences (KAS) Resolution In Support of Evolution At the annual business meeting of the KAS, on November 12, 2005, the KAS re- viewed and reaffirmed past resolutions in support of the teaching of Evolution and unanimously endorsed the American As- sociation for the Advancement of Scienc- es’s “Resolution on Intelligent Design Theory” The following resolution, already adopted by the Kentucky Academy of Science at the annual business meeting on November 14, 1981, was unanimously approved again at its annual business meeting on November 12, 2005: The Kentucky Academy of Science is op- posed to any attempt by legislative bodies to mandate specific content of science courses. The content of science courses should be determined by the standards of the scientific community. Science involves a continuing systematic inquiry into the man- ifold aspects of the biological and material world. It is based upon testable theories which may change with new data; it cannot include interpretations based on faith or re- ligious dogma. As scientists, we object to at- tempts to equate “scientific creationism” or “intelligent design” with evolution as sci- entific explanations of events. Teaching faith-based models implies that these views are equivalent alternatives among scientists; doing so would be misleading to students. “Scientific creationism” and “intelligent de- sign” are not equivalent to evolution. There is overwhelming acceptance by scientists of all disciplines that evolution (the descent of modern specifies of animals and plants from different ancestors that lived millions of years ago) is consistent with the weight of a vast amount of evidence. The understand- ing of the processes underlying evolution has provided the foundation upon which many of the tremendous advances in agri- culture and medicine and theoretical biol- ogy have been built. Differences among sci- entists over questions of how evolution was accomplished do not obscure the _ basic agreement that evolution has occurred. Most people who subscribe to religious views have developed belief systems that are compatible with evolution. There is a widespread consensus among theologians that biblical accounts of creation are mis- understood if they are treated as literal sci- entific explanations. We fully respect the re- ligious views of all persons but we object to attempts to require any religious teachings as science. We join the National Academy of Scienc- es, the American Association for the Ad- vancement of Science, and the academies of science in many other states in calling for the rejection of attempts to require the teaching of “scientific creationism” and “in- telligent design” as a scientific theory. It is further recommended that the Ken- tucky Academy of Science encourages its members and other professional scientific groups to give support and aid to those classroom teachers who present the subject matter of evolution fairly and encounter community objection. We also encourage administrators and individual teachers to oppose the inclusion of nonscientific con- cepts in the science classroom. Passed KAS Annual Business Meeting, No- vember 12, 1983. And, “A Resolution of the Kentucky Academy of Science In Regard to Omitting Evolution Terminology and Teaching in the Public Schools” (1999): Whereas the Kentucky Academy of Sci- ence, founded in 1914, is an organization that encompasses all of the accepted sci- entific fields, and Whereas the Scientific Method exempli- fies that search for Scientific Understand- ing, and 140 Press Release Whereas this methodology has consis- tently provided the means of questioning dogma, authoritarianism, and deliberate de- ception, by championing the spirit of inqui- ry based on testing, analysis, honest review, criticism and counter criticism and designs for further testing, and, Whereas the advancements of our under- standing of the interconnection of the phys- ical properties of our universe coupled with the life forms which together compose our biosphere clearly support that the evolu- tionary process has functioned and does function in the development, control, and survival of the earth’s living beings, and Whereas to deny the concepts of the known theoretical basis of the evolutionary process to the education arena of our public schools by avoiding or eliminating from the science curriculum any mention of the term evolution and evolutionary concepts would be an affront to an objective inquiry and the understanding of science, Thereby be it resolved that the Kentucky Academy of Science, in the strongest and most determined ways possible, deplores the decision to substitute “change over time” for “evolution” in the state teaching standards, urges that the original wording be reinstated, and decries any attempt to remove the teaching of basic evolutionary theory or any scientific concept that may be tested and examined in concert with the ba- sic scientific laws and principles that com- prise the Scientific Method, and further- more be it resolved that the public sup- ported education systems of the Common- wealth be enhanced with complete support of seeking knowledge by every means pos- sible commensurate with known principles of scientific theory, fact, and understanding. Adopted by the KAS Governing Board No- vember 6, 1999. Passed unanimously by KAS membership No- vember 6, 1999. Both resolutions reviewed and reaf- firmed by the KAS membership at the an- nual KAS business meeting on November 12, 2005. The KAS also voted to endorse the following AAAS Board Resolution: 141 AAAS Board Resolution on Intelligent Design Theory The contemporary theory of biological evo- lution is one of the most robust products of scientific inquiry. It is the foundation for re- search in many areas of biology as well as an essential element of science education. To be- come informed and responsible citizens in our contemporary technological world, students need to study the theories and empirical evi- dence central to current scientific understand- ing. Over the past several years proponents of so-called “intelligent design theory,” also known as ID, have challenged the accepted scientific theory of biological evolution. As part of this effort they have sought to intro- duce the teaching of “intelligent design theo- ry” into the science curricula of the public schools. The movement presents “intelligent design theory” to the public as a theoretical innovation, supported by scientific evidence, that offers a more adequate explanation for the origin of the diversity of living organisms than the current scientifically accepted theory of evolution. In response to this effort, indi- vidual scientists and philosophers of science have provided substantive critiques of “intel- ligent design,” demonstrating significant con- ceptual flaws in its formulation, a lack of cred- ible scientific evidence, and misrepresenta- tions of scientific facts. Recognizing that the “intelligent design the- ory” represents a challenge to the quality of science education, the Board of Directors of the AAAS unanimously adopts the following resolution: Whereas, ID proponents claim that con- temporary evolutionary theory is incapable of explaining the origin of the diversity of living organisms; Whereas, to date, the ID movement has failed to offer credible scientific evidence to support their claim that ID undermines the current scientifically accepted theory of evo- lution; Whereas, the ID movement has not pro- posed a scientific means of testing its claims; Therefore Be It Resolved, that the lack of scientific warrant for so-called “intelligent 142 design theory” makes it improper to include as a part of science education; Therefore Be It Further Resolved, that AAAS urges citizens across the nation to op- pose the establishment of policies that would permit the teaching of “intelligent design theory” as a part of the science cur- ricula of the public schools; Therefore Be It Further Resolved, that AAAS calls upon its members to assist those Journal of the Kentucky Academy of Science 66(2) the content of contemporary evolutionary theory and the inappropriateness of “intel- ligent design theory” as subject matter for science education; Therefore Be It Further Resolved, that AAAS encourages its affiliated societies to endorse this resolution and to communicate their support to appropriate parties at the federal, state and local levels of the govern- ment. engaged in overseeing science education policy to understand the nature of science, Approved by the AAAS Board of Directors on November 18, 2002. J. Ky. Acad. Sci. 66(2):143-145. 2005. INDEX TO VOLUME 66 Compiled by Ralph L. Thompson Abandoned limestone quarry, 24—34 ABBOTT, J. RICHARD, 24 ABERT, JAMES W.,, 3-16 Abstracts, 2004 KAS Annual Meet- ing, 57-65 Acer negundo, 26, 137 Acer saccharinum, 26, 42, 137 Acer saccharum, 26, 42 ACOSTA, CHARLES A., 65 Actinonaias pectorosa, 60 Adult Self-Report Scale (ASRS), 63 Agricultural Sciences abstracts, 57— 59 Aix sponsa, 60 ALL, JOHN, 61 Allegheny woodrat, 69 American Association for the Ad- vancement of Science (AAAS), 130-132 American mistletoe, 40 AMMONS, ANDY, 89, 94 AMONGE, AUGUSTINE, 59, 60 ANDERSON-HOAGLAND, ELIZABETH, 101 Anolis sagrai, 65 Antifungal materials in vitro, 118 ANTONIOUS, GEORGE F., 57, 58 Anthropology and Sociology ab- stracts, 59 Anuran larvae (tadpoles), 65 Appalachina sayana, 68 Archaeology, Chinese mining sites, 59 Artist, naturalist, land developer, and topographical engineer, 3 ASHER, WESLEY, 107 Asimina triloba, 58 Assessing water quality of two creeks, 50 Astronomy and space science work- shops, 63 Attention Deficit/Hyperactivity Dis- order (AD/HD), 63 Baccalaureate origins of doctorate recipients, 129-136 BARBOUR, THOMAS T., 60 BARNEY, ROBERT J., 57 BARTON, CHRIS, 101 BAUTISTA, DEBRA L., 17, 107 BEBE, FRED, 62 Berea College Forest, 35 BERRY, BOBBIE, 101 Beta vulgaris, 58 Biodegradable Dissolved Organic Carbon (BCOD) analyses, 50 Bivalvia: Unionidae, 60 Bluegill, 102 BOATENG, DADDY N., 57 Book Review: Plant Life of Ken- tucky, 66 BOSWELL, AYISA, 94 Botany and Microbiology abstracts, 59 Botrytis cinerea, 59, 120 BRAMWELL, FITZGERALD, B., 129 BRANDENBURG, DAVID M., 66 Brassica olearacea, 57 BRIGGS, JAMES A., 61 Broccoli, 57, 58 BROWN, ELINOR L., 129 Brown ADD Scales, 63 BUTHELEZI, THANDI, 59, 60 Cabbage looper, 57 Capsicum annuum, 58 CARPENTER, LISA, 107 Carya ovata, 42 Celtis occidentalis, 27, 42 Centrarchid fishes, 101 Cepaea nemoralis, 82-88 Cercaria, 89, 94, Chemistry abstracts, 59-60 CHERMAK, KEVIN A., 70 Chinese mining sites, 59 Chorus frogs, 65 Citrullus lanatus, 58 Clean Water Act (CWA), 61 Coal surface-mined areas, 24 Colletotrichum graminicola, 119 Competition between mycorrhiza, 60 Competition between wood ducks, 60 Computational investigations of enolase, 17-23 CORGAN, JAMES X., 62 Cornus drummondii, 27 CRAWFORD, NICHOLAS C., 61 CRAWFORD, NICK, 61 CRAWFORD, TODD J., 70 Creek, Ledbetter, 50 Creek, Panther, 50 Cryprogenia stegaria, 60 Ctenosaura similus, 65 Curcumene, 57 Curcurbita maxima, 58 Curcurbita pepo, 58 Cyclophane/anthracene, 59, 60 Cyclophane/9-fluorenone, 59, 60 Delta model, 109 Dichloromethane, 59, 60 143 DIEHL, TIFFANY, 63 Digenea: Azygiidae, 94 DigiScope 300 in middle school sci- ence, 64 Dimethoate insecticide residues, 57 Diplocarpon rosae, 120 Disabled postsecondary biology stu- dents, 64 Diseases of Warren County, 61 Dissolved Organic Carbon (DOC), 40 Dissolved Organic Matter (DOM), 40 DNA damage evaluation, 62 Doctorate recipients in Biological sciences, chemistry, and phys- ics, 129 DURTSCHE, RICHARD D., 44, 65 Dye tracing of groundwater through Russel Cave, 61 Eastern mistletoe, 137 Ecology and Environmental Science abstracts, 60-61 Effect of light wavelength and os- molality on cercariae, 94 Elevated spoil pile, 28 Elimia semicarinata, 89 Enhancement of plant disease resis- tance (EPDR), 118 Eponym of generic name Lindera, 44 ERVIN, CHRISTOPHER A., 64 Erythrocytes of farm workers, 62 ESTES, TERRI A., 70 Evolution resolution, 140 FAIRCHILD, JENNIFER L., 64 Fanshell mussel, 60 Festuca arundinacea, 27 Fire impact on Kentucky small mammals, 67—70 FLEMING, JONATHAN, 89, 94 Flora Wiksbergensis, 44 FORESTER, JOSEPH T., 70 Floristic survey, in Madison County, 35-39 Fraxinus americana, 26, 42, 137 FRISBIE, MALCOLM P.,, 64 G Protein Coupled Receptors (GPCRs), 107 GAR, JOSEPH D., 50 Gastropoda: Helicidae, 82 Gastropoda: Pleuroceridae, 89 Gekkonid lizards in Belize, 65 Generic name Lindera, 44 Geography abstracts, 61 Ginger, 57 144 Journal of the Kentucky Academy of Science 66(2) Gleditsia triacanthos, 42, 137 Glutathione peroxidase (GPX), 62 Glycolysis, 17 GOETZ, SCOTT M., 65 GOMELSKY, BORIS, 58 GRAY, BLAKNEY, 62 Guidelines for contributors, 71—72 HACKNEY, KAREN, 63 HACKNEY, RICHARD, 63 HAM, BRIAN, 61 HARDY, JONATHAN, 101 HEAD, JUSTIN A., 70 Health Sciences abstracts, 61-63 Hills of the Bluegrass, 41, 137 Host/guest interactions of cycloclo- phane molecules, 60 Host histopathology, 101 Host occurrence of eastern mistle- toe, 137-138 Host specificity of American mistle- toe, 40—43 Human Mu, Kappa, and Delta Opi- oid Receptors, 107, Hylidae, 65 Impact of fire on small mammals, 67-69 In memoriam: John W. Thieret, 73 Indian Fort Amphitheater, 35 Infections by trematodes, 101 Inner Bluegrass, 25, 41 Inquiry-based instruction, 64 Inquiry-based Science and Math in Appalachian Middle Schools (ISMAM), 64 Intelligent Design Theory, AAAS Board Resolution, 141-142 Intraspecific brood parasitism, pre- dation, stress levels of wood ducks, 60 Invasive species, 27 JOHNSON, PIERCE, JR., 60 JONES, RONALD L., 66 JORDAN CLARENCE E., 57 JOVANOVIC, BOJANA, 89, 94 Juglans nigra, 33, 40, 137 Juniperus virginiana, 26, 137 JUSTICE, TIFFANY D., 70 Kappa model, 109 Karst geomorphic features, 61 Karst stormwater discharges, 61 Kentucky Academy of Science, 2004 Abstracts, 57-65 Kentucky Academy of Science, 2005 Meeting, 70 Kentucky Academy of Science (KAS), Resolution in support of Evolution, 140-142 Kentucky Baccalaureate origins of doctorate recipients in Biologi- cal sciences, chemistry, and physics, 129-136 Kentucky Exotic Pest Plant Council, 27 Kentucky Lake, 50 Knobs-Lower Scioto Dissected Pla- teau, 36 Knobs-Norman Uplands, 41 KOCHHAR, TEJINDER S., 57 Land Between The Lakes, mining site, 59 Land snails, 68 LARDER, BRIDGET M.., 70 Lauraceae, 44 Ledbetter Watershed, 52 LEHMAN, AMELIA, 60 Lepomis gulosus, 102 Lepomis macrochirus, 102 Lepomis megalotis, 102 Life forms, 37 Life history of wood snails, 82 Light wavelength on cercariae, 95 LINDER, JOHAN, 45-49 Lindera, 44 LINDESTOLPE, JOHAN, 45 Liparis loeselii, 27 List of recent reviewers, 139 Lizard microhabitats in Belize, 65 Longear sunfish, 102 Lonicera maackii, 27 LOWE, JERMIAH D., 58 Lunch time intake, female sixth- graders, 62 Lunch time intake, male sixth-grad- ers, 62 Lycopersicon hirsutum, 57 Lymphocytes of farm workers, 62 Maclura pomifera, 42 Mammals, 67 MARLETTE, MARTHA A., 62 McGREGOR, MONTE A., 60 McINTOSH, AMY V., 64 Medical topography, 61 Mesic highwall talus slope, 27 MILLER, MELISSA A., 65 MIMS, STEVEN D., 58 Mining sites associated with over- seas Chinese, 59 Mistletoe, American, of Garrard County, 40-43 Mistletoe, Eastern, of Robertson County, 137-138 Mixed Mesophytic Forest, 67 Monilinia fructicola, 120 Morphine docking, 107 Mu model, 109 Mussels, freshwater, 60 Mycorrhiza, 60 Neotoma magister, 69 Nest boxes, clustered and isolated, 60 Nesting success of wood ducks, 60 Oak-Pine habitat, 67 Oak regeneration, 60 O'BRIEN, MARK, 59, 60 Octameric enolase enzyme, 17-23 Odocoileus virginianus, 68 Opioid receptor models, 107 Opioid sequences, 110 OSBORNE, MELISSA, 63 Osmolality of cercariae, 94 OSUNDE, ADESUWA, 62 OTIENO, TOM, 64 Outer Bluegrass, 41 Owens Farm, 59 Paddlefish, 58 PANEMANGALORE, MYNA, 62 Panther Watershed, 52 Paraquat, 123 PATTERSON, MATTHEW A., 57, 58 Pawpaw, 58 PEARSON, BROOKS C., 3 PENN, WESLEY, 17 Pepper, 58 Permanent pool, 27 Peromyscus leucopus, 69 Pestalotia sp., 120 Pesticides, 57, 62 Pheasant shell mussel, 60 Phoradendron leucarpum, in Gar- rard County, 40-43 Phoradendron leucarpum, in Rob- ertson County, 137-138 Physics and Astronomy abstracts, 63 Plant habitats, 24-34 Plant Life of Kentucky, 66 Plant-pathogenic fungi, 59, 118 Plant succession, 28 Platanus occidentalis, 26, 38, 137 POINDEXTER, DERICK B., 40 Polyodon spathula, 58 POMPER, KIRK W,, 58 PORTER, LARRY, 59 PORTER, LAWRENCE A., 118 POTEET, CATHERINE, 63 POWELL-McCOY, 64 Press release: KAS resolution in support of evolution, 140-141 Proterometra macrostoma, 89, 94, 101 Prunus serotina, 26, 40, 137 Pseudacris triseriata, 65 Psychology abstracts, 63-64 Purple lilliput, 60 Quarry flora, 24 Quercus muhlenbergii, 27, 42 Radish, 58 Raphanus sativus, 58 RAY, ZACHERY M., 58 Recycled waste as soil amendments, 57 Rediae of trematodes, 89 Refugium, 24, 29 REILLEY, SEAN P., 63 Reproductive behavior of rare and endangered freshwater mus- sels, 60 RIGGS, DEVIN L., 70 ROBERTS, AMANDA, 94 Robinia pseudoacacia, 26, 38, 40, 137 Robinson Forest, University of Ken- tucky, 67 Rodents (Muridae), 67 Root mean squared deviation, 19 ROESSLER, ROSE-MARIE, 59 Rosa multiflora, 28 ROSEN, RON, 89, 94, 101 Ruderal species, 36 Russel Cave National Monument, 61 SAKOFSKY, BRIAN D., 61 Salicylate, 59, 118 Salix exigua, 28 Salix nigra, 28 SAMS, ADRIANNE, 59 SARSHAD, AISHE, 89, 94 SCHAPKER, HEIDI T., 70 SCHELL, AMANDA, 94 SCHLOSSER, ANN M., 70 Science Education abstracts, 64-65 Scientists of Kentucky: James W. Abert, 3-16 SCOTT, ROGER, 63 SHEPARD, ADAM C., 60 SHIBER, JOHN G., 64 Shrews (Soricidae), 67 SHUPE, ANDREW E., 24 SIMONETTI, KAREN, 17 Index to Volume 66 Single Cell Gel Electrophoresis (SCGE), 63 Small Mammals, 67 Snails with rediae, 89 Sodium salicylate (SA), 118 SOTO, MARGARET, 64 Spatial analysis of Owens Farm, 58 Spiranthes lucida, 27 Squash, 58 State-listed rare species, 27 STEINMAN, KRISTOPHER, 63 STROBEL, NORM, 59 STROBEL, NORMAN E., 118 Subxeric quarry floor, 28 Survey of Earned Doctorates (SED), 130 Survey of mistletoe, 41, 137 Synergistic inhibition, 118 Swiss chard, 58 TEMPLETON, SUSAN B., 62 Test anxiety and perceptions of ac- ademic skills, 63 THIERET, JOHN W., 44, 73 THOMAS, KRISTIN L., 59 THOMPSON, RALPH L., 24, 35, 40, 73, 137 THROOP, EMILIFE, 89, 94 THUNBERG, KARL PETER, 44 TOBIN, BENJAMIN, 61 Tomato, 57 TOPE, AVINASH, 62 Toxolasma lividus, 60 Trematoda: Azygiidae, 89 Trichoplusia ni, 57 2-trideconone (hendecyl methylke- tone), 57 145 TYLER, RICO, 63 Typha latifolia, 28 Ulmus americana, 33, 40, 137 Vascular flora, in Clark County, 24— 34 VEERKAMP, HEATHER, 61 Viburnum molle, 59 Viscaceae, 40, 137 WANGMO, TENZIN, LOL Warmouth, 102 Water quality of two creeks, 50 Water quality testing experiment, 64 Watermelon, 58 WATKINS, MARCIA, 94 WECKMAN, JUDITH E., 59 WECKMAN, TIMOTHY J., 59 Western Mesophytic Forest, 26, 36, 41, 137 White-footed mouse, 70 White-tailed deer, 68 WHITSON, MAGGIE, 82 Wood ducks, 60 Wood snail, 82 Xeric vertical highwall, 27 Yeast enolase, 17 ZAKI, FADY, 89, 94 ZIMMERMAN ANDY, 61 Zingiber officinale, 57 Zingiberaceae, 57 Zingiberene, 57 Zoology abstracts, 65 =~ wv Poa, a: + Baas obits MMs, ROUTER.” SS, BETS ry Plant Lite Kentuck An titustrated Guide lo-the Vascular Ficrd PLANT LIFE OF KENTUCKY An Illustrated Guide to the Vascular Flora Ronald L. Jones Plant Life of Kentucky is the first comprehensive guide to all the ferns, flowering herbs, and woody plants of the state. This long-awaited work provides identification keys for Kentucky's 2,600 native and naturalized vascular plants, with notes on wildlife/numan uses, poisonous plants, and medicinal herbs. The common name, flowering period, habitat, distribution, rarity, and wetland status are given for each species, and about 80 percent are illustrated with line drawings. The inclusion of 250 additional species from outside the state broadens the regional coverage, and most plants occurring from northern Alabama to southern Ohio to the Mississippi River are examined, including nearly all the plants of western and central Tennessee. . The author also describes prehistoric and historical changes in the flora, natural regions and plant communities, significant botanists, current threats to plant life, and a plan for future studies. Plant Life of Kentucky is intended as a research tool for professionals in biology and related fields, and as a resource for students, amateur naturalists, and others interested in understanding and preserving our rich botanical heritage. $75.00 hardback wii John W. Thieret (1926-2005). Ralph L. Thompson. ................0cccececeeeess 73 Cepaea nemoralis (Gastropoda, Helicidae): The Invited Invader. Maggie WHHSOM oo oi nce Eee ana pind fase cecenua ons ab dinpeseides uaeteteUepeneeepiabmabaesdee eae itn ann 82 Location of Rediae of Proterometra macrostoma (Trematoda: Azygiidae) in the Snail Elimia semicarinata (Gastropoda: Pleuroceridae), and Daily Emergence of its Cercaria. Ronald Rosen, Jonathan Fleming, Bojana Jovanovic, Aishe Sarshad, Emilie Throop, Fady Zaki, and Andy AMMONS 60. ooo siosicccevcesodicedondnodduesethavensessqinscebdcsesduabacts sas dspmesioues sain nama 89 Effect of Light Wavelength and Osmolality on the Swimming of Cercariae of Proterometra macrostoma (Digenea: Azygiidae). Ronald Rosen, Andy Ammons, Ayisa Boswell, Amanda Roberts, Amanda Schell, Marcia Watkins, Jonathan Fleming, Bojana Jovanovic, Aishe Sarshad, Emilie Throop; and. Fady: ZAKI ciiccecc sos cdecentsindeses ives cessopeuweradivatwacs; Jennseus aaa 94 Natural and Experimental Infections of Centrarchid Fishes by the Digenetic Trematode Proterometra macrostoma: Detection of New Infections and Host Histopathology. Ronald Rosen, Elizabeth Anderson- Hoagland, Chris Barton, Bobbie Berry, Jonathan Hardy, and Tenzin Wangitio 55565 0b eee baw sega inlaw unade neuedaneheen auebawcnauers duevales dune aioe 101 Development of the Human Mu, Kappa, and Delta Opioid Receptors and Docking with Morphine. Debra L. Bautista, Wesley Asher, and Lisa CAPPONI voi si soees ok cteede dap eesd gn red eneleee ev as bbowbelude sn ecubcasuscsaiacs the cust eteg a aan 107 Salicylate Inhibits Growth of Plant-Pathogenic Fungi and Synergistically Enhances the Activity of Other Antifungal Materials In Vitro. Norman E. Strobel and: Lawrence A. (Porter: .i.0ccccccc ccs ci cceucbaasevcs ohscsenb aoe sbececsetvannatebe 118 Kentucky Baccalaureate Origins of Doctorate Recipients in the Biological Sciences, Chemistry, and Physics, 1978 through 2002. Fitzgerald B. Bramwell and Elinor L. Brown 1.62 s00.5003; Bcschocnssbecsavecoskcoucasdeuaabaeete aes 129 NOTES Host Occurrence of Eastern Mistletoe (Phoradendron leucarpum), Viscaceae in Robertson County, Kentucky. Ralph L. Thompson. ................20.seeee0es 137 NEWS List-of Recent Reviewers. oj. .0ascdsin ieee ek snc Gecs ahaevacacnedsthen ees tapes eueen in camme 139 Kentucky Academy of Sciences (KAS) Resolution in Support of Evolution .... 140 Index of Volume. 6G 56 iscc8 area aha ohare ek SU Ca a a ed 143