SDAS 2002 Vol 81

Proceedings of the South Dakota Academy of Science

Volume 81 2002

Published by the South Dakota Academy of Science Academy Founded November 22, 1915

Editor Kenneth F. Higgins

Co-Editor Steven R. Chipps

Terri Symens, Wildlife & Fisheries, SDSU provided secretarial assistance

Tom Holmlund, Graphic Designer

TABLE OF CONTENTS

Consolidated Minutes of the Eighty-Seventh Annual Meeting of the South Dakota Academy of Science ...... 1 Presidential Address: Reﬂections on Science in South Dakota and on

Vitamin B12. M. Steven McDowell ...... 11

Complete Senior Research Papers presented at The 87th Annual Meeting of the South Dakota Academy of Science Hydrogeology of the Homestake Mine. Perry H. Rahn and William M. Roggenthen ...... 19 Mycorrhizal Colonization as Impacted by Corn Hybrid. Marie-Laure A. Sauer, Diane H. Rickerl, Patricia K. Wieland, Courtenay Hoernemann, and W.B. Gordon...... 27 Retention and Survival Rates Associated With the Use of T-Bar Anchor Tags in Marking Yellow Perch. George D. Scholten, Daniel A. Isermann and David W. Willis ...... 35 Impact of Crop Harvest on Small Mammal Populations in Brookings County, South Dakota. Melissa K. Pinkert, Jonathan R. Meerbeek, George D. Scholten, and Jonathan A. Jenks ...... 39 Observations at the Lake Martin Rookery, Cypress Island Preserve, St. Martin Parish, Louisiana. James E. Martin...... 47 Juveline Marine Reptiles from the Late Cretaceous of the Antarctic Peninsula and their Relationships to Other Such Occurrences in Central South Dakota and Belgium. James E. Martin...... 53 Analysis of Wind Observations in the Black Hills Region. Srivisakha Skandaswamy Pokkunuri, Sameer Kumara Krishna Dronamraju, Abul R. Hasan, and Andrew G. Detwiler...... 59 Modeling Streamflow in Spearfish Creek, Black Hills, SD. Larry D. Stetler and John P. Knight...... 69 South Dakota Turfgrass Industry: Impacts and Value. Leo C. Schleicher...... 75 Chondrichthyes from the Upper Part of the Minnelusa Formation (Middle Pennsylvanian: Desmoinesian), Meade County, South Dakota. D.J. Cicimurri and M.D. Fahrenbach ...... 81 Influence of Age-0 Yellow Perch Abundance on Walleye Populations in Two Eastern South Dakota Lakes. Jonathan R. Meerbeek, Daniel A. Isermann, and David W. Willis ...... 93 Constitutive Expression of a 3-Deoxy-D-Arabino-Heptulosonate-7-Phosphate Synthase Gene in Alfalfa (Medicago sativa L.) Lines Differing in Morphology and Lignin Concentration. T. Liu, K.D. Kephart, and C.D. Carter ...... 101 Construction of Natural and Real Numbers from Generalized Order, Congruence and Continuity Axioms (A “Real Line” May Cross Itself). Richard P. Menzel...... 109 Initiation of Feeding During Hatchery Rearing of Landlocked Fall Chinook Salmon Fry. Michael E. Barnes, William A. Sayler, and Rick J. Cordes ...... 137 White-Tailed Deer Behavioral Response to Predator Feces. Christopher C. Swanson, Cory M. Kassube, and Jonathan A. Jenks ...... 143 Variation in Small Mammal Richness Among Ecotypes in Eastern South Dakota. David F. Terrall, Nick G. Cochran, and Jonathan A. Jenks...... 147 The Expanding Distribution of the Least Shrew, Cryptotis parva, in South Dakota. Douglas C. Backlund...... 153 Rare Earth Element Signatures of Fossil Vertebrates Compared with Lithostratigraphic Subdivisions of the Upper Cretaceous Pierre Shale, Central South Dakota. Doreena Patrick, D.E. Grandstaff, James E. Martin, and D.C. Parris ...... 161 3- Phosphonosulfonate(3-) O3PSO3 and Related Ions. Arlen Viste and Kari Lunder Stone ...... 181 Sets of Half-Reactions Which Yield Different Emf for an Overall Reaction, But Equivalent Standard Gibbs Free Energy and Equilibrium Constants. April Kay Bobbie and Arlen Viste...... 189 Application of ADF Amsterdam Density Functional Package to Relativistic 2+ Effects in Mercurous Ion Hg2 and Related Species. William J. Delfs and Arlen Viste...... 195 Estimating Summer Fawn Mortality of White-Tailed Deer Using Corpora Albicantia and Lactation Status. Robert G. Osborn, Beth A. Hippensteel, Jonathan A. Jenks, and Ted A. Benzon...... 201 Longitudinal Patterns in Fish Community Composition of Upper Missouri River Backwaters. Kipp A. Powell and Steven R. Chipps...... 211 Zooplankton Communities of Black Hills Reservoirs. Benjamin M. Holcomb and Steven R. Chipps...... 219 Fecal Egg and Oocyst Counts in Dogs and Cats from Animal Shelters from South Dakota. M.B. Hildreth, J.A. Bjordahl and S.R. Duimstra...... 227

Abstracts of Senior Research Papers presented at The 87th Annual Meeting of the South Dakota Academy of Science Good Wasps Gone Bad: Invasive Species in a Parasitoid Guild on a Native Legume. Paul J. Johnson and Arvid Boe...... 237 Comparative Morphology of Several Switchgrass Cultivars. Martine Zamy and Arvid Boe ...... 239 Polymorphism in the Agouti-Related Protein (agrp) Gene in Swine. Kent J. Donelan, Nels H. Granholm, Carl A. Westby, and Donald M. Marshall...... 241 Transgene Dispersal and Effects in Feed and Food from Roundup Ready Soybeans. C. Carter, S. Hansen, R. Scott, B. Turnipseed, L. Wrage, P. Krishnan, R. Thaler, and T. Cheesbrough ...... 243 Species Richness and Nesting Success of Neotropical Migrants in Natural River Corridors and Anthropogenic Woodlands in Southeastern South Dakota. Dale Gentry and David Swanson ...... 245 Isolation and Partial Characterization of the Hapliod State of Ustilago Maydis from Smutted Ears and Tassels of Field Corn (Zea mays) from Southeastern South Dakota. Jodie Linneweber and Donna Hazelwood ...... 247 Gene Expression Patterns in Segmenting Radials of Zebrafish, Danio rerio. Patricia L. Crotwell and Paula M. Mabee ...... 249 Relationships Among IKONOS Satellite Imagery, Airborne Scanning LIDAR Data, and Ground-Based Tree Inventory Data in a Ponderosa Pine Forest via Subpixel Interpretation. X. Chen, L. Vierling, D. Dykstra, W. Capehart, E. Rowell, and T. DeFelice ...... 251 Seasonal Variation on Brown Trout Diet in Rapid Creek, South Dakota. S. Martin Digler and Kerri T. Vierling...... 253 Urban Gradient Influence on Brown Trout Diet in Rapid Creek, South Dakota. Irene Cherni and Kerri T. Vierling ...... 255 An Evaluation of the Effects of Formalin and Hydrogen Peroxide Treatments on the Exterior Egg Membrane of Chinook Salmon (Onchorhyncus tshawytscha). Hans Stephenson, Mark Gabel, and Mike Barnes ...... 257 A More Biodegradable Quaternary Ammonium Compounds. Keith Hartman and Gary Earl ...... 259 Initial Examination of Possible Novel Macrocycles via CAChe. Ethan Englund, Arlen Viste, and Jetty Duffy-Matzner ...... 261 Synthesis of α,β-Unsaturated Carbaldehydes, Ketones And 1,4 Addition to α,β-Unsaturated Aldehydes. Jerrid Kruse, Andrea Votroubek, and Jetty Duffy-Maztner...... 263 Ridge Detection Using Artificial Neural Networks. Rishi Kishore and John M. Weiss...... 265 The Effects of Hyperbaric Environment-Induced pH Changes on the Generation Times of Pseudomonas aeruginosa. Contessa E. Creager, Sarah O. Hammar, Mitchell W. Jacobs, John Wayne Kalda, Spring J. Mayer, Brandon J. McCall, Jenny M. Volkert and William J. Soeffing...... 267 Observations on the Colonial Morphologies, Cell-Cell Adhesion Characteristics and Antibiotic Sensitivity of Pseudomonas aeruginosa in a Hyperbaric Environment. Brandon J. McCall, Sarah O. Hammar, Nicolas L. Strasser and William J. Soeffing...... 269 Nutritional values of Psoralea esculenta. Giani Perera and Neil Reese...... 271 Extraction, Purification, and Identification of Unique Protein in Root of (Psoralea esculanta). Marcus Braaten and R. Neil Reese ...... 273 Effects of Lethal Yellow and Mahogany Mutations on Reproduction in Female Mice. Trynda N. Oberg, Nels H. Granholm, and Maureen R. Diggins...... 275 Determination of Infective Gastrointestinal Nematode Juveniles on Spring Pastures . K. J. Mertz, M.B. Hildreth, and W.B. Epperson ...... 277 Determining the Feasibility of Restoring River Otters in South Dakota. Alyssa M. Kiesow and Charles Dieter...... 279 Comparative Analysis of Wing Beat Frequency and Eye Span in Stalk-Eyed Flies (Diptera: Diopsidae). John G. Swallow, Jilian M. Fazio, and Gerald S. Wilkinson ...... 281 Late Cretaceous Dinosaurs, Big Bend National Park, Texas. Julia T. Sankey ...... 283 Vertebrate Paleocology from the Late Cretaceous Hell Creek Formation, North Dakota: Surface Collection Versus Screen Washing. Georgia E. Knauss and Andrew A. Farke...... 285 Morphology of the Cornual Sinuses in Chasmosaurine Horned Dinosaurs (Ornithischia: Ceratopsidae). Andrew A. Farke ...... 287 Microvertebrate Fauna Recovered from the Hell Creek Formation (Late Cretaceous), Southwestern North Dakota. Marcus Ross and Tim Pranger ...... 289 Potential Effects of Genetically Modified Soybeans on Mammalian Fetal, Breast-fed Postnatal, Pubertal and Adult Development. Denise Brake and Donald P. Evenson ...... 291 Comparison of Two Techniques for the Detection of DNA Fragmentation in Sperm Chromatin: Sperm Chromatin Structure Assay and Comet. David Tritle, Lorna Jost, Kay Kasperson, and Donald P. Evenson ...... 293 Development of a GFP Construct to Determine the Localization of a Porcine Reproductive and Respiratory Syndrome Virus Protein. William D. Arndt, Susan Ropp, and David Benfield...... 295 Long-Term Plankton and Nutrient Changes in Prairie Lakes. Lois Haertel ...... 297 Plankton-Environment Association in Prairie Lakes. Lois Haertel...... 299 Use of Stable Isotopes to Document the Invasion of Tallgrass Prairie by Decidous Forest in Newton Hills State Park in Southeastern South Dakota. Craig Spencer, Matt Bukrey, Jason Smalley, Erik Wang, Dan Opheim, and Mike Chapman...... 301 The Utility of Aerial Photographs, Field Observations, and Invertebrate Fossils in the Construction of Geologic Maps. Frank J. Varriale and Alvis L. Lisenbee ...... 303 Observations of a Lightning Channel in a High Plains Thunderstorm. Tom Warner, John Helsdon, Andrew Detwiler, Qixu Mo, and Donna Kliche...... 305 A New Meandering Stream Complex from the Big Badlands of South Dakota. Robert W. Meredith and Jennifer Cavin...... 307 Using the Pentax Total Station and Arcview to Analyze the Fossil Accumulations Within the Conata Picnic Ground Excavation, Badlands National Park, South Dakota. Daniel B. Lien...... 309 Hydrolytic Stability Testing of Polymer Matrix Composites Reinforced with Fibers Having Novel Surface Treatments. H. Du, C. Struckman, D.H. He, W.M. Cross, J.J. Kellar, and D.A. Boyles...... 311 Forest Classification of the Black Hills, SD Using Decision Tree Analysis on IM and AVIRIS data. Sylvio Mannel, Doug Hua, and Maribeth Price...... 313 Pharmaceuticals, Binding Sites, and Pharmacological Effect. Rosina Barbara Flasch Hendrickson and Arlen Viste ...... 315 Nest Site Characteristics of Eastern Wild Turkey Females in Agricultural Landscapes of Northeastern South Dakota. Roger D. Shields and Lester D. Flake ...... 317 Correlation of Biochemical and Molecular Profiles for 35 Related Isolates of Methicillin Resistant Staphylococcus aureus. Amy J. Stephenson, Lona Haas, and Patricia M. Tille ...... 319 Biodiversity Assessment of The James River Basin. Chad Thury, Christy DeRoos, John Gullikson, Krisma Dewitt, James Sorenson and Patricia M. Tille ...... 321 Diversity, Density and Breeding Status of the Avifauna in the Schramm Addition at Ponca State Park, Nebraska. Eric T. Liknes ...... 323 Syntheses and Applications of the New Organoboron Reagents for Suzuki- Coupling Reaction. Jiangtao Hao and David A. Boyles ...... 325 Use of ArylDiazonium Tetrafluoroborates For Suzuki-Type Cross-Coupling Reactions with Diarylborinic Acids. Joseph Marshall, Jiangtao Hao, and David A. Boyles...... 327 Modified Suzuki Reaction Using Bisaryl Tosylates with Diarylborinic Acids. Celeste Mercado, Jiangtao Hao, and David A. Boyles ...... 329 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 1

CONSOLIDATED MINUTES OF THE EIGHTY-SEVENTH ANNUAL MEETING OF THE SOUTH DAKOTA ACADEMY OF SCIENCE APRIL 5, 2002 AUGUSTANA COLLEGE, SIOUX FALLS, SD

The Executive Council met at 8:00 a.m. Friday 5 April 2002 for a final check of plans for the day. President Charles Lamb opened the executive committee meeting, noted that a quorum was present, and thanked Ken Higgins for timely presentation of the 2001 Proceedings. Ken Higgins reported that the 2001 Proceedings has been printed and distributed to members. Because the funds in the proceedings account are sufficient to cover expenses and the abstracting services that Steve Chipps has contacted do not charge the Academy, Ken suggested that it would be possible to gift funds to the SDAS general fund to assist with expenses for the 2004 meeting in Chamberlain. Ken moved and Bill Soeffing seconded that the Treasurer will bill the Proceedings Account and the Proceedings account will gift back the sum of $2,000.00 or $3,000.00 to partially cover expenses for the 2004 annual meeting. The motion passed by voice vote. Krisma DeWitt reported that Mount Marty College has graciously accept- ed the invitation to become part of the rotation of hosting institutions for the Annual Meeting. The executive committee voted to allow the choice of meeting dates for the Annual Meeting of either Friday or Saturday will be the prerogative of the host institution. Neil Reese moved and Dave Swanson seconded that the procedure for selection of Fellows be modified to allow the format for nomination to consist of a brief letter of recommendation. Names of individuals for consideration will be forwarded to, or may originate with, the Executive Committee. The motion passed by unanimous voice vote. Bob Tatina moved and Steve Chipps seconded a motion that the President appoint one member of the executive committee, or ask for a member to volunteer to solicit nomination of Fellows at the Business Meeting. The motion passed by voice vote. Krisma DeWitt volunteered to contact each campus to solicit nominations of one or more Fellows. Ken Higgins reported that the By-laws need to be revisited and updated. A committee will be appointed. Ken Higgins moved and Neil Reese seconded that the position of Secre- tary-Treasurer be split into two positions. The motion passed by unanimous voice vote. The 2004 annual meeting will be held on the Missouri River at Cedar Shore Resort, Chamberlain, SD, and will have an emphasis on The Missouri River and/or Lewis and Clark. This meeting will not have a host institution, 2 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) but instead will have an Organizing Committee consisting of Ken Higgins, Chair, Miles Koppang, Bill Soeffing, and Donna Hazelwood. This meeting will consist of invited speakers Friday 2 April and paper sessions Saturday 3 April. The treasury of the Academy will support the cost of the keynote speaker to be selected by the committee. The Speaker Selection Committee includes Ken Higgins, Steve Chipps, Miles Koppang, Charles Lamb, Bill Soeff- ing, Bob Tatina, and Neil Reese. Jim Martin will arrange fieldtrips. In addition, Ken Higgins will be point person for the Program Committee. Registration for the Annual Meeting began at 7:30 a.m. this morning. Ken Higgins brought Terri Symens, and Mike Wanous has lined up Cheryl Holzapfel to assist with registration. The Nominating committee will have the following positions to fill, Sec- retary, Treasurer, Second Vice President, and two members-at-large. Bill So- effing and Mike Wanous will be going off as members-at-large. Kristel Bakker provided the Treasurer’s report. Audrey Gabel and Mark Gabel served as Auditing Committee. The CD at Dakotah Bank will be allowed to rollover for another term. The duties of Treasurer and the check- book were passed to Kristel 1 June 2001. The 2003 meeting of the Academy will be at South Dakota School of Mines and Technology and the 2004 meeting will be at Cedar Shore Resort Chamberlain SD. The executive committee will meet in Chamberlain, early Fall, most likely September. Ken Higgins provided the Proceedings report and had already passed out copies of the 2001 proceedings. Page costs are $25.00/page. The Academy is responsible for mailing copies to members and the Academy pays for the cost of mailing copies of the proceedings to participating Libraries. The cost of an issue of the Proceedings is currently $10.00 including postage. Ken suggested that the Academy provide honoraria in the following amounts to Terri Symens $200.00, Di Drake $100.00, and Cheryl Holzapfel $100.00. Neil requested $500.00 to pay for a student to assist with the Academy web site. Elections were held and officers for 2002-2003 are: members at large 2002-2004 Neil Reese and Gary Earl, Second vice-President Bob Tatina, Secre- tary Donna Hazelwood, and Treasurer Kristel Bakker. Audrey Gambel moved and Krisma DeWitt seconded acceptance of the nominations. The motion carried by voice vote.

Committee positions for 2002-2003 include: Membership Bob Tatina Maurine Diggins Bylaws Miles Koppang Bob Stoner Chuck Estee Resolutions to be ﬁlled Nominations to be ﬁlled Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 3

Andy Detweiler reported on the status of involvement of the Academy and Regional Science fairs. South Dakota hosts five regional science fairs, and the Academy could help by careful scheduling of Annual Meeting to not conflict with regional fairs, assisting with judging, donation of prizes especially at the middle school level, and providing to the five regional directors an e-mail list of members of the academy to allow for selection of judges of regional science fairs. Neil Reese moved and Mark Gabel seconded that the five regional directors be provided an e-mail list of members of the academy to allow for selection of judges of regional science fairs. The list to be distributed by the executive committee fall 2002. The motion passed by voice vote. Nels Granholm reported that a meeting on Bioethics will be held 1-7 June in Madison, Wisconsin. This meeting will address introduction of concepts of ethics in classrooms. Several items for consideration at the fall meeting of the Executive Com- mittee were discussed. 1) Ken pointed out that the by-laws were last updated in 1989 and need to be updated. 2) The 2004 meeting including examination of proposed speakers for the general session Friday 2 April. 3) Nomi- nation of individuals for Fellows Awards. 4) Recruitment of new members. 5) A new operations manual to replace the one that apparently has been lost. 6) Assisting the five regional science fairs, providing prizes at the middle school level. Miles and Andy will brainstorm and bring a report to the meeting. President Charles Lamb opened the meeting on Friday 5 April, Local Ar- rangements Committee Chair Milton Hanson welcomed all to campus, and Bruce A. Halverson, President of Augustana College, extended his welcome. In his address at the opening session, President-Elect Steven McDowell discussed Reflections on Science in South Dakota. Papers for the Academy began at 10:00 a.m. Friday 5 April, and continued throughout the day. Seventy- three papers were presented. The Keynote speaker, Joy B. Zedler, presented "Invasive Species in Wetlands: Unwanted Guests or Total Conquerors." One hundred sixteen individuals attended the annual meeting, 61 regular members and 55 associate members.

Respectfully submitted, Donna Hazelwood, DSU SDAS Secretary 4 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002)

SOUTH DAKOTA ACADEMY OF SCIENCE 2001-2002 EXECUTIVE COMMITTEE

PRESIDENT Steve McDowell, SDSM&T Chemistry 394-1229 [email protected]; FAX 394-1232

PRESIDENT-ELECT Miles Koppang, USD Chemistry 677-5011 [email protected]

FIRST VICE-PRESIDENT Andrew Detwiler, SDSM&T IAS 394-2291 [email protected]; FAX 605-394-6061

SECOND VICE-PRESIDENT Robert Tatina, DWU Biology 995-2712 [email protected]

CO-SECRETARY TREASURER Donna Hazelwood, DSU Natural Sciences 256-5187 [email protected]; FAX 256-5643

CO-SECRETARY TREASURER Kristel Bakker, DSU Natural Sciences 256-5182 [email protected]; FAX 256-5643

PROCEEDINGS EDITOR Kenneth F. Higgins, SDSU Wildlife 688-4779 [email protected]; FAX 688-4515

ASST. PROCEEDINGS EDITOR Steve Chipps, SDSU Wildlife 688-5467 [email protected]; FAX 688-4515

FIRST PAST PRESIDENT Charles Lamb, BHSU Biology 642-6026 [email protected]

SECOND PAST PRESIDENT Lenore Koczon, NSU Chemistry, University College, 626-2633 [email protected]; FAX 626-3317

MEMBERS-AT-LARGE

2001-2003 Krisma DeWitt, Mount Marty College Chemistry 668-1530 [email protected]

2001-2003 James Lefferts, Dakota Wesleyan University Chemistry [email protected]

2002-2004 Gary W. Earl, Augustana Chemistry [email protected]; FAX 357-9772

2002-2004 R. Neil Reese, SDSU Biology 688-4568 [email protected] Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 5

REPORT OF THE 2002 RESOLUTIONS COMMITTEE

The South Dakota Academy of Science wishes to thank Augustana Col- lege for hosting the 2002 meeting. In particular, we thank Bruce A. Halver- son, President of Augustana College, and are grateful to the local arrange- ments committee — Milton Hanson, Michael Wanous, Cheryl Holzapfel, Jay Smith, John Larkin, Maureen Diggins, Duane Weisshaar, Steve Shum, and Tim Sorenson and to the program committee — Ken Higgins, Steve Chipps, Michael Wanous, Terri Symens and Cheryl Holzapfel, for their hard work in producing an excellent meeting. A special thanks also to the several Augus- tana students for their assistance. Thank you to Charles Lamb for his dedication as President of the Acade- my for the past year. Thank you also to Steven McDowell, President-Elect, for his address on "Reﬂections of Science in South Dakota and on Vitamin

B12". The Academy wishes to thank Donna Hazelwood and Kristel Bakker for their service as Secretary-Treasurer over the past year. A special thanks goes to Editors Ken Higgins and Steve Chipps for their oversight of timely publication of the Proceedings. The Academy was also enlightened by the Public Lecture on "Invasive Species in Wetlands: Unwanted Guests or Total Conquerors?" by Dr. Joy B. Zedler, from the University of Wisconsin-Madison. Dr. Zedler was also a former Augustana College graduate.

Respectfully submitted, Gary Larson, Resolutions Committee

TREASURER’S REPORT

June 1, 2001 Starting Balance $11,306.64

Deposit 20.00

Withdrawals 124.38 Brookings Engraving 100.00 Terri Symens 100.00 Di Drake 81.00 AmericInn Conference Room 85.00 Cedar Shores Lunch 143.64 USD Biology Department (2001 SDAS)

Ending Balance: $10,692.62 (2001)

SDAS Treasurer, Kristel K. Bakker 6 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002)

EXECUTIVE SUMMARY, 2002 — PROCEEDINGS EDITORS

Volume 80 for 2001 totaled 457 pages and the production cost for 324 copies was $8,637.04. As of March 27, 2002, invoices for page and reprint charges totaled $11,060 for an estimated proﬁt of $2,194.02. To date $10,490 has been received and deposited in the Academy account for a proﬁt of $1,852.96. Our overall Proceedings account balance to date is $6,476.24. Copies have been mailed to all current members, all life members, all the State libraries and to eight indexing services.

Respectfully Submitted by: Kenneth F. Higgins, Editor and Steven R. Chipps, Assistant Editor For the S.D. Academy of Science Proceedings March 28, 2002

APRIL 5, 2002 BUSINESS MEETING REPORT – SDAS BY: K.F. HIGGINS, PROCEEDINGS EDITOR AND S.R. CHIPPS, ASSISTANT EDITOR

1. First, we want to thank those who helped us manage and produce the 2001 Proceedings: Terri Symens – Invoices, Mailing Labels, Correspondence and Entering Names and Titles for the Program Tom Holmlund – Proceedings Layout, Figure Scanning and Final Copy Formatting Donna Hazelwood – For Meeting Notes, etc. And several members who helped with obtaining member names and addresses at various Universities or Colleges 2. We’re happy to report that the 2001 Volume 80 of the Academy Proceed- ings have been published and distributed. It totaled 457 pages and costs were: $2,836.50 layout, $4,714.15 printing, and $1,086.39 reprints for a production cost of $8,637.04. 3. Finances for producing 324 copies of the Proceedings plus article reprints appear to be adequate, given the $25 rate per printed article or abstract page plus reprint costs. The use of the SDSU accounting system has been a positive action to date. Please refer to the Finance Summary Page for details of the account (see the green sheet). 4. The USD Library system is the repository and distributor of past volumes of Academy Proceedings. Joe Edelen is the contact person. They will provide storage, shelving and mailing of requested volumes, but the academy will have to handle invoices and collection of payments. He says most libraries sell extra copies of Proceedings for $5-10 per copy. 5. We have delivered copies of the 2001 Proceedings to all University Li- braries for the schools usually represented at the Academy meetings. Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 7

6. We have also submitted two copies of the 2000 and 2001 Proceedings to the contact for the National Academies — Tom Weaks, SASA Managing Editor, AcadSci, Inc., Box 141, Cumberland City, TN 37050. 7. We are gaining on the request to receive all abstracts and papers on electronic ﬁles (diskettes). We only had 2 or 3 submissions in 2001 that we had to scan or retype in Word or WordPerfect. We have over 70 papers and abstracts for 2002. 8. We recently increased the indexing coverage of the "Proceedings" by con- tacting numerous abstracting/indexing services. Six out of nine indexing services agreed to include "Proceedings of the South Dakota Academy of Sciences" in their Journals list. Table 1 lists these indexing services, the disciplines covered, contact information and web site access. All of the services are provided with a complimentary subscription to "Proceedings" and have been forwarded all back issues so they are up-to-date (e.g., through 2001). I checked several of these services to verify that Proceed- ings was listed in their Journals list and found that Proceedings is currently listed. AGRICOLA is reviewing recent issues and should let us know if they wish to include Proceedings in their Journal index. Several services were quite selective in the journals that they cover. Indexing services that decided not to include Proceedings are listed in Table 2. With this increase in coverage, we anticipate that more people will have access to information covered in Proceedings. To maintain this effort, we continue to look for appropriate indexing services and are open to suggestions from the membership for enhancing coverage of the Proceedings. Please forward questions/suggestions to Steve Chipps at [email protected] (605.688.5467).

Table 1. List of indexing services that include "Proceedings of the South Dakota Academy of Sciences".

Indexing Service Coverage Contact Web page access

Cambridge Scientiﬁc Aerospace Dr. Craig W. Emerson www.csa.com Abstracts Aquatic Sciences Director of Editorial Operations and Fisheries 7200 Wisconsin Ave. Sociological Abstracts Bethesda, MD 20814

GeoRef Geology/Physical Library www.agiweb.org/georef.html Sciences American Geological Institute 4220 King Street Alexandria, VA 22302-1502

National Information Wildlife Abstracts NISC Colorado www.nisc.com Services Corporation Attn: Luana Heikes (NISC) 1302 S. Shields Street, #A2-4 Fort Collins, CO 80521

Fisheries Abstracts NISC USA Attn: Debbie Durr 3100 Saint Paul St., Suite 806 Baltimore, MD 21218

Chemical Abstracts Chemistry Abstracts Margit Ballard www.cas.org Service Acquisitions Chemical Abstracts Service—CAS PO Box 3012 Columbus, OH 43210 8 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002)

Table 1 (continued). List of indexing services that include "Proceedings of the South Dakota Academy of Sciences".

Indexing Service Coverage Contact Web page access

State Academy of Proceedings of State Academies of www.acadsci.com Science Abstracts State Academies Science Abstracts AcadSci, Inc. PO Box 141 Cumberland City, TN 37050

Abstracts in Anthropology Baywood Publishing, Inc. www.baywood.com Anthropology Abstracts 26 Austin Ave. Box 337 Amityville, NY 11701

AGRICOLAa Agriculture Sally Sinn www.nal.usda.gov/ag98/ Agricola Journal Evaluation Panel Technical Services Division National Agricultural Library 10301 Baltimore Ave. Beltsville, MD 20705 a Proceedings are currently being reviewed by Journals Panel.

Table 2. Indexing services that were contacted but decided not to include Proceedings of the South Dakota Academy of Science.

Indexing Service Coverage Comment

INSPEC Physics/Engineering Covers primary, peer-reviewed journals

BIOSIS Biological Abstracts Too selective

Current Contents (ISI) Miscellaneous Too selective

PROCEEDINGS DISBURSEMENTS/RECEIVABLES Prepared by Di Drake, March 27, 2002

1996 1997 1998 1999 2000 2001 Total

Layout/Formatting 1,194.00 1,221.00 0.00 990.00 780.00 2,836.50 7,021.50

Publication 2,899.00 3,441.80 2,245.95 2,954.80 2,585.92 4,714.15 18,841.62

Reprints 677.88 1,034.67 1,526.17 299.75 823.57 1,086.39 5,448.43

Miscellaneous Printing 5.22 54.51 45.72 69.26 17.47 192.18

Supplies,Phone,Postage 37.70 104.88 160.56 126.55 211.47 641.16

TOTAL EXPENSES 4,813.80 5,856.86 3,978.40 4,244.55 4,385.30 8,865.98 32,144.89

TOTAL INVOICED 4,875.00 7,348.00 3,830.00 5,200.00 3,635.00 11,060.00 35,948.00

Proﬁt / Loss to date 61.20 1,491.14 -148.40 955.45 -750.30 2,194.02 3,803.11 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 9

CASH BALANCE IN SDSU — SOUTH DAKOTA ACADEMY OF SCIENCE PROCEEDINGS ACCOUNT Prepared by Di Drake, March 27, 2002

1996 1997 1998 1999 2000 2001 Total

Total Paid Expenses -4,813.80 -5,856.86 -3,978.40 -4,244.55 -4,385.30 -8,865.98 -32,144.89 Paid by SDAS Treasurer 502.58 4,244.55 4,747.13

Payments Received 4,600.00 7,334.00 2,990.00 4,640.00 3,010.00 10,490.00 33,064.00 Corrections Est 275.00 250.00 525.00 1,050.00 Paid to SDAS-Treasurer -240.00 -240.00

Balance of Account 288.78 1,477.14 -953.40 4,890.00 -850.30 1,624.02 6,476.24

Percent of Paid Invoices 94 100 78 89 83 95 92

Unpaid Invoices 275.00 14.00 565.00 310.00 100.00 570.00 2,074.00

Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 11

PRESIDENTIAL ADDRESS Reﬂections on Science in South Dakota

and on Vitamin B12

Address to the South Dakota Academy of Science April 5, 2002

Presented by M. Steven McDowell South Dakota School of Mines and Technology Rapid City, SD 57701-3995

As I reviewed the previous Presidential addresses from the past decade, I found two themes that dominate the addresses — comments on the state of science in South Dakota and beyond and presentations on scientiﬁc subjects of interest to the President of the Academy. I have chosen to speak with respect to both of these subjects.

SCIENCE AND CHANGE IN SOUTH DAKOTA

Change is an inevitable event. The only thing certain is uncertainty. If you do not like the weather in South Dakota, give it an hour and it will change. Clichéd sayings yet appropriate ones for all of us. As I sit and finalize my thoughts for this address, I have already had to change some of them, maybe many of them, even the ones that were written as recently as just a few days ago. A day does not seem to pass when what we knew yesterday is not what we know today, sometimes for the better, and sometimes for the worse. Science in South Dakota, whether it is related to its teaching or its research, has changed considerably over the twelve plus years during which I have lived and worked in the State of South Dakota. The changes have been at the local, statewide, and global levels. I, as many of you, have experienced these changes, which are, in large part, the result of the explosive growth and development of our electronic technologies. These changes are guaranteed to continue. We will soon be entering a new, possibly different, era of science and education in South Dakota. This era is one that will be mapped by a new official in the highest office of the state. For many of us, new officials at the local level will also map the direction within this new era. Personally, my institution, the South Dakota School of Mines and Technology (SDSM&T), is beginning to look at its next 10 to 20 years. Its executive officer, Dr. Richard Gowen, has given notification that he will be resigning during the summer of 2003, and now SDSM&T is considering its future. Other institutions in the state are facing similar situations as some of their high level administrators are considering career changes and moves. 12 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002)

I, as many of you, have experienced the highs and lows of education, particularly science education, in the state. I have met with satisfaction and frus- tration as I have performed my tasks and met my responsibilities as a member of one of the higher education institutions in the state. I have had to learn to adapt and change, sometimes as quickly as the proverbial overnight, to maintain a competency within my position. The delivery of education, perhaps as recently as 10 years ago, was for many of us, the standard classroom setting in which traditional lecture was likely employed to deliver the material to the student. The student was required to supplement this material by reading an accompanying textbook, and the student’s assessment was probably accomplished by traditional written examination. Since then, of course, many changes have occurred. Electronic methods of delivery and interaction have entered our classroom. The textbook is now complemented or even replaced by CD’s and websites. Students bring their laptops to class for downloading of materials as opposed to transcribing the material given during the lecture, and in turn, they upload their completed assignments. Group learning, team teaching, and interdisciplinary subjects have entered the classroom. Assessment likely still includes written examinations but has been expanded to include oral examinations, traditional student portfolios, and electronic student portfolios. Delivery of course content has also evolved. First, it was the satellite uplink, then the videotape, and now on-line streaming of lecture material. These changes have permitted the expansion of the boundaries of our institutions and our state. Articulation agreements and cooperativity agreements have created new centers of education and new op- portunities for students by which they can attain their higher education goals. These current trends are likely to be our future, especially the outsourcing of education to sites where traditional centers of learning do not exist. Change is now prevalent in the ways scientists conduct their research and share their discovery. Scientists by nature are solitary creatures and often pur- sue their scientific inquiry on an individual basis, sometimes with the help of a body of research assistants. But this is changing and also will continue to change. The speed at which data is collected, analyzed, and reported is increasing almost on a daily basis and the single scientist is a diminishing entity. Partnerships, collaborations, and research teams are developing between faculty within departments, between departments, between institutions, and between even institutions in different countries. Similar couplings are happening between academia and industry. A review of the current scientific literature re- veals a rapidly growing number of publications that are being submitted by multiple principal researchers and that are being submitted from multiple institutions. Within South Dakota, the Experimental Program to Stimulate Com- petitive Research (EPSCoR) and Small Business Innovation Research (SBIR) are two programs that have helped to foster the pursuit and establishment of many of these partnerships. A faculty member’s survival, tenure, and promotion at many institutions rely on the ability to conduct some level of research or schol- arly activity. Collaboration in research was at one time viewed with a negative attitude by evaluators of faculty performance, but now such teamwork is en- couraged if not mandated. Finally, scientists are faced with the reinvention of Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 13 themselves, often as the result of changes in their funding sources, and sometimes simply because of our inherent curiosity in the things that surround us. Research is being done still in the name of basic science and for the satisfaction of gaining knowledge, but more and more it is also being done for the sake of application, for profit, and for the support and benefit of the local and global communities. As a closing note on science in South Dakota, I earned a Ph.D. in Inor- ganic Chemistry. My graduate education prepared me for a life of research and scientific inquiry. A postdoctoral experience further prepared me for this en- deavor. However, I have often been the researcher last and everything else first, things for which I was never truly trained. It is hard to tell what it is that I am today, as I am the chair of my department, an educator of students, a mentor of graduate students, and an advisor of students and colleagues – I am an first administrator and a teacher, and then finally a researcher. But I am a scientist, and although what I was specifically prepared to do is not what I usually do each day, my training as a scientist did prepare me for the expected and frequently sudden changes that occur in my position and in my life. I believe I speak for many of us in that we are prepared for these changes because we are scientists. We are by nature the skeptic, and we are therefore always wary of the things that we observe, that we stand ready to always look at life with an open eye and mind, and that we respond quickly and correctly to the unexpected.

THE COBALAMINS – THE INORGANIC VITAMINS

Vitamins are vital chemical and biochemical species that are required in small or trace amounts to maintain a healthy life in the higher animals. The animals that require them do not make such species, nor do they store these vitamins in their bodies for extended periods of time. Commonly known vitamins include Vitamin C (ascorbic acid), Vitamin E (a-tocopherol), Vitamin B1 (thiamine hydrochloride), and Niacin (nicotinic acid). Unique among the vitamins is Vitamin B12, owing to the fact that it is considered to be inorganic in its composition and function. The other vitamins are considered organic in their composition and nature.

Vitamin B12 and its related compounds are collectively known as the cobalamins, a name that is derived from the shortening of the expression "cobalt vitamin". It is the presence of cobalt within these compounds that makes it inorganic, makes it unique among the vitamins, and permits it to perform its vital and life-sustaining transformations within the higher animals. Without the presence of the cobalt, the vitamin is non-functional and is of no worth to a living system.

In its biochemistry, Vitamin B12 does not function independently. As such, it is therefore biologically designated as a coenzyme, meaning that it acts in concert with other biological molecules to complete its designated function. As already mentioned, it is a species that is not produced by the animals that require it and plants also do not produce it. It is only produced by microorgan- 14 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) isms (anaerobic bacteria) that are typically found in the intestinal tract of animals. Humans and other animals acquire their daily dose of Vitamin B12 through a diet that includes products such as meat, milk, cheese, and eggs. Vitamins typically are classiﬁed as fat or water soluble, based upon where the vitamin is stored by the body. Vitamin B12 is one of the water-soluble vitamins and is widely found throughout the body in locations such as the blood, brain, milk, spleen, and kidney. A healthy person needs to ingest and uptake but a trace of the vitamin on a daily basis — only 10 µg. One of its most essential roles is in the production of healthy red blood cells. Deﬁciency of Vi- tamin B12 in the blood is responsible for the disease called known as pernicious anemia.

There are currently six forms of Vitamin B12 that can be isolated from biological sources. Prior to the late 1960s, there were only ﬁve forms that could be isolated. The ﬁrst to be isolated and characterized was cyanocobalamin, which is commonly given the name Vitamin B12. Standard multivitamins and

Vitamin B12 supplements contain the cyanocobalamin form. Other forms that were eventually isolated and characterized are adenosylcobalamin (coenzyme

B12), methylcobalamin, aquacobalamin, hydroxo(y)cobalamin, and most recently, sulfitocobalamin. The last is now considered to be technological arti- fact of modern food preserving practices. It is often found in fish products, which are frequently treated with sulfur dioxide (SO2) as a preservative. SO2 reacts with natural hydroxo(y)cobalamin in the fish to produce the sulfitocobalamin. Although all forms can be isolated from biological sources, only the adenosylcobalamin and the methylcobalamin are biochemically active and useful.

The history of Vitamin B12 begin in the mid-1920s when sufferers of the disease pernicious anemia were treated by a regular ingestion of 1 pound of raw (not cooked) liver, a dosage that varied with the severity of the patient’s condition. In 1948, the first form of Vitamin B12 — the anti-pernicious anemia factor — was isolated and crystallized from liver and its structure was determined in 1956. Unfortunately, what was isolated and later structurally characterized is an anomaly and it was not the truly active form of Vitamin B12. The biologically active forms, adenosylcobalamin and methylcobalamin, are destroyed upon exposure to light and are converted to aquacobalamin and hydroxo(y)cobalamin. The latter two species are in turn converted to cyanocobalamin upon exposure to cyanide ion, which was one of the reagents used in the original experiment to isolate the species from liver. The active forms were later isolated from both human and microbial material and clearly shown to be AdoB12 and MeB12 but the name Vitamin B12 had already been attached to the anomalous form of cyanocobalamin. Pernicious anemia, which has been mentioned several times, is a disease that results from insufficient levels of Vitamin B12 in the blood. This deficiency can be the result of lifestyle choice (vegetarian/ vegan, alcoholism), tempo- rary physiological changes (pregnancy), and biochemical breakdowns (inability to form the coenzyme B12, lack of enzymes and related species for uptake and transport). In most instances, supplements can overcome the deficiency; however, in the case of a lack of Intrinsic Factor, the glycoprotein responsible Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 15

for the uptake of Vitamin B12 from the intestine, supplements are usually inef- fective and direct delivery of Vitamin B12 into the blood is sometimes necessary.

Symptoms of a Vitamin B12 deﬁciency are manifested in the neurological and the physical. For the former, symptoms include apathy, irritability, memory defects, dementia, emotional instability, depression, and psychosis. The latter symptoms include fatigue, indigestion, burning sensation on tongue, pale or yellowish complexion, and gastric cancer. Most of the symptoms can be reversed by treatment with Vitamin B12 but without treatment, death is the even- tual outcome of a deﬁciency of the vitamin.

As a closing note on the cobalamins, exactly why does Vitamin B12 exist and what does it do. There are many reasons and I give just a few examples.

The active forms of Vitamin B12 are responsible for the production of healthy red blood cells and several important biochemical transformation functions. Adenosylcobalamin is a catalyst for numerous biological transformations of the isomerase and mutase type, reactions in which organic molecules are converted from less useful to more useful forms by simple rearrangement of their con- stituent nuclei. Methylcobalamin is a methyl transfer agent and among its many roles is its use as a detoxiﬁcation agent by anaerobic bacteria, where a need exists for the conversion of toxic elemental mercury to water-soluble methylmercury compounds. The latter compounds are readily eliminated by the bacteria whereas the elemental mercury is not and it remains within the organism.

CLOSING COMMENTS

I thank you for your time in listening to the thoughts of one scientist and educator in the State of South Dakota. I solicit your responses to my thoughts and welcome a dialogue on them. I would also like to thank Dr. Daniel L. Heglund for his valuable assistance in the preparation of this address.

Complete Senior Research Papers

presented at The 87th Annual Meeting

of the South Dakota Academy of Science

Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 19

HYDROGEOLOGY OF THE HOMESTAKE MINE

Perry H. Rahn and William M. Roggenthen Department of Geology and Geological Engineering South Dakota School of Mines and Technology Rapid City, SD 57701

ABSTRACT

Mining of gold ore at the Homestake Mine in Lead, South Dakota, ceased in December, 2001. Water normally pumped from as deep as 8,000 ft below the surface will begin to flood the mine if and when the pumps used to dewater it are turned off. We estimate the total volume of voids (drifts, shafts, and stopes) within the mine to be approximately 752 million ft3. At a constant rate of ground water infiltration estimated at 500 gpm, this void space would take approximately 21 years to fill. The dewatered zone of the Precambrian and Tertiary intrusive rocks may contain an additional void space of 35 million ft3, which would increase the time necessary for filling the mine to approximately 22 years.

INTRODUCTION

On December 14, 2001, the Homestake Mine ceased mining ore although pumping and closure activities continue to the present time. Since the discovery of the original lode in 1876, the mine produced about 10% of the nation’s gold. The mine includes an Open Cut, shown in Figure 1, but most of the mine is underground and contains a vast network of drifts, shafts, and volumes that were mined-out (Fig. 2). At 8,000 ft depth, it is one of the deepest mines in the world. The mine has about 63 levels and approximately 300 miles of horizontal drifts (Davis, 2001). An experiment to capture solar neutrinos using a 100,000 gallon chlorine detector has been operating at the 4850 ft level since 1965. Data from this experiment provided critical information about the neutrino flux through the earth, and the cessation of mining has led to proposals to expand the neutrino experiment. The expansion of the physics experiments would increase the types of neutrinos that could be detected and the efficiency of the detection (Bahcall et al., 2001). The experiments might be located as deep as the 7400 ft level, and the large volume of rock that would overlie the experiments would greatly decrease the cosmic ray background radiation and thus allow a host of particle physics experiments to be conducted. One concern, however, is the amount of water in the mine. If completely abandoned, the pumps used to dewater the mine would be turned off, and ground water would begin to flood the drifts and shafts. Construction of the proposed particle physics experiments will take considerable time and effort and will present some engineering challenges. The 20 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002)

Figure 1. Topographic map of Lead, South Dakota (from Deadwood South and Lead USGS 7.5’ topographic maps), showing location of Homestake Mine. The contour interval is 40 ft. Cross section A-A’ is shown in Figure 2. Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 21

Figure 2. Cross section of the Homestake Mine (modified from New York Times, 2001). Some drifts and shafts have been projected to this cross section. handling of the water inflow during the interim period between abandonment and the construction of the experiments will be an important factor. It is useful, therefore, to consider the scenario of complete abandonment, which would entail the shutting down of the pumps used to dewater the mine. Water presently entering the mine primarily comes from ground water sources resulting from natural precipitation and recharge from local streams. The average annual precipitation is approximately 28 in, of which approximately 8 in infiltrates and recharges the Madison Limestone (Driscoll and Carter, 2001); presumably recharge to the Precambrian rocks would be much less. Streams that could supply recharge include Whitewood Creek, Dead- wood Creek, and Grizzly Creek (Fig. 1). The main portion of the mine directly underlies Whitewood Creek but some of the shallower drifts underlie Dead- wood Creek. Whitewood Creek at Deadwood has an annual discharge of 27 cfs (Driscoll and Carter, 2001). Using data from similar Black Hills streams (Rahn and Davis, 1993), the average discharge for Deadwood Creek is approximately 2 cfs. While the mine operated, water discharged from the mine included naturally infiltrating ground water (approximately 500 gpm) and water contained in the tailings used to backfill the mined-out volumes (approximately 1,000 gpm) (Davis, 2001). The specific locations where ground water enters the mine are not known, although a likely collection and inflow point is in the open pit, which connects to the underground through a series of older workings. The combined discharge of Whitewood and Deadwood Creeks (29 cfs) greatly ex- ceeds the 500 gpm (equivalent to 1.1 cfs) now entering the mine. Therefore, a direct connection between these streams and the underground does not exist at the present time. In general, the Homestake Mine is considered a "dry" mine because, considering its vast size, relatively small amounts of ground water seep into the mine. The purpose of this paper is to examine the hydrogeology of the Homes- take Mine with particular emphasis on the rate of ground water that would fill 22 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) the mine upon abandonment. In order to determine the time required for water to fill the mine, it is necessary to determine the void space that is created by: (A) man-made openings created directly by the mining, and (B) the dewatered void space created by the cone of depression from the dewatering activities.

A. Volume of Voids Created by Mining

To determine the artiﬁcial void space in the mine, we assume that the void space is due to: (1) drifts, shafts, and (2) the mined volumes (stopes) that were back-ﬁlled with tailings:

(1) Drifts: We assume there are 300 miles of drifts and shafts, with a typical 10 ft X 10 ft cross-section. Therefore the volume = (5280 ft/mile)(300 miles) (10 ft X 10 ft) = 158 X 106 ft3.

(2) Mined volumes: The total gold production of the mine was approximately 40 million (troy) ounces (http://pubs.usgs.gov/gip/prospect1/goldgip .html). We assume the ore contained an average concentration of 0.3 ounces of gold per ton. There- fore the tonnage mined = 40 X 106 oz/0.3 oz/ton = 133 X 106 ton. In addition to the voids created by mining the ore (per se), "waste rock" was also removed. We assume the waste rock amounted to 25% of the volume of ore mined. Therefore, the total rock removed by mining activities in connection with ore removal = 125% (133 X 106 ton) = 167 X 106 ton. To convert tonnage (mass) to volume, we assume the rock density = 2.7 gm/cm3 = 168.5 lb/ft3 = 0.08425 ton/ft3. Therefore, the volume of mined out volumes (including the ore plus waste rock) = 167 X 106 ton/ 0.08425 ton/ft3 = 1,980 X 106 ft3. The mined-out volumes were back-ﬁlled with tailings (largely sand). We assume the porosity of these back-ﬁlled zones = 30%. Therefore, the void space in these zones = 0.30 (1,980 X 106 ft3) = 594 X 106 ft3. From the above, the total man-made void space at the Homestake Mine is: (1) Drifts ...... 158 X 106 ft3 (2) Mined volumes...... 594 X 106 ft3 Total ...... 752 X 106 ft3.

B. Volume of Voids Due to Dewatering

The extent of the cone of depression created by dewatering can be ap- proximated. No observation well surrounding the Homestake Mine is known which could be used to construct a cone of depression around the mine. How- ever, hydrologic information from nearby perennial streams and abandoned mines allow for a general interpretation of the cone of the depression. For example, Whitewood Creek and Deadwood Creek (Fig. l) seem to maintain their flow all year long. The Cutting Mine (near Central City on Fig. 1) is the water Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 23 supply for the town of Lead. Originally, water ran freely out of the portal of this abandoned mine, but now the water is just at the entrance (Fred Raubak, pers. comm.). In 1994 an exploratory drift was completed which extended northward from the 6800 level approximately 13,000 ft (Lalonde, pers. comm.). The drift encountered ground water reportedly under very high pressure. Based on this observation, the cone of depression had not extended into this region at the time of the construction of the drift. The cone of depression most likely is not symmetrical. The mine is developed in Precambrian metamorphic rocks with minor amounts of Tertiary in- trusives (Bachman and Caddey, 1990; Rogers, 1990). Jointing, foliation, and lithologic changes in the Precambrian rocks should influence the shape of the cone of depression. For example, joints measured from aerial photographs in Precambrian rocks show N 30º W and N 75º E trends (Lisenbee, 2000). Rahn and Johnson (2002) found that a N 20º W foliation in Precambrian metamorphic rocks near Nemo strongly influenced the direction of flow of contaminat- ed ground water. The N 20º W to N 30º W foliation extends throughout Pre- cambrian rocks in the Black Hills. As such, it would be expected that a cone of depression would be more extensive from the mine in this direction. Yet the Cutting Mine, located northwest of the mine, still contains water. It appears, therefore, that a large cone of depression from the Homestake Mine does not exist. Rather, in this low permeability rock, dewatering probably is restricted to the workings and the rock in the immediate vicinity of the mine (Fig. 2). Given the above discussion, a simplifying assumption is that only the mined area itself, the complex of shafts and drifts, has been dewatered during the 100+years history of the Homestake Mine. To calculate this volume, we assume a geometry of a truncated cone having a 15,000 ft diameter at the 8000 ft. level and extending upwards at a 45º angle to the surface. Thus the dewatered volume is 3.5 x 1012 ft3. The porosity of this dewatered zone can be only estimated. The mine is mostly within Precambrian metamorphic rocks of the Homestake Formation, a grunerite schist. Some fractures exist, and in the dewatered zone the gravitational water can drain out, leaving a void space. Fracture porosity and permeability in metamorphic rock decreases exponentially with depth (Davis and DeWeist, 1966). We judge that the average porosity of all the dewatered rocks would be less than 0.01 %. Of this, we estimate that only 10% of the void space would contain gravitational water that actually drained out due to dewatering, and the remaining 90% of the water would be retained due to capillary forces. Therefore, the volume of water removed is approximately 35 X 106 ft3. If the mine were to fill, water would resaturate these voids. It should be noted that if gravitational water occupied 50 % of the pore space, then the volume of water removed by dewatering would be increased by a factor of five. From the above, the total voids created by the mine workings and the dewatering is:

Mine workings...... 752 X 106 ft3 Cone of depression ...... 35 X 106 ft3 Total ...... 787 X 106 ft3. 24 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002)

Time Required to Fill the Mine

To determine the length of time required to fill the voids, we assume a constant natural ground water infiltration rate into the Homestake Mine of 500 gpm. Assuming a constant rate of fill, the length of time required to fill the Homestake Mine = 787 X 106 ft3/66.84 ft3/min = 11.7 X 106 min = 8176 days = 22.4 years. The rate that ground water fills any mine would not be constant with time. An analogy can be made to a pumped well. When the pump is turned on, the water level in the well drops quickly at first, but then gradually slows down until a nearly constant water level is obtained. When the well is turned off, the water level in the well quickly rises, but slows down as it gradually obtained the water level before the pump was turned on. The recovery curve is a mirror image of the drawdown curve (Driscoll, 1986). The drawdown and recovery applies to the well being pumped and the area immediately surrounding the well. For the Homestake Mine, it is not feasible to apply well pumping analogy using aquifer constants (transmissivity and storativity) to create a theoretical cone of depression and the mirror image curve for recovery Precambrian rocks in the Black Hills. Transmissivity can be estimated from specific capacity measurements. Galloway and Strobel (2000) measured the specific capacity for 71 shallow Precambrian wells, and they found an average of 51 gpm/ft of drawdown. The Precambrian rocks generally have low permeability (Rahn and Gries, 1973), but transmissivity values representative of the 8000 ft section are not available. Nevertheless, from typical well drawdown and recovery behavior, one would expect a gradual slowing of the rate of filling. Therefore we believe the 22-year figure obtained above using a constant 500 gpm recharge rate is a minimum time to fill.

CONCLUSION

If pumping of water ceased at the Homestake Mine, we believe it will take more than 22 years to flood the abandoned workings. Our calculations are based on limited data concerning the porosity of the dewatered zone, as well as unconfirmed data concerning the size of the mine and the ground water recharge rate. Additional work would be useful to more fully delineate the variation of porosity and permeability of the metamorphic rocks as a function of depth. As the water rises to its original (pre-mining) elevation, discharge from the workings would be expected to flow into Whitewood and Deadwood Creeks. The chemistry and the discharge rate from these workings in the future are not known. However, if any adverse environmental impacts from this water were to occur, they might be due to acidity and dissolved metals because the ore contains arsenopyrite and other sulfides, and it is known that the unregulated disposal of tailings in Whitewood Creek resulted in arsenic contamination of surface and ground water (Goddard, 1989; Rahn et al., 1996). Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 25

REFERENCES CITED

Bachman, R.L., and S.W. Caddey. 1990. The Homestake Mine, Lead, South Dakota: an overview. in Paterson, C.J., and A.L. Lisenbee, eds., Metalloge- ny of gold in the Black Hills, South Dakota: Society of Economic Geologists, Guidebook Series, 89-94. Bahcall, J., Barish, B., Calaprice, F., Conrad, J., Doe, P.J., Gaisser, T., Haxton, W., Lesko, K.T., Marshak, M., and Robinson, K. 2001. Underground Science (Bahcall Report), http://www.sns.ias.edu/~jnb/Laboratory/science.pdf Davis, R., 2001. Underground lab to mine secrets of the universe, Engineering Times. August/September 2001, 12-13. Davis, S.N., and R.J.M. DeWeist. 1966. Hydrogeology. John Wiley & Sons, New York. Driscoll, D.G., and J.M. Carter. 2001. Hydrologic conditions and budgets for the Black Hills of South Dakota, through water year 1998. U. S. Geological Sur- vey, Water-Resources Investigations Report 01-4226. Driscoll, F.G. ed. 1986. Groundwater and wells. Johnson Division, St. Paul, MN 55112. Galloway, J. M., and M.L. Strobel. 2000. Hydrologic characteristics of metamorphic rocks in the central core of the Black Hills South Dakota. in Stro- bel, M. L., and others, Hydrology of the Black Hills, South Dakota School of Mines and Technology, Bulletin No. 20, p. 87. Goddard, K. E., 1989. Composition, distribution, and hydrologic effects of con- taminated sediments resulting from the discharge of gold milling waste in- to Whitewood Creek at Lead and Deadwood, South Dakota. U.S. Geologi- cal Survey Water-Resources Investigations Report 87-4051, 76 p. Lisenbee, A.L. 2000. Structural anisotropy of the Laramide Black Hills region. in Strobel, M. L. and others, Hydrology of the Black Hills. South Dakota School of Mines and Technology, Bulletin No. 20, 88-97. New York Times. 2001. "Mix of Physics and Politics May Produce Lab in Mine", November 24, 2001. Rahn, P.H., and J.P. Gries. 1973. Large springs in the Black Hills, South Dako- ta and Wyoming. South Dakota Geological Survey, Report of Investigations No. 107, 46 p. Rahn, P.H., and A.D. Davis. 1993. Stream runoff from Black Hills watersheds. Proceedings, South Dakota Academy of Science, 72: 161-175. Rahn, P.H., A.D. Davis, and C.J. Webb and A.D. Nichols. 1996. Water quality impacts from mining in the Black Hills, South Dakota. Environmental Ge- ology, 27: 38-53. Rahn, P.H., and C.S. Johnson. 2002. Effects of anisotropic transmissivity on a contaminant plume at Nemo, South Dakota: Environmental and Engineer- ing Geoscience, 8:, 11-18. Rogers, H. 1990. Homestake Mine underground tour. in: Paterson and A.L. Lisenbee, eds., Metallogeny of gold in the Black Hills, Society of Economic Geologists, Guidebook Series, 7: 127.

Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 27

MYCORRHIZAL COLONIZATION AS IMPACTED BY CORN HYBRID

Marie-Laure A. Sauer, Diane H. Rickerl and Patricia K. Wieland South Dakota State University Brookings, SD 57007

Courtenay Hoernemann USDA

W.B. Gordon Kansas State University Irrigation Experiment Station Parsons, KS, 66939

ABSTRACT

Colonization of crop roots by mycorrhizal fungi can improve phosphorus uptake and thus reduce needs for phosphorus fertilizer application. Reduc- tions in fertilizer application decrease environmental risks from run-off and decrease farmers’ input costs. This study was conducted in 1996 and 1997 at the Kansas State University Irrigation Experiment Station near Bellville, Kansas to determine the effects of corn (Zea mays, L.) hybrid on mycorrhizal colonization. Twelve hybrids were selected for the study. A phosphorus starter fertilizer treatment was imposed as a split plot and the study was arranged in a randomized complete block design. Root samples were collected when corn reached the 6-leaf stage. Samples were washed, cut in to 1 cm lengths, cleared, and stained. The procedures of Giovannetti and Mosse were used to assess the percentage of root length colonized by hyphae, arbuscules, and total mycorrhizae. No differences, due to fertilizer or hybrid, in percent hyphae or percent arbuscules were found. Hyphae ranged from 60-90% and arbuscules ranged from 65-98%. The range in total mycorrhizal colonization (13-98%) suggested that hybrids varied in mycorrhizal colonization, but the variability was too high within each hybrid to result in statistically signiﬁcant differences between the hybrids.

Keywords

Mycorrhizae, corn hybrid, P fertilizer. 28 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002)

INTRODUCTION

Vesicular-arbuscular mycorrhizal (VAM) fungi are found in all kinds of habitat and have been shown to have a beneficial effect on crop growth, especially in low fertility soils (Black and Tinker, 1979). Soil disturbances (tillage) have been shown to decrease mycorrhizal colonization by reducing the in- noculum potential (mycelia and spores) in the soil (Anderson et al., 1987; Jasper et al., 1989; and Evans and Miller, 1988). Crop species also influences the species composition of mycorrhizae in the soil (Johnson et al., 1991). Public interest in alternative agricultural systems is growing. Practices are designed to protect agricultural resources and prevent environmental damage to the farm and off-site land, water, and air (Poincelot, 1990). Reduced tillage systems coupled with more efficient phosphorus fertilization can protect surface water from the effects of phosphorus run-off. Rapid mycorrhizal colonization of crop species, could reduce the need for phosphorus fertilization in early spring, when the risk of run-off is highest.

Objectives

The objectives of this study were: 1. To determine the effects of phosphorus starter fertilizer on phosphorus uptake and mycorrhizal colonization of corn (Zea mays, L.) hybrids; and 2. To determine if corn hybrids differed in mycorrhizal colonization.

METHODS

A ﬁeld study was conducted at the Kansas State University, Irrigation Ex- periment Field near Bellville, Kansas in 1996 and 1997. Twelve corn (Zea mays, L.) hybrids were planted as whole plots with phosphorus starter fertilizer as the split plot. The test was arranged as a randomized complete block with three replications. The twelve hybrids planted were Pioneer 3563, 3489, 3346, 3394; Cargill 6327, 7777; Dekalb 591, 626, 646; Northrup King 7333,6330; and ICI 8599. Split-plot fertilizer treatments were 0 fertilizer and 15 kg ha-1. Corn samples were collected at the V6 growth stage. Twenty corn plants per plot were randomly selected and carefully removed from the soil. Roots were washed, separated from top-growth at the soil line, and frozen until VAM analyses could be performed. To evaluate mycorrhizal infection, terminal feed- er roots were cut into 1 cm segments and 0.5 g of randomly selected segments were placed in plastic capsules, cleared in 10% KOH overnight and stained with trypan blue in lactoglycerin (Phillips and Hayman, 1970). The grid-inter- sect method of Giovannetti and Mosse (1980) was used to assess root colonization. Percentage root colonization by arbuscules, hyphae, and total (arbus- cule and/or hyphae) was calculated as:

Percentage root colonization = (VAM roots/total root intersects) X 100. Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 29

To assess the phosphorus content in the plant, plant top tissue samples were dried and ground to pass a 0.10 cm screen. A 0.25 g sub-sample of ground material was digested with sulfuric acid and hydrogen peroxide. phosphorus concentrations were measured with a dual-channel Technicon AutoAn- alyzer. Effects of starter fertilizer and hybrid on VAM colonization and phosphorus concentrations were compared using SAS Proc Mixed (Littell et al., 1996).

RESULTS

Plant phosphorus concentrations were significantly different due to starter fertilizer application, but not corn hybrid. There were no significant interactions between starter fertilizer and corn hybrid. In 1996 the mean P concentration was 0.30% in non-fertilized plots compared to 0.35% in fertilized plots (Fig. 1). In 1997 concentrations were 0.32% in non-fertilized and 0.35% in fertilized plots (Fig. 2). The range in P concentration by hybrid was 0.29 – 0.37% (Fig. 3) in 1996 and 0.32-0.36% (Fig. 4) in 1997. Pioneer 3489 consistently had the highest P concentration while Cargill 7777 was one of the lowest during both years of the study. The response to P fertilization and the range in concentrations are typical of early plant growth. No differences in mycorrhizal colonization, due to starter fertilizer or corn hybrid were found in either year. There were no significant interactions between starter fertilizer and corn hybrid. The percent colonization was slightly higher in non-fertilized plots than fertilized plots (Figs. 5 and 6), which is consistent with the literature (Arias et al., 1991). In 1996, the range in percent col-

Figure 1. Plant phosphorus concentration as inﬂuenced by starter fertilizer in 1996, Pr > F = 0.0001. 30 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002)

Figure 2. Plant phosphorus concentration as inﬂuenced by starter fertilizer in 1997, Pr > F = 0.0001.

Figure 3. Plant phosphorus concentration as inﬂuenced by corn hybrid in 1996, Pr > F = 0.0429. Corn hybrids: C = Cargill, D = Dekalb, I = ICI, N = NK, P = Pioneer. Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 31

Figure 4. Plant phosphorus concentration as inﬂuenced by corn hybrid in 1997, Pr > F = 0.6497. Corn hybrids: C = Cargill, D = Dekalb, I = ICI, N = NK, P = Pioneer.

Figure 5. Percentage of mycorrhizal colonization as inﬂuenced by starter fertilizer in 1996, Pr > F = 0.0833. 32 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002)

Figure 6. Percentage of mycorrhizal colonization as inﬂuenced by starter fertilizer in 1997, Pr > F = 0.9181.

Figure 7. Percentage of mycorrhizal colonization as inﬂuenced by corn hybrid in 1996, Pr > F = 0.4456. Corn hybrids: C = Cargill, D = Dekalb, I = ICI, N = NK, P = Pioneer. Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 33

Figure 8. Percentage of mycorrhizal colonization as inﬂuenced by corn hybrid in 1997, Pr > F = 0.5520. Corn hybrids: C = Cargill, D = Dekalb, I = ICI, N = NK, P = Pioneer. onization among hybrids was 57-85% and in 1997 it was 77-89%. The variability in colonization within hybrids was very high and probably prevented statistical signiﬁcance.

DISCUSSION

Several factors may have contributed to the lack of difference in mycorrhizal colonization among corn hybrids. The most obvious is the highly variable nature of colonization. If the study were to be repeated, it might be better to sample at a younger growth stage to detect early differences. The level of colonization in our samples from the V6 growth stage was quite high among all hybrids. Finding a site with low soil P concentrations would also be useful since high soil fertility can reduce the inﬂuence of mycorrhizae. The concept of using hybrid selection to reduce fertilizer needs may become more important as environmental concerns grow. Another approach would be to manipulate genes for improved mycorrhizal colonization and/or phosphorus uptake.

LITERATURE CITED

Anderson, E.L., P.D. Millner, and H.M. Kunishi. 1987. Maize root length density and mycorrhizal infection as inﬂuenced by tillage and soil phosphorus. J. Plant Nutr. 10:1349-1356. 34 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002)

Arias, I., I. Koomen, J.C. Dodd, R.P. White and D.S. Hayman. 1991. Growth responses of mycorrhizal and non-mycorrhizal tropical forage species to different levels of soil phosphate. Plant Soil.132:253-260. Black, R., and P.B. Tinker.1979. The development of endomycorrhizal root systems. II. Effect of agronomic factors and soil conditions on the development of vesicular-arbuscular mycorrhizal infection in barley and on endo- phyte spore density. New Phytol. 83:401-413. Evans, D. G., and M.H. Miller. 1988. Vesicular-arbuscular mychorrhizae and the soil-disturbance-induced reduction of nutrient absorption in maize. I. Causal relations. New Phytol. 110:67-74. Giovannetti, M., and B. Mosse. 1980. An evaluation of techniques for measuring vesicular arbuscular mycorrhizal infection in roots. New Phytol. 84:489- 500. Jasper, D.A., L.K. Abbott, and A. D. Robson. 1989. Hyphae of a vesicular-arbuscular mycorrhizal fungus maintain infectivity in dry soil, except when soil is disturbed. New Phytol. 112:101-107. Johnson, N.C., F.L. Pﬂeger, R.K. Crookston, S.R. Simmons and P.J. Copeland. 1991. Vesicular-arbuscular mycorrhizas respond to corn and soybean crop- ping history. New Phytol. 117:657-663. Littell, R.C., G.A. Miliken, W.W. Stroup, R.D. Wolﬁnger. 1996. SAS system for mixed models. Pp 53-63. Phillips, J.M., and D.S. Hayman. 1970. Improved procedures for clearing roots and staining parasitic and vesicular-arbuscular mycorrhizal fungi for rapid assessment of infection. Trans. Br. Mycol. Soc. 55:158-161. Poincelot, R. 1990. From the Editor. J. Sust. Agric. 1:1-3. Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 35

RETENTION AND SURVIVAL RATES ASSOCIATED WITH THE USE OF T-BAR ANCHOR TAGS IN MARKING YELLOW PERCH

George D. Scholten, Daniel A. Isermann and David W. Willis Department of Wildlife and Fisheries Sciences South Dakota State University Brookings, SD 57007

ABSTRACT

We assessed retention and short-term mortality rates associated with tagging yellow perch (Perca ﬂavescens) with t-bar anchor tags (FD-94 Floy® tags) by conducting 2- to 4-d net-pen trials. Yellow perch were captured from two South Dakota lakes with trap nets during October 2000 and 2001, when water temperatures ranged from 9º to 13º C. Retention and mortality rates for yellow perch (total length range: 151 - 266 mm) tagged from East 81 Slough during October of 2000 (n=19) were 100% and 5%, respectively. Retention and mortality rates for yellow perch (total length range: 265 - 325 mm) tagged from Pel- ican Lake during October of 2001 (n=22) were 95% and 0%, respectively. Based on average results, anchor tag retention (mean=97.7%) was high and tagging mortality (mean=2.6%) was low for yellow perch collected and tagged using this sampling approach. Yellow perch tagged in Pelican Lake were signiﬁcant- ly larger than yellow perch tagged in East 81 Slough (t = 13.929, df = 19, P < 0.001), but neither size-dependent tag retention (χ2 = 0.885, df = 1, P = 0.347), nor size-dependent tagging mortality (χ2 = 1.187, df = 1, P = 0.276) was observed.

INTRODUCTION

Since introduction in 1968 (Dell 1968), internal t-bar anchor tags have been extensively used to mark mass numbers of ﬁsh that need to be individually distinguishable upon recapture (Muoneke 1992; Pierce and Tomcko 1993; Pegg et al. 1996). To obtain unbiased data, tag retention must be accounted for in biological statistics derived from tagging studies. Several studies have shown t- bar anchor tag loss to be negligible (Cucin and Regier 1965; Curtis 1970; Sto- bo 1972; Bulak 1983; Franzin and McFarlane 1987; Pegg et al. 1996), while other studies reported relatively high rates of tag loss (Keller 1971; Rawstrom 1973; Wilbur and Duchrow 1973; Ebener and Copes 1982; Tranquilli and Childers 1982; Muoneke 1992). Mortality associated with tagging process must also be evaluated prior to drawing conclusions based on tagging study results (Bulak 1983; Curtis 1970; Stobo 1972; Tranquilli and Childers 1982). While proper tagging techniques can minimize mortality, inexperience or carelessness could cause increased mortality rates associated with the tagging process. 36 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002)

Severe underestimations of biological statistics for a population can result when tag loss and mortality associated with tagging are not measured (Dryer 1964; Ebener and Copes 1982). Few studies have assessed tag retention and tagging mortality in yellow perch (Perca ﬂavescens) (Stobo 1972). As part of a study designed to estimate yellow perch exploitation rates in eastern South Dakota glacial lakes, we determined initial tag retention and mortality rates associated with FD-94 Floy® t-bar anchor tags.

METHODS

Yellow perch were collected in overnight trap net sets when water temperatures ranged from 9º to 13º C. Yellow perch from East 81 Slough (Brook- ings County) were tagged during October 2000; perch tagged in Pelican Lake (Codington County) were collected during October 2001. Fish were removed from nets and placed in a holding tank supplied with fresh water prior to tagging. All tagged yellow perch were measured to the nearest mm (total length; TL). Tags (Floy® FD-94; 45 mm long with 10-mm anchor) were inserted at a 45° angle with a tagging gun (Avery-Dennison® Mark III) just under the dorsal fin. The tagging gun needle penetrated high on the body and was twisted 90° to allow the t-bar to become locked behind the interneural bones. Sub-samples of tagged yellow perch were held in floating net pens (1 m x 1 m x 10 m) for 2 - 4 d. Tag retention rates (% of fish retaining tags) and mortality rates (% of fish dead) were estimated for each trial.

RESULTS

Retention and mortality rates of yellow perch (TL range: 151 - 266 mm) tagged from East 81 Slough during October 2000 (n=19) were 100% and 5%, respectively. Retention and mortality rates of yellow perch (TL: 265 - 325 mm) tagged from Pelican Lake during October 2001 (n=22) were 95% and 0%, respectively. Yellow perch tagged in Pelican Lake were signiﬁcantly longer than yellow perch tagged in East 81 Slough (t = 13.929, df = 19, P < 0.001), but neither tag retention (χ2 = 0.885, df = 1, P = 0.347), nor tagging mortality (χ2 = 1.187, df = 1, P = 0.276) were signiﬁcantly different in the two trials.

DISCUSSION

Although Rawstrom (1973) observed high initial tag loss, our study supported numerous other studies that recorded high tag retention (Cucin and Regier 1965; Curtis 1970; Stobo 1972; Bulak 1983; Franzin and McFarlane 1987; Pegg et al. 1996). To achieve high tag retention it is important to lock the anchor bar of the tag behind the interneural bones. Tags can be lost if they are improperly inserted, if they rotate so the prongs are no longer perpendicular to the interneural spines, or the width of the tag is smaller than the distance between the interneural spines (Waldman et al. 1990). Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 37

The only tag shed in this study was from a 273-mm (TL) yellow perch from Pelican Lake. If the interneural spacing was larger than the width of the anchor (10 mm), the tag may have worked itself out; however, tags were retained in all perch larger than 273 mm (N = 20). Size-dependent t-bar anchor tag loss was seen in striped bass over 592 mm using t-bar anchor tags with a prong width of 10 mm (Waldman et al. 1990). We observed no size-dependent tag loss in yellow perch. Mortality associated with tagging is expected in all tagging studies but in our study tagging mortality was low, supporting previous work (Bulak 1983; Curtis 1970; Stobo 1972; Tranquilli and Childers 1982; Pierce and Tomcko 1993). We observed no evidence of size- dependent tagging mortality in yellow perch. Most tagging mortality is associated with handling during the tagging process, so if ﬁsh are tagged quickly and handled properly, mortality should be minimized. We found initial FD-94 ﬂoy t-bar anchor tag retention to be high in yellow perch. Yellow perch survival rates associated with using these tags was low. Al- though tag loss and mortality estimates were low, they must be accounted for in tagging studies designed to assess exploitation rates.

ACKNOWLEDGEMENTS

Partial funding for this project was provided by the South Dakota Depart- ment of Game, Fish, and Parks through Federal Aid in Sport Fish Restoration Project F-15-R, Study 1584. This manuscript was approved for publication by the South Dakota Agricultural Experiment Station as Journal Series No. 3321.

LITERATURE CITED

Bulak, J. S. 1983. Evaluation of floy anchor tags for short term mark-recapture studies with blueback herring. North American Journal of Fisheries Management 3:91-94. Cucin, D., and H. A. Regier. 1965. Effects of the floy anchor tag on growth and survival of brook trout (Salvelinus fontinalis). Journal of Fisheries Re- search Board of Canada 23:221-274. Curtis, T. A. 1970. Anadromous Fish Survey of the Santee and Cooper Rivers. Job Progress Report AFS 3-1. South Carolina Wildlife and Marine Resource Department, Columbia, South Carolina, USA. Dell, M. B. 1968. A new fish tag and rapid cartridge-fed applicator. Transac- tions of the American Fisheries Society 97:57-59. Dryer, W. R. 1964. Movements, growth and rate of recapture of whitefish tagged in the Apostle Islands area of Lake Superior. U.S. Fish and Wildlife Service Fishery Bulletin 63:611-618. Ebener, M. P., and F. A. Copes. 1982. Loss of floy anchor tags from lake whitefish. North American Journal of Fisheries Management 2:90-93 38 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002)

Franzin, W. G., and G. A. McFarlane. 1987. Comparison of floy anchor tags and fingerling tags for tagging white suckers. North American Journal of Fisheries Management 7:307-309. Keller, W. T. 1971. Floy tag retention by small brook trout. New York Fish and Game Journal 18:142-143. Muoneke, M. I. 1992. Loss of floy anchor tags from white bass. North Amer- ican Journal of Fisheries Management 12:819-824. Pegg, M. A., J. B. Layzer, and P. W. Bettoli. 1996. Angler exploitation of anchor-tagged saugers in the lower Tennessee River. North American Jour- nal of Fisheries Management 16:218-222. Pierce, R. B., and C. M. Tomcko. 1993. Tag loss and handling mortality for northern pike marked with plastic anchor tags. North American Journal of Fisheries Management 13:613-615. Rawstrom, R. R. 1973. Comparisons of disk dangler, trailer and internal anchor tags on three species of salmonids. California Fish and Game 59:266- 280. Stobo, W. T. 1972. The effects of dart tags on yellow perch. Transactions of the American Fisheries Society 101:365-366. Tranquilli, J. A., and W. F. Childers. 1982. Growth and survival of largemouth bass tagged with floy anchor tags. North American Journal of Fisheries Management 2:184-187. Waldman, J. R., D. J. Dunning, and M. T. Mattson. 1990. A morphological explanation for size dependent anchor tag loss from striped bass. Transac- tions of the American Fisheries Society 119:920-923. Wilbur, R. L., and R. M. Duchrow. 1973. Differential retention of five floy tags on largemouth bass (Micropterus salmoides) in hatchery ponds. Proceed- ings of the Annual Conference Southeastern Association of Game and Fish Commissioners 26(1972):407-413. Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 39

IMPACT OF CROP HARVEST ON SMALL MAMMAL POPULATIONS IN BROOKINGS COUNTY, SOUTH DAKOTA

Melissa K. Pinkert, Jonathan R. Meerbeek, George D. Scholten, and Jonathan A. Jenks Department of Wildlife and Fisheries Sciences South Dakota State University Brookings, SD 57007

ABSTRACT

In the Midwest, agricultural cropland provides the majority of habitat available to small mammals. In some regions of the Midwest, cropland comprises as much as 70% of the landscape. Importance of these agricultural habitats to small mammals pre- and post-harvest is not well documented. The distribution of small mammals was studied in shelterbelt, grassland, and adjacent cropland habitats pre- and post-harvest in Brookings County, South Dakota from 20 September (before crop harvest) to 15 November 2001 (after crop harvest). Deer mice (Peromyscus maniculatus) (n=30) and white-footed mice (Per- omyscus leucopus) (n= 29) were captured in relatively high numbers when compared to the other species: short-tailed shrews (Blarina brevicauda) (n=13), masked shrews (Sorex cinereus) (n=7), prairie voles (Microtus ochrogaster) (n=4), meadow voles (Microtus pennsylvanicus) (n=3), and northern grasshopper mice (Onychomys leucogaster) (n=2). Deer mice and white-footed mice were the only inhabitants of cropland. When compared to cropland, proportions of deer mice using the grassland and white-footed mice using the shelterbelt decreased after harvest. Competition among small mammals and predation risks from the short-tail shrew in grasslands and shelterbelts may force deer mice and white-footed mice into habitats such as cropland. Alter- natively, the abundance of waste grain after harvest may explain the increased use of cropland. The prairie vole demonstrated a strong use of grassland, while the white-footed mouse used shelterbelt habitats. Cropland habitats adjacent to shelterbelts and grasslands likely provide a relatively stable food source for small mammal populations.

Keywords

Crop harvest, deer mice, Peromyscus maniculatus, small mammals, South Dakota.

INTRODUCTION

The majority of habitat available to small mammals in the Midwest is cropland. Opportunistic feeders that do not rely on herbaceous ground cover, such as deer mice (Peromyscus maniculatus) are more likely to inhabit cultivated 40 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) cropland (Stallman and Best 1996). Manson and Stiles (1998) suggested that old cropland fields experience intense seed predation by white-footed mice (Peromyscus leucopus). Others species, such as house mice (Mus musculus), move into cultivated fields as cover becomes available (Stallman and Best 1996). Small mammals have been studied in wooded areas located along rivers and in shelterbelts in eastern South Dakota (Barnes and Linder 1982; Hallman and Butler 1992). Others have examined species diversity and abundance of small mammals utilizing cultivated fields in the Midwest, but few have studied the effects of crop harvest on habitat use (Stallman and Best 1996; Walker 1976). The purpose of this study was to determine the distribution of small mammals in habitats adjacent to and occupying pre- and post-harvested cropland. Study objectives were (1) to determine the small mammal abundance and species composition of a shelterbelt with an adjacent cornfield as well as grassland with an adjacent cornfield, (2) to determine small mammal habitat use and (3) determine if small mammals alter habitat use after crop fields are harvested.

STUDY AREA

To determine the impact crop harvest has on habitat use, a shelterbelt and a small grassland with adjacent cornfields were sampled. Both study areas were located in Brookings County, South Dakota. The shelterbelt/cornfield habitat was located at T 110 N, R 48 W, Section 26. The shelterbelt was about 2.3 ha in size and was composed of approximately 90% green ash (Fraxinus pennsylvanicus). The remaining 10% was made up of red cedars (Juniperus virginiana), cottonwood (Populus spp.), and tatarian honeysuckle (Lonicera tatarica). The understory was a mixture of brome grass (Bromus spp.) and green ash seedlings. The adjacent cornfield was 14.2 ha. The grassland/cornfield habitat was located at T 109 N, R 48 W, Section 2. The grassland was 28.3 ha of primarily switch grass (Panicum virgatum). The adjacent cornfield was 36.4 ha in size.

METHODS

Sampling periods began on 20 September 2001 (before crop harvest) and 15 November 2001 (after crop harvest). Seven x seven trap grids were randomly placed within habitat types on study areas. We assumed that each trap sampled a radius of 4.55-m; hence, traps were placed with a 9.09-m distance between each trap. Our approach was designed to effectively sample 0.4047 hectares (1 acre). Based on the recommendations of Stickel (1946), we used mouse-sized snap traps (Victor®, Woodstream Corporation, Lititz, PA) baited with a combination of rolled oats and peanut butter (Beer 1964). To minimize the possibility of catching drifting animals, traps were set for three consecutive nights for before and after crop harvest capture periods. In both sampling periods traps were set no more than two hours before dusk, Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 41 checked, and re-baited on the following two days, and checked and removed on the third day. For each captured animal the trap number was recorded and species was determined in the laboratory. Statistical analyses were conducted using an alpha level of 0.05. Chi- square tests (Bishop 1966) were used to test the hypothesis that small mammal use was equivalent among habitat types. For this portion of the analysis, small mammals captured in the cornﬁeld were classiﬁed based on adjacent habitat (grassland or shelterbelt). Relative abundance was calculated as the total number of individuals caught per 1000 trap nights: Total captures in habitat type Relative Abundance = X 1000 Total trap nights in habitat type

Change in proportion of captured individuals before and after crop harvest also was evaluated using chi-square analyses (SAS Institute 1996).

RESULTS

Eighty-eight small mammals were captured in 1176 trap nights. Deer mice (n=30) and white-footed mice (n=29) were the most frequently captured (Table 1). Other species captured included short-tailed shrews (Blarina brevicauda) (n=13), masked shrews (Sorex cinereus) (n=7), prairie voles (Microtus ochrogaster) (n=4), meadow voles (Microtus pennsylvanicus) (n=3), and northern grasshopper mice (Onychomys leucogaster) (n=2). With the exception of one grasshopper mouse, deer mice and white-footed mice were the only species captured in cornfields. Habitat use was evident with white-footed mice and prairie voles (Table 1). White-footed mice were captured more often in shelterbelt than other habitats, collective shrews were captured more often in shelter and grassland habitats than in cornfields, and prairie voles were only captured in grassland habitat (Table 1). Relative abundance of deer mice, masked shrews, prairie voles, and meadow voles was generally higher in grassland/cornfield habitat whereas relative abundance of white-footed mice, short-tailed shrews, and grasshopper mice was generally higher in shelterbelt/cornfield habitat (Table 2). There was no significant difference in the relative abundance of small mammals collected pre vs. post harvest in the grassland (χ2 < 0.001, df = 1, P = 0.939). However, relative abundance of small mammals collected in the shelterbelt was lower after harvest (χ2 = 6.377, df = 1, P = 0.012). In cornfields, deer mice and white-footed mice were the only species collected in sufficient numbers to test our hypothesis. Relative abundance of deer mice collected in the grassland was lower after harvest (χ2 = 11.397, df = 1, P = 0.001), but there was no significant difference in the proportion of deer mice collected in the shelterbelt after harvest (χ2 < 0.001, df = 1, P > 0.95). No white-footed mice were collected in the grassland; thus, comparisons could not be made. The proportion of white-footed mice collected in the shelterbelt was lower after harvest (χ2 = 4.202, df = 1, P = 0.040). 42 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002)

Table 1. Small mammals captured in shelterbelt/cornﬁeld and grassland/cornﬁeld habitats sampled with snap traps within Brookings County, South Dakota during the September–November 2001.

SHELTERBELT/CORNFIELD HABITAT

Pre-Harvest Post-Harvest Species Shelterbelt Cornﬁeld Shelterbelt Cornﬁeld Total

Deer mice (Peromyscus maniculatus) 424212 White footed mice (Peromyscus leucopus) 12 2 5 2 21 Grasshopper mice (Onychomys leucogaster) 00112 Short tailed shrew (Blarina brevicauda) 50409 Masked shrew (Sorex cinereus) 20002

Total 23 4 14 5 46

GRASSLAND/CORNFIELD HABITAT

Pre-Harvest Post-Harvest Species Grassland Cornﬁeld Grassland Cornﬁeld Total

Deer mice (Peromyscus maniculatus) 461718 White footed mice (Peromyscus leucopus) 03058 Short tailed shrew (Blarina brevicauda) 10304 Masked shrew (Sorex cinereus) 40105 Prairie vole (Microtus ochrogaster) 00404 Meadow vole (Microtus pennsylvanicus) 20103

Total 11 9 10 12 42

Table 2. Relative abundance (captures/1000 trap nights) of small mammals sampled with snap traps in two habitats adjacent to cornﬁelds within Brookings County, South Dakota during the fall of 2001.

Grassland/Cornﬁeld Shelterbelt/Cornﬁeld Species Relative Abundance Relative Abundance

Deer mouse (Peromyscus maniculatus) 30.61 20.41 White-footed mouse (Peromyscus leucopus) 13.61 35.71 Short-tailed shrew (Blarina brevicauda) 6.80 15.31 Masked Shrew (Sorex cinereus) 8.50 3.40 Prairie Vole (Microtus ochrogaster) 6.80 0.00 Meadow Vole (Microtus pennsylvanicus) 5.10 0.00 Grasshopper mouse (Onychomys leucogaster) 0.00 3.40 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 43

DISCUSSION

Deer mice and white-footed mice occurred most frequently in adjacent cornfields, indicating that this habitat is important to small mammals in eastern South Dakota. Species such as the deer mouse and white-footed mouse are adapted for such habitats (Stallman and Best 1996, Sietman et al. 1994). Stall- man and Best (1996) suggested that deer mice are the only permanent resident in cropland habitats because they can exist in areas without herbaceous ground cover by using extensive burrows, which reduces risk of predation. Deer mice used cropland habitats when adjacent to both the grassland and shelterbelt habitats. Deer mice in the grassland/cornfield habitat may have been affected by crop harvest because relative abundance in the grassland was lower after harvest. Deer mice in the shelterbelt/cornfield habitat were not significantly affected by crop harvest. Although the association of the white-footed mouse with woody vegetation is well documented (Sietman et al. 1994, Burt and Grossenheider 1964), the species also can occur in grasslands (Manson and Stiles 1998). White-footed mice exhibited a strong association for the shelterbelt habitat, but also occurred in cornfields. Corn stubble likely functioned structurally to allow co-oc- curance of white-footed mice with other small mammals. White-footed mice were not captured in switch grass habitat, but occurred frequently in the adjacent cornfield (before and after harvest). This suggests that white-footed mice avoid habitats with a large amount of herbaceous ground cover. Proportion of white-footed mice in the shelterbelt decreased after harvest, suggesting attrac- tion to post-harvest cropland. This may be the result of increased waste grain and/or competition, either through active exclusion or resource exploitation in the shelterbelt habitat (Manson and Stiles 1998). Shrews occurred in shelterbelt and grassland habitats, but were not captured in the adjacent cornfields. Romansky (1970) found that short-tailed shrews had an affinity for wooded areas. However, Burt and Grossenheider (1964) stated that short-tail shrews were not restricted to a particular habitat. Short-tail shrews can be an important predator on small animals, and can directly influence deer mouse populations (Eadie 1944). Masked shrews prefer moist habitats such as forests, open country, riparian, and upland habitats (Spencer and Pettus 1966, Yahner 1992, Walker 1976, Romanaky 1970). Hoff et al. (1989) captured masked shrews in marsh and grassland habitats but not in cornfield habitat. Shrew population estimates may be underestimated because snap traps are not as effective in catching these species as are live or pitfall traps (Fowle and Edwards 1954, Mengak and Guynn 1987, Briese and Smith 1974, Wiener and Smith 1972). If populations were underestimated, a risk of predation for deer and other mice could have affected distributions of small mammals. Voles (prairie and meadow) were associated with the grassland habitat. Prairie voles generally occur in open prairies with dense vegetation and a well- developed litter layer (Walker 1976, Heideman et al. 1983, Sietman et al. 1994). Meadow voles feed primarily on plant shoots, roots, and some monocot seeds (Manson and Stiles 1998) in associated grasslands (Grant 1971, Manson and 44 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002)

Stiles 1998, Morris 1969, Romansky 1970, Walker 1976). Few voles were captured in our study and no voles were captured in cornfield habitats. Deer mice and white-footed mice frequently used cornfields adjacent to shelterbelt and grassland habitats before and after harvest, suggesting they were permanent inhabitants of cultivated fields. Cropland habitats adjacent to shelterbelts and grasslands likely provide a relatively stable food source for small mammal populations. Because small mammals are an important food source for avian and mammalian predators, availability of small mammal populations likely affect predation rates on other species.

LITERATURE CITED

Barnes, T. G., and R. L. Linder. 1982. Small mammal occurrence in South Dakota shelterbelts and movements of Peromyscus maniculatus. Pro- ceedings South Dakota Academy of Science 61:56-63. Beer, J. R. 1964. Bait preferences of some small mammals. Journal of Mam- malogy 45:632-634. Bishop, O. N. 1966. Statistics for biology, Houghton Mifflin Company, New York, New York. 182pp. Briese, L. A., and M. H. Smith. 1974. Seasonal abundance and movement of nine species of small mammals. Journal of Mammalogy 55:615-629. Buchner, C. H. 1957. Population studies on small mammals of southeastern Manitoba. Journal of Mammalogy 38:87-97. Burt, W. H., and R. P. Grossenheider. 1964. A field guide to the mammals, Houghton Mifflin Company, Boston, MA. 284pp. Eadie, R. W. 1944. The short-tailed shrew and field mouse predation. Jour- nal of Mammalogy 25:359-364. Fowle, C. D., and R. Y. Edwards. 1954. The utility of breakback traps in population studies of small mammals. Journal of Wildlife Management 18:503- 508. Grant, P. R. 1971. The habitat preference of Microtus pennsylvanicus, and its relevance to the distribution of this species on islands. Journal of Mam- malogy 52:351-361. Hallman, K. A., and J. L. Butler. 1992. Small-mammal species diversity and abundance within woodland habitat patches along the Big Sioux River in Union County, South Dakota. Proceedings South Dakota Academy of Sci- ence 71:166-167. Heideman, P. D., J. B. Bowles, and K. R. Erickson. 1983. Habitat selection by small mammals on the shoreline of a flood control lake in South Central Iowa. Proceedings Iowa Academy of Science 90:93-97. Hoff, D. J., S. R. Moats, and R. P. Lampe. 1989. Habitat preference of small mammals during winter in northwestern Iowa. Proceedings Iowa Acade- my of Science 96:118. Manson, R. H., and E. W. Stiles. 1998. Links between microhabitat preferences and seed predation by small mammals in old fields. Oikos 82:37-50. Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 45

Mengak, M. T., and D. C. Guynn. 1987. Pitfalls and snap traps for sampling small mammals and hertpetofauna. American Midland Naturalist 118:284- 288. Morris, R. D. 1969. Competitive exclusion between Microtus and Clethriono- mys in the aspen parkland of Saskatchewan. Journal of Mammalogy 50:291-301. Romansky, J. W. 1970. Effects of population removal on the habitat of small mammals. New York Fish and Game Journal 17:102-109. SAS Institute. 1996. SAS System version 6.12. Cary, North Carolina. Sietman, B. E., W. B. Fothergill, and E. J. Finck. 1994. Effects of haying and old-field succession on small mammals in tall grass prairie. American Mid- land Naturalist 131:1-8. Spencer, A. W., and D. Pettus. 1966. Habitat preferences of five sympatric species of long-tailed shrews. Ecology 47:677-683. Stallman, H. R, and L. B. Best. 1996. Small-mammal use of an experimental strip intercropping system in northeastern Iowa. American Midland Natu- ralist 135:266-273. Stickle, L. F. 1946. Experimental analysis of methods for measuring small mammal populations. Journal of Wildlife Management 10:150-159. Walker, J. A. 1976. Relative abundance and habitat preference of some small mammals in southeastern Nebraska. Transactions Nebraska Academy of Science 3:74-77. Wiener, J. G., and M. H. Smith. 1972. Relative efficiencies of four small mammal traps. Journal of Mammalogy 53:868-873. Yahner, R. H. 1992. Dynamics of a small mammal community in a fragmented forest. American Midland Naturalist 127:381-391.

Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 47

OBSERVATIONS AT THE LAKE MARTIN ROOKERY, CYPRESS ISLAND PRESERVE, ST. MARTIN PARISH, LOUISIANA

James E. Martin Museum of Geology South Dakota School of Mines and Technology Rapid City, SD 57701

ABSTRACT

Within the southeastern portion of the Lake Martin cypress-tupelo swamp is a large rookery where a variety of avian types nest from late January through the spring. A sabbatical leave afforded the opportunity to observe the rookery during the nesting season of 2002. The primary goal was to observe the changes in wildlife in order to make comparisons with paleontological occurrences at higher latitudes. Utilization of the uniformitarian principle resulted in a number of ancillary zoological observations recorded herein. A procession of events occurred at Lake Martin from January to April, 2002. During early January when leaves had been lost and cooler conditions pre- vailed, Great (American) and Snowy Egrets, Great and Little Blue Herons, An- hinga, and White Ibis foraged, usually as individuals or in small flocks. Dur- ing a warm spell in late January, the first birds to begin mating/nesting behavior were the Great Egrets. They displayed and nested principally in cypress and water tupelo trees 5-7 meters above water level; none nested in the higher reaches of the canopy or near water level. Great Blue Herons and Anhin- ga were the next to congregate in early February. The herons took residence in the treetops, whereas the Anhinga normally nested 3-5 meters below and approximately 6 meters above the egrets. In early March, the Little Blue Herons began to congregate at the Lake, followed by Cattle Egrets in mid- March. Both taxa began nesting in the lower reaches of the buttonbush, some only a meter above water level. The last major nesting group was the Roseate Spoonbill, which arrived in the third week in March, and began building nests at the same level as the American Egrets. This is also when the first hatchlings of the American Egrets were observed, as well as the initiation of mating behavior of the American alligator. By the end of March, all major large water birds had engaged in nesting behavior, and some American Egrets were ready to fledge. The persistence of the rookery is dependent, among other parameters, upon the vegetation, isolation by water, a rich feeding area, and probably the prevalence of alligators. More alligators occur in the rookery than anywhere else in Lake Martin. Alligators do feed upon the birds, but they also aid in the restriction of other terrestrial predators such as raccoons and opossums from the nesting area. 48 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002)

INTRODUCTION

A major rookery for larger water birds occurs in St. Martin Parish, southern Louisiana, at Lake Martin, which lies between the towns of Layfayette and Breaux Bridge. Estimates of 20 to 30,000 wading birds occur within this 600- acre cypress-tupelo swamp known as Lake Martin. Lake Martin lies within the Cypress Island Preserve, an 8600-acre preserve donated to the Nature Conser- vancy by Texaco. Observations of the procession of waterfowl from January through March were made in the southwestern portion of the Lake. In order to maintain consistency, observations were confined to this smaller area of easy access. Observations were recorded along with temperature, cloud cover, and wind, as these parameters appeared to affect wildlife behavior. Unfortunately, daily observations were not possible due to other commitments; nevertheless, the major processional events were noted. The Lake was normally checked twice a day, once at midday and again at dusk when birds returned to roost. These observations are not comprehensive but provide insight into the major zoological procession during a specific interval at a specific area. Lake Martin originated through flooding scour within the Atchafalaya-Mis- sissippi River floodplains. The result was a marshy depression that was deep- ened up to four meters (averaging slightly over a meter) in 1952 when a dirt levee was built around the marsh by the Louisiana Department of Public Works. The deeper water provided the necessary parameters for a rookery, and according to local informants, nesting began in the late 1980’s. Over 200 species of birds have been observed at Lake Martin. Of these, this study concentrated on larger water birds, including the following major groups: herons, egrets, grebes, anghina, ibis, spoonbills, some ducks, and gallinules (Table 1). Observations of other taxa such as turtles, American alligators, and nutria (Myocastor coypus) were also conducted.

Table 1. Avian Taxa Observed at the Lake Martin Cypress-Tupelo Swamp.

Cormorants Anhinga anhinga, Anhinga

Herons Ardea herodias, Great Blue Heron Hydranassa tricolor, Louisiana (Tricolored) Heron Florida coerulea, Little Blue Heron Nycticorax nycticorax, Black Crowned Night Heron

Egrets Casmerodius albus, Great (American) Egret Leucophoyx thula, Snowy Egret Bubulcus ibis, Cattle Egret

Ibis Guara alba, White Ibis

Spoonbill Ajaia ajaja, Roseate Spoonbill

Ducks Aix sponsa, Wood Duck

Gallinules and Coots Gallinula chloropus, Florida Gallinule Fulica americana, American Coot Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 49

TIMELINE

January

The later part of December and the first three weeks of January were cool (~12C), and little activity occurred at the Lake, except for that of a few nutria. Thereafter, a warm spell (22-27C) persisted until the end of the month. At the beginning of the warm interval (1/21/02), approximately 30 Great (American) Egrets arrived in the southeastern portion of Lake Martin. The egrets were solitary but began taking up scattered nesting sites at a horizon of 5 meters above the water level in the cypress-tupelo stand. At the same time, a few White Ibis appeared. Only three days later (1/23), twice as many Great Egrets were in the rookery, and the males began their display. A solitary Great Blue Heron appeared, as well as 10 Snowy Egrets. The first three American alligators of the year were also observed. By the last two days of the month, the zenith of the warm spell resulted in hundreds of Great Egrets, most of which were paired, mating, and engaged in nest building. A flock of 18 Snowy Egrets were observed foraging, and the first pair of Florida Gallinules was observed. Nine adult and two juvenile alligators were sunning themselves, as well as numerous turtles and a water moccasin.

February

The first 12 days of the month were very cool (4-9C), so no alligators or turtles were observed, although a few nutria remained active. One day (2/2) was warmer (16C), and five Great Blue Herons (including 2 pairs) and a second pair of Gallinules were observed. A few White Ibis were noted, including a gray-brown juvenile. By the first week of February, hundreds of Snowy Egrets and ~50 White Ibis were roosting in the buttonbush just north of the principal nesting area of the Great Egrets, although some Great Egrets established nests north of the roost. Nineteen Little Blue Herons were vying for roosting sites, principally on the periphery of the Snowy Egret/White Ibis roost. The first pair of Tricolored Herons and numerous nutria were active. By the end of the first week of February, six Great Blue Herons were observed nesting at the treetops west of the roost. Anhinga began pairing and built initial nests 1-3 meters below the Great Blue Heron nests, well above the Great Egrets. Two pair of Black Crowned Night Herons were noted foraging, and a flock of nearly 40 American Coots was among the cypress trees within the Great Egret rookery. At the roost, many more White Ibis arrived so the proportions of Snowy Egrets and White Ibis were nearly the same of approximately 1000 birds; only 45 Little Blue Herons were counted in the roost. By the end of the cold spell, most of the ~500 pairs of Great Egrets were incubating eggs, 55 Anhinga and 14 herons were nesting. Hundreds of White Ibis foraged in the area during the day (many less Snowy Egrets, usually 15-30 individuals, foraged along the eastern margin of the Lake). Although nearly freezing at night, daytime temperatures reached 16C by 2/13 and warmed into the upper teens and lower twenties during the remain- 50 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) der of the month. The moderate warm spell brought the return of alligators and turtles. Additional Gallinules (5 pairs), the first observed pair of Wood Ducks, and additional flocks of American Coots appeared. A huge flock (>100 individuals) of yellow-beaked grebes (Holboell’s Grebe) was observed feeding on the open waters of the Lake.

March

Another cool interval occurred during the first part of March. During this interval, the northern roost was abandoned, although many Little Blue Herons took up residence. The Snowy Egrets had moved south in the buttonbush within the major portion of the Great Egret rookery. By the second week of March, they were joined by equal numbers of Little Blue Herons; the White Ibis were now roosting high up in the cypress-tupelo trees. The first Roseate Spoonbills arrived on 3/18. By the third week in March, the major nesting birds had arrived, and the Great Egrets who had arrived in January now had hatchlings. By this time, the Spoonbills numbered 51 individuals, most were solitary, but one pair was observed constructing a nest. The last nesting bird of the study, the Cattle Egret, was observed in the southern portion of the rookery on 3/21. The alligators were relatively abundant, and all large males that hitherto had been solitary, were now accompanied by smaller females. The American Coots were no longer in flocks but found in discrete pairs. By this time, the Little Blue Herons outnumbered the Snowy Egrets at the southern roost. During the remainder of the month, a warming trend with temperatures up to 25C occurred. By the end of the month, 71 pairs of Spoonbills were counted in the study area where they nested at the same level as the Great Egrets. The Night Herons were nesting just above the Great Egrets, over 500 pairs of Little Blue Herons and nearly 100 pairs of Cattle Egrets were building nests within the buttonbush, some less than a meter above water level. Interesting- ly, the Snowy Egrets, which had been observed at the Lake in January, were building nests at the end of the month, competing for space in the buttonbush with Cattle Egrets and the very abundant Little Blue Herons, but many were found near the base of the bushes. The Great Egrets were ready to fledge when the Spoonbills began their nesting. Overall, all major wading birds had begun the nesting process by the end of the month, and some had young ready to fly.

OBSERVATIONS AND INTERPRETATONS

The nesting levels of various avian species differs. The Great Egrets first colonized the horizon 4-7 meters above the water level in the cypress-tupelo trees. The herons nested at the highest reaches of the trees, and the Anhinga nested 3-5 meters below, but well above (~6 meters) the Great Egrets. The night herons nested about a meter above the Great Egrets. The only taxon to nest at about the same level as the Great Egrets were the Roseate Spoonbills, Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 51 and they appeared very late when the Great Egrets were about to fledge. The Snowy Egrets, Cattle Egrets, and Little Blue Herons nested in the lower buttonbush, less than 1 meter to 2.5 meters above water level. Great Egrets oc- casionally nested at the highest reaches of the buttonbush that was at about the same level as the other Great Egrets in the cypress-tupelo stands. There- fore, most species nested at discrete horizons, although those in the buttonbush competed for any space in the short vertical distance above the water. The original northern buttonbush roost was dominated by hundreds of Snowy Egrets initially with few White Ibis and Little Blue Herons. The White Ibis became much more abundant until nearly equal numbers of Snowy Egrets and ibis dominated the roost. Relatively few Little Blue Herons were found interspersed, but mostly on the periphery of the roost. The Snowy Egrets roost- ed lowest on the buttonbush, whereas the White Ibis were higher. Juvenile ibis are brown and become patchy until they finally are white except for the black tipped wings. Interestingly each night, the darker juveniles were at the highest reaches of the roost, and the white birds were down among the branches. The darker birds would be better camouflaged at night, and the adults certainly show dominance forcing the juveniles to the top of the branches. The highest occurring birds would be vulnerable to night predators, such as Great Horned Owls, which I observed at the Lake. Eventually, Little Blue Herons nested within this roost, and the Snowy Egrets and White Ibis moved to a roost farther south. In the southern roost, Snowy Egrets and Little Blue Herons dominated, and eventually began nesting. The White Ibis were forced out and would roost higher in the cypress-tupelo trees; many juvenile ibis were observed roosting in this manner. Later, Cattle Egrets appeared and also began nesting in the buttonbush. The occurrence of the rookery at Lake Martin may be explained by the cypress-tupelo forest environment, which provides ample forage and nesting materials. The rookery is surrounded by water, affording protection for nesting birds from most predators. Of course, the relative abundance of alligators results in some loss of birds, but these creatures also reduce the numbers of nocturnal mammalian predators such as raccoons and opossums that wreck hav- oc upon nesting birds.

SUMMARY

During 2002, a progression of wading birds arrived to nest at the Lake Mar- tin rookery. The ﬁrst appearing birds were the Great Egrets, followed by herons and anhingas, followed by Night Herons, with Little Blue Herons, Snowy Egrets, Roseate Spoonbills and Cattle Egrets beginning to nest at about the same time. Each species nested at discrete horizons, although Little Blue Herons, Snowy Egrets, and Cattle Egrets competed for space within the 1-2.5 meter interval above the water level in the buttonbush, with the Snowy Egrets normally the lowest. The Spoonbills began nesting at approximately the same level as the Great Egrets, although by the time the Spoonbills arrived, the Great Egret hatchlings were ready to ﬂedge. 52 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002)

Initially, a major roosting area of thousands of Snowy Egrets and White Ibis was established north of the Great Egret rookery on the eastern margin of the Lake. However, it later was moved to the south within the Great Egret rookery. The old roost was taken over by Little Blue Herons that eventually began nesting. The new roost was dominated by Snowy Egrets and Little Blue Herons; the White Ibis were excluded from the low buttonbush and were forced up into the surrounding cypress-tupelo forest. The roost eventually became a nesting area for Snowy Egrets, Little Blue Herons, and Cattle Egrets. The rookery is the result of a number of parameters, including cypress, tupelo, and buttonbush isolated by water, a rich feeding area, and probably the prevalence of alligators. More alligators occur in the rookery than anywhere else in Lake Martin. Alligators do feed upon the birds, but they also aid in the restriction of other nocturnal predators such as raccoons and opossums from the nesting area.

ACKNOWLEDGEMENTS

I wish to thank Lena A. Martin for introducing me to Lake Martin, for her assistance at Lake Martin, for making bird counts, and supporting the project in many other ways. Mr. Robert Smith of Breaux Bridge, LA, provided local information and made observations when other commitments prevented the author from doing so. Finally, the SD School of Mines kindly allowed the author sabbatical leave, providing time for these observations. The project began as a study for comparisons with Late Cretaceous environments, and these observations were made as a result of this undertaking. National Science Foundation grant number 0087972 provided some publication funds. Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 53

JUVENILE MARINE REPTILES FROM THE LATE CRETACEOUS OF THE ANTARCTIC PENINSULA AND THEIR RELATIONSHIPS TO OTHER SUCH OCCURRENCES IN CENTRAL SOUTH DAKOTA AND BELGIUM

James E. Martin Museum of Geology South Dakota School of Mines and Technology Rapid City, SD 57701

ABSTRACT

The Late Cretaceous deposits of Antarctica, South Dakota, and Belgium,have produced higher concentrations of juvenile marine reptiles, both isolated elements and associated skeletons, compared to elsewhere in the world. These concentrations of mosasaurs and plesiosaurs occurred in shallow marine waters, relatively near to shore. In contrast, the deeper, open water areas have produced few juvenile marine reptiles. These concentrations may have been sites where unique preservation occurred or where marine reptiles came into the shallows for birthing and where the young remained until they reached sufﬁcient size and ability to navigate and survive in more open waters.

INTRODUCTION

Recent joint United States-Argentinean expeditions to Vega Island east of the Antarctic Peninsula resulted in the collection of numerous elements indicative of juvenile mosasaurs and plesiosaurs. These expeditions sponsored by the US National Science Foundation (OPP 9815231 & 0087972) and the In- stituto Antartico Argentino were concentrated in the Cape Lamb and Sandwich Bluff members, Lopez de Bertodano Formation. These Maastrichtian sediments were deposited in very shallow marine environments where even terrestrial wading bird and dinosaur elements have been found (Case et al., 2000). From these deposits, partial skeletons and numerous isolated postcranial elements, particularly vertebrae, are attributable to mosasaurs and plesiosaurs, giant marine reptilian groups of the Late Cretaceous. In Antarctica, the plesiosaur specimens vastly outnumber those of the mosasaurs, unlike the situation in central South Dakota or Belgium. For many years, relatively few elements of juvenile marine reptiles were known from the Late Cretaceous deposits of North America, one of the most proliﬁc producers of mosasaurs and plesiosaurs in the world. A concentration of juvenile mosasaurs and even babies has been discovered in central South Dakota along the Big Bend of the Missouri River from the DeGrey and lower 54 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002)

Verendrye members of the Pierre Shale, a black marine shale deposited in the Late Cretaceous (Campanian) North American Epicontinental Seaway. The seaway was continental and therefore shallow, with some estimated depth ranges in the Big Bend area from 30 to 70 meters. In the Big Bend area, mosasaurs are exceedingly abundant. In contrast, only a few fragmentary plesiosaurs have been found in these deposits, so their ontogenetic stages are indetermi- nate at this time. Interestingly, in western South Dakota, hundreds of mosasaur specimens and many plesiosaurs have been collected by our parties from slightly older Campanian deposits of the Pierre Shale, but very few juveniles are known. This region represents deeper, open-water conditions compared to those in central South Dakota which were nearer to the eastern margin of the Epicontinental Seaway. In the late nineteenth century and early twentieth centuries, extensive quarry operations were undertaken in the Late Cretaceous (type Maastrichtian) deposits in Belgium and the Netherlands. In Belgium, mining potassium-rich deposits for fertilizer resulted in approximately 50 partial mosasaur skeletons but only few plesiosaur elements for the National Museum of Belgium. These specimens were derived from shallow depths of 30-50 meters. Although the specimens had reposed in the Museum for approximately a century, the abundance of juvenile individuals was seldom noted.

JUVENILE MATERIAL

Antarctica

Most of the juvenile marine reptile material from Antarctica consists of juvenile partial plesiosaur skeletons, one mosasaur skull, and isolated vertebrae. Many vertebrae are very small, poorly ossiﬁed, and one plesiosaur vertebra even retains the notochordal canal. Two types of juvenile mosasaur vertebrae occur, those with fused haemel arches suggesting the Mosasaurinae or Hal- isaurinae and those with articulated haemals suggesting the Tylosaurinae or Plioplatecarpinae (=Russellosaurinae). Another mosasaur specimen represents a partial skull of a juvenile tylosaurine mosasaur. Adult specimens indicate that the Mosasaurinae are represented by three genera and four species, whereas the Russellosaurinae by two genera and two species (Martin et al., in press and subsequent observations). In addition to the mosasaurs, numerous plesiosaur specimens have been found on Vega Island. Two types of plesiosaurs occur in Antarctica; one is a typical elasmosaurid and the other appears to be a highly derived elasmosaurid. Of the six partial elasmosaurid skeletons noted during our expeditions, three were juveniles and four of eight elasmosaurid skeletons from nearby Seymour Island listed in Chatterjee and Small (1989) represent juveniles or young adults. Only a single skeleton of the derived elasmosaurid (Aristonectes) has been found, and it too is a juvenile individual (Gas- parini et al., in press). Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 55

South Dakota

In the Big Bend area, partial skeletons of both Mosasaurinae and Plio- platecarpinae young have been observed, including a mother Plioplatecarpus containing at least two babies (Bell et al., 1996). Both adult and juvenile mosasaurs are found in abundance, and enough young specimens have been found through our efforts in the last decade that a growth series now exists for Plioplatecarpus tympaniticus and Mosasaurus missouriensis. Only a few vertebrae and two fragmentary plesiosaurs have been found whereas hundreds of mosasaur specimens have been secured. One of the plesiosaurs appears to be juvenile. In western South Dakota around the Black Hills, numerous specimens of mosasaurs and plesiosaurs have been found; however, only one juvenile mosasaur skull (Plioplatecarpinae) and one baby plesiosaur skeleton (Martin, 1994) have been secured.

Belgium

Belgium has been famous for the diversity and abundance of mosasaur specimens from the type Maastricht units. Most of these specimens were described in a series of papers by Dollo (e.g. 1882, 1889) and later reviewed by Lingham-Solier (e.g. 1992, 1993). These papers were principally descriptive, so ontogenetic stage was not emphasized. The Maastricht units are best known for the occurrences of Mosasaurus hoffmanni, the ﬁrst known mosasaur. The type specimen and most others in the literature are very large, and generally, the taxon has been considered as a giant species. Investigations of specimens in Brussels indicate that at least two juveniles are in the collections. These collections also contain numerous specimens of a smaller species, M. lemmonieri; most of these specimens are juvenile or young adults. Nearly all of the specimens of the Plioplatecarpinae are juveniles; the two specimens of the Ty- losaurinae are adults. Of the specimens of Prognathodon, one of three is juvenile. Therefore, although assemblages of mosasaurs from the type Maastricht area have been generally considered adults, many specimens in fact represent juveniles or young adults.

SUMMARY

In Antarctica, South Dakota, and Belgium, a relatively high concentration of juvenile specimens is known compared to elsewhere in the world. In each case, these concentrations are in relatively shallow marine waters, relatively near to shore. In contrast, the deeper, open water area of western South Dako- ta has produced only two juveniles of hundreds of marine reptile specimens. The Antarctic, South Dakota, and Belgian juvenile concentrations were found in very shallow marine sediments, but such concentrations are not found everywhere in similar depositional environments. Therefore, these concentrations may have been sites where unique preservation occurred or where ma- 56 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) rine reptiles came into the shallows for birthing and where the young remained until they reached sufﬁcient size and ability to navigate and survive in more open waters.

ACKNOWLEDGEMENTS

I wish to sincerely thank the Ofﬁce of Polar Programs under the direction of Dr. Scott Borg, without whom this work could not have been undertaken; this investigation was supported by grants OPP 9815231 and 0087972. The Antarctic Institute of Argentina, their personnel, and the personnel of the Ar- gentinean National Museum, La Plata, contributed immensely to the success of the Antarctic project. Nathalie Bardet of the Paris Museum of Natural History; Pierre Bulfynak, Anne Dhondt, Pascual Godefoid, and Jean-Jacques Blarivacq of the Belgium National Museum; Douwe th de Graaf, John Jagt, and Anne Schulp of the Museum of Natural History, Maastricht, kindly allowed me to view collections from the type Maastricht. The Crow Creek Tribe kindly allowed collection and investigations of the fossils on their lands in South Dako- ta, as did the US Corps of Engineers, who aided in ﬁeld excavations and supported curational activities, resulting in the recovery and preservation of juvenile mosasaurs.

REFERENCES

Bell, G.L., Jr., Sheldon, A., Lambe, J. and Martin, J.E., 1996, First direct evidence of live birth in Mosasauridae (Squamata): exceptional preservation in the Cretaceous Pierre Shale of South Dakota: Jour. Vert. Paleont., Abstr., 16(3 suppl.):21-22A. Case, J.A., Martin, J.E., Chaney, D.S., Reguero, M., Marenssi, S.A., Santillana, S.M. and Woodburne, M.O., 2000, The First Duck-billed Dinosaur (Family Hadrosauridae) from Antarctica: Jour. Vert. Paleont., 20(3):612-614. Chatterjee, S. and Small, B.J., 1989. New plesiosaurs from the Upper Cretaceous of Antarctica, in Crame, J.A. (Ed.) Origins and Evolution of the Antarctic Biota, Geol. Soc. London, Spec. Publ., 47:197-215. Dollo, L., 1882. Note sur l’osteologie des Mosasauridae: Bull. Mus. Royal d’His- toire Nat. de Belgique 1:55-65. Dollo, L., 1889. Premiere note sur les mosasauriens de Mesvin: Bull. de la so- ciete Belge de Geologie, de Paleontologie et Hydrologie, 3:271-304. Gasparini, Z., Bardet, N., Martin, J.E. and Fernandez, M., in press, Systematical review of Aristonectes and Mortuneria (Plesiosauria) from the latest Creta- ceous of South America and Antarctica: Jour. Vert. Paleont. Lingham-Soliar, T., 1992. The tylosaurine mosasaurs (Reptilia, Mosasauridae) from the Upper Cretaceous of Europe and Africa: Bull. Royal Inst. Nat. Sci. Belgium, Ear. Sci., 62:171-194. Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 57

Lingham-Soliar, T., 1993. A mosasaur bares its teeth, pp. 445-458: in Sargeant, W.A.S (Ed.) Fossil Vertebrates: Faunas and Concepts, Special Issue, Hal- stead Mem. Vol., Part IV, Modern Geology, 18(4):419-534.Lingham-Solier, Martin, J.E., 1994, A baby plesiosaur from the Late Cretaceous Pierre Shale, Fall River County, South Dakota: Jour. Vert. Paleont., 14(3 suppl.):35A. Martin, J.E., Bell, G.L., Jr., Case, J.A., Chaney, D.S., Fernandez, M.S., Gasparini, Z., Reguero, M., and Woodburne, M.O., in press, Mosasaurs (Reptilia) from the Late Cretaceous of the Antarctic Peninsula: New Zealand Jour. Geol. Geophy.

Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 59

ANALYSIS OF WIND OBSERVATIONS IN THE BLACK HILLS REGION

Srivisakha Skandaswamy Pokkunuri, Sameer Kumara Krishna Dronamraju, and Abul R. Hasan Department of Electrical and Computer Engineering and Andrew G. Detwiler Institute of Atmospheric Sciences South Dakota School of Mines and Technology Rapid City, SD 57701

ABSTRACT

The potential for wind energy production was assessed for 4 specially in- strumented sites in the region of the Black Hills of South Dakota. More than 3 years of observations were obtained at 75 and 100 ft AGL at these sites, from 1995-1998. A site to the north of the main body of the Black Hills, north of Stur- gis and southwest of Bear Butte, has significantly higher mean winds than the other sites, although it is also the site at the lowest terrain elevation. Monthly mean winds at 100ft AGL at this site were more than 14 mph while mean winds were near 12 mph at 3 other sites in the higher elevations of the Black Hills region. The frequency distribution of wind speeds has a broader multi-modal form at the most windy site compared to the other sites, with significantly more frequent occurrence of winds greater than 30 mph compared to the other sites. The strongest winds at all sites were typically northwesterly, although strong winds also occur from the southwest, south, and southeast in fall, winter, and spring, at most sites. At all sites, summer was typically the season with the lightest winds. There was weak correlation between monthly mean wind speeds at the Rapid City Regional Airport and at the three sites around the northern and central Black Hills. There was no correlation between monthly mean wind speeds at Rapid City Regional Airport and the site at Four Corners, Wyoming on the western border of the Black Hills, about 100 km west of the airport. These results suggest that examination of the 100+ year record of winds at the official observing sites in and around Rapid City may yield a valid estimate of the variability of winds at sites like those involved in this study as long as they are in the central or eastern portion of the Black Hills. Rapid City winds will not be as good a predictor of winds on the western side of the Black Hills.

Keywords

Wind, meteorology, renewable energy 60 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002)

INTRODUCTION

Hasan and Hughes (1995) discuss almost one year of wind observations at four sites in the Black Hills region of western South Dakota. These investiga- tors installed instrumentation at these sites with support from the Black Hills Power and Light Company (now called Black Hills Power) to evaluate the potential for commercial generation of electric power using wind turbines at these sites and other sites with similar exposure in the Black Hills region. All sites were chosen based on their exposure, proximity to transmission lines, and the permission and support of landowners. The locations of the sites are indicated on a regional map in Figure 1. Three of the four sites, one near Four Corners, Wyoming, another on the Clinton Ranch, and a third on the Sanders ranch, are all located on relatively high terrain in a roughly west-east line extending across the central Black Hills. The fourth site is located on grazing land then owned by the Homestake Mining Company 14 miles north of the Sturgis, on lower terrain to the north of the Black Hills. Bear Butte rises about 4 miles east northeast of this Homestake site.

Figure 1. Contour map of the Black Hills region of South Dakota with Black Hills Power and Light Company wind survey sites indicated. Contour interval is 200 m, and spatial resolution is 50 m. Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 61

Wind speed observations were obtained at 2 levels at each site, 100ft (30 m) and 70 ft (21 m) above ground level (AGL). Wind direction was observed only at the 100ft (30 m) level. Data obtained beginning in September 1994, and continuing through July 1995, were discussed in the initial report by Hasan and Hughes. Although all 4 sites were located in open areas where high mean wind speeds were expected, based on low-spatial-resolution national wind energy surveys, only one site, the Homestake site on the northern ﬂank of the Black Hills, actually turned out to have high enough and steady enough winds for economical wind turbine power generation. We examine here several years of additional data to see whether or not the climatological characteristics of wind at these sites, estimated based on the ﬁrst year of observations, remained similar over a several year period. Equipment failures occurred over these subsequent years, and the last good quality data from the last functioning site were obtained in the spring of 1999. Wind vanes were more prone to failure than anemometers. Hail from thunderstorms is known to have been responsible for some of the instrument losses. Other weather hazards (lightning, winter icing, etc.) may have been responsible for other losses. In this discussion we will focus mainly on the meteorological analysis of the wind observations, with some emphasis on those characteristics of wind that contribute to a more useable wind-driven turbine electric power-generat- ing site.

REVIEW OF FIRST YEAR’S DATA

Towers and instrumentation were installed in the summer and fall seasons, 1994. Data were recorded on-site using battery-powered data loggers. Magnet- ic data cards were exchanged and batteries replaced during roughly bi-monthly site visits by project personnel. An analysis of the ﬁrst year of data was performed during fall, 1995. (Hasan and Hughes, 1995) reported mean wind speeds at 100 ft (30 m) AGL, based on the 11 to 12 months of data available at that time from each site, are shown in Table 1. Monthly mean wind speeds varied from 10 to 18 Table 1. mph. Variation from Site Mean Wind speed (mph) month-to-month was great- est at the Homestake site Clinton 12.3 and least at the Four Cor- Sanders 11.9 ners site. At most sites, for Four Corners 11.9 this period, the maximum Homestake 14.3 monthly means occurred in the fall (October or Novem- ber) and spring (March and April), within lower means in early winter (De- cember) and late summer (July and August). For comparison, the annual mean wind speed at the ofﬁcial wind observing height of 33 ft (10 m) AGL at the 62 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002)

Rapid City Regional Airport is 11 mph, based on observations from 1930-1996 (National Climatic Data Center, 1998). Monthly means at the airport vary from 10 to 13 mph, with highest monthly means in March and April, and lowest means in July and August. The (non-zero) wind speed frequency distribution was monomodal at 3 of the sites, with most frequent wind speeds at these sites between 9 to 12 mph, and a peak frequency for the mode wind speed of 6 to 7% of all 15 min observing periods. At these 3 sites, the wind speed was < 1 mph for 5 to 8% of the time, and wind speeds greater than 20 mph were observed no more than 2% of the time. At the 4th site, the Homestake site, the peak in the wind speed frequency distribution was much broader and extending to greater speeds. There was a broad peak in the distribution, with each integer wind speed value between 7 and 22 mph being observed more than 3% of the time. At all 4 sites, the dominant winds were from the SW and NW quadrants. Winds were most frequently from the NW at all sites except Clinton, where the most frequent wind direction was SW. The Four Corners wind vane data show an unrealistically narrow peak from the NW in the direction distribution. This wind vane apparently was not functioning properly as early as this ﬁrst season of data. All sites are on cleared land that is elevated above the surroundings in most directions. There is a small knob of land located about 300 m south of the Clinton site that might account for the dearth of strong winds from the south at that site. The Sanders site sits on a prominent knob just east of a south- north valley through which runs US Route 385. Channeling of winds by this valley may account for the relatively more frequent southerly winds observed at this site compared to the others. The strongest winds at all sites were most frequently from the north and west quadrants. The weakest winds were from the east. We next compare these published analyses based on the nearly complete ﬁrst year of data from each site, to an analysis of the entire record, extending to 4 years for some parameters at some sites.

ANALYSIS OF COMPLETE DATA SET

An analysis of the complete data set was performed using the Winsite software package (Second Wind, Inc., Somerville, MA. URL: http://www.sec- ondwind.com). In the present analysis data were broken down seasonally, and wind roses and wind speed frequency spectra were computed for each site for the ‘seasons’ January-March (‘winter’), April-June (‘spring’), July-September (‘summer’), and October-December (‘fall’). Although seasonality of the wind regime over the Black Hills may not correspond exactly to these calendar periods each year, the aim is to capture at least some of the seasonal variability. Hasan and Hughes (1995) note that overall, for the period fall, 1994, through summer, 1995, fall and spring were windier than summer. In the following analysis we look in more detail at how much variation there is from ‘season’ to ‘season’, and from one year to the next in a given ‘season’. Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 63

WIND DIRECTION DISTRIBUTION BY SEASON

The Four Corners wind direction data appear to be unreliable, even during the ﬁrst year, indicating an unrealistically narrow peak frequency from the northwest. From April 1996, onward, the wind direction is indicated as always exactly from the northwest. This appears to be due to a failure in the recording of the wind vane position, although as even as late as the spring, 1999, the wind vane appeared to be working properly mechanically. The Homestake wind direction data also developed problems early in the study, becoming unrealistic during summer, 1995. During the period of reliable data, there is a prominent peak in frequency and magnitude of northwesterly winds, from December 1994, through March 1995. In spring, 1995, there were some frequent and strong winds from the southeast, as well as from the northwest, associated with several strong low-pressure systems that passed near or over the Black Hills region during April of that year. At the Sanders site, good data are available from winter, 1995, through summer, 1996. The most frequent and strongest winds were from the northwest, although spring and summer in both 1995 and 1996 also show frequent winds from the south and southeast that generally are not quite as strong as the northwest winds. It is probable that these spring and summer south and southeasterly winds result from synoptic-scale low pressure areas moving through or around the region in the this time period. At the Clinton site good direction data are available from fall, 1994, through summer 1997. Westerly winds dominate during this period from fall, through winter, and spring. Summers generally are dominated by winds from the southeast through west quadrants, although summer, 1997, also shows a prominent northwesterly wind peak. Winds from the southeast are not as strong at this site as they are at the Sanders and Homestake sites. The Clinton site is more centrally located in the Black Hills and often does not see winds as strong as those at the Sanders and Homestake sites (Sanders near the eastern edge, Homestake somewhat north of the northern edge of the Black Hills) when springtime low-pressure areas migrate by the region.

WIND SPEED FREQUENCY DISTRIBUTIONS BY SEASON

Wind speed data were generally more reliable than wind direction data. The 3 higher elevation sites had similar mean wind speeds, while the Homes- take site on lower elevation land to the north of the Black Hills (see Figure 1), had a signiﬁcantly higher mean wind speed. Data generally are available from fall, 1994, or winter, 1995, through winter, 1998. The wind speed at 100 ft (33 m) AGL will be the basis for the following discussion. At Four Corners the seasonal frequency distributions typically were monomodal. An example from winter, 1995, is shown in Figure 2. In this season, the peak value of 12 mph was observed 6.7% of the time. Over the entire period of record the peak seasonal median wind speeds varied between 10 and 13 mph and this peak value was observed from 6 to 8% of the time. Winds 64 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002)

greater than 20 mph were least frequent during summer. Near-zero winds were generally least frequent in the spring, but there was much year-to-year variability in the frequency of very light winds in a given season. Outside of the tenden- cy for winds to be generally lighter in the summer, Figure 2. Frequency distribution of mean wind we found in all seasons, at over 15 minute intervals at the Four Corners site for Winter, 1995. Data are from the 100 ft. (30m.) this site, that there was AGL level. about as much variability from year-to-year in a given season, as there was within a year from season-to-season. Of course, the seasons do not necessarily change at precise 3-month intervals at the same times each year, and some of the ‘seasonal’ variability is probably due to variation in the timing and length of the true meteorological seasons, fall, winter, spring, and summer. Figure 3. Frequency distribution of mean wind Seasonal trends similar over 15 minute intervals at the Clinton Ranch site for Fall, 1995. Data are from the 100 ft. (30m.) AGL to those at the Four Cor- level. ners site were observed at the Sanders Ranch and Clinton Ranch sites. At these latter two sites, the months with higher median winds speeds tended to have broader maxima in their speed frequency distribution, and a longer tail towards high wind speeds, compared to the Four Cor- ners data. An example from the Clinton Ranch from the Fall, 1995, is shown in Fig- ure 3. The winds at the Figure 4. Frequency distribution of mean wind over 15 minute intervals at the Homestake site for Sanders Ranch and Clinton Spring, 1996. Data are from the 100 ft. (30m.) AGL Ranch sites were more vari- level. able from season-to-season Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 65 than at Four Corners, with seasonal median wind speeds ranging from 7-16 mph at the Sanders site and 8-17 mph at the Clinton site. The Homestake site was the windiest site, with much broader maxima in the seasonal speed frequency distributions compared to the other sites. These speed frequency distributions almost always exhibited multiple modes along with much fatter tails extending towards higher speeds. An example from Spring, 1996, is shown in Figure 4. The lowest mode typically was in the 7-10 mph range, and an upper mode typically was between 13 and 21 mph. As at the other sites, winds were lightest in the summer, and speed frequency distributions tended to be more monomodal in summer than during the other 3 seasons. Higher winds were almost always observed at the Homestake site in the fall through spring seasons, compared to the other sites. The frequency distributions, though broader at the Homestake site, were somewhat less variable in shape than at the other sites during fall through spring.

SEASONAL MEAN WINDS

Monthly mean winds at 100 ft (30 m) AGL at the four wind survey sites were computed and compared to those at 33 ft (10 m) AGL Rapid City Regional Airport. The monthly mean winds at the airport are almost always lower than at any of the four sites. The primary reason is the fact that the wind speed measurement is made at a lower height AGL at the airport at the other sites, but geographic location differences may also play some role. The Homestake site generally has the highest monthly mean winds among the 5 sites, consistent with Table 1 in which it also had the highest mean winds over the entire first year of the study, and Table 2, in which it had the highest mean wind over the entire period of study. Although the wind speeds at the study sites cannot be directly compared to those at the airport due to the different heights at which observations were made, it is possible to compare deviations from the long-term mean at each study site to the corresponding deviation at the airport. Correlation coefficients for each series, correlating monthly mean wind speed at each study site with monthly mean wind speed at the airport, for the period January, 1995, through February, 1998, are listed in Table 2. Correlation coefficients range from 0.01 to 0.39, with the best correlation between airport and Homestake site wind speeds, and the worst correlation between airport and Four Corners site wind speeds. All sites except Four Corners show larger amplitude of variability from month-to-month and season-to-season compared to the airport.

Table 2.

Site Airport Clinton Four Corners Sanders Homestake

Mean Wind (mph) 10.4 12.2 11.8 12.3 14.5

Correlation Coefﬁcient 1 0.31 0.01 0.32 0.39 66 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002)

The Four Corners site is on the western edge of the Black Hills, while the other sites are along the eastern and northern periphery (Sanders, Rapid City Airport, and Homestake) or in the east central (Clinton) Black Hills. Geo- graphic distance as well as location on opposite sides of a significant terrain feature, the Black Hills, probably account for the independence of wind variability between the Four Corners site and the airport. Table 2, based on more than three years of data, shows essentially the same relative ranking between mean wind speeds at the four Black Hills Pow- er study sites as was found by Hasan and Hughes (1995) from an analysis of just the first year of data. Mean wind speed over the whole period at the Home- stake site is more than 2 mph faster than mean winds at the other 3 sites, which are similar in magnitude to each other. This small difference translates to a significant difference in wind energy class as defined in Hasan and Hughes (1995). The Clinton, Four Corners and Sanders sites are Class 2, while the Homestake site is Class 4. Based on the mean wind speed of 10.4 mph at 33 ft (10 m) AGL at the Rapid City Airport, the class definitions given by Hasan and Hughes (1995), suggest that the Rapid City Airport site also would be a Class 2 wind energy site.

DISCUSSION AND CONCLUSIONS

This analysis confirms the relative rankings for wind energy production between the four Black Hills Power sites deduced in a study of wind energy potentials by Hasan and Hughes (1995). The earlier study looked at the first year of observations, while the present study examined more than 3 years of observations (including the same first year). The Homestake site, north of Sturgis and southwest of Bear Butte, has significantly higher mean winds than the other sites. Mean winds at 100ft (30 m) AGL are more than 14 mph at the Home- stake site while they are near 12 mph at the other sites. The frequency distribution of wind speeds has a broader multi-modal form at the Homestake site compare to the other sites, with a significantly more frequent occurrence of winds greater than 30 mph compared to the other sites. Wind direction data from the Four Corners site were found to be unreliable for the entire period, while at the other sites the direction data became unreliable after about the first year. The strongest winds at all sites with valid wind direction observations are most frequently northwesterly, although strong winds also are common from the southwest, south, and southeast in fall, winter, and spring, at most sites. At all sites, summer was typically the season with the lightest winds. There was weak correlation between monthly mean wind speeds at the Rapid City Regional Airport and at the three sites around the northern and eastern fringe of, and in the east central, Black Hills. There was no correlation between monthly mean wind speeds at Rapid City Regional Airport and the Four Corners, Wyoming, site on the western border of the Black Hills. The variability in monthly and seasonal mean winds at the 100 ft (30 m) level at the northern and eastern sites was greater than the variability at 33 ft (10 m) at the air- Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 67 port. The wind speed at 100 ft (30 m) AGL at Four Corners was less variable than at all other sites, including the airport. These results suggest that examination of the 100+ year record of winds at the various official observing sites in and around Rapid City used since the late 1800’s may yield a rough estimate of the variability on century time scales of winds at sites like those involved in this study as long as they are on the same side of the Black Hills as Rapid City. Rapid City winds will not be a good predictor of winds on the western side of the Black Hills. When estimating wind energy potential over broad areas, it often is assumed that winds will be generally higher over higher terrain. This study shows that this in not true in the Black Hills region. Other factors, including interaction between the northern flank of the Black Hills and strong northwest winds from cold season storms, and possibly some channeling of these winds between the northern flank of the Black Hills and the neighboring Bear Butte, can lead to higher mean winds at lower elevations. The higher mean speed results from the appearance of a second mode in the wind speed frequency distribution at higher speeds, while at the other sites the frequency distribution is monomodal with a much lower frequency of higher wind speeds. One of the problems with generation of electric power using wind-driven turbines is the variability of the winds, and the frequent lack of sufficient wind at any one site to generate useable power. This study suggests that a power distribution grid covering an area equivalent to or larger in size compared to the Black Hills region, with wind generation sites distributed over the entire grid area, may be able to depend on having some wind-generated power from at least some of the sites much of the time. In comparison, a distribution system with most of its wind-generation capacity located at one or a few sites will have more intermittent power contributions from wind-driven turbines.

REFERENCES

Hasan, A. R., and W. L. Hughes, 1995: Black Hills Power Wind Study. Depart- ment of Electrical Engineering, South Dakota School of Mines and Tech- nology, Rapid City, SD 57701. 25 pp. + appendices. National Climatic Data Center, 1998. Climatic Wind Data for the United States. National Climatic Data Center, 151 Patton Avenue, Room 120, Asheville, NC 28801-5001. http://www.ncdc.noaa.gov . np.

ACKNOWLEDGEMENTS

We thank Don Martinez from Black Hills Power Corporation for sharing with us the data collected by his company and facilitating its analysis. We also thank the landowners for facilitating these observations. The Institute of At- mospheric Sciences and Department of Electrical Engineering at the South Dakota School of Mines and Technology supported this analysis effort. Dr. William Capehart produced Figure 1.

Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 69

MODELING STREAMFLOW IN SPEARFISH CREEK, BLACK HILLS, SD

Larry D. Stetler and John P. Knight Dept. of Geology & Geological Engineering SD School of Mines & Technology Rapid City, SD 57701

ABSTRACT

Spearfish Creek flows in a narrow, incised canyon in the northern Black Hills which causes high velocity flood flows to impinge on channel banks and bottom. Scour holes, channel widening, damaged fish habitat, damage to, or collapse of, existing roadways and bridges could result. Thus, it would be desirable to predict locations where increased bank and/or channel stress would be expected under varying flow conditions. An approximately 1.0 mi reach, between Hydro No. 2 and Maurice intake, was surveyed using a laser-digital transit. Channel cross-sections were measured every 100 ft producing a detailed topographic map of the channel, banks, and limited floodplains. Flow modeling was performed using dis- charges that were based on current stream-flow and known historic flood flows. The stream was modeled as segments where inflow to each segment occurred at the upstream boundary. The segment containing Squaw Creek tributary had two inflow boundaries. Results indicted high stress at numerous points along the entire stream channel as a function of river channel morphology. Areas of concern included sections that contained bends or narrowing of the channel. Where the highway was closest to the potential channel scour zones, damage might be expected under severe flow conditions causing immediate damage to the fishery. These results will allow highway engineers and fisheries personnel to maximize efforts to sustain existing roadways and to provide and maintain quality fisheries within Spearfish Canyon.

Keywords

Spearfish Creek, flow modeling, fish habitat

INTRODUCTION

Naturally reproducing trout in Spearﬁsh Creek are limited in spawning success to the amount and location of available and suitable habitat. The purpose of this study was to 1) model potential effects of increased discharge on ﬁsh habitat, 2) determine behavior of substrate (scour and deposition of sediment), and 3) identify potential locations of increased bank stress. The study was conducted with the cooperation of the SD Dept. of Game, Fish, and Parks. 70 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002)

The study area was located approximately 10 miles south of the city of Spearfish on Spearfish Creek between Homestake Mining Co. (now Barric Min- ing Co.) Hydroelectric Power Plant Number Two (Hydro No. 2) and Maurice intake, a diversion tunnel supplying Hydro No. 1 in Spearfish (Fig. 1). Physi- cal descriptions of local geology, morphology, and stream character are given by Stetler and Sieverding (2001) and Sieverding (2001). Two streams flow through the study area. Spearfish Creek (the larger of the two) originates from springs emanating from the Mississippian Madison Limestone and flows northward within the confines of Spearfish Canyon. Squaw Creek (the smaller tributary stream) enters Spearfish Creek about 1/3 mile below Hydro No. 2 and sources from springs and seeps within the Cambrian Deadwood formation aquifer east of Spearfish Canyon and flows westward. Spearfish Creek is a low sinuosity single channel within the study reach Figure 1. Study site location map. The small (Mussetter 1997), although square (at 90° bend) is the location of data shown it does meander through in Figures 3 and 4. the bedrock of the canyon through which it flows. State Highway 14A, located immediately to the west of the main channel, is stabilized by using gabion structures (wire mesh baskets filled with cobbles and boulders) to armor the road side channel bank which effectively constrains the creek from natural meandering and bank adjustments during high flows. Thus, increased shear stress generated during high flows would primarily be directed toward the gabions and the channel bottom promoting scour. Potential loss to fish habitat under these conditions would be high. In addition, under extreme flow events, gabions could easily be washed out causing bank and possible road failure, both of which would exacerbate loss of fish habitat. Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 71

STUDY METHODS AND RESULTS

A 3-D numerical flow model, Surfacewater Modeling System, was used to evaluate potential effects of various flow conditions on the banks and substrate of Spearfish Creek. The model was run on a topographic base map (Fig. 2) that was prepared for this study from detailed surveys of the channel, banks, floodplains, and surrounding areas. Head loss along the main channel of Spearfish Creek was 80ft (4463 ft at Hydro No. 2 to 4383 ft at Maurice intake). Model limitations for maximum head loss of <25 ft forced the study reach to be segmented for flow analysis. Figure 3 is an example segment (see Fig. 1 Figure 2. Topographic model generated from de- for location) showing the tailed survey. Main channel indicated is Spearfish designation of material Creek. Squaw Creek enters from the east. properties. For the study reach, 3 material types were designated: 1) channel, 2) vegetated bank, and 3) engineered bank. Each of these materials were assigned an erodibility factor (Manning roughness coeffi- cient) with in-channel erosion also controlled through an eddy viscosity value (transverse flow). Adjustment of parameter values were based on actual physical conditions at the stream and from published data. Flow modeling was Figure 3. Modeled segment from northern part of the study area. Each segment modeled is defined performed using the mean by erodibility factors as indicated by the patterns. average discharge (~60 cfs) and a medium flood event discharge (2000 cfs). Re- sults from the 2000 cfs event for the segment shown in Fig. 3 are shown in Fig. 4. Topography is not shown in this figure. Inflow of 2000 cfs is shown along the upstream boundary and the downstream water elevation is shown as 4000 72 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002)

ft elevation. The arrows represent flow magnitude vectors, the larger the ar- row, the higher the velocity. At the upstream edge, flow is spread across the entire segment and is flowing out of the channel. Ve- locities out of the channel are ~1.5 ft/s and range from 6-7.5 ft/s in the main channel. Zero’s mark the edge of the out-of-channel Figure 4. Model results for segment shown in flow. Erosion on the vege- Figure 3. Length of arrows relate to velocity mag- tated bank is low due to nitude. Note the increase in velocity directed to- the velocity slowing from ward the west bank. the vegetation. In the main channel, flow impinging on the outer, west-side bank, increases in velocity, and thus, shear stress as well. Velocities of up to 12 ft/s are shown and would effectively erode banks and engineered gabion structures. A similar magnitude flood occurred in the canyon in 1994 and undercut and eroded gabion structures in this corner and washed them into the stream. They were removed in 2000 and the corner was re-engineered.

SUMMARY

These results form a preliminary assessment of channel and bank erosion for Spearfish Creek. They indicate areas of concern exist from both a fisheries and a highway standpoint. During increased flow, increased bank stress (and the potential for bank failures) exists at several points along the highway side of the channel. Many of these banks are protected by gabion structures which in effect, transfers the stress to the channel bottom which can reduce or eliminate fish spawning habitat in these areas. In extreme events, complete failure of the highway side bank is possible by under-washing of the gabions. Results from future model runs at different flow rates can assist in identification of these locations.

ACKNOWLEDGEMENTS

Funding and support for this project were provided by the South Dakota Department of Game, Fish and Parks, through Federal Aid in Sport Fish Restoration funds (Federal Aid Project F-15-R, Study 1588), and by the South Dakota School of Mines and Technology. Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 73

LITERATURE CITED

Mussetter Engineering, Inc. 1997. Geomorphic and Sediment Transport Study; Spearfish Creek Between Hydro No. 2 and Maurice Intake Dam. Prepared for the SD Dept. Game, Fish, & Parks, 2 plts, 59p. Sieverding, H.L. 2001. Environmental controls on fish spawning habitat in Spearfish Creek between Hydro No. 2 and Maurice intake. MS Thesis, SDSM&T, 67p. Stetler, L.D. and H.L. Sieverding. 2001. Environmental controls on fish spawning habitat in Spearfish Creek, Black Hills, SD. Proc. SD Acd. Sci., 80:109- 117.

Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 75

SOUTH DAKOTA TURFGRASS INDUSTRY: IMPACTS AND VALUE

Leo C. Schleicher Department of Horticulture, Forestry, Landscape and Parks South Dakota State University Brookings, SD 57007

ABSTRACT

Turfgrass is the preferred ground cover in modern American culture. Tur- fgrass enhances the quality of life, is a viable economic component, and serves as a valuable employment provider. The economic impact of the turf industry has been quantiﬁed in other states, but little information is available for South Dakota. The objective of this study was to compile data from other states as a basis for estimating the economic impact of the South Dakota turf industry. Based on estimates from Georgia, Indiana, Iowa, Kentucky, Missouri, North Carolina, Pennsylvania and Wisconsin, annual turfgrass expenditures in South Dakota are predicted to be greater than $500 million. The value of South Dakota turfgrass as an economic agricultural contributor ranks third behind corn and soybeans, valued at $690 and $681 million, respectively, in 2000.

INTRODUCTION

Turfgrass is ubiquitous in modern American culture as a component of the urban and suburban landscape, outdoor sports and recreation, land management, and beautification. Turfgrass is valued for aesthetic quality as well as functional use (Beard, 1973). Beard and Green (1994) summarized the benefits of turfgrass into three major classifications: functional, recreational, and aesthetic. Functional benefits include reduction of soil erosion, dust, noise, glare, air pollution, and high temperatures. Low cost turfgrass surfaces enhance recreational activities and reduce sports injuries. The aesthetic value of turf adds to the beauty, mental health, community pride, and quality of life. Additionally, the turf industry is a major economic contributor and employment provider. Turfgrass is the preferred ground cover for home lawns, commercial and industrial sites, parks, playgrounds, athletic fields, cemeteries, golf courses, and roadsides. The turf industry provides employment in professional turf management; the service sector; fertilizer, chemical and pesticide production and distribution; sod and seed production; equipment and irrigation manufacturing and distribution; and retail product outlets such as lawn and garden centers (Watson et al., 1992). The overall value of the U.S. turf industry in 1992 was estimated at $25 billion annually (Watson et al., 1992). Recent estimates increased the overall value to 35 to $45 billion (Agric. Res. Serv., 2002; Hortic. and Landscape Archi- tecture Dep., 2002). Vinlove and Torla (1995) estimated total home lawn area 76 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) in the U.S. at 5.7 to 10.5 million ha. According to the National Golf Founda- tion, there are 26.7 million U.S. golfers playing on 17,108 golf courses, of which 524 opened last year (National Golf Foundation, 2001). South Dakota was listed among the top 4 of 50 states in percentage of residents > 12 years of age who play golf. South Dakota has one of the highest golf course per capita ratios in the U.S. with approximately 120 golf courses; and all cities with population > 1,000 have a golf course (personal research; U.S. Bureau of the Cen- sus, 2000). The economic impact of the turf industry in some states has recently been quantified from detailed survey data (Pennsylvania Agric. Statistics Serv., 1990; Fleishman-Hillard, 1999; Murphy et al., 1999; Iowa Agric. Statistics Serv., 2001). Turf industry estimates from other states are available from university websites; however, data used to derive estimates are often not provided nor are sources referenced (Hallonquist, 1998; Eclov, 2000; Rodekohr, 2000; Fisher, 2001; Dep. of Hortic., 2002; Schoettle, 2002). Because turf industry data in South Dakota is limited, existing data from other states were used as a basis for estimating the economic impact of the South Dakota turf industry. The objective of this study was to estimate annual turfgrass expenditures in South Dakota without conducting a detailed survey.

METHODS

Simple linear regression was used to predict annual turfgrass expenditures in South Dakota. Georgia, Indiana, Iowa, Kentucky, Missouri, North Carolina, Pennsylvania, and Wisconsin were used as the sample population, with total developed land as the independent variable and annual turfgrass expenditures as the dependent variable. Natl. Resources Inventory (NRI) data were used to determine relative turfgrass area among comparison states (Natl. Resources In- ventory, 2000a; 2000b).

RESULTS AND DISCUSSION

Land usage, as defined by the NRI, was categorized into pasture, forest, crop, federal, developed, and other land (Natl. Resources Inventory, 2000a). Turfgrass is classified as developed land. Developed land, calculated as a percent of total land, was considerably less in South Dakota than other states. Al- though South Dakota total land area was greater, there were < 0.4 million ha of developed land in South Dakota compared to 0.7 million ha in Iowa and 1.0 million ha in Missouri. Developed land is sub-classified as large urban and built-up, small built- up, or rural transportation land (Natl. Resources Inventory, 2000b). Percent rural transportation land, e.g., roadside turf, was greater in South Dakota compared to other states, but percent large urban and built-up land was least among comparison states (Table 1). Simple linear regression indicated a positive relationship between total developed land and annual turfgrass expenditures (r2=0.73, p=0.05) among the sample population states (Fig. 1). Based on South Dakota’s 388,679 developed Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 77

Table1. Developed land usage and annual expenditures attributed to turfgrass production, maintenance, and industry.

DEVELOPED LAND†

Large Urban Small Rural Annual Turfgrass State and built-up built-up Transportation Total Expenditures Reference

ha x 104 $ x 109 MO 62.25 5.16 34.55 101.95 1.30 Fleishman-Hillard Res. 1999) WI 64.46 6.95 26.52 97.93 1.00 Dep. of Hortic. (2002) IA 30.66 2.17 36.11 68.94 1.07 Iowa Agric. Statistics Serv. (2001) KY 38.09 2.17 38.30 78.57 0.50 Eclov, M. (2000) GA 126.89 8.60 24.79 160.27 1.56 Rodekohr, J.L. (2000) PA 127.08 17.70 16.54 161.32 1.50 Murphy et al. (1999) NC 117.79 15.82 22.58 156.18 2.00 Fisher, A. (2001) IN 65.34 6.40 19.80 91.55 1.00 Schoettle, A. (2002) SD 13.12 0.96 24.79 38.87 0.51‡

† National Resources Inventory, 2000b. ‡ Predicted annual turfgrass expenditures.

2.5

Y = 1.30exp8 + 9.79exp2X 2.0 r2 = 0.73, p = 0.05 NC ) 9

GA 1.5 PA

IA 1.0 INWI

0.5 KY Annual turfgrass expenditures ($ x 10 expenditures turfgrass Annual

0.0 0 5 10 15 20 Developed hectares ( x 10 5) Figure 1. Relationship between developed hectares and annual turfgrass expenditures among sample population of eight states. 78 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) hectares, the predicted value of the state’s turfgrass expenditures is $511 ± 255 million annually. The value of turfgrass to South Dakota’s economy appears to be comparable to the major agricultural crops. According to the South Dakota Agric. Statistics Serv. (2001), the value of crop production in 2000 for corn, soybeans, and hay was $690, $681, and $402 million, respectively. The predicted annual value of $511 million for South Dakota turfgrass would rank third, behind corn and soybeans. Similar ﬁndings were reported in other states. Turfgrass was listed as the fourth major crop in Missouri, following soybeans, hay, and corn (Fleishman-Hillard, 1999). According to the College of Agricultural Sci- ences and Natural Resources at Oklahoma State University (Hortic. and Land- scape Architecture Dep., 2002), turfgrass is the third-largest agricultural crop in Oklahoma. In Indiana, turfgrass is the third major crop, following corn and soybeans (Schoettle, 2002). In summary, South Dakota turfgrass is valuable as a major contributor to the state’s economy in addition to the aesthetic, functional, and recreational beneﬁts it provides.

LITERATURE CITED

Agric. Res. Serv. 2002. Priority turf research needs: document one—initial feed- back product for customers. Turf customer workshop. Dallas, TX, Jan. 22- 25, 2002. Beard, J.B. 1973. Turfgrass science and culture. Prentice-Hall, Englewood Cliffs, NJ. Beard, J.B. and R.L. Green. 1994. The role of turfgrasses in environmental protection and their benefits to humans. J. Environ. Qual. 23:452-460. Dep. of Hortic. 2002. O.J. Noer research and education facility [Online]. Univ. of Wisconsin. Available at http://www.hort.wisc.edu/uwturf/pages/ojno- er.htm (verified 10 June 2002). Eclov, M. 2000. UK College of Ag reaches urban audiences through turf topics [Online]. Univ. of Kentucky website. Available at http://www.ca.uky.edu/age/news/2000/Feb/turf2.htm (verified 11 June 2001). Fisher, A. 2001. Approaching the green. Perspectives on line [Online]. College of Agric. and Life Sci., North Carolina Univ. Available at http://www.cals.ncsu.edu/agcomm/magazine/spring01/approach.htm (verified 10 June 2002). Fleishman-Hillard Research. 1999. Turfgrass in Missouri. Fleishman-Hillard, Inc., St. Louis, MO. Hortic. and Landscape Architecture Dep. 2002. Huffine presidential distinguished scholarships—brochure [Online]. Oklahoma State Univ. Available at http://osu-ns03.cis.okstate.edu/tools/webtools.nsf/Images/hortschol- ar/$FILE/huffine.pdf (verified 11 June 2002). Iowa Agric. Statistics Serv. 2001. Iowa’s turfgrass industry. Natl. Agric. Statistics Serv., Des Moines, IA. Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 79

Murphy, B., C. Hayes, M.S. Phipps, R. Bosecker, K. Neas, and H. Smith. 1999. 1999 North Carolina Turfgrass Survey. North Carolina Agric. Statistics Serv., North Carolina Dep. of Agric. and Consumer Serv., Raleigh, NC. National Golf Foundation. 2001. Golf facilities in the U.S. 2001 edition. Jupiter, FL. Partially available on-line at http://www.ngf.org/faq/ (verified 11 June 2002). Natl. Resources Inventory. 2000a. Broad land cover/use, by state, 1997 [Online]. Available at http://ftp.nhq.nrcs.usda.gov/pub/outgoing/land/data/m5150d.txt (verified 10 June 2002). Natl. Resources Inventory. 2000b. Developed land, by state, 1997 [Online]. Available at http://ftp.nhq.nrcs.usda.gov/pub/outgoing/land/data/m5803d.txt. (verified 10 June 2002). Pennsylvania Agric. Statistics Serv. 1990. Pennsylvania 1989 Turfgrass Survey. Pennsylvania Dep. of Agric., Harrisburg, PA. Rodekohr, J.I. 2000. The beauty of tough turf. UGA Research Reporter, Univer- sity of Georgia 29(1): Summer 2000. Schoettle, A. 2002. Turfgrass research center turns course maintenance into a science. Indianapolis Business J. 22 (52): Mar. 4-10. South Dakota Agric. Statistics Serv. 2001. Crop data downloads 1882-2000 [On- line]. South Dakota Dep. of Agric. Available at http://www.nass.usda.gov/sd/dnlds/dnld_c.htm (verified 12 June 2002). U.S. Bur. of the Census. 2000. Profile of general demographic characteristics: 2000 [Online]. Available at http://censtats.census.gov/cgi-bin/pct/pctPro- file.pl (verified 10 June 2002). Vinlove, F.K. and R. F. Torla. 1995. Comparative estimations of U.S. home lawn area. J. Turfgrass Manage. 1(1):83-97. Watson, J.R., Kaerwer, H.E., and D.P. Martin. 1992. The turfgrass industry. In D.V. Waddington et al. (ed.) Turfgrass. Amer. Soc. of Agron., Madison, WI.

Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 81

CHONDRICHTHYES FROM THE UPPER PART OF THE MINNELUSA FORMATION (MIDDLE PENNSYLVANIAN: DESMOINESIAN), MEADE COUNTY, SOUTH DAKOTA

D.J. Cicimurri Bob Campbell Geology Museum Clemson University Clemson, SC 29634

M.D. Fahrenbach Geological Survey Program South Dakota Department of Environment and Natural Resources Rapid City, SD 57702

ABSTRACT

Numerous teeth, denticles, and scales of selachians and holocephalans have been recovered from an outcropping of the upper Minnelusa Formation in Meade County, South Dakota. Specimens include the denticles Petrodus patelliformis and Listracanthus histrix, the scale Holmesella quadrata, and teeth of Caseodus aff. C. eatoni, Edestus sp., cf. "Cladodus" sp., and Janassa sp. Associated conodont elements of Idiognathodus “delicatus” and Idioprioniodus conjunctus indicate a middle Pennsylvanian (Desmoinesian) age. A similar chondrichthyan assemblage has been reported from temporally equivalent rocks of Arkansas, Kansas, Missouri, Illinois, Oklahoma, and Iowa.

Keywords

Minnelusa Formation, Pennsylvanian, Chondrichthyes, South Dakota

INTRODUCTION

The material presented in this report was obtained during ﬁeld trips conducted by the Department of Geology and Geological Engineering, South Dakota School of Mines and Technology, Rapid City, South Dakota. Matrix was collected to recover conodont fossils, although it had been known that fossil vertebrates occurred at the locality (Elder 1993). The outcrop is located on the north side of Little Elk Creek, approximately 2.5 km west of Piedmont, and approximately 30 km north of Rapid City (Fig. 1). Fossils were recovered from green and gray silty shale referred to as the Petrodus II bed by Elder (1993) (Fig. 2). This shale occurs approximately 0.7 m above the Petrodus I bed (Elder 1993) and consists of 1-2.5 cm of light green 82 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002)

shale underlying 10–15 cm of dark gray shale. The shale unit is slightly calcareous, blocky, and moderately indurated, becoming soft and fissile where weathered. The upper and lower contacts are wavy and irregular. Selachian fossils have been found throughout the green shale, with concentrations in de- pressions along the lower contact of the green shale and an underlying dolomite. Fossils appear to be less abundant in the overlying dark gray shale. The lower, irregular contact represents a disconformity between the shale and dolomite, and concentrations of fossils may represent a lag deposit. The light green color is due to Figure 1. Generalized geologic map of the Black Hills region showing surface outcrops of the Min- secondary reduction of iron nelusa Formation (stippled area). The large star in- resulting from water move- dicates the Little Elk Creek locality. Modified from ment through fractures and Darton (1951) and Elder (1993). along bedding contacts. Unfortunately, the Petrodus II shale bed is now difficult to sample because weathering and previous collecting efforts for conodonts have resulted in the shale being removed up to 0.75 m into the outcrop, with the adjacent limestone and dolomitic limestone beds restricting access.

GEOLOGIC SETTING

Elder (1993) proposed that during periods of higher sea level, the remains of deceased animals fell to the bottom and were worked by higher energy currents, which would account for the lag deposit that is thought to be present at the base of the Petrodus II bed. However, the teeth and denticles do not show signs of abrasion that would be expected if they were subjected to such actions. Based on associated lithologies, it is believed that the Petrodus shale beds were deposited under reducing conditions in a semi-restricted lagoonal environment, and in close proximity to tidal ﬂats. The presence of delicate Listra- Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 83

Figure 2. Stratigraphic section of a portion of the Minnelusa For- mation exposed at Little Elk Creek, Meade County, South Dakota. See following for lithologic descriptions: 13) Limestone, 90-120 cm thick, with abundant coarse, rounded, quartz sand. Contains many hollow and calcite-lined vugs and boxwork structures. Thin chert lenses and sub-angular chert clasts occur near the base. Very irregular upper contact. 12) Limestone, 17.5-22.5 cm thick, light gray to brown, very fine-grained. Contains abundant chert lenses and sub-angular chert clasts; minor quartz sand. Flat upper contact. 11) Limestone, 5 cm thick, white to beige, very fine grained, chalky. Contains lenses of coarse, well-rounded sand and sub-angular chert clasts; some lenses cut into the subjacent unit. Moderately irregular upper contact. 10) Dolomite, 32.5 cm thick, light beige, medium- to coarse- grained. Contains abundant moldic texture and calcite-lined vugs to 20 cm diameter. 9) Limestone, 32.5 cm thick, white to whitish gray, very fine grained. Contains very fine moldic texture. A 0.5-1 cm thick green shale occurs along the irregular upper contact. Massive with angular, blocky weathering. 8) Shale and silty shale, 20 cm thick, black to gray with thin, reddish to maroon laminae and associated limonite nodules to 1 cm diameter, especially in the upper half. Fissile. Thin purple lamina- tions also present. Lower 4 cm of unit is greenish yellow fissile shale, thickest in low spots along the irregular lower contact. Irregular, wavy upper contact. Equivalent to the Petrodus II bed of El- der (1993). 7) Dolomitic limestone, 7.5-30 cm thick, beige to tan, fine- to medium-grained. Contains moldic texture of crinoid columnals and brachiopod fragments. Lobate and irregular, hollow to calcite lined vugs to 30 cm diameter occur throughout. Irregular, wavy upper contact. Massive. 6) Limestone, 40 cm thick, beige to orange gray, very fine grained. Contains planar stromatolites with crenulated laminae weathering in relief. Irregular, wavy upper contact. Flaggy, angular weathering. 5) Shale, 5 cm thick, gray with purple staining. Moderately irregular upper contact. Fissile to flaggy weathering. Equivalent to the Petrodus I bed of Elder (1993). 4) Dolomitic limestone, 35 cm thick, upper half beige, lower half grayish white, very fine grained. Fine planar stromatolites occur in the upper half; lacks fine laminae in the lower half. Stylolite surfaces separate some beds. Thin orange chert lenses and sub-angular chert clasts occur along the moderately irregular upper and lower contacts. 3) Dolomitic limestone, 25 cm thick, beige to light tan, very fine grained. Fine planar stromatolites occur in the upper 5-7.5 cm. Upper contact moderately irregular. 2) Shale, 5 cm thick, gray with maroon staining. Fissile. Irregular upper contact. 1) Dolomitic limestone, 15 cm thick, gray with maroon staining, very fine-grained. Massive, dense. Moderately flat upper contact, lower contact covered. 84 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) canthus denticles and slightly disturbed conodont assemblages in the shale beds indicate very low energy conditions, with minimal current action or periods of quiescence. The fragile spines and conodont assemblages would not have survived intact if subjected to strong currents or reworking. The shale beds probably represent periods of very slow clastic deposition between cycles of carbonate deposition, possibly related to cyclothemic sequences or a maximum flooding episode during a highstand system tract.

METHODS

Matrix was disaggregated using a solution of 1% acetic acid in water. The concentrate was placed in water to remove calcium acetate residue, then screened through 5 mm, 3 mm, and 1 mm mesh sieves. The remaining material was dried and sorted through a binocular microscope. Specimens that had been broken were repaired with thinned Butvar (B-76 in acetone). The classiﬁcation scheme follows Zangerl (1981). All referred specimens are housed in the Bob Campbell Geology Museum (BCGM), Clemson Univer-

Figure 3. A: Listracanthus histrix denticle, BCGM 2935; B: Petrodus patelliformis denticle, BCGM 4425, dorsal view; C and D: Holmesella quadrata scales, BCGM 4747 and BCGM 4748, dorsal views with anterior at right. Scale lines = 1mm. Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 85 sity, South Carolina. Additional specimens are housed at the Museum of Ge- ology (SDSM), South Dakota School of Mines and Technology. Precise sample locality information is on ﬁle at both institutions. The specimens are the prop- erty of the U.S. Forest Service.

SYSTEMATIC PALEONTOLOGY

Chondrichthyes Huxley, 1880 Holocephali Bonaparte, 1832-41 Cochliodontiformes Obruchev, 1953 Menaspidae Woodward, 1891 Listracanthus histrix Newberry and Worthen, 1870 Figure 3, A

Referred specimens: BCGM 2934, isolated denticle in matrix; BCGM 2935, isolated denticle in matrix. Additional specimens: SDSM 26057, one denticle; SDSM 26058, one denticle with four associated Petrodus. Description: BCGM 2934 and BCGM 2935 are compound denticles consisting of an elongate base supporting six or seven curved, needle-like spines arranged in a row, forming a comb-like structure. Both specimens are preserved as impressions on the matrix, with some of the original material remaining. Discussion: Zangerl (1981) stated that Listracanthus is a common component of Upper Carboniferous rocks in the Midwestern United States, where it is often associated with Petrodus. Chorn and Reavis (1978) and Zangerl (1981) noted that Listracanthus and Petrodus denticles occurred on different types of chondrichthyans, and were therefore merely form genera. Listracanthus-type denticles were described by Patterson (1965) and Moy-Thomas (1936) as occurring on the dorsal surface of Deltoptychius (illustrated by Stahl 1999).

Euselachii Hay, 1902 Hybodontoidea Zangerl, 1981 Hybodontidae Owen 1846 Petrodus patelliformis M’Coy, 1848 Figure 3, B

Referred specimens: BCGM 2932, 170 incomplete denticles; BCGM 2933, 90 denticle fragments; BCGM 2950, 30 denticles; BCGM 4425, one denticle. Additional specimens: SDSM 26051, one denticle; SDSM 26052, one denticle, SDSM 26053, 125 denticles. Description: This type of denticle ranges in size from 2-11 mm in diameter, and 1–9 mm in height. They are circular to elliptical in outline. The crown of small to medium-sized denticles has a conical cross-section, pointed, smooth apices, and coarse radiating ridges that bifurcate basally. Larger denticles have fewer but more massive bifurcating ridges that coalesce at the apex, forming a large rounded surface. The base of the denticle extends outwards a short distance from the crown, and the attachment surface may be ﬂat, weakly convex, or concave. 86 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002)

Discussion: According to Zidek (1973), North American occurrences of Petro- dus were typically referred to P. occidentalis. When originally described, it was with some reservation that P. occidentalis was considered as being distinct from P. patelliformis M'Coy 1848 (Newberry and Worthen 1866). We assign the Minnelusa specimens to P. patelliformis because this speciﬁc designation has seniority over P. occidentalis. Petrodus is the dominant vertebrate fossil at the Little Elk Creek locality. This type of denticle morphology is not necessarily unique to a single taxon, and for this reason Chorn and Reavis (1978) considered the generic designation a nomen dubium. Petrodus-like denticles were described by Patterson (1965) as occurring on the body of Deltoptychius, but Zangerl (1981) suggested that Petrodus was a single, very large taxon whose skin was covered ex- clusively with this type of denticle. Chorn and Reavis (1978) described a mat of Petrodus (a cluster of articulated denticles) in which smaller denticles were intermingled with much larger ones, and the diversity in size of the denticles therefore does not necessarily indicate populations containing juvenile and adult individuals.

Protacrodontoidea Zangerl, 1981 Tamiobatidae Glickman, 1964 Holmesella quadrata Gunnel, 1931 Figure 3, C and D

Referred specimens: BCGM 2931, 8 scales; BCGM 4747, scale; BCGM 4748, scale. Description: This type of scale typically has a rhombohedral shape, although one specimen having a hexagonal outline was collected. The long axis ranges from 2–4 mm and the short axis ranges from 1.5–3 mm. The height varies between one and two millimeters. The crown is very low, ﬂat, or weakly concave, and highly ornamented with tubercles and/or longitudinal ridges. The base of the scale is much thicker and wider than the crown, having a convex basal surface. Concentric zonation is visible on the basal surface of some specimens. Discussion: The scales described above are identical in morphology to scales of Holmesella in the collections of the Field Museum of Natural History (FMNH), Chicago (PF 2367). Zangerl (1981) noted that this type of scale is common in black carbonaceous shales of the Midwestern United States. Mer- tiniene (1999) described the histology of Holmesella as consisting of a primor- dial odontode surrounded by later odontodes. The odontodes are composed of mesodentine, whereas the base consists of acellular bone tissue. As with Listracanthus and Petrodus, this type of scale may not be characteristic of any one selachian taxon and can be considered a form genus (Mer- tiniene 1999). Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 87

Eugeneodontida Zangerl, 1981 Edestoidea Hay, 1930 Edestidae Jaekel, 1899 Edestus sp. Agassiz, 1833 Figure 4, A

Referred specimen: BCGM 2938, symphyseal tooth with associated root fragment. Description: The tooth has a very laterally compressed, labially inclined crown with nearly flat faces. Cutting edges have large, apically directed serrations that become finer toward the apex. These larger serrations are themselves more finely serrated. The root is fragmentary but appears to have been elongated, extending posterior- ly. Discussion: BCGM 2938 is part of a symphyseal tooth whorl, which was an unusual feature of several Pa- leozoic shark taxa Figure 4. A: Edestus sp. symphyseal tooth, BCGM (Zangerl,1981). The basal 2938, lateral view with anterior at right; B: cf. edge of the root of BCGM "Cladodus" sp. tooth fragment, BCGM 2936, lin- 2938 forms a sharp ridge gual view; C: Janassa sp. incomplete tooth, BCGM that would sit in a dorsal 2940, labial view; D: Janassa sp. incomplete tooth, BCGM 2939, labial view. Scale lines = 1 mm in B- groove in the root of the D, 5 mm in A. preceding tooth, a characteristic of some edestoid sharks (particularly Edestus) rather than agassizodontid sharks. When viewed anteriorly, agassizodontid symphyseal teeth are bifurcated basally (Zangerl 1981). BCGM 2938 is not as heavily serrated as specimens of Edestus minor and E. heinrichi in the FMNH collection (UC 2092 and UC 14345 respectively), but the weak serrations and small size probably reflects the anterior position of the tooth in the whorl (i.e. older part). Zangerl (1981) reported this taxon from Upper Carboniferous (Pennsylvanian) rocks of the United States, parts of Europe, and Russia. 88 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002)

Caseodontoidea Zangerl, 1981 Caseodontidae Zangerl, 1981 Caseodus aff. C. eatoni Zangerl, 1981 Figure 5, A - D

Referred specimens: BCGM 2942, 16 tooth fragments; BCGM 2943, incomplete anterior tooth; BCGM 2944, incomplete anterior tooth; BCGM 2945, incomplete anterior tooth; BCGM 2946, lateral tooth; BCGM 2947, incomplete lateral tooth; BCGM 2948, incomplete posterior tooth, BCGM 2949, incomplete posterior tooth. Description: Anterolateral teeth are mesiodistally elongated with low crown. There is a central cusp, as well as one or two indis- tinct lateral cups. The labial crown face is convex, the lingual face is ﬂat to convex, with crenulations occurring on both the labial and lingual faces. Large longitudinal protuberances are found on the labial Figure 5. Caseodus aff. C. eatoni - A: BCGM 2493, face, with corresponding incomplete anterior tooth, lingual view; B, BCGM smaller protuberances on 2944, anterior tooth, lingual view; C, BCGM 2946, the lingual face. An unser- incomplete lateral tooth, lingual view; D, BCGM rated cutting edge is con- 2948, posterior tooth, lingual view. Scale lines = 5 mm in A and C, 1 mm in B and D. tinuous along the entire mesiodistal width of the crown, with smaller transverse cutting edges extending out onto the labial and lingual buttresses. An unusual feature is the alter- nating nature of short vertical ridges that are perpendicular to the cutting edges (see Fig. 5A). The ends of the crown are rounded and there is a weak central bulge on the labial face. Lateral teeth are similar to anterior teeth except that the main cusp is lower, situated more distally on the crown, and distally inclined. Posterior teeth are very small, having a very low, straight crown with rounded ends. There are no cusps, and the cutting edge is unserrated and Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 89 straight across the crown. The labial and lingual crown faces are only slightly convex, and crenulation is much reduced. Labial protuberances are only weakly developed, and lingual protuberances are nearly absent. Though roots are incomplete, they are nearly as long as the crown, labio- lingually compressed, with numerous vertical ridges and grooves. Discussion: Unfortunately all of the teeth are incomplete, making precise taxonomic assignments difﬁcult. However, they are similar to several euge- neodontid genera, namely Agassizodus, Bobbodus, Campodus, and Caseodus. In Agassizodus, the crowns are arched (Zangerl 1981, Case 1982), which is a feature not seen in the teeth of our sample. The Minnelusa specimens have smaller labial buttresses, and the vertical ridges on the cutting edge are stronger than in Bobbodus (Zangerl 1981, Schultze and West 1996). In Campodus, the labial and lingual buttresses are nearly the same size, giving the teeth a symmetrical appearance in occlusal view (Zangerl 1981). The teeth in our sample have much larger labial buttresses compared to those on the lingual face. Based on these comparisons, we assign our specimens to the genus Caseodus. Of the two species, Caseodus eatoni and C. basalis, the large lateral teeth in our sample compare most closely with the ornamented teeth of C. eatoni. Both taxa are known from the Desmoinesian Series and could theoretically occur together (Zangerl 1981). However, we regard the smaller, simpler teeth in our sample as distal lateral and posterior teeth of C. eatoni, rather than C. basalis.

Petalodontida Zangerl, 1981 Janassidae Munster, 1839 Janassa sp. Figure 4, C and D

Referred specimens: BCGM 2939, incomplete tooth; BCGM 2940, incomplete tooth; BCGM 2941, incomplete tooth. Description: The teeth are very labio-lingually compressed with a weakly convex labial face and slightly concave lingual face. There may be some lingual flexure of the crown. The cutting edge is smooth and weakly to strongly convex. Fine longitudinal striations are found on the lingual face, and there are very short longitudinal grooves on the labial face that are restricted to an area just below the cutting edge. Discussion: Schaumberg (1979) presented a ray-like reconstruction of Janas- sa, with a ventral mouth and large, expanded pectoral fins. Zangerl (1981) reported Janassa from Permian rocks of Germany and Greenland, and documented a Mississippian to Permian range for the taxon in the United States and Britain. The teeth in our sample compare favorably with those of Janassa bi- tuminosa figured by Case (1982) from Upper Pennsylvanian rocks of Nebras- ka. However, with such a small sample size (n=3) and incomplete specimens, we hesitate to make a more specific identification. 90 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002)

Symmoriida Zangerl, 1981 cf. "Cladodus" sp. Figure 4, B

Referred specimens: BCGM 2936, incomplete central cusp; BCGM 2937, incomplete tooth. Additional specimens: SDSM 26054, tooth base with two tooth fragments; SDSM 26055, incomplete tooth. Description: BCGM 2936 is an incomplete central cusp having a nearly flat labial face and very convex lingual face. The lower portion of the labial face possesses very fine longitudinal striations, whereas coarse striations extend nearly the entire height of the lingual face. Cutting edges are smooth. BCGM 2937 is a tooth fragment preserving part of the root and the bases of two lateral cusplets. The cusplets have an elliptical cross-section and several longitudinal striations on the labial face. The root is shelf-like and extends lingually beyond the lateral cusplets. Discussion: Zangerl (1981) considered Cladodus a nomen dubium because several distinct Paleozoic shark taxa possessed a cladodont dentition. Our specimens are so fragmentary that it is difficult to make even a generic identification.

DISCUSSION

The present chondrichthyan assemblage of the Minnelusa Formation consists of at least seven taxa including Listracanthus, Petrodus, Holmesella, Edes- tus, Caseodus, Janassa, and "Cladodus". We have also found small fragments of fin spines that are distinctly channeled and could be referrable to Acondy- lacanthus St. John and Worthen, 1875 or Ctenacanthus Agassiz, 1835. More complete material is needed for a precise taxonomic assignment. Elder (1993) illustrated a tooth that was tentatively identified as Cranodus. Unfortunately the specimen could not be located, and the illustration is not clear enough to confirm the identification. Semi-articulated conodont assemblages and the good preservation of delicate Listracanthus denticles indicate that deposition of the Petrodus II bed took place in relatively quiet water. The apparent lag deposit at the base of this bed may be due to scouring or dissolution of the underlying dolomite beds during an initial transgression, followed by very slow deposition of fine-grained sediment and selachian remains as sea level rose. The sandstone-shale-carbonate "cycles" at the Little Elk Creek locality may be related to widespread cyclothemic deposition occurring during the Pennsylvanian. Conodants associated with the chondrichthyan remains, such as Idioprioniodus conjunctus, Neog- nathodus roundyi-dilatus plexus, and Idiognathodus “delicatus” morphotypes, indicate a Middle Pennsylvanian, middle to late Desmoinesian age (Rowe and Fox 1997). Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 91

ACKNOLWEDGEMENTS

We thank the U.S. Forest Service for allowing us to study the fossils described in this report, and James Foxx for providing information pertaining to associated conodonts. Rainer Zangerl was kind enough to examine the teeth and denticles, and Marjorie DeCocker illustrated the specimens seen in Fig- ures 3, 4, and 5.

REFERENCES

Case, G.R. 1982. A Pictorial Guide to Fossils. Van Nostrand Reinhold Company, New York, 514 p. Chorn, J., and E.A. Reavis. 1978. Affinities of the chondrichthyan organ-genera Listracanthus and Petrodus. University of Kansas Contributions to Paleon- tology 89:4-9. Darton, N.H. 1951. Geologic Map of South Dakota. U.S. Geological Survey. Elder, R.A. 1993. The first occurrence of vertebrates, represented by the Paleo- zoic sharks Petrodus, Listracanthus, Cladodus, and Cranodus (?) from the Pennsylvanian-Permian Minnelusa Formation near Piedmont, South Dako- ta. Proceedings of the South Dakota Academy of Science 72:221-233. Gries, J.P., and J.E. Martin. 1985. Composite section of the Paleozoic and Meso- zoic strata in the Black Hills and surrounding areas. 261-292 in F.J. Rich (ed.), Geology of the Black Hills, South Dakota and Wyoming, Geological Society of America Field Trip Guidebook, Rocky Mountain Section. Mertiniene, R. 1999. Structure of scales of the "Holmesella" morphological type, Ichthyolith Issues Special Publication 5:34. Moy-Thomas, J.A. 1936. The structure and affinities of the fossil elasmobranch fishes from the Lower Carboniferous rocks of Glencartholm, Eskdale. Pro- ceedings of the Zoological Society of London 1936:761-788. Newberry, J.S., and A.H. Worthen. 1866. Description of vertebrates. Geological Survey of Illinois 2(1):10-134. Patterson, C. 1965. The phylogeny of chimaeroids. Transactions of the Royal Philosophical Society of London 249:101-219. Rowe, B.L. and J.E. Fox. 1997. Conodont assemblages and element morphotypes relating to depositional environments of the Pennsylvanian Min- nelusa Formation, Black Hills, South Dakota (abstract). 95 in Geological Society of America, 1997 annual meeting, Abstracts with Programs—Geo- logical Society of America 29(6). Schaumberg, G. 1979. Neue Kenntnisse uber die Anatomie von Janassa bitu- minosa (Schlotheim), Holocephali, Chondrichthyes aus dem Permischen Kupferschiefer. Palaontologie Z. 5(3 and 4):334-346. Schultze, H., and R.R. West. 1996. An eugeniodontid elasmobranch from the late Paleozoic of Kansas. Journal of Paleontology 70(1):162-165. St. John, O.H. and Worthen, A.H. 1875. Description of fossil fishes. Geological Survey of Illinois 6(2):245-488. Stahl, B.J. 1999. Chondrichthyes III: Holocephali, In: Schultze, H.P. (ed.), Hand- book of Paleoichthyology 3C, Gustav Fischer Verlag, 164 p. 92 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002)

Zangerl, R. 1981. Chondrichthyes I: Paleozoic Elasmobranchii, In: Schultze, H.P. (ed.), Handbook of Paleoichthyology 3A, Gustav Fischer Verlag, 115 p. Zidek, J. 1973. Oklahoma Paleoichthyology Part II: Elasmobranchii (Cladodus, Minute Elements of Cladoselachian Derivation, Dittodus, and Petrodus). Oklahoma Geology Notes 33:87-103. Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 93

INFLUENCE OF AGE-0 YELLOW PERCH ABUNDANCE ON WALLEYE POPULATIONS IN TWO EASTERN SOUTH DAKOTA LAKES

Jonathan R. Meerbeek, Daniel A. Isermann, and David W. Willis Department of Wildlife and Fisheries Sciences, South Dakota State University, Brookings, SD 57007

ABSTRACT

Eastern South Dakota lakes typically range from mesotrophic to hypereutrophic based on the trophic state index, and prey supplies, whether they be macroinvertebrates or fishes, tend to be abundant. Thus, biologists often believe that quality of sport fishing is more closely linked to interannual variation in recruitment (i.e., year-class strength) than to prey availability for sport fishes. Here, we document two case histories that challenge this premise. We noted significant (P ≤ 0.002) increases in age-0 yellow perch (Perca flavescens), a common prey for walleye (Stizostedion vitreum), abundance between 2000 and 2001 in Sinai and Waubay lakes. For example, larval yellow perch abundance (number/100 m3) was 723 times higher in 2001 than 2000 in Lake Sinai and 46 times higher in Lake Waubay. Similarly, juvenile yellow perch abundance as indexed by seine hauls increased by a factor of 251 in Waubay Lake and a factor of 385 in Lake Sinai between 2000 and 2001. In conjunction with these increases in age-0 yellow perch as available prey, walleye condition factors significantly (P ≤ 0.004) increased in 2001 compared to 2000 for walleyes ≤51 cm in both lakes. In addition, age-0 walleye growth rates increased, but not always significantly. Based on these two examples, we suggest that prey availability may be more important to sport fish population structure and dynamics in eastern South Dakota lakes than previously believed.

INTRODUCTION

The natural lakes of glacial origin in eastern South Dakota range from mesoeutrophic to hypereutrophic, based on the Carlson trophic state index (Stueven and Stewart 1996). Thus, biologists often suspect that the lakes provide an abundant food supply (either invertebrates or fishes) to support fish communities. For example, chironomid densities have been measured as high as 55.6 million/ha (Lott 1991). Various life stages of walleye (Stizostedion vitreum), from larvae to adults, often consume a large proportion of zooplankton and macroinvertebrates when invertebrate densities are sufficiently high (Isaak et al. 1993; Blackwell et al. 1999; Starostka 1999). Several studies have related prey abundance to walleye recruitment, growth, and condition (Forney 1977; Knight et al. 1984; Marwitz and Hubert 94 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002)

1997; Porath and Peters 1997). The abundance of juvenile yellow perch (Per- ca flavescens) has frequently been related to walleye recruitment and growth (Forney 1974, 1976; Madenjian 1991; Marwitz and Hubert 1997). Annual variability in juvenile walleye growth was directly related to size of available yellow perch (Madenjian 1991; Marwitz and Hubert 1997). Previous studies suggest that management of walleye populations should be based solely on prey production rather than increased recruitment or stocking because intraspecific competition among individuals limits walleye growth and survival (Forney 1976; Swenson 1977). Inter-annual variation in recruitment (i.e., strong and weak year classes) and annual high food productivity has previously been ac- cepted as the determinant of sport fish quality in eastern South Dakota glacial lakes. However, the effect of age-0 yellow perch abundance on walleye recruitment and growth has not been assessed. Thus, the objective of this study was to determine the relationship between age-0 yellow perch abundance and walleye growth and condition in two South Dakota lakes.

METHODS

Larval yellow perch were collected at approximately weekly intervals during late May and early June of 2000 and 2001 using a 1,000-µm mesh conical ichthyoplankton net with a 0.75-m diameter circular steel frame and a 500-ml bucket. Nets were towed from a 10-m lead in a semicircular pattern for 2-5 min at an average speed of 0.5 m/s. A flowmeter (General Oceanics, model 2030R) mounted in the center of the frame estimated the amount of water volume fil- tered. Larval yellow perch catches (number/100 m3) were indexed on the date of highest abundance. Sampling sites in both Waubay Lake (20 sites/day) and Lake Sinai (10 sites/day) were random, and remained fixed throughout the study. Samples were preserved in a 10% formalin solution and were taken to the lab for examination. A dissecting microscope and the criteria provided by Auer (1982) were used to identify larval fishes. During the daylight hours of August 2000 and 2001, a 15.2-m bag seine with a 6-mm bar mesh was used to index juvenile yellow perch abundance (20 hauls at Sinai and 40 hauls at Waubay). Catch per unit effort (CPUE) was defined as the mean number of age-0 yellow perch caught per 90º arc. Randomly selected fixed sites were sampled each year. Juvenile yellow perch were counted and then released. Walleye populations were sampled in 2000 and 2001 using experimental gill nets during mid-July at Lake Sinai and mid to late August at Waubay Lake. Experimental monofilament gill nets were composed of six 7.6 m panels (13, 19, 25, 32, 38, and 51 mm bar measure). Lengths and weights of all walleyes were measured at Waubay Lake, while only a subsample was measured at Lake Sinai. Walleye condition was analyzed using relative weight (Wr; fish weight divided by a "standard" weight for a fish of that length and then multiplied by 100) using the standard weight equation provided by Murphy et al. (1990). Mean Wr values were compared between years at each lake using length categories recommended by Gabelhouse (1984; <25 cm, 25-37 cm, 38-50 cm, and 51-62 cm). Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 95

Night electrofishing using pulsed-DC current was used to sample juvenile walleyes in October of both years in each lake. Scales were collected from walleyes to verify those designated as age 0. Mean length at time of capture for age-0 walleyes was compared between years by water. All comparisons were conducted with SAS software (SAS Institute 1988) with a significance level of α = 0.05. Yellow perch larval and juvenile abundance data did not meet the assumption of homogeneity of variance; peak larval abundance and seining CPUE were compared using a Wilcoxon signed rank test for paired sample data (SAS NPAR1WAY procedure). Two factor analysis of variance (ANOVA) for unbalanced data (GLM procedure) was used to compare mean Wr values with year and length group as main effects. When the two-way ANOVA exhibited a significant interaction between year and length group, mean Wr was compared between years by length group using a the TTEST procedure and the appropriate test for equal or unequal variances. Similarly, age-0 walleye growth (i.e., mean length at time of fall capture) was initially compared between years and lakes using a two-way ANOVA, but the interaction term was significant. Thus, we again used the TTEST procedure to compare age-0 growth between years by lake.

RESULTS

Substantial differences in interannual variation in age-0 yellow perch abundance were observed between 2000 and 2001 (Table 1). Highest larval yellow perch abundance occurred on 21 May 2000 and 24 May 2001 in Lake Sinai, and

Table 1. Yellow perch (YEP) abundance indices and mean length at time of capture for walleyes (WAE) collected by fall night electroﬁshing from Lakes Sinai and Waubay, South Dakota. Larval abundance was estimated using an ichthyoplankton trawl during late May/early June, while juvenile relative abundance was indexed using a bag seine in August. N = units of sampling effort; CPUE = catch per unit effort; SE = standard error the mean.

Index Lake Year N Mean SE

Larval YEP abundance Sinai 2000 10 1.9/100 m3 0.8 2001 10 3,097.9/100 m3 2,914.5 Waubay 2000 20 4.8/100 m3 1.5 2001 20 224.1/100 m3 47.9

Juvenile YEP CPUE Sinai 2000 20 2.7/haul 1.3 2001 20 1,019.2/haul 931.5 Waubay 2000 40 0.3/haul 0.1 2001 39 225.5/haul 105.0

Mean length age-0 WAE Sinai 2000 10 162.3 mm 2.0 2001 78 171.0 mm 2.9 Waubay 2000 14 153.6 mm 3.0 2001 6 173.5 mm 12.2 96 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002)

25 May 2000 and 6 June 2001 in Waubay Lake. Larval abundance was significantly higher in 2001 than in 2000 in both Sinai (S = -26.5, P = 0.004) and Waubay (S = -105.0, P = 0.0001) lakes, increasing 723 times between 2000 and 2001 in Lake Sinai, and by a factor of 46 between 2000 and 2001 in Waubay Lake. Similarly, juvenile yellow perch abundance was significantly higher in 2001 than in 2000 for both Sinai (S = -105.0, P = 0.0001) and Waubay (S = - 144.5, P = 0.0001) lake seine samples. Mean CPUE for juveniles increased by a factor of 385 in Lake Sinai and 251 in Waubay Lake between 2000 and 2001. Condition indices were higher for walleyes in 2001 than in 2000 for most length groups in both lakes (Table 2). In Lake Sinai, there was no significant interaction between year and length group (F = 0.95, df = 2, P = 0.39). The two-way ANOVA indicated that mean Wr values were significantly higher in 2001 than 2000 (F = 8.97, df = 1, P = 0.004), while there was no difference among length groups (F = 0.10, df = 3, P = 0.96). For Waubay Lake walleyes, we found a significant interaction between year and length group (F = 16.65, df = 3, P = 0.0001); thus, we compared mean Wr values by length group using a one-way ANOVA. Mean Wr values for <25 cm (t = -5.41, df = 70, P = 0.0001), 25-37 cm (t = -8.0, df = 28.7, P = 0.0001), and 38-50 cm (t = -4.68, df = 27.2, P = 0.0001) walleyes were significantly higher in 2001 than 2000. Only for the largest length group, 51-62 cm, was there no difference in mean Wr between years (t = -0.24, df = 32, P = 0.81). Although sample size by length group often was low (Table 2), especially in Lake Sinai, condition indices were consistently higher in 2001 than 2000 for walleyes ≤51 cm.

Table 2. Mean Wr by length group for walleyes collected from Lake Sinai on 10-11 July of 2000 and 2001, and from Waubay Lake on 15-18 August 2000 and 21-24 August 2001. SE = standard error of the mean.

2000 2001

Lake Length group (cm) N Mean Wr (SE) N Mean Wr (SE)

Sinai <25 43 84 (2.1) 5 97 (1.2) 25-37 2 79 (4.5) 50 98 (0.9) 38-50 3 88 (1.7) 3 93 (2.5) 51-62 2 87 (2.0) 0

Waubay <25 70 85 (0.4) 2 99 (4.0) 25-38 29 84 (1.4) 229 95 (0.2) 38-51 44 85 (0.4) 25 93 (1.7) 51-62 20 89 (0.9) 14 89 (1.7)

In Lake Sinai, age-0 walleye growth was significantly faster (t = -2.49, df = 56.4, P = 0.02) in 2001 than in 2000 (Table 1). In Waubay Lake, age-0 walleye growth was not significantly different (t = 01.57, df = 5.6, P = 0.17) between years; however, the mean length of captured fish was 20 mm longer in 2001 than in 2000. Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 97

DISCUSSION

Yellow perch recruitment (i.e., year-class strength) in South Dakota glacial lakes tends to be inconsistent. Several environmental (water levels, spawning habitat, water temperature, wind, currents) and biological (food availability and size, predation rates) variables are important in determining the success or failure of year classes (Nelson and Walburg 1977; Whiteside et al. 1985; Hartman and Margraf 1993). Our data, along with data from six other eastern South Dakota lakes (Anderson et al. 1998), suggest that the number of age-0 yellow perch caught per seine haul in the summer was positively related to the number of larval perch collected in ichthyoplankton tow nets. This potential ability for fishery biologists to index yellow perch recruitment through larval abundance (mid-May to early June) allows for 1) an understanding of the available prey for predators, as well as 2) the possibility of implementing stocking programs as suggested by Anderson et al. (1998) to enhance perch year-class strength during years of expected low recruitment. When age-0 yellow perch are abundant, walleyes often extensively prey on them, which can allow enhanced survival of young walleye (Forney 1974). Hartman and Margraf (1993) found that even in large systems such as Lake Erie, intense walleye predation on age-0 yellow perch could cause low recruitment of yellow perch. In Enemy Swim Lake, South Dakota, walleye food habit analysis suggested that even where macroinvertebrate densities were high, yellow perch remained the dominant food item selected by walleye (Blackwell et al. 1999). Conversely, in Lake Thompson, invertebrates composed a substantial portion of walleye diets when yellow perch densities were low (Isaak et al. 1993). Condition indices (Wr) for walleyes in Sinai and Waubay lakes were significantly higher in 2001 than 2000 and there were indications that growth also was faster in 2001, perhaps because prey supply (i.e., age-0 yellow perch abundance) was significantly higher in 2001. Strong year classes of walleye may not necessarily result in quality fishing if low prey abundance results in slow walleye growth. Evidence is accumulating that suggests Wr may best serve as an indicator of prey abundance (Wege and Anderson 1978; Liao et al. 1995; Marwitz and Hubert 1997; Porath and Peters 1997). Thus, the increases in walleye condition between 2000 and 2001 may indicate that the increased abundance of age-0 yellow perch between years resulted in increased walleye predation and thus improved body condition. We were surprised at the apparent effect of fish prey abundance (i.e., age- 0 yellow perch abundance) on walleye growth and condition. Admittedly, we cannot be certain of cause and effect for these two case histories; some un- measured factor may have been responsible for the increase in walleye population indices. However, our pre-conceived notion that food supply (whether it be fish prey or invertebrate prey) is always abundant for walleyes in mesotrophic to hypereutrhopic eastern South Dakota glacial lakes apparently may be incorrect. At the least, we should re-examine this concept. 98 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002)

ACKNOWLEDGEMENTS

We would to acknowledge walleye data collection by South Dakota De- partment of Game, Fish and Parks (SDGFP) personnel D. O. Lucchesi, T. R. St. Sauver, B. G. Blackwell, and M. J. Hubers. We appreciate their gracious permission to utilize these data. Manuscript review was provided by C. P. Pauk- ert. Partial funding for this project was provided by SDGFP through Federal Aid in Sport Fish Restoration Project F-15-R, Study 1584. This manuscript was approved for publication by the South Dakota Agricultural Experiment Station as Journal Series No. 3320.

REFERENCES

Anderson, M. R., S. J. Fisher, and D. W. Willis. 1998. Relationship between larval and juvenile yellow perch abundance in eastern South Dakota glacial lakes. North American Journal of Fisheries Management 18:989-991. Auer, N. A. 1982. Identification of larval fishes of the Great Lakes basin with emphasis on the Lake Michigan drainage. Great Lakes Fish Commission, Special Publication 82-3, Ann Arbor, Michigan. Blackwell, B. G., C. A. Soupir, and M. L. Brown. 1999. Seasonal diets of walleye and diet overlap with other top-level predators in two South Dakota lakes. South Dakota Department of Game, Fish and Parks, Fisheries Divi- sion Report 99-23, Pierre. Forney, J. L. 1974. Interactions between yellow perch abundance and walleye predation, and survival of alternate prey from Oneida Lake, New York. Transactions of the American Fisheries Society 103:15-24. Forney, J. L. 1976. Year-class formation in the walleye (Stizostedion vitreum vitreum) population of Oneida Lake, New York, 1963-73. Journal of the Fisheries Research Board of Canada 33:783-792. Forney, J. L. 1977. Reconstruction of yellow perch (Perca flavescens) cohorts from examination of walleye (Stizostedion vitreum vitreum) stomachs. Journal of the Fisheries Research Board of Canada 34:925-932. Gabelhouse, D. W., Jr. 1984. A length-categorization system to assess fish stocks. North American Journal of Fisheries Management 4:273-285. Hartman K. J., and F. J. Margraf. 1993. Evidence of predatory control of yellow perch (Perca flavescens) recruitment in Lake Erie, U.S.A. Journal of Fish Biology 43:109-119. Isaak, L. C., R. M. Neumann, and D. W. Willis. 1993. Food habits of walleyes in Lake Thompson, South Dakota. The Prairie Naturalist 25:325-330. Knight, R. L., F. J. Margraf, and R. F. Carline. 1984. Piscivory by walleyes and yellow perch in western Lake Erie. Transactions of the American Fisheries Society 113:677-693. Liao, H., C. L. Pierce, D. H. Wahl, J. B. Rasmussen, and W. C. Leggett. 1995. Relative weight (Wr) as a field assessment tool: relationships with growth, prey biomass, and environmental conditions. Transactions of the Ameri- can Fisheries Society 124:387-400. Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 99

Lott, J. P. 1991. Food habits of yellow perch in eastern South Dakota natural lakes. M.S. Thesis. South Dakota State University, Brookings. Madenjian, C. P. 1991. Limits to growth of young-of-the-year walleye (Sti- zostedion vitreum vitreum): an individual-based model perspective. Cana- dian Journal of Fisheries and Aquatic Sciences 48:1492-1499. Marwitz, T. D., and W. A. Hubert. 1997. Trends in relative weight of walleye stocks in Wyoming reservoirs. North American Journal of Fisheries Man- agement 17:44-53. Murphy, B. R., M. L. Brown, and T. A. Springer. 1990. Evaluation of the relative weight (Wr) index, with new applications to walleye. North American Journal of Fisheries Management 10:85-97. Nelson, W. R., and C. H. Walburg. 1977. Population dynamics of yellow perch (Perca flavescens), sauger (Stizostedion canadense), and walleye (S. vitreum) in four main stem Missouri River reservoirs. Journal of the Fisheries Research Board of Canada 34:1748-1763. Porath, M. T., and E. J. Peters. 1997. Use of walleye relative weights (Wr) to assess prey availability. North American Journal of Fisheries Management 17:628-637. SAS Institute. 1988. SAT/STAT user’s guide. SAS Institute, Cary, North Caroli- na. Starostka, A. B. 1999. Food habits and diet overlap of age-1 and older walleye and white bass in Lake Poinsett, South Dakota. M.S. Thesis. South Dakota State University, Brookings. Stueven, E., and W. C. Stewart. 1996. 1995 South Dakota lakes assessment final report. South Dakota Department of Environment and Natural Re- sources, Pierre. Swenson, W. A. 1977. Food consumption of walleye Stizostedion vitreum vitreum and sauger S. canadense in relation to food availability and physical conditions in Lake of the Woods, Minnesota, Shagawa Lake, and western Lake Superior. Journal of the Fisheries Research Board of Canada 34:1643- 1654. Wege, G. J., and R. O. Anderson. 1978. Relative weight (Wr): a new index of condition for largemouth bass. Pages 79-91 in G.D. Novinger and J.G. Dil- lard, editors. New approaches to the management of small impoundments. North Central Division, American Fisheries Society, Special Publication No. 5, Bethesda, Maryland. Whiteside, M. C., C. M. Swindoll, and W. L. Doolittle. 1985. Factors affecting the early life history of yellow perch, Perca flavescens. Environmental Bi- ology of Fishes 12:47-56.

Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 101

CONSTITUTIVE EXPRESSION OF A 3-DEOXY-D-ARABINO-HEPTULOSONATE- 7-PHOSPHATE SYNTHASE GENE IN ALFALFA (MEDICAGO SATIVA L.) LINES DIFFERING IN MORPHOLOGY AND LIGNIN CONCENTRATION

T. Liu Department of Molecular Pharmacology State University of New York/Stony Brook Stony Brook NY 11794

K.D. Kephart and C.D. Carter Department of Plant Science South Dakota State University Brookings, SD 57007

ABSTRACT

A partial cDNA for 3-deoxy-D-arabino-heptulosonate-7-phosphate (DAHP) synthase, the ﬁrst enzyme of the shikimate pathway, was isolated and used as a probe to examine DAHP synthase gene expression in alfalfa (Medicago sativa L.) lines divergently selected for morphology and lignin content. High lignin (HL) lines have lower leaf to stem ratios and consequently a higher proportion of lignin in the total herbage than do low lignin (LL) lines. However, LL lines have higher stem cell wall lignin concentrations than HL lines. Thus, we predicted that higher levels of DAHP synthase expression would be observed in LL lines. However, DAHP synthase mRNA levels were higher in HL lines than in LL lines. The relatively enhanced DAHP synthase expression in HL lines may be associated with increased biosynthesis of shikimate and/or phenylpropanoid-derived compounds other than lignin.

Keywords

Medicago sativa L., alfalfa, lignin, DAHP synthase, shikimate pathway

INTRODUCTION

Digestibility of alfalfa herbage is limited by lignin content (Buxton and Hornstein, 1986). Leaves have much less lignin and are more digestible than stems. Thus, herbage digestibility is affected by the proportion of leaves to stems as well as by the cell wall composition and digestibility of stems or leaves alone (Albrecht et al., 1987; Buxton and Hornstein, 1986; Kephart et al., 1990). 102 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002)

Kephart et al. (1989, 1990) analyzed morphological characters and lignin content of two classes of alfalfa lines developed by divergent selection for high vs. low total herbage lignin concentration. The low-lignin (LL) lines had higher leaf to stem ratios and shorter stems than high-lignin (HL) lines. HL and LL lines did not differ in number of branches or number of internodes, but HL internodes were longer than LL internodes. LL herbage was more digestible than HL herbage, largely due to the dilution effect of the higher leaf to stem ratio in LL lines. However, the concentration of lignin in stem cell walls was actually higher in LL lines than in HL lines (Kephart et al., 1990). Lignin is a polymer of monolignol precursors synthesized via the shikimate and phenylpropanoid pathways (Campbell and Sederoff, 1996; Grisebach, 1981; Herrmann, 1995a,b; Whetten and Sederoff, 1995). The shikimate pathway produces chorismate, from which are derived the aromatic amino acids phenylalanine (PHE), tyrosine (TYR) and tryptophan. Monolignols are subsequently derived from PHE and TYR via the phenylpropanoid pathway. Various branches of the shikimate and phenylpropanoid pathways lead to the synthesis of many other secondary plant compounds including flavonoids, flavonols, isoflavonoids, anthocyanins, phenolics, quinones and tannins. The first enzyme of the shikimate pathway is 3-deoxy-D-arabino-heptulosonate-7-phosphate (DAHP) synthase (Herrmann,1995a, 1995b). DAHP synthase activity and gene expression are regulated by tissue specific and developmental effects (Gorlach et al., 1993; Graziana and Boudet, 1980; Jones et al., 1995; Rothe et al., 1976; Suzich et al., 1984) and environmental factors including light (Henstrand et al., 1992), physical wounding and pathogen attack (Dy- er et al., 1989; Henstrand et al., 1992; Keith et al., 1991), exposure to the herbicide glyphosate (Pinto et al., 1988), and Ca2+ ions and jasmonate-related compounds (Suzuki et al., 1995). DAHP synthase cDNAs have been isolated from potato (Dyer et al., 1990; Zhao and Herrmann, 1992), tobacco (Wang et al., 1991), Arabidopsis (Keith et al., 1991) and tomato (Gorlach et al.,1993). In most plant species, at least two copies of the DAHP synthase gene are present, and the isoforms differ substantially in their 5’ and 3’ regions and in expression. To determine whether the differences in stem cell-wall lignin concentration between HL and LL lines are related to differences in DAHP synthase gene expression, we have isolated an alfalfa partial cDNA for DAHP synthesis and compared DAHP synthase mRNA levels in HL and LL lines.

MATERIALS AND METHODS

Plant material

The alfalfa plants used in this study were derived from `Saranac AR' following divergent selection for herbage lignin concentration (Kephart et al., 1989). Total above-ground herbage (minus the basal 1" of the main stem) of each plant was collected from six plants of ﬁeld-grown HL and LL lines at early-bloom stage. Leaves were removed and discarded and stems and petioles were collected and frozen in liquid nitrogen. HL plants were H-5, H-7, and H- 9. LL plants were L-4, L-6, and L-8. Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 103

Isolation of alfalfa partial cDNA

RNA was extracted and fractionated by gel electrophoresis (Sambrook et al., 1989). Total RNA (1 ug) was reverse transcribed in vitro using SuperScript Reverse Transcriptase (BRL). The cDNA pool was amplified by polymerase chain reaction (PCR) using DNA Taq polymerase (Promega). The degenerate primers were based on the conserved regions of known DAHP synthase sequences from other species (Gorlach et al., 1993). PCR amplification was conducted in a Perkin-Elmer Model 480 thermal cycler to 30 cycles using 94C/30 sec to denature, 44C/60 sec for annealing, and 72C/60 sec for extension. The final extension step was 7 min. The plasmid pLW3-210, provided by K. M. Her- rmann (Purdue University), carrying a potato DAHP synthase cDNA insert, was used as a positive control in the PCR reactions. The products resulting from PCR amplification of the alfalfa cDNAs were separated on an 0.8% agarose gel and DNA fragments of about 875 bp were isolated and subcloned into EcoR I cleaved-pCR II (Invitrogen) and transformed into E. coli INαF’ (Invitrogen). The cDNA insert (MsDHS1) was sequenced at the Iowa State University Molec- ular Biology facility. Sequence comparisons were performed with the Gene- Jocky II program.

Isolation of alfalfa genomic DNA

Genomic DNA was extracted from stems and petioles of H-7 according to Rogers and Bendich (1994) with some modiﬁcations. Genomic DNA (10 ug/reaction) was restriction digested with EcoRI, KpnI, PstI or PstI plus KpnI at 37ºC for 60 min, and fractionated on an 0.8% agarose gel (Sambrook et al., 1989). The DNA was transferred to nylon membrane (Nytran) and hybridized to the 32P-dCTP-labeled MsDHS1 cDNA as described below.

Isolation of RNA

For analysis of DAHP synthase gene expression, alfalfa RNA was extracted, separated by formaldehyde-agarose gel electrophoresis, and transferred to nylon membranes (Ausubel et al., 1990; Sambrook et al., 1989). The Northern blots were hybridized as described below.

Northern and genomic Southern hybridizations

Prehybridizations were performed at 42ºC for 1.5 h with gentle shaking in a solution containing 50% (v/v) formamide, 6x SSPE (0.9 M NaOH, 60 mM NaH2PO4-H2O, 6 mM EDTA), 5x Denhardt's solution (0.1% Ficoll, 0.1% polyvinylpyrrolidone, 0.1% BSA), 1% (w/v) SDS, and 200 mg/ml denatured salmon sperm DNA. The MsDHS1 cDNA insert was isolated by restriction digestion and gel electrophoresis, and the insert was cut and extracted from the gel (Sambrook et al., 1989), and labeled with 32P-dCTP by random priming using a Prime-a-Gene Kit (Promega). Hybridizations were carried out at 42ºC in the same solution plus the 32P-labeled MsDHS1 probe (2 ng/ml) for 18 to 22 h 104 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) with gentle agitation. The blots were washed three times with 2x SSPE (0.3 M

NaOH, 20 mM NaH2PO4-H2O, 2 mM EDTA), 0.1% SDS for 5 min at RT, and twice with 0.2x SSPE (30 mM NaOH, 2 mM NaH2PO4-H2O, 0.2 mM EDTA), 0.1% SDS for 30 min at 55ºC. Hybridized blot membranes were imaged using a Bio- Rad PhosphorImager. Quantitative data were obtained from PhosphorImager scans using Molecular Analyst software. The Northern blot membrane was stripped by immersion in 0.1% SDS at 95-100C, and then re-hybridized with an 18S rRNA cDNA insert from soybean clone pSR1.2B3 (provided by R. Meagher, University of Georgia) to quantify RNA loaded in each lane. DAHP synthase mRNA levels obtained for each sample were normalized against the 18S rRNA values.

RESULTS AND DISCUSSION

The sequence of the 875 bp alfalfa cDNA (GenBank Accession Number U50544) showed sequence similarities of 89%, 88%, 87%, and 87% to AtDHS2, LeDHS2, NtDAHPS-1, and StshKA, respectively, and slightly lower similarities of 84%, 86%, and 86% to AtDHS1, LeDHS1, and StshKB, respectively. The alfalfa cDNA, designated MsDHS1, corresponds to base pairs 472 to 1330 of potato StshkB, and appears to be a partial cDNA for alfalfa DAHP synthase. Alfalfa genomic DNA was restriction digested with EcoRI, KpnI, PstI, or KpnI plus PstI, and hybridized to the alfalfa partial DAHP synthase cDNA, MsDHS1, to determine the gene copy number (Fig. 1). PstI does not have a cutting site within the cDNA and produced a single band (lane 1, Fig. 1). EcoRI and KpnI each have a single restriction site within the Ms- DHS1 cDNA, and each digestion produced two hybridizing bands (lanes 2 and 3, Fig. 1). The combined KpnI plus PstI cleav- age produced two fragments that hybridized to the probe (lane 4, Fig. 1). The results indicate that only one copy of the DAHP synthase gene is present in the alfalfa genome, or that the copies are highly conserved. Two copies of DAHP synthase genes occur in Arabidopsis, potato and tomato genomes; these isoforms belong to two different gene families and differ in expression and in se- Figure 1. Southern blot of alfal- quence, especially in the 5’ and 3’ regions fa genomic DNA. DNA samples (Hermann, 1995). (10 ug each) were digested with DAHP synthase gene expression in the restriction enzymes PstI (P), HL and LL lines was analyzed by hybridiza- XhoI (X), EcoRI (E) or a combi- tion of HL and LL RNA to the MsDHS1 nation of PstI and XhoI (P/X), and hybridized with 32P-labeled cDNA. The results are shown in Figures 2-4. MsDHS1. Molecular marker The level of DAHP synthase mRNA was sub- sizes (Kb) are shown at the left. stantially greater in stems of HL lines than in Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 105 stems of LL lines. (Figure 3 shows that the control hybridizations to an 18S RNA probe did not differ between HL and LL lines, indicating that equivalent amounts of RNA from each line were loaded onto the gel. Figure 4 compares DAHP synthase mRNA levels adjusted for 18S rRNA levels.) Although total herbage lignin content is lower in LL lines than in HL lines, this Figure 2. Alfalfa DAHP synthase mRNA levels in 3 is due primarily to the HL plants (H5, H7, H9) and 3 LL plants (L4, L6, L8). greater proportion of leaves Total RNA (10 ug/lane) was isolated from each in LL herbage. Stem cell line, separated by formaldehyde gel electrophore- wall lignin concentration, sis, transferred to nylon membrane and hybridized to 32P-labeled MsDHS1. Counts were visualized by however, is in fact higher in BioRad PhosphorImager scan and analyzed using LL lines (Kephart et al., Molecular Analyst software. A: Northern blot. 1989, 1990). If the higher B: Densitometry proﬁle. stem cell wall lignin concentration of LL lines were due to coordinately elevated transcription of shikimate and phenylpropanoid pathway genes in LL lines, then DAHP synthase expression might be expected to be greater in LL lines. However, our results suggest the reverse; i.e., alfalfa DAHP synthase mRNA levels were higher in HL lines than in LL lines. Lignin deposition has been shown to contribute to resistance to injury or Figure 3. Alfalfa 18S rRNA levels in 3 HL plants pathogen attack, and other (H5, H7, H9) and 3 LL plants (L4, L6, L8). The same phenylpropanoid com- membrane used in Figure 4 was stripped and re- hybridized with a 32P-labeled 18S rRNA probe. pounds have been shown Counts were visualized by BioRad PhosphorIm- to act as phytoalexins in in- ager scan and analyzed using Molecular Analyst duced resistance responses software. A: Northern blot. B: Densitometry pro- (Paiva et al., 1991; Cramer ﬁle. et al. 1989; Dixon and Pai- va, 1995). Constitutive lev- 106 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002)

els of antimicrobial compounds also contribute to pathogen resistance and differ among genotypes and environmental conditions (Dornbos et al., 1990; Pennypacker et al., 1994; Silverman et al., 1995). HL alfalfa lines are more vigor- ous and more resistant to pathogens in the ﬁeld than LL lines (Buxton and Figure 4. Normalization of DAHP synthase RNA Casler, 1993). The greater levels relative to 18S rRNA levels, in 3 HL plants (H5, H7, H9) and 3 LL plants (L4, L6, L8). pathogen resistance, lower cell wall lignin content, and higher DAHP synthase gene expression of HL lines may be associated with increased biosynthesis of shikimate- or phenylpropanoid-derived compounds other than lignin in HL lines, or with a diversion of phenylpropanoid precursors away from lignin biosynthesis in favor of increased production of phenylpropanoid phytoalexins or other secondary metabolites associated with constitutive or induced pathogen resistance.

ACKNOWLEDGMENTS

We thank T. Cheesbrough (Biology/Microbiology, South Dakota State Uni- versity) for assistance with sequence and image analysis software, and K. Her- rmann (Biochemistry, Purdue University) for providing the potato DAHP synthase probe.

LITERATURE CITED

Ausubel FM (Ed) (1994) Current Protocols in Molecular Biology. John Wiley and Sons, Inc., NY. Buxton DR, Casler MD. 1993. Environmental and genetic effects on cell wall composition and digestibility. In: Forage Cell Wall Structure and Digestibil- ity, ASA-CSSA-SSSA, Madison WI. Campbell MM, Sederoff RR (1996) Variation in lignin content and composition: Mechanisms of control with and implications for the genetic improvement of plants. Plant Physiol 110:3-13 Cramer CL, Edwards K, Dron M, Liand X, Dildine SL, Bolwell GP, Dixon RA, Lamb CJ, Schuch WW. 1989. Phenylalanine ammonia-lyase gene organiza- tion and structure. Plant Mol Biol 12: 367-383 Dixon RA, Paiva NL. 1995. Stress-induced phenylpropnoid metabolism. Plant Cell 7: 1085-1097. Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 107

Dornbos DL Jr, Spencer GF, Miller RW. 1990. Medicarpin delays alfalfa seed ger- mination and seedling growth. Crop Sci 30: 162-166. Dyer WE, Weaver LM, Zhao J, Kuhn DN, Weller SC, Herrmann KM (1990) A cDNA encoding 3-deoxy-D-arabino-heptulosonate 7-phosphate synthase from Solanum tuberosum L. J Biol Chem 265: 1608-1614 Dyer WE, Henstrand JM, Handa AK, Herrmann KM (1989) Wounding induces the first enzyme of the shikimate pathway in Solanaceae. Proc Natl Acad Sci USA 86: 7370-7373 Gorlach J, Beck A, Henstrand JM, Handa AK, Herrmann KM, Schmid J, Amrhein N (1993) Differential expression of tomato (Lycopersicon esculentum L.) genes encoding shikimate pathway isoenzymes. I. 3-deoxy-D-arabino- heptulosonate 7-phosphate synthase. Plant Mol Biol 23: 697-706 Graziana A, Boudet AM (1980) 3-Deoxy-D-arabino-heptulosonate 7-phosphate synthase from Zea mays: general properties and regulation by tryptophan. Plant Cell Physiol 21: 793-802 Grisebach H (1981) Lignins. In Stumpf PK, Conn EE, eds, The Biochemistry of Plants Vol 7, Academic Press, New York, pp 457-478 Henstrand JM, McCue KF, Brink K, Handa AK, Herrmann KM, Conn EE (1992) Light and fungal elicitor induce 3-deoxy-D-arabino-heptulosonate 7-phosphate (DAHP) synthase mRNA in suspension cultured cells of parsley (Pet- roselinum crispum L.). Plant Physiol 98: 761-763 Herrmann KM (1995a) The shikimate pathway as an entry to aromatic secondary metabolism. Plant Physiol 107: 7-12 Herrmann KM (1995b) The shikimate pathway: early steps in the biosynthesis of aromatic compounds. The Plant Cell 7: 907-919 Jones JD, Henstrand JM, Handa AK, Herrmann KM, Weller SC (1995) Impaired wound induction of 3-deoxy-D-arabino-heptulosonate 7-phosphate (DAHP) synthase and altered stem development in transgenic potato plant expressing a DAHP synthase antisense construct. Plant Physiol 108: 1413- 1421 Keith B, Dong X, Ausubel FM, Fink GR (1991) Differential induction of 3-deoxy-D-arabino-heptulosonate 7-phosphate synthase genes in Arabidopsis thaliana by wounding and pathogen attack. Proc Nat Acad Sci (USA) 88: 8821-8825 Kephart KD, Buxton DR, Hill RR Jr (1989) Morphology of alfalfa divergently selected for herbage lignin concentration. Crop Sci 29: 778-782 Kephart KD, Buxton DR, Hill RR Jr (1990) Digestibility and cell-wall components of alfalfa following selection for divergent herbage lignin concentration. Crop Sci 30: 207-212 Paiva NL, Edwards R, Sun Y, Hrazdina G, Dixon RA. 1991. Stress responses in alfalfa (Medicago sativa L.) II. Molecular cloning and expression of alfalfa isoflavone reductase, a key enzyme of isoflavonoid phytoalexin biosynthesis. Plant Mol Biol 17: 653-667. Pennypacker BW, Knievel DP, Risius ML, Leath KT. 1994. Photosynthetic pho- ton flux density X pathogen interaction in growth of alfalfa infected with Verticillium albo-atrum. Phytopathol 84: 1350-1358. 108 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002)

Pinto JEBP, Dyer WE, Weller SC, Herrmann KM (1988) Glyohosate induces 3- deoxy-D-arabino-heptulosonate 7-phosphate synthase in potato (Solanum tuberosum L.) cells grown in suspension culture. Plant Physiol 87: 891-893 Rogers SO, Bendich AJ (1994) Extraction of total cellular DNA from plants, algae and fungi. In Gelvin SB, Schilperoort RA eds, Plant Molecular Biology Manual, 2nd Ed, Kluwer Acad Publ, Belgium, pp D1: 1-8 Rothe GM, Maurer W, Mielke C (1976) A study on 3-deoxy-D-arabino-heptu- losonic acid 7-phosphate synthase in higher plants. The existence of three isozymes in Pisum sativum. Ber Dtsch Bot Ges 89: 163-173 Sambrook J, Fritsch EF, Maniatis T (1989) Molecular Cloning: A Laboratory Man- ual, 2nd Ed. Cold Spring Harbor Lab Press, New York Silverman P, Seskar M, Kanter D, Schweizer P, Metraux JP, Raskin I. 1995. Sali- cylic acid in rice: Biosynthesis, conjugation, and possible role. Plant Phys- iol 108: 633-639. Suzich JA, Ranjeva R, Hasegawa PM, Herrmann KM (1984) Regulation of the shikimate pathway of carrot cells in suspension culture. Plant Physiol 75: 369-371 Suzuki K, Fukuda Y, Shinshi H (1995) Studies on elicitor-signal transduction leading to deferential expression of defense genes in cultured tobacco cells. Plant Cell Physiol 36: 281-289 Wang Y, Herrmann KM, Weller SC, Goldsbrough PB (1991) Clonic and nucleotide sequence of a complementary DNA encoding 3-deoxy-D-arabino- heptulosonate 7-phosphate synthase from tobacco. Plant Physiol 97: 847- 848 Whetten R, Sederoff R (1995) Lignin biosynthesis. Plant Cell 7: 1001-1013 Zhao J, Herrmann KM (1992) Cloning and sequencing of a second cDNA encoding 3-deoxy-D-arabino-heptulosonate 7-phosphate synthase from Solanum tuberosum L. Plant Physiol 100: 1075-1076 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 109

CONSTRUCTION OF NATURAL AND REAL NUMBERS FROM GENERALIZED ORDER, CONGRUENCE AND CONTINUITY AXIOMS (A “REAL LINE” MAY CROSS ITSELF)

Richard P. Menzel Box 254 Martin, SD 57551 110 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 111 112 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 113 114 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 115 116 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 117 118 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 119 120 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 121 122 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 123 124 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 125 126 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 127 128 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 129 130 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 131 132 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 133 134 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 135 136 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 137

INITIATION OF FEEDING DURING HATCHERY REARING OF LANDLOCKED FALL CHINOOK SALMON FRY

Michael E. Barnes, William A. Sayler, and Rick J. Cordes South Dakota Department of Game, Fish and Parks McNenny State Fish Hatchery Spearﬁsh, SD 57783

ABSTRACT

Hatchery feeding of landlocked fall chinook salmon (Oncorhynchus tshawytscha) fry was initiated at two different stages of development to determine the time to start feeding for optimal fish growth. Despite initially weighing less, fry started on feed at the first visible indication of free-swimming (approximately 20% yolk sac remaining), were significantly heavier after 28 rearing days than fry started on feed at complete yolk sac absorption. Total tank weights were also significantly heavier in the fry started on feed the earliest. No differences in fry mortality were observed between the two feed initiation regimes. For maximum growth, initiating feeding of landlocked fall chinook salmon at the first observation of free-swimming fry is recommended, unless there are overriding fish health concerns.

Keywords

Oncorhynchus tshawytscha, chinook salmon, salmonid fry, hatchery feeding, swim-up

INTRODUCTION

Although salmonid fry will begin feeding prior to complete yolk sac absorption (Knight 1963), there are considerable differences of opinion on when to initiate feeding of chinook salmon (Oncorhynchus tshawytscha) and other salmonids during hatchery rearing. The most common practice is to start feeding at fry swim-up, when fry have inflated their swim bladders and absorbed enough yolk sac to maintain a position in the water column (Piper et al. 1982). Needham (1988) also suggested first feeding when fry could swim upright, but then suggested species specific differences in the timing of initial feeding. He gave as a guide initiating feeding of rainbow trout (Oncorhynchus mykiss) fry at 50% of yolk sac remaining, as opposed to only 20% of the yolk sac remaining for initial feeding of Atlantic salmon (Salmo salar). The Alaska Department of Fish and Game (1983) fish culture manual rec- ommends making food available as soon as any salmonid fry swim-up is observed. Contrarily, delayed feeding until near complete yolk-sac absorption 138 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) has been suggested to promote efficient growth and transition to exogenous feeding in rainbow trout (Twongo and MacCrimmon 1976; MacCrimmon and Twongo 1980). Heming et al. (1982) identified an optimal time frame to initiate feeding chinook salmon fry based on incubation water temperatures. Their work was conducted with anadromous salmon in their native range. However, landlocked chinook salmon exhibit substantially different reproductive characteristics, such as egg size, than salmon in their native range (Barnes et al. 2000), which likely make the time frame identified by Heming et al. (1982) unusable with landlocked salmon. Landlocked chinook salmon fry obtained as eggs from Lake Oahe, South Dakota are typically maintained in vertical-tray incubators, moved to rearing tanks just prior to complete absorption of the external yolk sac (button-up), and subsequently started on feed when an estimated 30% of the fry are free- swimming. The impact of the starting feeding at this time on fish growth is uncertain (Palmer et al. 1951; Twongo and MacCrimmon 1976; MacCrimmon and Twongo 1980; Heming et al. 1982). The objective of this study was to determine the effect on the hatchery growth and survival of landlocked chinook salmon fry if feeding is initiated at the first sign of fry swim-up, as compared to initiating feeding at complete yolk sac absorption.

METHODS

Fry used for this experiment were obtained as eggs during spawning of chinook salmon from Lake Oahe at Whitlocks Spawning Station on October 27, 1999. After fertilization, eggs were pooled during water-hardening and subsequent transport to McNenny State Fish Hatchery, Spearﬁsh, South Dakota. Eggs were incubated in Heath (Flex-a-lite Consolidated, Tacoma, Washington) incubator trays. Fry were removed from the same incubator tray on two dates (January 1 and January 10) corresponding to the ﬁrst sign of free-swimming fry (approximately 20% of the yolk sac remaining) and near-complete yolk sac absorption. At each date, each of three 100-L cylindrical tanks received 200 fry. Total tank weights were recorded to the nearest g. In addition, 30 fry were individually weighed to the nearest mg. Well water (11º C; total hardness as

CaCO3, 360 mg/L; alkalinity as CaCO3, 210 mg/L; pH, 7.6; total dissolved solids, 390 mg/L) was used both in the incubator trays and tanks. Flows in each tank were set at 20 L/min throughout the experiment. Hourly hand feeding with #2 BioDiet Starter (Bio-Oregon, Inc., Warrenton, Oregon) was initiated the day after fry were placed in the tanks. Fry were fed to satiation, as indicated by the presence of wasted feed on the bottom of each tank. All food dispensed was recorded to the nearest g. Tanks were cleaned daily to eliminate excess food and ﬁsh waste, and any mortalities were removed and recorded at that time. On January 29, 2000, total tank weights were again recorded to the nearest g, and 20 ﬁsh from each tank were individually weighed to the nearest mg. Individual fry weight data were analyzed using Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 139

Student’s t-tests and tank data were analyzed using the Kruskal-Wallis test. Sta- tistical testing was performed using the SPSS (9.0) statistical analysis program (SPSS 1999), with signiﬁcance pre-determined at P ≤ 0.05.

RESULTS

At the start of the experiment, fry with 20% of their yolk sac remaining weighed significantly less (P = 0.006) than fry which had completely absorbed their visible yolk sacs (Table 1). Despite weighing less at the time of initial feed, at the end of the experiment mean weights from fry started on feed with 20% of their yolk sac remaining were 80 mg more than those fry started on feed 10 days later (P = 0.0001). The improvement in growth of the fry started on feed at the earliest date was also evident in total tank weights and weight gained per tank (Table 2). Mean weights in the tanks started on feed at the first sign of fry swim-up were significantly heavier, by an average of 14 g, than mean weights from the tanks of fry started on feed at complete yolk sac absorption. Weight gain per tank was significantly different, and over 20% greater, in the fry started on feed the earliest. Mean tank mortality ranged from 1 to 7% and was not significantly different between the two feed initiation treatments.

Table 1. Mean (± SE) weights (mg) from chinook salmon fry started on feed either at the ﬁrst observance of free swimming fry (20% yolk sac remaining) or at complete yolk sac absorption. Means in a row are all signiﬁcantly different at P = 0.05.

Weight N First free swimming fry Complete yolk sac absorption

Start 30 257 ± 6 z 285 ± 7 y

End 60 718 ± 16 z 636 ± 15 y

Table 2. Data (mean ± SE) from tanks of chinook salmon fry started on feed either at the ﬁrst observance of free swimming fry (20% yolk sac remaining) or at complete yolk sac absorption (N = 3). Means in a row followed by the same letter are not signiﬁcantly different at P = 0.05.

First free swimming fry Complete yolk sac absorption

Start weight (g) 56 59 End weight (g) 151 + 5 z 137 + 3 y Gain (g) 95 + 5 z 78 + 3 y Food fed (g) 265 + 7 z 281 + 15 z Mortality 8 + 2 z 6 + 4 z 140 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002)

DISCUSSION

Heming et al. (1982) stated that it would not be beneficial to initiate feeding of chinook salmon fry before 82 incubation days at 11ºC, the point at which 50% of fry in their study were first able to ingest food. In contrast, our earliest feeding started at 67 incubation days, and complete yolk sac absorption occurred on incubation day 76. Heming et al. (1982) also recorded wet weights of approximately 600 mg from non-feeding salmon fry, more than double the weights of fry that we obtained. Obviously the sizes and developmental rates we observed with landlocked fry from Lake Oahe are substantially different than those of chinook salmon in their native range. We believe this is due to the differences in egg size (Barnes et al. 2000), but it could also be due to other, as yet unidentified, environmental factors (Bagenal 1978; Donaldson 1990). Because of these differences, the quantitative models developed by Heming et al. (1982) to identify the time at which initial food presentation to chinook salmon fry should occur do not apply to landlocked chinook salmon fry from Lake Oahe and possibly other freshwater bodies of water. Our results validate the suggestion of Palmer et al. (1951) that salmon should be fed when they are first capable of free swimming if maximum growth is to be attained. The amount of yolk remaining at this stage that they describe appears to be very similar to that which we observed in the early feeding salmon fry from our experiment. Initiating feeding prior to visible yolk sac absorption was also suggested by Heming et al. (1982). Although they found that the optimal time for initial feeding did not coincide with any easily recognizable development stage of chinook salmon fry development, they indicated that such an time occurs well prior to complete yolk sac absorption when fry are reared at 11ºC. Species or strain (domestic vs. wild) differences may explain the disagreement between our results and those reported for rainbow trout (Twongo and MacCrimmon 1976; MacCrimmon and Twongo 1980). Using our smaller experimental tanks, we observed no difference in mortality between fry started on feed at two different developmental times. Wast- ed feed and fecal material were regularly and easily removed. Starting fry on feed prior to complete swim-up may be a concern in hatcheries with a history of bacterial (environmental) gill disease, particularly if rearing units used for feed initiation are difficult to clean (Post 1987; Warren 1991).

RECOMMENDATION

We recommend initiating feeding of landlocked chinook salmon lots at the ﬁrst occurrence of any free-swimming fry, unless there are overriding ﬁsh health concerns. Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 141

ACKNOWLEDGEMENTS

We thank the reference librarians at the South Dakota State Library for their assistance with literature procurement, Kody Steinbrecher and Fritz Fonck for assistance during ﬁsh culture.

LITERATURE CITED

Alaska Department of Fish and Game. 1983. Fish culture manual. Alaska De- partment of Fish and Game, Juneau. Bagenal, T. B. 1978. Aspects of fish fecundity. Pages 75-101 in S. D. Gerk- ing, editor. Freshwater fish production. Wiley and Sons, New York. Barnes, M. E., R. P. Hanten, R. J. Cordes, W. A. Sayler, and J. Carreiro. 2000. Reproductive performance of inland fall chinook salmon. North American Journal of Aquaculture 62:203-211. Donaldson, E. M. 1990. Reproductive indices as measures of the effects of environmental stressors in fish. Pages 109-122 in S. M. Adams, editor. Bio- logical indicators of stress in fish. American Fisheries Society Symposium 8, Bethesda, Maryland. Heming, T. A., J. E. McInerney, and D. F. Alderdice. 1982. Effect of temperature on initial feeding in alevins of chinook salmon (Oncorhynchus tshawytscha). Canadian Journal of Fisheries and Aquatic Sciences 39:1554- 1562. Knight, A. E. 1963. The embryonic and larval development of the rainbow trout. Transactions of the American Fisheries Society 92:344-355. MacCrimmon, H. R., and T. K. Twongo. 1980. Ontogeny of feeding behaviour in hatchery-reared rainbow trout, Salmo gairdneri Richardson. Canadian Journal of Zoology 58:20-26. Needham, T. 1988. Salmon smolt production. Pages 87-116 in L. M. Laird and T. Needham, editors. Salmon and trout farming. Wiley, New York. Palmer, D. D., H. E. Johnson, L. A. Robinson, and R. E. Burrows. 1951. The effect of retardation of the initial feeding on the growth and survival of salmon fingerlings. Progressive Fish-Culturist 13:55-62. Piper, R. G., I. B. McElwain, L. E. Orme, J. P. McCraren, L. G. Fowler, and J. R. Leonard. 1982. Fish hatchery management. U.S. Fish and Wildlife Ser- vice, Washington, D.C. Post, G. 1987. Textbook of fish health. T. F. H. Publications, Neptune City, New Jersey. SPSS. 1999. SPSS Base 9.0. SPSS, Inc. Chicago, Illinois. Twongo, T. K., and H. R. MacCrimmon. 1976. Significance of the timing of initial feeding in hatchery rainbow trout, Salmo gairdneri. Journal of the Fisheries Research Board of Canada 33:1914-1921. Warren, J. W. 1991. Diseases of hatchery fish. U.S. Fish and Wildlife Service. Portland, Oregon.

Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 143

WHITE-TAILED DEER BEHAVIORAL RESPONSE TO PREDATOR FECES

Christopher C. Swanson, Cory M. Kassube, and Jonathan A. Jenks Department of Wildlife and Fisheries Sciences South Dakota State University Brookings, SD 57007

ABSTRACT

White-tailed deer (Odocoileus virginianus dakotensis) rely upon their ability to recognize predators. Avoidance of predators can be due to innate or learned behaviors. We tested the hypothesis that white-tailed deer fawns raised in a controlled setting with no exposure to native predators would have the innate ability to recognize predator feces via an alert response. To test our hypothesis, fawns were released into a pen in which predator feces or a control were placed. Predator feces evaluated included gray wolf (Canis lupus), mountain lion (Puma concolor), coyote (Canis latrans), and Bengal tiger (Panthera tigris tigris). Plaster-of-Paris formed to a shape similar to feces and eastern cottontail rabbit (Sylvilagus ﬂoridanus) feces were used as controls. Presence of an alert response to feces was documented during 180-second trials. Vigilance behavior of fawns to gray wolf, mountain lion, and coyote feces was higher (P < 0.10) than that for cottontail rabbit, Bengal tiger, and the control. White-tailed deer fawns demonstrated an innate behavioral response to the presence of feces of native predators.

Keywords

Behavior, feces, Odocoileus virginianus, predators, white-tailed deer

INTRODUCTION

White-tailed deer fawns have long had the ability to escape predation. Yet ungulate predation remains highest during the fawning season (Gese 1999). Fawns rely on cryptic coloration, minimum activity, and scent glands that lack development to avoid predation (Carl and Robbins 1988). Recognition of on- coming predators by olfactory and auditory senses enables fawns to "freeze" to avoid predation. Prey species have the capability to retain predator recognition for thousands of years after predation has been diminished (Berger 1999). Thus, ungulates may be born with the predisposed notion of what poses a threat. How- ever, no information is available on innate responses to predators of fawns without prior experience to predation. We tested the hypothesis that white- tailed deer fawns raised in a controlled setting with no exposure to native predators would have the innate ability to recognize predator feces via an alert 144 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) response. Because innate responses may be more important than learned behavior to survival of neonates, we predicted that white-tailed deer fawns with no exposure to predators would exhibit higher vigilance to predator feces than to controls.

STUDY AREA

We observed white-tailed deer fawns during October 2001 at the South Dakota State University Wildlife Research Facility in Brookings, South Dakota. The facility is located approximately 1.5 km north of South Dakota State Uni- versity in Brookings County (R-50-W, T-110-N). The 305-cm tall enclosure re- stricts predators and receives little disturbance other than human intervention. Captive white-tailed deer fawns were separated from adult females after consuming colostrum and hand raised to imprint fawns to the presence of humans.

METHODS

Trials were conducted in which white-tailed deer fawns were removed from a holding pen and released in an individual pen where they were observed for 180 seconds. Five randomly selected white-tailed deer fawns from a pool of 10 fawns were placed in the pen (one per trial) and observed during trials. North American predator feces including gray wolf, mountain lion, and coyote were used in trials. In addition, Bengal tiger feces, which represented a non-native predator, were included in experiments. Plaster of Paris molded into the shape of feces was used as a control and eastern cottontail feces were used as a biological control. Feces from predators or a control (random assignment) were placed in the pen before introducing a fawn and the fawn observed for an 180-second time frame (Berger 1999). Change in posture and movements were recorded from a ground blind. We determined the percent vigilance of each fawn to the feces by recording the amount of time the fawn was in alert posture per 180-s interval. Percent vigilance of fawns to feces was compared using Analysis of Variance. We used SYSTAT (SPSS Inc. 2001) for all analyses. Alpha was set at P = 0.10 for analyses.

RESULTS

Mean percent vigilance varied from 8.75 (Bengal tiger) to 27.50 % (gray wolf) (Table 1). Percent vigilance was similar for controls and predator feces

(F5,22 = 1.14, P = 0.369). However, percent vigilance of fawns to feces of native predators (gray wolf, mountain lion, coyote) differed (F2,25 = 3.013, P = 0.067) from controls (Plaster of Paris, cottontail) and nonnative predators (Bengal tiger). Response of fawns to gray wolf, mountain lion, and coyote feces was over twice (24.16 ± 1.82) the response as that of controls and Bengal tiger feces (9.79 ± 0.55). Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 145

Table 1. Percent vigilance of captive white-tailed deer fawns to North American predator feces (gray wolf, mountain lion, coyote), non-native predator feces (Bengal tiger), Plaster of Paris control feces, and biological control feces (cottontail rabbit), October 2001.

Treatment Mean1 SE2 N3

Control 10.62 5.67 8 Gray Wolf 27.50 8.02 4 Mountain lion 21.25 8.02 4 Coyote 23.75 8.02 4 Bengal Tiger 8.75 8.02 4 Eastern Cottontail 10.00 8.02 4

1LS Mean = average vigilance per trial. 2SE = standard error of the mean. 3N = number of trials per experiment.

DISCUSSION

In ecological communities where predators remove large numbers of prey from populations, miscues related to non-vigilant behavior to predators could prove fatal (Berger 1999). Coyote and native carnivores can affect juvenile survival (Berger et al. 1999). Naïve prey may be killed in areas of predator de- population because they lack recognition of cues that indicate predator presence (Berger et al. 2001). Berger (1999) suggested that when predation no longer exists in an ecological community the behavioral response by prey might altogether be absent. Conversely, risk of predation is thought to increase with the presence of predators (Magurran 1992). Berger et al. (2001) implied that prey in high-density predator populations were more vigilant due to the increased risk of attack. Despite studies indicating that response to predators is related to presence of predators, fawns reacted to predator feces with an alert and nervous response. Fawns displayed increased vigilance for gray wolf, mountain lion, and coyote feces. Increase in vigilance was likely due to the fact that gray wolf, mountain lion, and coyote are native predators to white-tailed deer in North America. White-tailed deer may maintain an innate response to predator cues because they recognize the feces as prevalent in the ecological community. Eastern cottontail feces received low response because this animal coincides with white-tailed deer interacting in a docile manner. Moreover, Bengal tiger was once a native predator but feces likely received low response because of its prolonged absence from North America. In the wild, fawns also may learn to avoid predators from their mother’s knowledge of predator impact. Ozoga and Verme (1986) indicated that as white-tailed deer females increase in maternal age and experience they are more likely to avoid predation and protect fawns. Consequently, predation might involve innate and learned response by fawns. Fawns may develop acute awareness to cues that trigger alert response to recognize danger. Our data indicate that survival of fawns involves innate responses to predators that 146 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) currently occur in North America. However, fawn alert response likely bene- ﬁts from experience and doe-fawn interaction.

ACKNOWLEDGEMENTS

We would like to thank the South Dakota State University Wildlife Research Facility and Kevin Montieth and Aaron Montieth for their assistance at the facility. We would also like to thank the Bramble Park Zoo, Watertown, South Dakota for providing feces used in trials. We thank D. M. Fecske for comments made on a previous draft of this manuscript.

LITERATURE CITED Berger, J. 1999. Anthropogenic extinction of top carnivores and interspeciﬁc animal behaviour: implications of the rapid decoupling of a web involving wolves, bears, moose and ravens. Proceedings Royal Society of London 266:2261-2267. Berger, J., J. W. Testa, T. Roffe, and S. Monfort, S. 1999. Conservation en- docrinology: a noninvasive tool to understand relationships between car- nivore colonization and ecological carrying capacity. Conservation Biology 13: 980-989. Berger, J., J. E. Swenson, and I. Persson. 2001. Recolonizing carnivores and naïve prey: conservation lessons from Pleistocene extinctions. Science 291:1036-1039. Carl, G. R., and C. T. Robbins. 1988. The energetic cost of predator avoidance in neonatal ungulates: hiding versus following. Canadian Journal of Zool- ogy 66: 239-246. Gese, E. 1999. Threat of predation: do ungulates behave aggressively towards different members of a coyote pack? Canadian Journal of Zoology 77:499- 503. Magurran, A. E., B. H. Seghers, G. R. Carvalho, and P. W. Shaw. 1992. Be- havioural consequences of an artiﬁcial introduction of guppies (Poecilia reticulata) in N. Trinidad: evidence for the evolution of anti-predator behaviour in the wild. Proceedings Royal Society of London 248:117-122 Mech, D. L. 2000. A record large wolf, Canis lupus, pack in Minnesota. Cana- dian Field-Naturalist 114:504-505. Ozoga, J. J., and L. J. Verme. 1986. Relation of maternal age to fawn rearing success in white-tailed deer. Journal of Wildlife Management 50:480-486. SPSS. 2000. SYSTAT 10. SPSS Science and Marketing Department. Chicago, Illi- nois. Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 147

VARIATION IN SMALL MAMMAL RICHNESS AMONG ECOTYPES IN EASTERN SOUTH DAKOTA

David F. Terrall, Nick G. Cochran, and Jonathan A. Jenks Department of Wildlife and Fisheries Sciences South Dakota State University Brookings, SD 57007

ABSTRACT

Small mammal diversity and distribution was assessed for three ecotypes at Oakwood Lakes Game Production Area in eastern South Dakota. Ecotypes were an eastern red cedar (Juniperus virginiana) stand, native grassland consisting mainly of big bluestem (Andropogon gerardii), Indian grass (Sorghas- trum nutans), and switchgrass (Panicium virgatum), and a cultivated corn (Zea mays) food plot. Deer mice (Peromyscus maniculatus) (38%), short- tailed shrews (Blarina brevicauda) (38%), masked shrews (Sorex cinereus) (12%), meadow voles (Microtus pennsylvanicus) (6%), a meadow jumping mouse (Zapus hudsonius) (3%), and a thirteen-lined ground squirrel (Citellus tridecemlineatus) (3%) were trapped using snap, pitfall, and small live traps. Species were combined into mice (i.e., deer mice and meadow voles) and shrews (i.e., short-tailed shrews and masked shrews) to test the hypothesis that distribution of small mammals differed across ecotypes. No signiﬁcant difference in species diversity (P=0.118) or distribution (P= 0.700) was found among ecotypes. Results indicate that necessary ecological components exist in non- native, human induced habitats to support populations of small mammals.

Keywords

Blarina brevicauda, deer mouse, eastern South Dakota, ecotypes, masked shrew, Peromyscus maniculatus, short-tailed shrew, small mammals, Sorex cinerus

INTRODUCTION

Small mammals provide an important wildlife component among habitats of North America (Gibbons 1988). In eastern South Dakota, habitat use and distribution of small mammals is poorly documented except for a few studies associated with wetlands and shelterbelts (Lindell 1971, Searls 1974, Barnes and Lindner 1982). Information on small mammal use of food plots and cultivated agricultural ﬁelds also is lacking. Small mammals can act as a buffer against game bird predation and inﬂuence predator population dynamics (Wilson and Bromley 2001,Schwartz and Schwartz 1981). 148 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002)

Habitat variability can have a profound affect on the distribution of mammals by providing an increased number of available niches (Adler 1988, Sieg 1988). Small mammal abundance can be inﬂuenced by changes in habitat such as losses of food and cover associated with tree harvest and the cultivation of herbaceous vegetation following disturbances, thus making population estimates and species diversity difﬁcult to measure (Szaro et al. 1988, Waters and Zabel 1998). The purpose of this study was to examine the diversity and distribution of small mammals among three ecotypes in eastern South Dakota. We hypothesized that native grasslands would have higher species richness and abundance than other (i.e., coniferous woodland and cultivated food plots) un- natural ecotypes.

STUDY AREA

The study area was a Game Production Area (GPA) managed by the South Dakota Department of Game, Fish and Parks located in Brookings County, in eastern South Dakota. Location of the plots were Township 111 N, Range 51 W in the south one half of Section 8 between Lake Tetonkaha and Lake Oak- wood. Elevations for the study site ranged from 494 to 500 m. The physio- graphic description of the area is the Central Prairie Coteau region (Westin and Malo 1978). Soil zone is cool moist prairie with average annual temperatures of 5ºC to 7.2ºC and average annual rainfall between 45.6 cm to 55.2 cm (West- in and Malo 1978). The Oakwood GPA surrounding the study site was a mixture of coniferous forest, grasslands, and a cultivated food plot. Coniferous forest was dominated by eastern red cedar. Grasslands were composed of native grasses consisting of big bluestem, Indian grass, and switchgrass intermixed with Kentucky bluegrass (Poa praetensis), smooth bromegrass (Bromus iner- mis), sweetclover (Melilotus spp.), prairie rose (Rosa arkansana), alfalfa (Med- icago sativa), green sagewort (Certemisia drucunculus), and Canada golden- rod (Solidago canadensis). The cultivated food plot was primarily corn intermixed with yellow foxtail (Seteria glauca) and green foxtail (Seteria viridis). Scientiﬁc nomenclature follows that of Johnson and Larson (1999).

METHODS

Three cover types were chosen for the study; a stand of eastern red cedar, a native grass seeding, and a cultivated unharvested corn food plot. One, ﬁve by ﬁve trapping grid was set per ecotype, with stations set 5 m apart for a total area of 625 m2. Four trap types were placed in a square pattern with even distribution throughout the grid (Fig. 1). Trap types were; 1) 11 snap traps, 2) 5 pit-fall traps (#10 tin cans with 8 cm by 1 m tarp fence on both sides), 3) 3 large live traps (38 cm x 38 cm x 91 cm) and 4) 6 small live traps (8 cm x 8 cm x 30 cm). Traps were baited with a mixture of peanut butter, raisins, beef fat, and oatmeal (Giles 1969) and checked daily. Species frequencies for each habitat were divided by the trap nights per grid and multiplied by 1000. Anal- ysis of Variance (ANOVA) was used to test for effects of species and ecotype. Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 149

SN SL LL SN P

SL SN SN P SL

SN SL P SN SN

LL P SN SL LL

PSNSLSNSN

Figure 1. Outline of trap (SN=snap trap, SL=small live trap, P=pitfall trap, LL=large live trap) placement in ecotypes at Oakwood Lakes Game Production Area, November 2001.

RESULTS

Traps were set from 3 to 16 November 2001 for a total of 13 nights. The large live traps were removed from analyses because those traps produced no captures. Twenty-two traps were used in the analysis for each grid for a total of 286 (snap traps = 143, pitfall traps = 65, small live traps = 78) trap nights per grid. A total of 858 (snap traps = 429, pitfall traps = 195, small live traps = 234) trap nights were analyzed for all three grids. Six species of small mammals were captured during the study; deer mice (n=13), short-tailed shrews (n=13), masked shrews (n=4), meadow voles (n=2), a thirteen-lined ground squirrel (n=1), and a meadow jumping mouse (n=1). The meadow jumping mouse and thirteen-lined ground squirrel were removed from the analysis because only one individual of each species was captured during the study. Species richness was similar across ecotypes with the grassland and corn ecotypes each having four species, and the cedar stand having three species (Table 1). Species were combined into mice species (deer mice and meadow voles), and shrew species (short-tailed shrews and masked shrews) in analyses. No signiﬁcant difference was noted in species types (mouse vs. shrew) (F1,2=7.000,

P=0.118) by ecotype. In addition, no signiﬁcant difference (F2,2=0.429, P=0.700) was found in captures/trap night for ecotypes. 150 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002)

Table 1. Species and species (per trap night) captured in ecotypes at Oakwood Lakes Game Production Area, November 2001.

Habitat Species Number Number/trap nights Captured x 1000

Cedar Deer mouse 5 17.5 Short-tailed shrew 2 7.0 Meadow jumping mouse 1 3.5

Grassland Short-tailed shrew 7 24.5 Masked shrew 3 10.5 Meadow vole 1 3.5 Thirteen-lined ground squirrel 1 3.5

Corn Deer mouse 8 28.0 Short-tailed shrew 4 14.0 Masked shrew 1 3.5 Meadow vole 1 3.5

Table 2. Captures by trap type used in ecotypes at Oakwood Lakes Game Production Area, November 2001.

Trap Type Species Number Captured

Snap Trap Deer mouse 13 Short-tailed shrew 9 Masked shrew 1 Meadow vole 2

Pitfall trap Short-tailed shrew 4 Masked shrew 3

Small live trap Meadow jumping mouse 1 Thirteen-lined ground squirrel 1

DISCUSSION

The relative distribution of small mammal species captured was similar across habitats. Seig (1988) found similar results showing average numbers of small mammals distributed among grassland and woodland habitats. Seig (1988) captured deer mice in both woodland and grassland types with 66% of the total deer mice captures in woodlands and 48% of captures in grasslands. Results differed from our study in that no deer mice were captured in grassland areas. Deer mice were however, captured in corn and represented 62% of the total deer mice captured. Short-tailed shrews were captured in all ecotypes with 54% of captures occurring in grassland. Adler (1988) documented a positive association of white-footed mice (Peromyscus leucopus) and woody vegetation and a negative association with herbaceous vegetation, which paralleled our study. Adler (1988) also found that meadow voles were positively Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 151 associated with herbaceous grassland habitats but avoided woody habitats. Those ﬁndings were similar to our study because meadow voles were present in the grassland and corn but absent from the cedar ecotype. Effectiveness of various trap types in this study paralleled those of Szaro et al. (1988). Pitfall traps were only effective on shrew species. Szaro et al. (1988) theorized that deer mice might escape by jumping out of pitfalls traps. The most effective trap type however, was the snap trap with greater numbers (74%) and species captured (67%). Small mammals are a signiﬁcant and important ecological wildlife component in any ecosystem. Research and management of these species has not kept pace with those of more popular wildlife species. Schwartz and Schwartz (1981) indicated that small mammals could reduce predation on game birds by acting as an alternate prey species for small predators. Wilson and Bromely (2001) also found a functional response by arctic foxes (Alopex lagopus) to high lemming density; foxes spent less time foraging in goose nesting sites during years of high lemming cycles. Small mammals can have both positive and negative effects on plant communities and soil quality (Schwartz and Schwartz 1981, Hayward and Philipson 1979). Tunnels and burrows from small mammals help to aerate soils and reduce runoff, while plant herbivory can affect seed dispersal, community structure, and reduce crop yields (Schwartz and Schwartz 1881). Results of this study indicate that necessary ecological components exist in non-native, human induced habitats to support populations of small mammals. The large amount of agricultural land in eastern South Dako- ta warrants inclusion of these areas into future studies of small mammal populations.

ACKNOWLEDGMENTS

Support for this study was provided by South Dakota State University. We thank the South Dakota Department of Game Fish and Parks for providing the use of the study area and some large traps.

LITERATURE CITED

Adler, H.A. 1988. The role of habitat structure in organizing small mammal populations and communities. Proceedings of a symposium on the management of amphibians, reptiles, and small mammals in North America. Pages 289-299. U.S. Forest Service General Technical Report. RM-166. Barnes, T.G., and R.L. Linder. 1982. Small mammal occurrence in South Dako- ta shelterbelts and movements of Peromyscus maniculatus. Proceedings of the South Dakota Academy of Science. 61:56-63. Gibbons, J.W. 1988. The management of amphibians, reptiles, and small mammals in North America: the need for an environmental attitude adjustment. Proceedings of a symposium on the management of amphibians, reptiles, and small mammals in North America. Pages 4-9. U.S. Forest Service Gen- eral Technical Report. RM-166. 152 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002)

Giles, R.H. Jr. 1969. Wildlife Management Techniques. Third edition. The Wildlife Society, Washington, D.C. Hayward, G.F. and J. Phillipson. 1979. Community structure and functional role of small mammals in ecosystems. Pages 135-211 in D. M. Stoddard, editor. Ecology of small mammals. John Wiley and Sons, New York. Johnson, J.R., and G.E. Larson. 1999. Grassland plants of South Dakota and the Northern Great Plains. South Dakota State University, College of Agri- cultural and Biological Sciences, Ag Communications, Brookings, South Dakota. Lindell, J.R. 1971. Small mammal distribution in relation to marshland vegetation types in southeastern South Dakota. Thesis, University of South Dako- ta, Vermillion. Schwartz, C.W., and E.R. Schwartz. 1981. The wild mammals of Missouri. Uni- versity of Missouri Press, Columbia. Searls, D.A. 1974. Influence of vegetation on the distribution of small mammals on a waterfowl production area. Thesis, South Dakota State University, Brookings. Sieg, C.H. 1988. The value of Rocky Mountain juniper (Juniperus scopulorum) woodlands in South Dakota as small mammal habitat. Proceedings of a symposium on the management of amphibians, reptiles, and small mammals in North America. Pages 328-332. U.S. Forest Service General Techni- cal Report. RM-166. Szaro, R.C., L.H. Simmons, and S.C. Belfit. 1988. Comparative effectiveness of pitfalls and live-traps in measuring small mammal community structure. Proceedings of a symposium on the management of amphibians, reptiles, and small mammals in North America. Pages 282-288. U.S. Forest Service General Technical Report. RM-166. Waters, J.R., and C.J. Zabel. 1998. Abundances of small mammals in fir forests in Northeastern California. Journal of Mammalogy. 79(4):1244-1253. Westin, F.C., and D.D. Malo. 1978. Soils of South Dakota. Plant Science De- partment, Agricultural Experiment Station, South Dakota State University, Brookings, South Dakota. Wilson, D.J., and R.G. Bromley. 2001. Functional and numerical responses of predators to cyclic lemming abundance: effects on loss of goose nests. Canadian Journal of Zoology. 79(3):525-532. Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 153

THE EXPANDING DISTRIBUTION OF THE LEAST SHREW, CRYPTOTIS PARVA, IN SOUTH DAKOTA

Douglas C. Backlund South Dakota Natural Heritage Program South Dakota Dept. of Game, Fish and Parks Pierre, SD 57501

ABSTRACT

Westward range expansions of least shrew, Cryptotis parva, have been reported for New Mexico, Texas, Kansas, Nebraska, and Colorado. A westward range expansion is now reported for South Dakota with the ﬁrst documenta- tion of a northward range expansion. Prior to 1992, only two specimen records of C. parva were reported from South Dakota. One specimen was collected in 1932 near Okreek in Todd County by T. J. Turner (USNM #250199). The second specimen was collected near Cottonwood in Jackson County in 1954 by J. S. Findley. Since 1992, small mammal trapping in western South Dakota has extended the known range of the least shrew considerably north and west, in- to Dewey and Ziebach counties to the north and into Fall River County, near the Wyoming border, to the west. Other specimens have been taken from near Pierre in Hughes County and from Wind Cave National Park in Custer County. A lack of historical specimen records from elsewhere in South Dakota, particularly the lack of records from the Black Hills region, suggests that the species is colonizing habitat in a northward and westward direction.

INTRODUCTION

In the Northern Great Plains, C. parva, a short-tailed shrew, is easily iden- tiﬁed, unlike the species of long-tailed shrews in the genus Sorex. Blarina brevicauda is the only other species of short-tailed shrew known to occur in South Dakota. C. parva is much smaller than B. brevicauda. B. brevicauda has lead- gray pelage while C. parva is brownish-gray with lighter underparts. In addition, Cryptotis has only 30 teeth (large incisors 1/1, unicuspids 4/1, large premolars 1/1, molars 3/3), whereas Sorex and Blarina have 32 teeth (large incisors 1/1, unicuspids 5/1, large premolars 1/1, molars 3/3). C. parva is the most widespread species in a genus of about twelve extant species (Choate, 1970). Most species of Cryptotis inhabit forests of tropical and subtropical Cen- tral and South America, but C. parva, the only species found north of Mexico, usually inhabits grasslands with average annual precipitation approaching or greater than 50 cm (Porter, 1978; Hafner and Shuster, 1996). Until 1992 C. parva was reported from only two locations in South Dakota, both in the south- central area of the state (Fig. 1). Small mammal trapping in grassland and riparian habitats of central and western South Dakota has resulted in specimen 154 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002)

records of C. parva north and west of the reported range (Fig. 2). At the same time, extensive trapping in eastern South Dakota has not detected C. parva.

METHODS

Small mammal trapping in western and central Figure 1. Historic specimen records of Cryptotis South Dakota has docu- parva in South Dakota. mented new range extensions of C. parva. A search of the literature, the South Dakota Natural Heritage Database, and personal communication with mammalogists and wildlife biologists are combined in this paper to present the current distribution pattern of C. parva in South Dakota. Interpretation of the new distribution records is based on other research re- Figure 2. Historic and new specimen records for sults and the fossil record Cryptotis parva in South Dakota. of the late Pleistocene and Holocene. The author has deposited specimens from Hughes County, Ziebach County, Dewey County and Fall River County at Uni- versity of Kansas Natural History Museum, KU155934, KU155933, KU155932, KU155931, KU155935, KU146018, KU146017, KU146016, KU155936.

DISCUSSION

Prior to 1992, only two specimen records of C. parva were known from South Dakota. One specimen was collected in 1932 near Okreek in Todd County by T. J. Turner (USNM #250199). The second specimen was collected near Cottonwood in Jackson County in 1954 by J. S. Findley (1956). Based on the lack of records, the species was thought to be rare in South Dakota and was one of the original rare species tracked by the South Dakota Natural Her- itage Program (Houtcooper, et al., 1985). Since 1992, small mammal trapping in western South Dakota has extended the known range of the least shrew considerably northward and westward. The author has collected specimens in Dewey and Ziebach counties to the Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 155 north and in Fall River County, near the Wyoming border, to the west. Many specimens have been taken from near Pierre in Hughes County (Backlund, 1995). One specimen was collected at Wind Cave National Park in Custer County in 1999 (Jeremy Duckwitz, personal communication). Figure 3. Late Pleistocene and Holocene Distribu- Eleven specimens of C. par- tion of Cryptotis Parva (closed circles, from Faun- va were taken in 2000 on R. map Database, Illinois State Museum) and modern E.(Ted) Turner’s Bad River distribution (black line; from Hall, 1981 Mammals Ranches in Stanley and of North America. Jones counties (Kevin Hon- ness, personal communication). These are significant range extensions beyond the known historical distribution. Range expansions have been reported in Texas (Jones, 1993), New Mexico (Hoditschek et al., 1985), Nebraska (Benedict et al., 2000), Kansas, and Colorado (Choate and Reed, 1988). The new South Dakota records could be the result of a natural range expansion or range extensions that reflect increased trapping effort. While other researchers have suggested that irrigation increased available habitat for range expansion (Hafner and Shuster, 1996; Armstrong, 1972), none of the collection sites in South Dakota were affected by irrigation. Habitat at recent South Dakota collection sites ranged from dense riparian vegetation to dry prairie. One specimen in Ziebach County was taken in a dry woody draw. All of the specimens were taken from relatively undis- turbed habitats. Hafner and Shuster (1996) reported that one of three peripheral populations in eastern New Mexico represented a relic population from the late Wis- consinan, based on a unique allozymic complement and cranial morphology, while the other two populations were thought to be recent dispersals. Bene- dict et al. (2000) speculated that C. parva taken from the Pine Ridge area of northwestern Nebraska differed sufficiently from populations in eastern Ne- braska to indicate vicariance but did not rule out recent dispersal. Harris (1985) speculated that bones of Cryptotis from the Little Boxelder Cave, in the foothills of the Laramie Mountains in southeastern Wyoming were from populations that spread west during the pluvial pre-Altithermal or, alternatively, that the bones could have come from populations of Cryptotis forced to higher elevations during the warmer and drier Altithermal. The Black Hills function as refugia for many species, isolated from the more arid and warmer climate of the surrounding grasslands. Small mammalian species such as Clethrionomys gapperi, Microtus longicaudus, and Za- pus hudsonius persisted in the Black Hills during climatic extremes of the late Pleistocene and Holocene to the present (Turner, 1974). Jones et al. (1983) placed Cryptotis parva in the eastern faunal element. Other flightless, small 156 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) mammal species in the eastern faunal element include Peromyscus leucopus and Synaptomys cooperi. Peromyscus leucopus is extant in the Black Hills but Synaptomys cooperi appears to be extipated (Turner, 1974). S. cooperi remains have been recovered from the Beaver Creek Holocene site in Wind Cave Na- tional Park (Rachel Benton, personal communication). S. cooperi is a moist grassland species that is now sympatric with C. parva across a broad range in the east-central United States. A grassland species such as C. parva could have persisted during climatic extremes as a relic population in the Black Hills region, utilizing riparian areas and wet meadows around the periphery of the Black Hills. However, as with S. cooperi, there is no evidence that vicariance occurred in the Black Hills (Turner, 1974). The Cryptotis remains from Little Boxelder Cave, Converse County, Wyoming indicate a Holocene distribution (Harris, 1985) that would have included the Black Hills region (Figure 3). Both species were probably extirpated from this region during the Holocene, possibly during the Altithermal period. Turner (1974) does not include C. parva as a species of unverified occurrence or as a species of uncertain status in the Black Hills region. Turner considered the least shrew to be completely absent from the area. Cinq-Mars et al. (1979) used approximately 144 pitfall traps in Jackson and Haakon counties during the summer of 1970. Although four specimens of the rare Sorex nanus were taken, no C. parva were detected. The 1990’s collections in western South Dakota, including specimens from Wind Cave National Park, Fall River Coun- ty, Hughes County, Ziebach County and Dewey County indicate the species is in fact expanding northward and westward. The efforts of Cinq-Mars et al. (1979) and Turner (1974) indicate that C. parva has not been merely overlooked. However, the Jackson County specimen taken by Findley in 1954 is somewhat of an enigma. Findley’s record was based on a specimen consisting of the rostrum and one mandible with the covering skin. Findley stated "To my knowledge this species has not been recorded in the Great Plains northwest of a line connecting Yuma County in northeastern Colorado with the area of the type locality in Washington County, Ne- braska." Findley was not aware of the Todd County specimen collected by T. J. Turner in 1932. Recent small mammal trapping in eastern South Dakota has not detected C. parva. There are no historical records. Skadsen conducted intensive pitfall trapping in Day and Marshall counties in 1997, catching more than one hundred shrews of the genus Sorex, including eighteen Sorex hoyi, a species once considered rare in South Dakota (Dennis Skadsen, personal communication). Mullican used pitfall traps in a variety of locations in eastern South Dakota, capturing many shrews, including S. hoyi (Mullican, 1993). King et al. (1999) trapped shrews in Union County in 1994 and 1995, also capturing S. hoyi. Pit- fall traps were used at Ordway Prairie to capture shrews in 1983, including S. hoyi and the second state record for Sorex arcticus (Gruebele and Steuter, 1988). Rinker trapped small mammals for many years in the Vermillion area of southeastern South Dakota (George Rinker, personal communication). None of these sampling efforts detected C. parva in eastern South Dakota. Other small mammal sampling efforts are too numerous to list but are cited in Higgins et Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 157 al. (1999). The apparent absence of remnant populations in eastern South Dakota is further evidence that C. parva was not historically present. The species is expanding in the western South Dakota but is apparently unable to colonize favorable habitat in eastern South Dakota, possibly due to the barrier created by extensive conversion of upland grassland to agriculture. Loss of grassland habitat in eastern South Dakota may be a barrier to dispersal in that region of the state even though the higher annual precipitation in southeastern South Dakota indicates optimum habitat in remaining grasslands. The Missouri River may act as a barrier. It is interesting that the only C. parva specimens collected from the eastern side of the Missouri River were taken near Pierre, near the Oahe Dam (Backlund, 1995). The dam may provide a dispersal route across the river allowing recent colonization of this area. More small mammal sampling is needed along the Missouri River to determine the actual distribution of C. parva. Competitive exclusion due the presence of other shrew species, such as Sorex haydeni, Sorex cinereus, S. hoyi, S. arcticus, and B. brevicauda may also be a factor limiting the distribution of C. parva in eastern South Dakota and the Black Hills. These five species are found in eastern South Dakota in varying abundance, becoming much less common to completely absent in most of western South Dakota. B. brevicauda and S. haydeni are widespread and often abundant in eastern South Dakota. In the Black Hills, S. haydeni is the dominant Soricid and appears to be the only Soricid present in most of the Black Hills. Sorex merriami (Mullican, 1994) and Sorex nanus are known from grasslands and scrub habitats of western South Dakota, but are rare and local. The drier western grasslands may be suitable habitat for C. parva but not optimal for S. haydeni or B. brevicauda, species that become more abundant to the east. Porter (1978) also speculated that competition with Sorex species may inhibit dispersal and colonization of C. parva. The construction of thousands of small stock dams in western South Dako- ta could provide new habitat, free of Soricid competitors, for C. parva. These stock dams may provide "stepping stone" habitats for C. parva to advance westward and northward and refuge for survival in drier years. It is likely that range expansions occur during years of higher precipitation or warmer winters followed by retreats during less optimum years. Small populations become isolated in refugia until favorable conditions return. Such a pattern could result in periods of time when the species is very difficult to detect followed by periods of abundance and expansion.

CONCLUSIONS

I suggest that the South Dakota distribution of Cryptotis parva is expanding north and west. C. parva may be taking advantage of new habitat created by the construction of thousands of stock dams in western South Dakota. Ex- pansion and colonization into eastern South Dakota may be limited on a landscape scale by the Missouri River, loss of grassland habitat, and the presence of Soricid competitors in the remaining suitable habitat. It is likely that a gen- 158 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) eral trend of increasing annual precipitation and and temperature is contribut- ing to this expansion. Westward range expansions have been reported along the entire western periphery of the range of C. parva. This is the ﬁrst report of a northward range expansion. I predict new state records for C. parva in eastern Wyoming and south-central North Dakota in the near future.

LITERATURE CITED

Armstrong, D. M. 1972. Distribution of Mammals in Colorado. Monograph Mu- seum of Natural History, University of Kansas 3: 1-415. Backlund, D. C. 1995. New Records for the Dwarf Shrew, Pygmy Shrew, and Least Shrew in South Dakota. Prairie Naturalist 27(1): 63-64. Benedict, R. A., H. H. Genoways, and P. W. Freeman. 2000. Shifting Distribu- tional Patterns of Mammals in Nebraska. Transactions of the Nebraska Academy of Sciences 26: 55-84. Choate, J. R. 1970. Systematics and zoogeography of Middle American shrews of the genus Cryptotis. Univ. Kansas Publ. Mus. Nat. Hist. 19:195-317. Choate, J. R. and K. M. Reed. 1988. Least Shrew, Cryptotis parva, in South- western Kansas and Southeastern Colorado. Southwestern Naturalist 33:361-362. Cinq-Mars, R. J., R. S. Hoffman, and J. K. Jones, Jr., 1979. New Records of the Dwarf Shrew (Sorex nanus) in South Dakota. Prairie Naturalist 11:7-9 Findley, J. S. 1956. Distribution of Some South Dakota Mammals. Natural His- tory Miscellanea. Chicago Academy of Sciences. No. 155. Gruebele, M. J. and A. A. Steuter. 1988. South Dakota Records of Pygmy and Arctic Shrews: Response to Fire. Prairie Naturalist 20: 95-98. Hafner, D. J. and C. J. Shuster. 1996. Historical Biogeography of Western Pe- ripheral Isolates of the Least Shrew, Cryptotis parva. Journal of Mammalo- gy 77(2): 536-545. Harris, A. H., R. A. Smartt, and W. R. Smartt. 1973. Cryptotis parva from the Pleistocene of New Mexico. Journal of Mammalogy 54: 512-513. Harris, A. H. 1985. Late Pleistocene Vertebrate Paleoecology of the West. Univ. of Texas Press, Austin. 293 pp. Higgins, K. F., A. E. Gabbert, T. R. Cooper, and J. J. Higgins. 1999. An Anno- tated Bibliography: Published Articles (1861-1998) about the Wild Mam- mals of South Dakota. Dept. of Wildlife and Fisheries, SDSU, Brookings, S.D. B731. 99 pp. Hoditschek, B., J. F. Cully, Jr., T. L. Best, and C. Painter. 1985. Least Shrew (Cryptotis parva) in New Mexico. Southwestern Naturalist 30(4):600-601. Houtcooper, W. C., D. J. Ode, J. A. Pearson, and G. M. Vandel III. 1985. Rare Animals and Plants of South Dakota. Prairie Naturalist 17(3): 143-165. Jones, J. K., D. M Armstrong, R. S. Hoffman, and C. Jones. 1983. Mammals of the Northern Great Plains. University of Nebraska Press, Lincoln and Lon- don. 371 pp. Jones, J. K., Jr., R. W. Manning, F. D. Yancey II, and C. Jones. 1993. Records of Five Species of Small Mammals from Western Texas. Texas Journal of Sci- ence 45:104-105. Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 159

King, C. B., G. M. Wilson, and P. D. Sudman. 1999. New Records of the Pygmy Shrew in Southeast South Dakota. Prairie Naturalist 31(2): 115-117. Mullican, T. R., 1993. Distribution of the Pygmy Shrew in South Dakota. Prairie Naturalist 24(4):257-259. Mullican, T. R. 1994. First Record of Merriam’s Shrew from South Dakota. Prairie Naturalist 26(2):173. Porter, L. S. W. 1978. Pleistocene Pluvial Climates as Indicated by Present Day Climatic Parameters of Cryptotis parva and Microtus mexicanus. Journal of Mammalogy 59(2):330-338. Turner, R. W. 1974. Mammals of the Black Hills of South Dakota and Wyoming. University of Kansas, Museum of Natural History. Misc. Publication No. 60. University of Kansas, Lawrence. 178 pp.

Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 161

RARE EARTH ELEMENT SIGNATURES OF FOSSIL VERTEBRATES COMPARED WITH LITHOSTRATIGRAPHIC SUBDIVISIONS OF THE UPPER CRETACEOUS PIERRE SHALE, CENTRAL SOUTH DAKOTA

Doreena Patrick Department of Geology Temple University Philadelphia, PA 19122

James E. Martin Museum of Geology South Dakota School of Mines and Technology, Rapid City, South Dakota 57701

D.C. Parris Natural History Bureau New Jersey State Museum Trenton, NJ 08625

D.E. Grandstaff Department of Geology Temple University Philadelphia, PA 19122

Keywords

Rare earth elements, Pierre Shale, South Dakota, mosasaur, Verendrye, Cre- taceous, Hornerstown, Sharon Springs, paleoenvironments, New Jersey

ABSTRACT

Rare earth elements were measured in mosasaur bones collected from ﬁve members (Sharon Springs, Gregory, Crow Creek, DeGrey, and Verendrye) of the upper Cretaceous Pierre Shale at localities near the Missouri River in Brule, Buffalo, Hughes and Hyde counties. Fossils from each member of the Pierre Shale have different REE signatures. Signatures of fossils from individual members may be distinctive over wide areas; fossils from the Verendrye Member have REE signatures that are consistent over 250 square kilometers. Fossils from the Sharon Springs Member have distinctive REE signatures that may be further subdivided into three superposed groups that correspond with the upper, middle, and lower Sharon Springs Member. Because REE signatures may differ 162 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) among stratigraphic units, fossil bones eroded from their stratigraphic context may be assigned to their proper depositional unit based on REE signature comparisons. We interpret changes in REE signatures among members as resulting from differences in mixing between two end members: oxygenated and anoxic seawaters. If differences in mixing are interpreted as depth differences, the lower Sharon Springs Member was deposited in deep, anoxic water. Water depths decrease in the middle and upper Sharon Springs, but the overlying Gregory Member was deposited in shallow water. The overlying Crow Creek, DeGrey, and Verendrye members were then deposited in progressively deeper marine waters, but not as deep as the lower Sharon Springs. These interpretations are generally consistent with those based on faunal diversity and eustatic sea level curves. REE signatures in fossils from the Campanian Verendrye Member were also compared with those of similar fossils from the Maastrichtian Navesink For- mation of New Jersey. Although the signatures differ, they are sufﬁciently similar so as to indicate similar degrees of mixing of oxygenated and anoxic deep waters and suggest similar water depths.

INTRODUCTION

REE concentrations and neodymium isotopes in fossil bones and teeth have been used to infer paleo-redox conditions in marine waters (Wright et al., 1987), detect reworking of fossils (e.g., Trueman and Benton, 1997; Trueman, 1999; Staron et al., 2001), and to accomplish or test paleoenvironmental (Gi- rard and Albarède, 1996; Reynard et al., 1999) and paleogeographic reconstructions (Wright et al., 2002). Many of these and other studies have examined stratigraphic variations of REE and have attempted to infer paleoenvironmental or paleoredox conditions and their variations in time and space as well as chemical nature and origin of water masses producing those signatures. The total Rare Earth Element concentrations (ΣREE) in modern bones and teeth are generally less than 20 ppm (Chenery et al., 1996; Staron et al., 2001; Patrick et al., 2001). However, ΣREE in fossil bones may be greater than 1,000 to 10,000 ppm (e.g., Arrhenius et al., 1957, and Table 1). Therefore, more than ca. 95% of REE in fossil bone is diagenetically incorporated into bone post mortem. Because most REE are introduced post mortem, REE signatures in fossils do not reﬂect the diet, trophic level, or phylogenetic position of the organism. Different osteological materials from a single organism may have different concentrations of REE; however, the signatures in all of the bones are essentially the same within analytical error (Patrick et al., 2001, in preparation). Research indicates that REE and possibly other trace elements are incorporated within 3,000 to 10,000 years after deposition (Grandstaff et al., 2001; Patrick et al., 2001, in preparation; Millard and Hedges, 1995, 1996) during early diagenetic recrystallization of the bone apatite. After incorporation into the osteological material, the REE signature in fossils is apparently stable and serves as a record of depositional or early diagenetic conditions (Wright et al., 1987; Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 163 lable, - Not determined, Sh Spr = Sharon Springs Member (degree N) (degree W) 1 Plioplatycarpus2 Plioplatycarpus3 Verendrye Plioplatycarpus5 Verendrye 43.9335 Plioplatycarpus Verendrye 43.9335 99.5119 Verendrye 43.9335 99.5119 43.9335 99.5119 - 99.5119 - 214.7 - 31.27 55.70 - 128.2 161.1 8.11 27.70 21.70 42.68 32.67 6.20 85.56 5.91 7.34 17.56 34.94 23.55 1.70 4.29 5.40 4.62 9.43 25.05 36.04 1.13 1.50 3.89 7.95 6.51 9.92 26.50 23.09 1.02 2.20 6.31 3.24 6.95 19.30 20.31 6.67 1.65 2.78 3.24 0.98 17.43 5.04 5.94 2.85 0.71 0.98 4.51 0.76 7256 Tylosaurine65 Tylosaur (?) Lower Sh Spr53 Lower Sh Spr Mosasaur 43.701262 43.6967 Tylosaur Middle Sh Spr 99.401264 Mosasaur 43.7669 99.4119 Upper Sh Spr59 Mosasaur 9.31 Upper Sh Spr60 43.7782 99.4326 Mosasaur 13.86 18.18 43.7811 Gregory 23.23 Mosasaur 99.4298 3.11 10.04 Crow Creek 99.3989 3.65 19.15 19.64 12.04 DeGrey 19.98 * 4.60 3.34 * 47.79 13.24 4.79 19.14 1.06 44.32 1.7821 1.11 * 4.74 1.64 Plioplatycarpus 4.15 15.4236 * * 4.10 Plioplatycarpus 1.12 Verendrye 6.5933 3.51 0.49 0.47 Plioplatycarpus Verendrye 4.4534 1.37 0.75 2.50 43.9035 * 4953.2 Plioplatycarpus 2.38 469.9 Verendrye35 0.54 0.58 4987.5 3.73 0.40 43.9035 99.5384 825.6 Plioplatycarpus Verendrye 469.9 0.37 2.73 3.58 44.1841 0.55 0.92 22.82 172.2 99.5384 1932.2 Verendrye 0.86 0.42 44.1841 0.31 4.25 0.11 328.0 73.49 269.2 99.7289 - 0.11 44.1841 257.2 113.0 0.94 13.57 17.57 2.17 1.00 0.74 99.7289 50.84 422.3 731.7 0.67 214.4 69.06 7.47 0.11 0.48 3.26 0.09 99.7289 53.60 7.47 123.3 354.4 14.97 184.7 40.25 0.08 1.29 1.86 0.49 1060.4 221.1 46.22 28.31 291.4 4.15 208.0 3.61 321.8 0.11 0.24 48.59 3.44 194.0 37.29 6.27 318.3 23.50 25.71 1176.7 11.25 41.45 1.46 168.3 0.54 40.04 1.50 3.36 184.3 7.62 61.65 35.63 9.38 164.8 1251.5 0.31 25.49 8.19 28.43 8.80 221.6 34.22 8.30 51.96 6.63 4.61 1.31 61.08 8.13 46.63 7.53 21.47 30.32 13.32 8.64 46.23 6.74 51.22 3.06 5.34 40.26 1.92 11.35 6.97 47.45 18.53 5.71 34.41 10.35 5.79 49.40 3.30 35.85 31.65 4.87 10.86 0.83 5.56 33.47 4.58 30.62 5.19 4.78 30.35 4.74 0.87 31.36 4.68 4.85 Number Specimen Unit Latitude Longitude La Ce Pr Nd Sm Eu Gd Tb Dy Ho Er Tm Yb Lu Table 1.Table of the Pierre Shale. selected members from REE in fossils REE concentrations in ppm, * South Dakota School of Mines and Technology Specimen, primary Latitude and Longitude data not avai REE concentrations in ppm, * South Dakota School of Mines and Technology 164 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002)

Henderson et al., 1983; Trueman, 1999). REE signatures in fossils from different stratigraphic units may differ, providing unique "fingerprints" of variations in their depositional or early diagenetic environments. In this paper we present results of a preliminary study of REEs in mosasaur fossils from five Campanian members (Sharon Springs, Gregory, Crow Creek, DeGrey, and Verendrye) from the lower part of the upper Cretaceous (Cam- panian to Maastrichtian) Pierre Shale at localities near the Missouri River in Brule, Buffalo, Hughes and Hyde counties, central South Dakota. This study was conducted to determine whether REE signatures in vertebrate fossils vary stratigraphically from one member to another in the Pierre Shale. We have also investigated whether REE signatures are similar laterally within in a single member. Where REE in fossils differ stratigraphically they may be used to distinguish fossils from different units. Because the REE signatures reflect the original composition and source of water masses from which the REE were incorporated, we have attempted to draw inferences about paleoenvironmental conditions and the sources of marine waters in this part of the Western Interior Seaway. We introduce the use of ternary diagrams for interpreting variations of REE signatures in fossils and their paleoenvironmental implications.

STRATIGRAPHY OF THE PIERRE SHALE

The Pierre Shale, one of the best known lithologic units of the Western In- terior, was deposited during the upper Cretaceous (Campanian and Maas- trichtian Stages) and is primarily a sequence of organic-rich black and grey shales, and interbedded bentonites, most of which formed under relatively anoxic marine conditions. Originally named the Fort Pierre Group (Meek and Hayden, 1862), it is extensively exposed in South Dakota. Within central South Dakota, members recognized by Crandell (1958) have gained general acceptance and were used in the research reported here. Validity of the subdivision has been confirmed by Hanczaryk (2002), whose comprehensive field studies were performed in association with our own. Eight members were recognized by Crandell (1958) of which only the five lower members are pertinent to our work thus far. In ascending order, these are the Sharon Springs, Gregory, Crow Creek, DeGrey, and Verendrye members. All are widespread within the region, and the Sharon Springs has been recognized throughout the Northern Plains. The stratigraphic column of the Pierre Shale in central South Dakota is shown in Figure 1. The Sharon Springs Member has been described in numerous reports not only in central South Dakota, but also in the vicinity of the Black Hills and elsewhere. The member consists largely of dark fissile shale, which weathers blue- grey, with many bentonite beds, and has abundant pyrite and gypsum derived from recent pyrite weathering. Vertebrate and invertebrate fossils are abundant, including mosasaurs, plesiosaurs, diving birds, and large fish. Great variations of its thickness have been reported, ranging from four to six meters in outcrops to as much as thirty meters recognized by some drillers (Crandell, 1958). With- in the present study areas, complete sections are uncommon but thicknesses Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 165 should be closer to the higher figure. Three superposed lithostratigraphic units have been noted (Martin, 1996) and are in- formally termed lower, middle, and upper Sharon Springs until they are formally named. The Sharon Springs Member contains moderately abundant iron Figure 1. Stratigraphy of the upper Cretaceous phosphate (vivianite) nod- Pierre Shale in central South Dakota (after ules. Hanczaryk, 2002). The Gregory Member is approximately 23 m thick in the Ft. Thompson section (Hanczaryk, 2002) and consists primarily of grey claystone and marl with a few bentonites. Much of its bentonitic content appears to have been redeposited and dispersed, which is probably responsible for its characteristic "popcorn" weathering surface. The unit has yielded significant numbers of invertebrate (Baculites gregoryensis) and a few vertebrate fossils. The Crow Creek Member is a calcareous siltstone, locally sandy, which is distinguishable from the other members by a light grey to yellow-brown color. In addition to being readily recognized by surface appearance, it often supports a distinctive botanical cover. The member is essentially unfossiliferous, and only a single mosasaur sample from this member was available for REE analysis. The Crow Creek is 1.2 meters thick in the Fort Thompson section and is never a very thick unit. The DeGrey Member measured 10.1 meters in the Fort Thompson section and its lower portion is gray shale and interspersed with abundant iron-manganese concretions. It has yielded numerous vertebrate and invertebrate fossils. The upper portion of the member is less conspicuously concretionary and is mostly silver-gray shale with some bentonites. The Verendrye Member is primarily silver-gray shale within the study area, but only the lower 17 m is exposed in the Fort Thompson section. Vertebrate and invertebrate fossils are common. Within the primary study area a number of marker units are recognizable, giving additional accuracy to lithostratigraphic measurements. The relatively thin Crow Creek Member itself is one such stratum. Although not of such limited thickness, the iron-manganese concretionary levels of the DeGrey Member are readily recognized. A thin, but persistent, red stratum in the lower Verendrye Member provides a reliable reference datum essentially throughout the study area. Also, a recognizable terrace-forming concretionary stratum occurs about 20 meters above the DeGrey-Verendrye contact. Because a complete section of the Verendrye Member has not been found in the area, these marker beds are of considerable stratigraphic value. 166 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002)

Geochronological determinations of the Campanian Sharon Springs- Verendrye interval are of the order of ten million years, ranging from 81-71 Ma B.P. (Hanczaryk, 2002). In places, the lower part of the Crow Creek Member contains shocked quartz and feldspar grains which may originate from the 74.1 ± 0.1 Ma Manson impact structure (Izett et al., 1998). The biostratigraphic zonation of the Pierre Shale for this region, based on ammonites, is from the Bac- ulites obtusus Zone (oldest) through the Baculites reesidei Zone (youngest), a span of thirteen zones. The entire formation is interpreted to be of marine origin and appears to represent an ideal paleoenvironment for mosasaurs, highly aquatic lizards of the Late Cretaceous. The total thickness (Fig. 1) of this part of the Pierre Shale in central South Dakota, from the base of the Sharon Springs to the top of the Verendrye Mem- ber, is approximately 55 meters (Hanczaryk, 2002; Hanczaryk et al., 1999). In central South Dakota, the 17 meter-thick Verendrye Member is incomplete and has been truncated by an unconformity. In other areas of central South Dako- ta where the Verendrye is conformable with overlying Virgin Creek Member, the thickness of the Verendrye is about 46 meters. Similarly, the complete Sharon Springs section is probably about 30 meters. Therefore, the total thickness, compensating for unconformities, is about 110 meters. The time required for deposition of these members of the Pierre Shale was approximately 10 million years (Hanczaryk, 2002). Therefore, the average depositional rate is approximately 1.1 cm/1000 years.

METHODS

Mosasaur fossil sample numbers and localities are given in Table 1. Sam- ple preparation techniques follow those in Staron et al. (2001). For some specimens, small fragments that could be used for destructive analysis were available. These bone samples were immersed in water or a dilute acid solution in an ultrasonic cleaner and then dried. Other specimens were sampled by drilling into an intact bone using a Dremel variable speed electric drill. Be- tween samples the drill was cleaned with trace metal grade dilute (5%) nitric acid or trace metal grade acetone and rinsed in distilled water. Matrix and secondary minerals were removed by handpicking. An ultrasonic technique removed more of the matrix and secondary carbonate from the bone. Samples were then rinsed with distilled water and dried. Cleaned bone fragments were mechanically crushed in an acid-washed mortar and pestle. Approximately 0.2 gram of powder was weighed for each sample. Each sample was dissolved in

3 mL Ultrex-grade HNO3 and diluted to 50 mL with 5% HNO3. When only smaller masses of bone were available, the initial solution was diluted to smaller volumes in order to preserve the mass/volume ratio. Samples were diluted to appropriate levels with 5% ultra-pure HNO3, and an internal standard containing 10 ppb 145Nd and 2 ppb 171Yb was added. Samples were analyzed using a Finnigan MAT Element/1 Inductively Coupled Plasma – Mass Spectrometer (ICP-MS). Analytical procedures follow those in Field and Sherrell (1998). Re- sults of analyses are given in Table 1. The coefﬁcient of variation for most REE Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 167 is less than + 5% of the analyzed value. Analytical results presented in Figures 2–7 have been normalized relative to the North American Shale Composite (NASC) (Gromet et al., 1984).

RESULTS

Results of REE analyses of fossil bones are given in Table 1. REE signatures (patterns of REE concentration vs. atomic number) in bones from a single fossil are essentially identical within analytical error (Patrick et al., 2001, in preparation). REE signatures of fossils within individual members also appear nearly identical. For example, Figure 2 shows REE signatures in bones from mosasaur fossils collected from the Verendrye Mem- ber at four sites. These REE signatures are light rare earth (LREE) depleted with a small negative Cerium anomaly. REE concentrations in various fossils differ, in part due to differences in bone type. Cortical bone tends to have higher concentrations than trabec- ular bone from the same or associated fossils (Patrick et al., 2001; in preparation). However, the signatures (relative concentrations of Figure 2. NASC-normalized REE signatures in REE) are essentially identi- mosasaur fossils from the Verendrye Member of cal. Verendrye mosasaur the upper Cretaceous Pierre Shale. Although REE fossils were collected at concentrations vary in different fossils, REE sig- three different localities natures in all of the fossils are the same. Fossils containing REE in this diagram were collected over an approximately 250 over an area of more than 250 km2 in central km2 area of central South South Dakota. This indicates that, at least under Dakota. Therefore, at least some conditions, REE signatures from a strati- in some marine environ- graphic unit may be similar over extensive areas. ments, REE signatures may have great lateral extent. However, REE signatures in fossils from neighboring stratigraphic units may be signiﬁcantly different. Figure 3 illustrates examples of REE signatures that indicate that fossils from various parts of the Pierre Shale have different REE signatures. In many cases the differences can be visually distinguished. Signatures in Figure 3 vary from Middle REE (MREE) enriched in the lower Sharon Springs, to Heavy REE (HREE) enriched in the Gregory Member, and to slightly Light REE (LREE) depleted in the Verendrye Member. To conﬁrm the visual impression (Fig. 3), REE data were analyzed by Dis- criminant Analysis using NCSS statistical software (Hintze, 1997). Discriminant 168 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002)

Analysis is a statistical technique (Davis, 1986) which ﬁnds linear combinations of variables (REE ratios) that produces the maximum or optimal separation between the deﬁned groups (members). Discriminant Analysis was conducted using NASC-normalized REE ra-

tios (e.g. GdN/YbN, Fig. 4). Ratios were used, rather than concentrations, to remove the effect of differ- Figure 3. NASC-normalized REE signatures in mosasaur fossils from selected members of the ences in REE concentra- Pierre Shale. REE signatures from different strati- tions between different graphic units are visibly and statistically different. types of osteological materials and different localities. Discriminant Analysis was conducted using all REE ratios and by stepwise regression to select the best ratios. Results of Discriminant Analysis showed that the REE signatures (ratios) in the Verendrye and Sharon Springs members, for which several analyses are available, were statistically different. For example, linear combinations of

NdN/TmN and DyN/LuN ratios allowed accurate classification of the units. If REE signatures in fossils from different members or stratigraphic units are statistically different, then these differences may be used to establish or limit the provenience of fossils from unknown or questioned localities. In some cases, significant fossils may be found in float or field notes for fossils in museum collections may be incomplete or have become lost. In such cases the scientific value of the fossil may be limited by the lack of stratigraphic information. Comparison of the REE signature of the fossil with REE signatures of fossils from candidate stratigraphic units may allow the provenience of the fossil to be established or limited. The differences within and between members may be illustrated by plotting ratios of various REEs as a function of stratigraphy. GdN/YbN from the various members are shown in Figure 4. These data show that GdN/YbN ratios differ from one member to another and indicate that different members can be distinguished based on REE signatures and ratios. Gd and Yb also show relations between the MREE and HREE elements. The GdN/YbN ratio is high in the lower Sharon Springs Member, indicating MREE enrichment, decreases to a minimum in the Gregory Member, and then increases slightly to the Verendrye Member. The Verendrye Member shows a consistent REE pattern throughout the member, whereas the Sharon Springs displays significant REE variations within the member. The Sharon Springs Member may be further sub-divided into three parts, based on REE signatures. The subdivisions generally coincide with previously identified units by one of us (Martin) of upper, middle and lower Sharon Springs. Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 169

Another way of visual- izing variations in REE patterns is by use of a triangular or ternary diagram. Fig- ure 5 shows a ternary diagram with NASC-normalized Yb, Gd, and Nd (YbN,

GdN, and NdN) at the ver- tices. Yb is a representative HREE, Gd a MREE, and Nd a LREE. Further, Nd, Gd, and Yb are even-numbered elements, and therefore Figure 4. Shale-normalized Gd/Yb ratios (GdN/YbN) have higher concentrations in mosasaur fossils from selected members of the according to the Oddo- Pierre Shale. Harkins effect, and thus have generally more reliable analytical data. The LREEs, Ce and La, were not used because Ce, unlike most other REE, has two valence states and is subject to effects of oxidation and reduction and La which, although a member of the REE, often has slightly different chemical variations than the others. Other REE combinations were tested, particularly substi- tuting Pr for Nd, but do not Figure 5. Triangular Diagram of NASC-normalized greatly alter the patterns. values of Yb (a heavy rare earth [HREE]), Gd The ternary diagram al- (MREE) and Pr (LREE) in fossils from the Pierre lows the basic shape of the Shale. Flat, shale-like REE patterns will plot in the REE pattern to be repre- middle of the diagram, samples plotting toward the Yb vertex will be HREE-enriched, whereas sented. NASC-normalized N samples plotting toward the PrN vertex will be samples that plot in the LREE- and those plotting toward the GdN vertex middle of the diagram (33% MREE-enriched. Mosasaur samples plot along a YbN, 33% GdN, and 33% straight trend line (from lower Sharon Springs to NdN) have equal amounts Gregory), which may result from mixing of two end-member waters, one enriched in MREE and of these elements and will LREE and the other enriched in HREE. Coastal have a ﬂat, "shale-like" REE seawater may represent the HREE-enriched water patterns (near Verendrye and deep, anoxic bottom water the MREE and signature, Fig. 3). REE in LREE enriched end-members. shale samples from the Pierre Shale do indeed plot 170 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002)

near the center of the diagram. Samples plotting toward YbN will be enriched in the HREE and signatures will have a positive slope (e.g., Gregory, Fig. 3), those plotting near NdN are enriched in LREE with a negative slope, and those plotting near GdN are enriched in MREE and have a bell shape (e.g., lower Sharon Springs, Fig. 3). Thus fossils from the lower Sharon Springs are relatively enriched in MREE, whereas those from the Gregory are enriched in HREE and relatively depleted in the other end-members. REE data for fossils from the various members lie near a straight line between the Gregory and lower Sharon Springs Members (Fig. 5). This indicates that, as a ﬁrst approximation, REE data can be explained by mixing or evolution between a Yb-rich (HREE) end-member near the Gregory Member and a relatively Gd-rich (MREE, with subordinate LREE Nd) end-member near the lower Sharon Springs. The amount of variation in REE signatures in the Pierre is as great as that in some terrestrial units. The REE variations in the fossils should be related to variations in marine waters from which the REE were obtained.

FACTORS INFLUENCING REE SIGNATURES IN FOSSILS

A long debate continues about the source of REE, the mode of REE uptake by fossils, and the relationship between the REE signature of the fossil and the REE composition of the water. For example, some workers have proposed that fossils obtain their REE primarily from surface waters, including sea bottom waters (Wright et al., 1987). In this case REE signatures in marine fossils may be used to infer paleo-oceanographic conditions. However, others have argued that, depending on sedimentation rate, REE may be derived more from ground or pore waters (e.g. Elderﬁeld and Pagett, 1986). In this case REE signatures will more likely reﬂect early diagenetic conditions. Previous workers have suggested that REE signatures measured in vertebrate fossils may be affected by a number of factors, including: mechanism of REE incorporation into fossils, REE composition and speciation of water providing trace elements, redox state, sedimentation rate, as well as late diagenetic alteration.

Incorporation mechanisms

Reynard et al. (1999) suggested that REE could be incorporated into apatite by two mechanisms (1) equilibrium crystal-chemical fractionation between apatite and water and (2) adsorption and apatite surface reactions. They suggested that some "bell shaped" REE signatures, such as those commonly found in Paleozoic ichthyoliths or conodonts (e.g., Wright et al., 1987; Girard and Al- barède, 1996; Reynard et al., 1999), could arise from equilibrium fractionation between seawater and the biogenic apatite. Reynard et al. (1999) calculated relative distribution coefﬁcients or fractionation factors between water and apatite. Although their equilibrium fractionation model could qualitatively explain the "bell shaped" patterns, it could not quantitatively explain all of the REE ra- Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 171 tios, nor explain other patterns. Therefore, REE incorporation in fossil bone apatite results primarily from surface reactions between apatite and water. Incor- poration may be controlled by sorption of REE on the apatite surface, followed by incorporation during diagenetic apatite recrystallization (Trueman, 1999; Armstrong et al., 2001).

Water Composition and Speciation

Concentrations and signatures of REE have been measured in a variety of natural waters (Johannesson and Xiaoping, 1997; Piepgras and Jacobson, 1992); however, many factors controlling REE in natural waters are still a matter of debate. The aqueous properties of the REE change systematically across the series. For example, there is a systematic increase in the strength of REE-carbonate ion complexes from light to heavy REE (Haas et al., 1995). This leads to fractionation between the REE in natural waters. REE concentrations from some representative natural waters are plotted on a Nd-Gd-Yb ternary (Fig. 6). REE concentrations in seawater are very low and are strongly HREE enriched (Piepgras and Jacobsen, 1992). Thus, seawater signatures (grey circles) plot toward the Yb end of the Figure 6. NdN-GdN-YbN Ternary of REE signatures ternary (Fig. 6, data from from selected natural waters. Data from: Baue et Westerlund and Ohman, al. (1997), DeBaar et al. (1988), Elderﬁeld et al. 1992). The HREE enrich- (1990), German and Elderﬁeld (1989), Hoyle et al. ment is due to enhanced 91984), Johannesson and Xiaoping (1997), Piep- solubility of HREE in rela- gras and Jacobsen (1992), Sholkovitz et al. (1992) and Westerlund and Ohman (1992). tively high pH and alkaline seawater, produced by preferential HREE-carbonate complexing, and sorption of LREE by hydrous ferric oxides (HFO) and tests of planktonic organisms (Piepgras and Jacobsen, 1992). The slight variations between oceans appear to be consistent with variations between deep and shallow ocean water and oceanic circulation patterns (e.g., Piepgras and Jacobsen, 1992). These open marine waters also have large negative cerium anomalies as a result of strong sorption of oxidized Ce4+ on HFO. 172 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002)

In contrast, river water signatures (e.g. Elderfield et al., 1990) are relatively enriched in MREE and LREE, and often exhibit bell-shaped signatures. These data plot toward the upper left of the triangular diagram. REE signatures in estuaries (not plotted in Fig. 6) are variable and intermediate between river waters and sea waters (Hoyle et al., 1984; Elderfield et al., 1990; Sholkovitz and Szymczak, 2000). Coastal sea waters tend to have somewhat higher REE concentrations than does open ocean sea water. Their signatures (open circles) are somewhat variable but, except in the area of estuaries, dissolved REE tend to plot in the same area as do open ocean waters. Although most open marine waters and coastal sea waters have similar REE signatures, anoxic or suboxic marine bottom or pore waters are more LREE and MREE enriched (German and Elderfield, 1989; Sholkovitz et al., 1992) and tend to plot more toward the upper left of Figure 6 than do oxic marine waters. The LREE and MREE enrichment is probably due to desorption of LREE or dissolution of REE-bearing HFO under anoxic conditions (Sholkovitz et al., 1992; Bau et al., 1997). These anoxic waters also have small cerium anomalies, in contrast with overlying oxygenated marine waters.

Sedimentation rate and timing of REE uptake

Sedimentation rate may affect the source of waters (surface vs. pore or ground waters) providing REE. Researchers such as Bernat (1975) and Elder- ﬁeld and Pagett (1986) have suggested that REE and other trace elements are incorporated into fossils over relatively short periods of time, perhaps as little as 3,000 – 10,000 years (Patrick et al., 2001). If a fossil incorporates REE only at the sediment-water interface (zero sedimentation rate) then its REE signature must be most closely related to the REE composition of surface (sea bottom) water. However, if the fossil becomes deeply buried during REE uptake, then the REE signature in the fossil will result from a combination of surface and pore water REE compositions, depending on the rate and timing of incorporation. Sholkovitz et al. (1992) and others have shown that, with increasing depth, submarine pore waters become more enriched in REE and that the LREE/HREE ratio increases with depth as LREEs are preferentially released or desorbed from solid phases. Therefore, in a marine system, fossils incorporat- ing REE in areas of slow sedimentation could have lower LREE/HREE ratios, reﬂecting the low LREE/HREE ratio in seawater, whereas higher sedimentation rates should produce signatures with higher LREE/HREE values. Marine waters typically have strong negative Ce anomalies; however, anoxic bottom waters, pore waters, and some coastal seawaters have only small anomalies. The source of REEs incorporated into bone and the timing of incorporation may be important in determining the resulting REE signature, including the size of any Ce anomaly.

Diagenesis

Most studies (see discussion in Trueman, 1999) have suggested that once REE are incorporated into the bone, they are retained and provide a stable sig- Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 173 nal reflecting the depositional or diagenetic environment from which they were incorporated. Apatite crystals in living organisms are very small, their lattices have low degrees of crystallinity (Person et al., 1995), and they contain carbonate, sodium, and other species which increase the solubility of the bone material. During fossilization, and while the REE are being taken up and incorporated into the bone, the bone apatite recrystallizes. Fossil apatite contains larger crystals which have more highly crystalline lattices. Incorporation of REE, fluoride, and other trace elements may also lower the solubility of the resulting fossilized material. Growth of larger, more highly crystalline materials, which contain REE, fluoride and other elements, reduces the solubility and potential rate of dissolution of the fossil materials. Thus the material is much less likely to be affected by dissolution and remobilization or alteration of REE. REE signatures will be altered only by dissolution and re-precipitation of the apatite, by solid state diffusion, or recrystallization during metamorphism (Armstrong et al., 2001).

DISCUSSION AND PALEOENVIRONMENTAL INTERPRETATION

Changes in REE signatures in vertebrate fossils from the various members (Figs. 3–5) should be explainable in terms of variations in REE concentrations in marine or pore waters, such as those shown in Figure 6. REE patterns in fossils may not be identical with those of waters from which the REE were obtained. They may differ from water compositions due to crystal-chemical effects (preferential incorporation of MREE, Reynard et al., 1999), differences in aqueous REE speciation, and relative efficiencies of adsorption of various REEs. However, for marine waters these factors should be relatively constant because pH and compositional variations in sea water and sea bottom waters are relatively small. Therefore, relative changes in fossil REE signatures should reflect similar relative changes in the compositions of natural waters from which they were obtained (see also Armstrong et al., 2001). If the average sedimentation rate in the Pierre Shale is approximately 1.1 cm/1000 years and the period of REE uptake less than ca. 3,000 to 10,000 years (Patrick et al., 2001), then the fossil would not be deeply buried and REE signatures probably most closely reflect composition of ocean bottom waters. REE signatures in fossils are different from those in the surrounding sediments, which have essentially flat patterns. This tends to support incorporation of REE largely from the sea bottom water. Changes in REE signatures in fossils from these members of the Pierre Shale might be explained either by (1) mixing between LREE/MREE enriched estuarine/river water (upper left in Fig. 6) and HREE enriched oxic open marine water (lower right in Fig. 6) end-members, or (2) by mixing between LREE/MREE-enriched anoxic water (also toward the upper left in Figure 6) and HREE-enriched oxic open marine water. There is no lithological evidence that the Pierre Shale in central South Dakota was deposited in an estuarine environment. As indicated above, the Pierre Shale consists of laterally extensive organic-rich, largely unbioturbated, 174 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) black and grey shales, silts, and bentonites. This suggests deposition in relatively anoxic conditions, with low deposition rates and relatively remote from estuaries. Therefore, the main trend observed in Figure 5 appears to be most consistent with incorporation of REE from mixtures between oxygenated open sea water and anoxic or suboxic marine water masses. For example, REE signatures in fossils from the lower Sharon Springs Member, which are relatively enriched in LREE and MREE (Fig. 5), and have low Ce anomalies (Fig. 3), are more similar to signatures in anoxic or seasonally anoxic marine waters such as the Cariaco Trench (DeBaar et al., 1988) or the Eastern Mediterranean (Bau et al., 1997). This also seems consistent with the nature of the black, pyrite-rich shale of the lower Sharon Springs. In contrast, REE signatures in fossils from the Gregory Member are highly HREE (YbN) enriched. Such signatures are more similar to those of open seawater. REE signatures in the overlying Crow Creek, DeGrey, and Verendrye Members become progressively more MREE and LREE enriched (Figs. 2 and 3), indicating progressively greater mixing with anoxic waters. Such changes in mixing between oxic and anoxic or suboxic marine waters in one locality might occur in three ways: (1) changes in water depth, (2) changes in organic productivity, and (3) changes in upwelling or intensity of marine currents bringing anoxic waters toward the surface. Although changes in marine currents, storms, or organic productivity cannot be ruled out, many researchers have proposed eustatic or relative changes in sea level during this period (Haq et al., 1987; Stoffer et al., 1998; Hanczaryk et al., 1999). If the data are interpreted primarily in terms of changes in relative water depth, then the lower Sharon Springs fossils were deposited in relatively deep water. REE patterns in the middle and upper Sharon Springs suggest shallowing and fossils in the overlying Gregory Member may have been deposited in relatively shallow water. Overlying members are then deposited in progressively deeper marine environments, but not as deep as the Sharon Springs. These results are generally consistent with paleoenvironmental interpretations by Hanczaryk (2002) and sea level variations proposed by Haq et al. (1987). Hanczaryk (2002) interpreted biofacies in members of the Pierre Shale as indicating shallow depths, perhaps 100 meters, for the Gregory, depths less than 100 meters in the Crow Creek, 100+ meters for the lower DeGrey, 200 meters for the upper DeGrey and 200 meters for the DeGrey-Verendrye boundary. Hanczaryk (2002) did not investigate the Sharon Springs Member; however, the Sharon Springs appears to have been deposited in deeper waters. Haq et al. (1987) proposed a general long-term eustatic shallowing between 81 Ma (Sharon Springs) and 71 Ma (Verendrye), with a short-term, abrupt shallowing about 75 Ma, generally consistent with HREE enriched Gregory and Crow Creek fossils. A short-term period of shallower waters around 80 Ma is not recorded in these fossils or in the lower Sharon Springs lithology, but might occur below the lowest Sharon Springs section preserved in central South Dakota. Although the REE signatures of mosasaur bones appear to correlate generally with members of the Pierre Shale, not enough samples have been obtained to demonstrate how closely the members and REE signatures correspond. More extensive sampling and better stratigraphic control will probably Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 175 show that REE signatures do vary within members, much as they do within the Sharon Springs.

COMPARISON OF REE IN THE VERENDRYE MEMBER WITH THE NAVESINK AND HORNERSTOWN FORMATIONS, NEW JERSEY

REE signatures in mosasaur fossils from the Pierre Shale may be compared (Fig. 7) with those from the Navesink and Hornerstown formations (Staron et al., 2001). The Navesink and Hornerstown formations consist of slowly deposited, olive-green to gray, pelletized glauconitic sand. These formations are thought to represent mid-shelf marine deposits. The glauconitic sediments were probably deposited under relatively anoxic conditions. The Navesink For- mation is Late Cretaceous (Maastrichtian) in age. The overlying Hornerstown Formation may span the Cretaceous-Tertiary boundary (Staron et al., 2001); its Main Fossiliferous Layer (MFL) is correlated with the K-T boundary and yields both vertebrate and invertebrate fossils. Because of some reworking of fossils between the Navesink and overlying Hornerstown Formation and MFL, only bones having cortical material have been used for comparison. These bones are less likely to have been reworked. These bones also have REE signatures that are characteristic of their formations and consistent with in-situ fossil deposition. The Figure 7. Comparison of YbN,PrN, and GdN in REE Verendrye and signatures in mosasaur fossils from the Verendrye Navesink/Hornerstown are Member of the Pierre Shale with the Navesink and not the same age. The Hornerstown Formations (upper Cretaceous Verendrye Member is Cam- [Maastrichtian] and lowest Paleocene [Danian]), at panian whereas the New Inversand, New Jersey. The Hornerstown Forma- tion contains a bone bed, the Main Fossiliferous Jersey formations are Maas- Layer (MFL), which is also shown in the Figure. trichtian to Danian. These Relative REE enrichments are similar in the data are plotted only for a Verendrye, Navesink, and Hornerstown Forma- general comparison of pa- tions. This suggests similar degrees of mixing of leoenvironments. Data near surface and anoxic bottom waters and may indicate similar depths of formation for these for- from the two sites are plot- mations. ted in a triangular diagram (Fig. 7). The degree of MREE and LREE enrichment is similar for the two 176 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) units. This suggests that, if the mixing end-members are similar, the Navesink and Hornerstown formations may have been deposited similar conditions of surface water/anoxic bottom water mixing.

CONCLUSIONS

Rare Earth Element signatures are essentially identical in bones of fossil mosasaurs from individual members or units of the Pierre Shale. However, REE signatures in fossils from different members are different and can be distinguished by statistical methods, such as Discriminant Analysis. Such unique REE signatures may enable paleontologists to determine the provenience of fossils of unknown or questioned origin, or fossils for which there are insufﬁcient cu- ratorial data. Ternary diagrams show that REE data for members of the Pierre Shale lie along a two-component mixing line. This line is interpreted as most consistent with mixing of oxygenated marine waters having HREE enrichment with anoxic or suboxic marine water having MREE and moderate LREE enrichment. If the REE trends are interpreted as being related to ocean depth, then the lower part of the Sharon Springs Member, the lowest member of the Pierre Shale, was deposited in relatively deep anoxic sea bottom waters. REE signatures in the lower, middle, and upper Sharon Springs members are different. The upper Sharon Springs member is more HREE enriched, possibly consistent with decreased depths and regression. The overlying Gregory Member was apparently deposited in shallow, possibly coastal waters. Overlying members (Crow Creek, DeGrey, and Verendrye) were deposited in progressively deeper water. These results are generally consistent with previous paleoenvironmental conclusions based on analysis of biofacies (Hanczaryk, 2002) and sea level curves of Haq et al. (1987). REE signatures from the Verendrye Member of the Pierre Shale were compared with signatures from the Maastrichtian Navesink and Maastrichtian-Dani- an Hornerstown formations. Based on similarities in LREE enrichments and assuming that the mixing end-members have similar REE signatures, the two sites were deposited under similar degrees of mixing of oxic and anoxic marine waters.

ACKNOWLEDGMENTS

The authors wish to thank the U.S. Army Corps of Engineers for their on- going support of ﬁeld, laboratory, and research investigation at the South Dakota School of Mines and Technology. We also thank the Crow Creek Sioux Tribe for permission to collect on their land. Paul Field and Louise Bolge aided ICP-MS analyses. Temple University provided partial support for this research with a grant from the Research Incentive Fund. This study represents part of a M.S. thesis by the ﬁrst author. Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 177

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Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 181

3- PHOSPHONOSULFONATE(3-) O3PSO3 AND RELATED IONS

Arlen Viste and Kari Lunder Stone Department of Chemistry Augustana College Sioux Falls, SD 57197

ABSTRACT

2- 4- The ions dithionate S2O6 and hypophosphate P2O6 are well known. How- 3- ever the isoelectronic phosphonosulfonate(3-) ion O3PSO3 is not. Neither is the - isoelectronic O3SClO3 ion. Previous experimental study of the oxidation of thio- 3- 3- phosphate ion O3PS suggested the formation of O3PSO3 but the conclusion remains tentative. (Viste 1991) In the present study, HF/6-31G and HF/6- 31G(d) molecular orbital calculations were carried out for each of the four ions, both in the gas phase and solvated in aqueous solution. With Jaguar 4, geometries were optimized for both gas phase and solvated structures for the four ions. In the gas phase in HF/6-31G, Jaguar 4 and Gaussian (94 or 98W) show 4- 2- - 3- stable structures for P2O6 , S2O6 , and O3SClO3 , while gas phase O3PSO3 is cal- - 2- culated to dissociate to planar PO3 and pyramidal SO3 . (Schrödinger 2000, 3- Gaussian 1999) However solvated (aqueous) O3PSO3 is calculated by Jaguar 4 to be stable in HF/6-31G, though with a slightly long S-P bond length of 258 2- pm, compared with 230 pm for S-S in solvated S2O6 and 219 pm for P-P in sol- 4- 4- vated P2O6 . In HF/6-31G solvation also shortens the central X-Y bond in P2O6 2- - and S2O6 but lengthens it in O3SClO3 . Next in HF/6-31G(d) and DFT B3LYP/6- 31G(d), all four ions are calculated to be stable both in the gas phase and in solution. Vibrational frequencies have been calculated in HF/6-31G(d) and DFT B3LYP/6-31G(d) for the normal modes of vibration of the four ions, both gas phase and solvated. Gas phase infrared and Raman intensities have been calculated in HF/6-31G(d). Thus calculations indicate that the phosphonosul- 3- fonate (3-) ion O3PSO3 is stabilized by solvation in HF/6-31G, while all four ions are stable in HF/6-31G(d) and DFT B3LYP/6-31G(d), both gas phase and solvated. Further investigation of protonated species is planned.

Keywords

Phosphonosulfonate(3-), vibrations, Jaguar, Gaussian

INTRODUCTION

3- 3- phosphate ion O3PS suggested the formation of O3PSO3 but the conclusion remains tentative. (Viste 1991) In the present study, HF/6-31G and HF/6- 31G(d) molecular orbital calculations were carried out for each of the four ions, both in the gas phase and solvated in aqueous solution. Structure and vibrations have been investigated through ab initio and DFT molecular orbital calculations, using Jaguar 4 and Gaussian 94 computational chemistry software

COMPUTATIONAL METHODS

Jaguar 4.0 software was used under SuSE 6.4 Linux on a PC. Gaussian 94 software was run under Aix 4.3 on an IBM RS/6000, through a WebMO interface. (Schrödinger 2000, Gaussian 2001, WebMO 2001).

RESULTS AND DISCUSSION

Preliminary calculations carried out at the 6-31G level are summarized in 3- Table 1. Note that in the gas phase at this level, the S-P bond of SPO6 breaks, - but is stabilized in aqueous solution, while SClO6 has a longer calculated bond length in solution.

Table 1. 6-31G calculations.

4- 3- 2- - Geometry P2O6 SPO6 S2O6 SClO6

6-31G gas Failed

X-Y, Å 2.525 18.197 2.359 2.871

X-O, Å 1.650 1.576 1.636 1.582

Y-O, Å 1.685 1.948

X-Y-O, deg 108.6 110.89 105.11 105.25

O-X-Y, deg 90.39 96.76

6-31G solvated (aq)

X-Y, Å 2.194 2.578 2.297 3.973

X-O, Å 1.610 1.571 1.622 1.569

Y-O, Å 1.641 1.851 X-Y-O, deg 107 112 104 113 O-X-Y, deg 96 89

** X=lower, Y=higher atomic number

Next, calculations were carried out at the 6-31G* level, also known as 6- 31G(d). The results are presented in Table 2. Note that the addition of d or- 3- bitals in the calculation has stabilized SPO6 in the gas phase. In solution, cal- 4- 3- 2- culated X-Y bond lengths are slightly shorter for P2O6 , SPO6 , and S2O6 , but - longer for SClO6 . Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 183

Table 2. 6-31G* Calculations in Gaussian 94, Jaguar 4.

4- 3- 2- - Geometry P2O6 SPO6 S2O6 SClO6

6-31G* gas

X-Y, Å 2.315 2.257 2.115 2.848

X-O, Å ** 1.550 1.509 1.455 1.408

Y-O, Å 1.490 1.473

X-Y-O, deg ** 108.4 109.5 105.3 109.3

O-X-Y, deg 103.1 93.8

6-31G* solvated (aq)

X-Y, Å 2.128 2.120 2.075 3.52

X-O, Å 1.521 1.498 1.449 1.404

Y-O, Å 1.469 1.481 X-Y-O, deg 107 108 105 111 O-X-Y, deg 103 89

** X=lower, Y=higher atomic number

Next, DFT (Density Functional Theory) calculations were carried out, at the 6-31G*/B3LYP level. Results are presented in Table 3.

Table 3. 6-31G*/B3LYP (DFT) calculations.

4- 3- 2- - Geometry P2O6 SPO6 S2O6 SClO6

6-31G*/B3LYP gas

X-Y, Å 2.508 2.432 2.255 2.557

X-O, Å 1.582 1.540 1.496 1.47

Y-O, Å 1.533 1.504

X-Y-O, deg 108 109.9 105.4 107

O-X-Y, deg 102.7 108

6-31G*/B3LYP solvated (aq)

X-Y, Å 2.141 2.161 2.165 2.840

X-O, Å 1.545 1.524 1.486 1.453

Y-O, Å 1.504 1.520 X-Y-O, deg 107 108 105 110 O-X-Y, deg 103 92

** X=lower, Y=higher atomic number

4- For comparison, a literature crystal structure containing P2O6 provides these experimental bond length, in Na4P2O6.10H2O. (Emmerson 1973) P-P 2.201 Å, P-O 1.530-1.538 Å, P-P-O 101.5-113.1 deg. The 6-31G* solvated (aq) and the 6-31G*/B3LYP solvated (aq) calculated results are in quite good agreement with these experimental bond lengths and bond angles. 184 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002)

Vibrational calculations were made both for gas phase and solvated species. Table 4 presents results for selected vibrations, calculated in 6-31G*, gas and solvated(aq).

Table 4. Selected Vibrations in 6-31G*, Gaussian 94, Jaguar 4

4- 3- 2- - P2O6 SPO6 S2O6 SClO6

Vibrations (cm-1) A1g, A2u (D3d) and A1 (C3v) 6-31G* gas X-Y stretch (A1g, A1) ** 255 246 317 54 X-Y-O, Y-X-O bend (A2u, A1) 625 618 669 524 X-Y-O, Y-X-O bend (A1g, A1) 713 741 806 655 X-O, Y-O stretch (A2u, A1) 925 1008 1071 1010 X-Y, X-O, Y-O stretch (A1g, A1) 1038 1118 1189 1198

6-31G* solvated (aq) X-Y stretch (A1g, A1) 355 319 366 58 X-Y-O, Y-X-O bend (A2u, A1) 638 549 711 643 X-Y-O, Y-X-O bend (A1g, A1) 835 783 898 775 X-O, Y-O stretch (A2u, A1) 1066 1055 1161 1135 X-Y, X-O, Y-O stretch (A1g, A1) 1201 1186 1292 1274

** X=lower, Y=higher atomic number

Of these selected normal modes of vibration in D3d symmetry, A1g is Ra- man active and A2u is infrared active. In C3v symmetry, A1 is symmetry al- 3- lowed in both Raman and infrared, but calculated intensity patterns in SPO6 4- 2- are still fairly similar to those of P2O6 and S2O6 .

Vibrational calculations were also made in DFT for both gas phase and solvated species. Table 5 presents results for selected vibrations, calculated in 6- 31G*/B3LYP, gas and solvated(aq). Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 185

Table 5. Selected Vibrations in 6-31G*, Gaussian 94, Jaguar 4

4- 3- 2- - P2O6 SPO6 S2O6 SClO6

Vibrations (cm-1) A1g, A2u (D3d) and A1 (C3v) 6-31G*/B3LYP gas X-Y stretch (A1g, A1) 184 138 272 175 X-Y-O, Y-X-O bend (A2u, A1) 673 525 757 730 X-Y-O, Y-X-O bend (A1g, A1) 721 619 830 782 X-O, Y-O stretch (A2u, A1) 928 903 1075 1062 X-Y, X-O, Y-O stretch (A1g, A1) 1005 966 1154 1102

6-31G*/B3LYP solvated (aq) X-Y stretch (A1g, A1) 306 276 281 120 skew X-Y-O, Y-X-O bend (A2u, A1) 548 500 584 527 X-Y-O, Y-X-O bend (A1g, A1) 723 677 728 647 X-O, Y-O stretch (A2u, A1) 949 956 993 972 X-Y, X-O, Y-O stretch (A1g, A1) 1064 1057 1093 1036

For comparison, experimental vibrational frequencies are summarized in Table 6. (Palmer 1961) The comparison between calculated and observed vibrational frequencies is quite good, particularly for 6-31G*/B3LYP solvated (aq).

Table 6. Comparison with Experiment (Palmer 1961) for Selected Vibrations

4- 3- 2- - P2O6 SPO6 S2O6 SClO6

Vibrations (cm-1) A1g, A2u (D3d) and A1 (C3v) X-X stretch (A1g) 275 293 X-X-O bend (A2u) 562 577 X-X-O bend (A1g) 670 710 X-O stretch (A2u) 942 1000 X-Y, X-O stretch (A1g) 1062 1102

3- Figure 1 shows the electron density with electrostatic potential for SPO6 , calculated in Gaussian 94 6-31G*, as represented by Molekel. (Gaussian 1999, Molekel 2001)

3- Fig. 1. SPO6 electron density with electrostatic potential. 186 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002)

CONCLUSION

3- Calculations indicate that the phosphonosulfonate (3-) ion O3PSO3 is sta- 4- 3- 2- bilized by solvation in HF/6-31G, while all four ions (P2O6 , SPO6 , S2O6 , and - SClO6 ) are stable in HF/6-31G(d) and DFT B3LYP/6-31G(d), both gas phase and solvated. The comparison between calculated frequencies and experimental vibrational frequencies observed by Palmer is quite good, particularly for 6- 31G*/B3LYP solvated (aq). (Palmer 1961) Further investigation of protonated species is planned.

Notes

1. Kari Lunder participated in the beginning of this work as an undergraduate at Augustana College (SD). Kari Lunder Stone is currently a graduate student at Pennsylvania State University. 2. This work was presented in a poster session at the Finnish Symposium on Quantum Chemistry, An International Conference, Kuusamo, Finland, June 11-17, 2001. http://www.chem.helsinki.ﬁ/Research/Kuusamo/ http://inst.augie.edu/~viste/Kuusamo/

LITERATURE CITED

Anderson, Wayne. 2001. Jagconvert1.1.tcl. Bloomsburg University, PA. http://facstaff.bloomu.edu/wpa/ Emmerson, D. S. and D. E. C. Corbridge. 1973. “The crystal structure of tetra-

sodium hypophosphate decahydrate Na4P2O6.10H2O.” Phosphorus. 3: 131. Gaussian, Inc. 2001. Gaussian 94 and 98 software. http://www.gaussian.com/ Laaksonen, Leif. 2000. gOpenMol software. http://www.csc.ﬁ/~laaksone/gopenmol/gopenmol.html Molekel. 2001. Advanced Interactive 3D-Graphics for Molecular Sciences. http://www.cscs.ch/molekel/molekel.html Palmer, W. G. 1961. The vibrational spectra and structures of dithionate, hypophosphate, and related ions. J. Chem. Soc. 1552-1562. Schrödinger, Inc. 2000. Jaguar 4 software. http://www.schrodinger.com/ Viste, Arlen and Jeanne Pfaff. 1991. Oxidation of Thiophosphate. Proc. S.D. Acad. Sci.. 70: 262-263. Abstract. WebMO. 2000. Hope College. Holland, MI. http://www.chem.hope.edu/webmo/ WebMO. 2001. Department of Chemistry, Augustana College, Sioux Falls, SD. http://inst.augie.edu/~spartan/ Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 187

ACKNOWLEDGMENTS

We thank Hope College for their public-spirited development and provi- sion of the WebMO software. Thanks also to Augustana College and its De- partment of Chemistry for software and hardware support.

Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 189

SETS OF HALF-REACTIONS WHICH YIELD DIFFERENT Emf FOR AN OVERALL REACTION, BUT EQUIVALENT STANDARD GIBBS FREE ENERGY AND EQUILIBRIUM CONSTANTS

April Kay Bobbie and Arlen Viste Department of Chemistry Augustana College Sioux Falls, SD 57197

ABSTRACT

The purpose of this project was to investigate sets of half-reactions that produce the same cell reaction, different cell emf values (Eºcell), but similar val- ∆ ues of nEºcell, standard Gibbs free energy ( Gº) and equilibrium constants (Keq). For several ions, sets of half-reactions were determined and cell emf values, nEºcell, equilibrium constants, and standard Gibbs free energy were calculated. For example, the realization that the following two sets of mercury half-reactions give different Eºcell values, but (within roundoff error) equivalent values ∆ for nEºcell, Gº, and Keq, prompted the current investigation:

2+ - 2 Hg = Hg2 + 2 e Eº = -0.7960 V 2+ - 2+ 2 Hg + 2 e = Hg2 Eº = +0.9110 V 2+ 2+ = 2 Hg + 2 Hg = 2 Hg2 Eºcell +0.115 V 2+ 2+ Hg + Hg = Hg2 (n=1) n Eºcell = +0.115 V

∆ -1 Gº = -11.1 kJ mol Keq = 87.7

2+ - 2 Hg = Hg2 + 2 e Eº = -0.7960 V Hg2+ + 2 e- = Hg Eº = +0.8535 V 2+ 2+ Hg + Hg = Hg2 Eºcell = +0.0575 V

(n=2) n Eºcell = +0.115 V

∆ Gº = -11.1 kJ mol-1 Keq = 87.7

The standard Gibbs free energy and equilibrium constant calculated were found to be equivalent for the two sets of half-reactions. The same cell reaction is produced by both sets of half-reactions. Each set differs in the number of electrons transferred and the cell emf value, but agrees in their product, nEºcell. (Atkins 1998, Bard 1985, Latimer 1952) 190 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002)

Keywords

Emf, cell potential, half-reaction, Gibbs free energy, equilibrium constant

INTRODUCTION

Standard electrode potentials (redox potentials, oxidation-reduction potentials) provide information about the behavior of chemical reactions. This project investigated sets of half-reactions that produce the same cell reaction, different cell emf values (Eºcell), but similar values of nEºcell, standard Gibbs free ∆ energy ( Gº) and equilibrium constants (Keq). For mercury, chromium, iron, sulfur, chlorine, copper, and indium, sets of half-reactions were determined and cell emf values, nEºcell, equilibrium constants, and standard Gibbs free energy were calculated. Eº values for half-reactions are tabulated in several sources. (Atkins 1998, Bard 1985, Latimer 1952) Where the NBS Tables in- ∆ ∆ clude Gºf for the species in a half-reaction, Eº = - Gº/nF, and this was used as the preferred value. (Wagman 1982) Where the NBS Tables were incomplete, the source of the Eº used is referenced.

METHODS OF CALCULATION

For all calculations: F = 96485.3 C/mol R = 8.31451 J/mol*K T = 298.15 K ∆Gº = -nFEº nFEº/RT Keq = e

RESULTS

Mercury: (Wagman 1982)

2+ - 2 Hg = Hg2 + 2 e Eº = -0.7960 V 2+ - 2+ 2 Hg + 2 e = Hg2 Eº = +0.9110 V 2+ 2+ 2 Hg + 2 Hg = 2 Hg2 Eºcell = +0.115 V 2+ 2+ Hg + Hg = Hg2 (n=1) n Eºcell = +0.115 V

∆ -1 Gº = -11.1 kJ mol Keq = 87.7

2+ - 2 Hg = Hg2 + 2 e Eº = -0.7960 V Hg2+ + 2 e- = Hg Eº = +0.8535 V 2+ 2+ Hg + Hg = Hg2 Eºcell = +0.0575 V

(n=2) n Eºcell = +0.115 V

∆ -1 Gº = -11.1 kJ mol Keq = 87.7 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 191

Chromium: (Atkins 1998)

3 (Cr2+ + 2 e- = Cr) Eº = -0.91 V 2 (Cr = Cr3+ + 3 e-) Eº = +0.74 V 2+ 3+ 3 Cr = 2 Cr + Cr Eºcell = -0.17 V

(n=6) n Eºcell = -1.02 V

∆ -1 ≈ -1 -18 Gº = 98.4 kJ mol 100 kJ mol Keq = 5.74x10

Cr3+ + 3 e- = Cr Eº = -0.74 V 3 (Cr2+ = Cr3+ + e-) Eº = +0.41 V 2+ 3+ 3 Cr = 2 Cr + Cr (n=3) Eºcell = -0.33 V

n Eºcell = -0.99 V

∆ -1 ≈ -1 -17 Gº = 95.5 kJ mol 100 kJ mol Keq = 1.84x10

Iron: (Wagman 1982)

3 (Fe2+ + 2 e- = Fe) Eº = -0.409 V 2 (Fe = Fe3+ + 3 e-) Eº = +0.049 V 2+ 3+ 3 Fe = 2Fe + Fe Eºcell = -0.360 V

(n=6) n Eºcell = -2.16 V

∆ -1 Gº = 208 kJ mol Keq = 3.23x10-37

Fe3+ + 3 e- = Fe Eº = +0.049 V 3 (Fe2+ = Fe3+ + e-) Eº = -0.769 V 2+ 3+ 3 Fe = 2Fe + Fe (n=3) Eºcell = -0.72 V

n Eºcell = -2.16 V

∆ -1 -37 Gº = 208 kJ mol Keq = 3.23x10 192 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002)

Sulfur: (Wagman 1982)

2- - + - 2 (S2 + 2 e + 2 H = 2 HS ) Eº = +0.287 V - + - 2- 3 HS = 3 H + 4 e + S3 Eº = -0.097 V 2- + - + 2- 2 S2 + 4 H = HS + 3 H + S3 Eºcell = +0.190 V 2- + - 2- 2 S2 + H = HS + S3 (n=4) n Eºcell = +0.760 V

∆ -1 12 Gº = -73.3 kJ mol Keq = 6.91x10

2- - + - S2 + 2 e + 2 H = 2 HS Eº = +0.287 V 2- 2- 3 S2 = 2 e- + 2 S3 Eº = +0.473 V 2- + - 2- 4 S2 + 2 H = 2 HS + 2 S3 Eºcell = +0.760 V 2- + - 2- 2 S2 + H = HS + S3 (n=1) n Eºcell = +0.760 V

∆ -1 12 Gº = -73.3 kJ mol Keq = 6.91x10

Chlorine: (Wagman 1982)

- - - - ClO3 + H2O + 2 e = ClO2 + 2 OH Eº = +0.295 V - - 2 (ClO2 = ClO2(g) + e ) Eº = -1.071 V - - - ClO3 + ClO2 + H2O = 2 ClO2(g) + 2 OH Eºcell = -0.776 V

(n=2) n Eºcell = -1.552 V

∆ -1 -27 Gº = 150 kJ mol Keq = 6.03x10

- - - ClO3 + H2O + e = ClO2(g) + 2 OH Eº = -0.481 V - - ClO2 = ClO2(g) + e Eº = -1.071 V - - - ClO3 + ClO2 + H2O = 2 ClO2(g) + 2 OH Eºcell = -1.552 V

(n=1) n Eºcell = -1.552 V

∆ -1 -27 Gº = 150 kJ mol Keq = 6.03x10 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 193

Examples where the number of electrons is equal

Copper: (Wagman 1982)

2 (Cu+ + e- = Cu) Eº = +0.520 V Cu = Cu2+ + 2 e- Eº = -0.340 V + 2+ 2 Cu = Cu + Cu Eºcell = +0.180 V

Cu2+ + 2 e- = Cu Eº = +0.340 V 2 (Cu+ = Cu2+ + e-) Eº = -0.160 V + 2+ 2 Cu = Cu + Cu Eºcell = +0.180 V

n=2 n Eºcell = +0.360 V

∆ -1 6 Gº = -34.7 kJ mol Keq = 1.22x10

Indium: (Wagman 1982)

2 (In2+ + e- = In+) Eº = -0.400 V In+ = In3+ + 2 e- Eº = +0.445 V 2+ 3+ + 2 In = In + In Eºcell = +0.045 V

In3+ + 2 e- = In+ Eº = -0.445 V 2 (In2+ = In3+ + e-) Eº = +0.490 V 2+ 3+ + 2 In = In + In Eºcell = +0.045 V

n=2 n Eºcell = +0.090 V

∆ -1 Gº = -8.68 kJ mol Keq = 33.1

DISCUSSION

For mercury, chromium, iron, sulfur, chlorine, copper, and indium, the cal- ∆ culated Gº and Keq are equal for both sets of half-reactions. The cell reaction was the same for each set of half-reactions. For all of the ions, except for copper and indium, the number of electrons transferred in each set of half-reactions was different. Regardless of this difference, the standard Gibbs free energy, equilibrium constant, and nEºcell were equivalent. Use of Eº data from sources other than the NBS Tables tended to lead to modest round-off error. Note added in proof: a related discussion is provided by Cooke and Willis (Cooke 1996). 194 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002)

CONCLUSION

For each set of the two sets of half-reactions, the standard Gibbs free energy and equilibrium constant calculated were found to be equivalent. The same cell reaction is produced by both sets of half-reactions. Each set differs in the number of electrons transferred and the cell emf value, but agrees in their product, nEºcell. Using Eº values based on the NBS Tables tended to give the most consistent results.

LITERATURE CITED

Atkins, Peter W. 1998. Physical Chemistry, 6th Ed. Freeman, New York. Bard, Allen J., Roger Parsons, and Joseph Jordan. 1985. Standard Potentials in Aqueous Solution. Marcel Dekker, New York. Cooke, Ron C., and Grover C. Willies, 1996. Which Eº is it? Chemical principles revisited. J. Chem. Ed. 73:450. Latimer, Wendell M. 1952. Oxidation Potentials, 2nd Ed. Prentice-Hall, Engle- wood Cliffs, N.J. Wagman, Donald D., et al. 1982. The NBS Tables of Chemical Thermodynam- ic Properties. Journal of Physical and Chemical Reference Data. 11: Sup- plement 2.

ACKNOWLEDGMENT

Useful conversations with Jennifer L. Engelbrecht (Augustana College) are gratefully acknowledged. Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 195

APPLICATION OF ADF AMSTERDAM DENSITY FUNCTIONAL PACKAGE TO RELATIVISTIC EFFECTS 2+ IN MERCUROUS ION Hg2 AND RELATED SPECIES

William J. Delfs and Arlen Viste Department of Chemistry Augustana College Sioux Falls, SD 57197

ABSTRACT

A useful review of relativistic effects in chemistry is provided by Pyykkö. (Pyykkö 1988) For a number of years, at the undergraduate level at Augustana College this topic has been treated brieﬂy in Advanced Inorganic Chemistry, and sometimes in Quantum Chemistry. The work reported here was undertaken for undergraduate instructional purposes, to compare relativistic and nonrelativistic calculations on some relatively simple systems, involving gold and mercury, where relativistic effects are known to often be signiﬁcant. The quantum chemistry used for this purpose was ADF, the Amsterdam Density Func- tional package. (Te Velde 2001, SCM 2001) The software was installed on a PC under Mandrake 8.0 Linux. (Mandrake 2001) Examples included with the ADF package include AuH and Au2, treated relativistically. The present work con-

ﬁrmed and compared those two with nonrelativistic treatments of AuH and Au2. + +2 Relativistic and nonrelativistic calculations for HgH and Hg2 were carried out as well. Results were compared with Huber and Herzberg (Huber 1979) for the +2 ﬁrst three, and with X-ray crystallographic results for compounds of Hg2 . (Wyckoff 1982) Satisfactory agreement was obtained between relativistic ADF calculations of bond lengths and experiment. The results illustrate the concept of relativistic contraction, since the calculated bond lengths were shorter for +2 relativistic than for nonrelativistic calculations. (Pyykkö 1988) For Hg2 this work found nonrelativistic 2.94 Å, relativistic 2.69 Å. For comparison, crystallographic literature gives 2.69 Å in Hg2I2 or 2.58 in Hg2Br2. (Wyckoff 1982) ADF calculations were carried out for AuHg+. ADF also calculated frequencies, IR intensities, and Raman intensities for each of these species.

Keywords

ADF, relativistic quantum chemistry, mercurous ion

INTRODUCTION

A useful review of relativistic effects in chemistry is provided by Pyykkö. (Pyykkö 1988) For a number of years, at the undergraduate level at Augustana 196 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002)

College this topic has been treated briefly in Advanced Inorganic Chemistry, and sometimes in Quantum Chemistry. The work reported here was undertaken was undertaken for undergraduate instructional purposes, to compare relativistic and nonrelativistic calculations on some relatively simple systems, involving gold and mercury, where relativistic effects are known to often be significant. Au and Hg+ have the electron configuration [Xe] 4f14 5d10 5s1. Single 2+ + + bonded species exist such as Au2 , Hg2 , AuH, HgH , and perhaps AuHg .

COMPUTATIONAL METHODS

The quantum chemistry used for this purpose was ADF, the Amsterdam Density Functional program. (SCM 2001) The software was installed on a PC under Mandrake 8.0 Linux. (Mandrake 2001) Examples included with the ADF package include AuH and Au2, treated relativistically.

RESULTS AND DISCUSSION

Examples included with the ADF package include AuH and Au2, treated relativistically. The present work conﬁrmed and compared those two with nonrelativistic treatments of AuH and Au2, with results shown in Table 1.

Table 1. Bond Length Results for Au2 and AuH

ADF Relativistic ADF Nonrelativistic Experiment (Huber 1979)

Au2 Bond length 2.522 Å (ZORA) 2.708 Å 2.4719 Å

AuH Bond length 1.479 Å (RelGO) 1.702 Å 1.52385 Å

Relativistic and nonrelativistic calculations were carried out for HgH+ as shown in Table 2.

Table 2. Bond Length Results for HgH+

ADF Relativistic ADF Nonrelativistic Experiment (Huber 1979)

+ HgH Bond length 1.583 Å 1.759 Å 1.5944 Å Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 197

Figure 1 shows HgH+ from relativistic ADF, represented in Molekel 4.1 with a Connolly surface. (Molekel 2002)

Figure 1. Relativistic HgH+ with Connolly surface, represented in Molekel

+2 Relativistic and nonrelativistic calculations were carried out for Hg2 as shown in Table 3.

2+ Table 3. Bond Length Results for Hg2

ADF Relativistic ADF Nonrelativistic Experiment (Wyckoff 1982)

2+ Hg2 Bond length 2.707 Å (RelGO) 2.935 Å 2.69 Å in Hg2I2 or

2.58 in Hg2Br2

Thus satisfactory agreement was obtained between relativistic ADF calculations of bond lengths and experiment.

+ Table 4. Stretching Frequency Results for Au2, AuH, and HgH

ADF Relativistic ADF Nonrelativistic Experiment (Huber 1979)

-1 Au2 frequency, cm 175 134 190.9

Au2 IR intensity 0 0 symmetry forbidden

Au2 Raman intensity 148 259 symmetry allowed AuH frequency, cm-1 2260 1720 2305.01 AuH IR intensity 3 28 symmetry allowed AuH Raman intensity 1121 1874 symmetry allowed HgH+ frequency, cm-1 1963 1688 2027.7 HgH+ IR intensity 166 88 symmetry allowed HgH+ Raman intensity 1503 3023 symmetry allowed 198 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002)

2+ Table 5. Hg-Hg Stretching Frequency Results for Hg2

ADF Relativistic ADF Nonrelativistic Experiment (Gager 1966)

2+ -1 Hg2 frequency, cm 113 (RelGO) 97 169 (aqueous) 2+ Hg2 IR intensity 0 0 symmetry forbidden 2+ Hg2 Raman intensity 109 172 symmetry allowed 2+ -1 Hg2 frequency, cm 119 (ZORA) 169 (aqueous) 2+ Hg2 IR intensity 0 symmetry forbidden 2+ Hg2 Raman intensity 104 symmetry allowed

ADF calculations for AuHg+ are presented in Table 6. Although experimental values apparently are not available, relativistic CCSD(T)+g(MP2) calculations of Wesendrup are listed for comparison. (Wesendrup 1999)

Table 6. Some ADF Results for AuHg+

ADF Relativistic ADF Nonrelativistic Lit Rel CCSD(T)

Bond length 2.53 Å 2.78 Å 2.562 Å Vibrational frequency 157 cm-1 116 165 cm-1 IR intensity 0.5 0.5 Raman intensity 69 90

CONCLUSION

The results illustrate the concept of relativistic contraction, since the calculated bond lengths were shorter for relativistic than for nonrelativistic calcula- 2+ tions. (Pyykkö 1988) Theoretical studies of the mercurous ion Hg2 include that of Neisler and Pitzer. (Neisler 1987) In the present study, bond lengths and vibrational frequencies calculated in ADF were in reasonable agreement with ex- 2+ -1 periment. The frequency calculated for Hg2 (113 or 119 cm by two ADF methods in the gas phase) is somewhat lower than the experimental value (169 cm-1) in aqueous solution. This may result in part from solvation effects for this dipositive cation.

ACKNOWLEDGMENTS

Thanks to Augustana College and its Department of Chemistry for software and hardware support.

LITERATURE CITED

Gager, H. M., J. Lewis, and M. J. Ware. 1966. Metal-metal stretching frequencies in Raman spectra. Chem. Commun. 616-617. Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 199

Huber, K. P. and G. Gerzberg. 1979. Molecular Spectra and Molecular Structure. IV. Constants of Diatomic Molecules. Van Nostrand Reinhold Co. p. 50, 56, 318. Mandrake. 2001. Mandrake Linux. http://www.linux-mandrake.com/en/ Molekel. 2002. Advanced Interactive 3D-Graphics for Molecular Sciences. http://www.cscs.ch/molekel/molekel.html 2+ Neisler, R. P. and K. S. Pitzer. 1987. The dipositive dimeric ion Hg2 : A theoretical study. J. Phys. Chem. 91:1084-1087. Pyykkö, Pekka. 1988. Relativistic Effects in Structural Chemistry. Chem. Rev. 88: 563-594. SCM. 2001. ADF Program System. Scientiﬁc Computing & Modelling NV. http://www.scm.com/ Te Velde, G., F. M. Bickelhaupt, E. J. Baerends, C. Fonseca Guerra, S. J. A. Van Gisbergen, J. G. Snijders, and T. Ziegler. 2001. Chemistry with ADF. J. Comp. Chem. 22:931-967. Wesendrup, Ralf, Jon K. Laerdal, and Peter Schwerdtfeger. 1999. Relativistic effects in gold chemistry. VI. Coupled cluster calculations for the isoelec- - + tronic series AuPt , Au2, and AuHg . J. Chem. Phys. 110:9457-9462. Wyckoff, Ralph W. G. 1982. Crystal Structures. Volume 1, second ed. Robert E. Krieger Publishing Co., p. 152.

Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 201

ESTIMATING SUMMER FAWN MORTALITY OF WHITE-TAILED DEER USING CORPORA ALBICANTIA AND LACTATION STATUS

Robert G. Osborn, Beth A. Hippensteel, and Jonathan A. Jenks Department of Wildlife & Fisheries Sciences South Dakota State University Brookings, SD 57007

Ted A. Benzon South Dakota Department of Game, Fish and Parks Rapid City, SD 57702

ABSTRACT

Most fawn mortality occurs during the first 2-3 months of life, but it is difficult to quantify. Estimating fawn mortality from radio telemetry studies is time consuming and expensive. The purpose of this study was to develop a rapid method for estimating summer fawn mortality using corpora albicantia and evidence of lactation (CAL), to identify sources of bias influencing these estimates, and to compare CAL estimates to those determined via radio telemetry. Twenty-five adult female white-tailed deer (Odocoileus virginianus) were collected each August in 1993, 1994, and 1995 from the Black Hills of South Dako- ta. Lactation status (i.e., lactating or non-lactating) was recorded and the number of corpora albicantia in each pair of ovaries were counted. Mortality estimated from CAL data in 1993 (0.1% < 8.7% < 31.7%; 95% CI), 1994 (2.1% < 14.8% < 37.2%), and 1995 (0.8% < 10.3% < 30.6%) were less (P = 0.012) than those calculated from radio telemetry data in 1993 (5.1% < 33.3% < 52.9%), 1994 (0.2% < 25.0% < 60.2), and 1995 (8.8% < 41.2% < 54.6%). The inability of the CAL method to detect the death of offspring when one or more litter mates survive was the bias which most influenced mortality estimates. Esti- mates of summer mortality from CAL data may be a useful index for ungulates which typically produce a single offspring. Better estimates of partial litter loss are necessary before this approach can be extended to species producing more than 1 offspring per litter.

Keywords

Black Hills, corpora albicantia, corpora lutea, fawn mortality, lactation, Odocoileus virginianus, ovary, South Dakota, summer mortality, white-tailed deer. 202 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002)

INTRODUCTION

Mortality is one of the major factors influencing population dynamics. Consequently, a great deal of time, effort and resources are expended to identify causes of mortality. Additionally, mortality rates vary with age. For ungulates, mortality is highest during the first year of life, with much of that mortality occurring within the first month of life (Cook et al. 1971, Huegel et al. 1985, Nelson and Woolf 1987). This source of mortality, however, is particularly difficult to quantify because the relative speed with which small bodies are scavenged and/or decompose (Cook et al. 1971). Most fawn mortality studies rely on data collected from radio-collared fawns. Although reliable, radio telemetry studies are time consuming and expensive. A physiological approach to determining fawn mortality is possible but has received less attention. Robinette et al. (1955) used corpora lutea and udder development to assess fawn mortality among mule deer (Odocoileus hemionus). Mule deer that possessed corpora lutea but lacked udder development were considered to have suffered prenatal fawn mortality, or at least to have lost their fawn(s) prior to any appreciable udder development (Robi- nette et al. 1955). While this method of evaluating prenatal mortality works well for first time breeders, it is less useful after initial development occurs and the udder loses its virginal appearance. Harder and Kirkpatrick (1994) suggested that lactation indices combined with measures of reproductive rate may improve estimates of net natality. Navarre (1993) used reproductive rate data from ovaries combined with lactation status data collected from hunter killed animals to estimate postnatal mortality of roe deer (Capreolus capreolus) in France. In the United States, hunting seasons of deer generally extend into Novem- ber and December. By this time many females that produced fawns may have already stopped lactating (Scanlon and Urbston 1978, Woolf and Harder 1979). Furthermore, waiting till this time of year increases the likelihood that developing ovules will compress and obscure corpora albicantia from the past breeding season, potentially leading to an underestimation of productivity (Golley 1957, Teer et al. 1965). Consequently, it is unlikely that using hunter killed animals would provide an accurate estimate of juvenile mortality of deer in the United States. The purpose of this study was to refine and evaluate a physiological method of estimating fawn mortality during the first 3 months of life and to compare these results with fawn mortality data from a radio telemetry study, which was conducted concurrently. We also identified sources of bias that may influence this procedure and examined the extent to which they may affect accuracy. If practical, a rapid, physiological approach to quantifying summer mortality rate might prove useful for identifying areas with differing mortality. Radio telemetry studies could then be used to gather more detailed information. Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 203

STUDY AREA

The study was conducted in the northern and central Black Hills, South Dakota. The northern Black Hills (NBH) study area ranged in latitude and longitude from 441 12´ N to 441 25´ N and 1031 24´ W to 1041 04´ W, respectively. The central Black Hills (CBH) study area ranged in latitude and longitude from 431 52´ N to 441 15´ N and 1031 22´ W to 1041 07´ W, respectively. Osborn (1994) and Hippensteel (2000) described the northern and central regions of the Black Hills.

METHODS

Seventy-five adult female white-tailed deer (O. virginianus) were collected from summer ranges in the Black Hills. Deer collections began mid-August and were completed within 10 days in 1993, 1994, and 1995. Twenty-five deer were collected each year. Deer collections were approved by the South Dako- ta Department of Game, Fish and Parks, met the guidelines of the Institution- al Animal Care and Use Committee at South Dakota State University, and followed the animal research guidelines established by The American Society of Mammalogists (Ad Hoc Committee for Animal Care and Use 1998). During necropsy, ovaries were removed and stored in 10% formalin until they could be examined. Lactation status (i.e., lactating or non-lactating) was determined by expressing milk from teats. In the lab, ovaries were sectioned and degenerating corpora lutea (i.e., corpora albicanta [CA]) were counted using the macroscopic method (Cheatum 1949). Based on CA and lactation status data, deer were placed in 1 of 3 categories: class 1), no CA and non-lactating (i.e., non-breeders); class 2), 1 or more CA and lactating (i.e., successfully bred and fawn(s) still alive at the time of collection); and class 3), 1 or more CA but non-lactating (i.e., successfully bred but lost fawn(s) prior to collection). Capture of fawns for the radio telemetry study began 1 June 1993, 1994, and 1995 and continued until at least 20 fawns were captured or until fawns could no longer be captured. Captured fawns were sexed and fitted with Telonics (Telonics, Mesa, AZ) MOD-300 radio transmitters with whip antennas attached to CB-6 expandable break-away collars. Collars and transmitters of a brown color were used to match fawn pelage. Handlers wore vinyl gloves during capture and placement of collars. Radio transmitters were equipped with motion-sensing mercury switches that accelerated pulse rates if fawns were in- active for > 4 hr. Mortality was determined by a change in transmitter pulse rate and was verified by field observation. For the purposes of this paper, summer fawn mortality was defined as the number of fawns dying between parturition and the time when deer collections began (i.e, mid-August). For deer collections, mortality was calculated as the number of CA found in the ovaries of class 3 deer divided by the total number of CA present in the ovaries of both class 2 and class 3 deer (i.e., all deer that bred the previous fall). For the telemetry portion of this study, mortality 204 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) was calculated by dividing the number of fawns that died by the total number of fawns collared. Ninety-five percent simultaneous confidence intervals for proportional data were calculated for mortality rates using the Bailey method (Cherry 1996). Comparisons of mortality estimates between methods were conducted via t-tests (Zar 1996) on ranked data (Conover and Iman 1981) using SYSTAT (Wilkinson 1990).

RESULTS

Of the 75 deer collected in the Black Hills, data from 13 were incomplete and were excluded from analysis. In 1993, 4 animals were placed in class 1, 14 in class 2, and 2 in class 3. The ovaries of class 2 animals contained a total of 21 CA and 2 CA were found in the ovaries of class 3 animals. In 1994, there were 2 class 1, 15 class 2 (23 CA), and 2 class 3 (4 CA)deer. In 1995, there were 4 class 1, 17 class 2 (26 CA), and 2 class 3 (3 CA) deer. Twenty-six fawns were captured and ﬁtted with radio transmitters in 1993. Five of these fawns died within 48 hr of capture. These deaths were attributed to abandonment by the mother and were not considered in mortality calculations. Additionally, 6 fawns lost collars and also were not considered in mortality calculations. Of the 15 fawns used for mortality calculations, 5 died during 1 June - 15 August. In 1994, 10 fawns were captured but 2 lost collars and were excluded from mortality calculations. Two of the remaining 8 fawns died in the summer of 1994. In 1995, 21 fawns were captured. A total of 4 fawns was excluded from calculations, 1 due to abandonment, 2 because of collar loss, and 1 as a result of collar failure. Seven of the remaining 17 fawns died in the summer of 1995. Using the CA/lactation (CAL) method, summer fawn mortality was calculated at 0.1% < 8.7% < 31.7% (95% CI; note that the Bailey method does not necessarily produce symmetric intervals) in 1993, 2.1% < 14.8% < 37.2% in 1994, and 0.8% < 10.3 < 30.6% in 1995. Mortality rates calculated from telemetry data were 5.1% < 33.3% < 52.9% in 1993, 0.2% < 25.0% < 60.2% in 1994, and 8.8% < 41.2% < 54.6% in 1995. The CAL method yielded lower (t = -3.647, df = 4, P = 0.021) mortality estimates than radio telemetry data. Mortality rates estimates from the CAL method were 5, 41, and 75% of those from telemetry for 1993, 1994, and 1995, respectively.

DISCUSSION

We identiﬁed 10 sources of bias that could affect the accuracy of the CAL method of calculating summer mortality. Because of the importance associated with understanding biases inﬂuencing any method, a detailed account of each source of bias is provided below. Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 205

Polyovulation and Identical Twinning

If a single ova produces more than 1 fetus, CAL mortality will be biased downward. Instances of polyovulation and identical twinning have been documented for mule deer (Robinette and Gashwiler 1950) and white-tailed deer (Hesselton et al. 1964, Hesselton 1967, and Jocobson et al. 1979), but appear to be relatively uncommon occurrences. Robbinette and Gashwiler (1950) reported that approximately 0.6% of mule deer examined produced identical twins. Jacobson et al. (1979) reported that approximately 1.5% of white-tailed deer examined had more CA than embryos as a result of 1 or the other of these processes.

Prenatal Mortality

Mortality estimates from the CAL method will be biased upward if all eggs expelled during the estrus of conception are not fertilized. Because fertilization rates among deer are high, corpora lutea are generally a good index to the number of fetuses (Harder and Kirkpatrick 1994). Among white-tailed deer, comparisons of corpora lutea numbers to fetus numbers indicate ova/fetal survival rates between 84% and 90% at midgestation (Roseberry and Klimstra 1970, Jacobson et al. 1979, Woolf and Harder 1979, Mundinger 1981, Teer 1984). If fetuses are reabsorbed, aborted, or stillborn, CAL mortality will be biased upward. Reabsorption and abortion rates are more difﬁcult to quantify but appear to be low. Among mule deer, Robinette and Gashwiler (1950) noted a 0.9% reabsorption rate and Robinette et al. (1955) reported a 1.45% reabsorption/abortion rate. Data on mortality rates from midgestation to parturition are scarce, but Robinette et al. (1955) predicted an additional 4.4% during the latter part of gestation. While prenatal losses are not inconsequential, it is possible to quantify the level of impact these sources of bias may have upon CAL estimates using supporting reproductive data collected in the ﬁeld. For example, NBH deer collected during late February and early March of 1993 had 1.56 corpora lutea/female and 1.56 fetuses/female, indicating that all ova were fertilized and no fawns had been aborted by mid-pregnancy (R. G. Osborn, South Dakota State University, Unpubl. data). Additionally, there was no indication that fetuses were being reabsorbed. In the CBH, during 1994, there were 1.30 corpora lutea/female and 1.30 fetuses/female (B. A. Hippensteel, South Dakota State University, Unpubl. data). In the CBH during 1995, however, there were 1.50 fetuses/female and 1.55 corpora lutea/female. These discrepancies could have been the result of reabsorption or abortion. One female had 2 corpora lutea but only 1 fetus, a second female had 1 corpora lutea but no fetuses. If these discrepancies were the result of reabsorption or abortion, they occurred early in pregnancy because no signs of fetal reabsorption were noted during necropsy (B. A. Hippensteel, South Dakota State University, Unpubl. data). Therefore, the amount of error introduced by prenatal mortality during this study appears 206 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) to have been minimal. But, this source of error may not always be inconsequential and should re-evaluated on a case by case basis.

Unable to Differentiate CA of Estrus from CA of Pregnancy

If CA formed from corpora lutea of estrus cannot be distinguished from CA formed from corpora lutea of pregnancy, mortality rates will be biased upward. Although Cheatum (1949) reported that corpora lutea of estrus did not become pigmented as they degenerated, Golley (1957) found 2 large pigmented scars and 10 smaller pigmented scars in the ovaries of a captive black-tailed deer (O. h. columbianus) that had never been bred. After correction based on the size and color characteristics of CA described by Cheatum (1949), Golley (1957) reported that CA counts still overestimated the incidence of ovulation by 18%. Teer et al. (1965) found no reliable method to distinguish between CA formed from corpora lutea of pregnancy and pigmented scars from other sources. As with prenatal mortality, the impact of this source of bias can be assessed using reproductive data. Number of CA recorded during this study ranged from 0 to 3, with overall averages of 1.15, 1.32, and 1.26 CA per female in 1993, 1994, and 1995, respectively. These numbers were comparable with known white-tailed deer reproductive rates in the NBH in 1992 (1.12 fetuses/female) and 1993 (1.53 fetuses/female) (Osborn 1994), and for the CBH in 1995 (1.30 fetuses/female) and 1996 (1.36 fetuses/female) (Hippensteel 2000). Therefore, while it is possible pigmented scars other than those formed from corpora lutea of pregnancy were included in CA counts, it does not appear to have been a large source of error for the Black Hills data set.

CA Persistence

Mortality rates will be biased upward if it is not possible to distinguish CA scars of the most recent breeding season from those that are 2 or more years old. Cheatum (1949) reported CA scars degenerated and were not reliable for more than 8 months after parturition. However, CA persistence for 2 breeding seasons or longer has been documented for black-tailed deer (Golley 1957), white-tailed deer (Teer et al. 1965), and moose (Alces alces; Simkin 1965). Teer et al. (1965) reported that the number of CA present in ovaries increased with age in white-tailed deer. Simkin (1965) reported similar ﬁndings for moose. Using the macroscopic method it is difﬁcult to accurately distinguish between CA from the current breeding season and those of past breeding seasons; however, it is possible microscopically (Mansell 1971). Although deer collected in the Black Hills during this study ranged in age from 1 to 12 years (deer were aged via cementum annuli), the number of CA ranged from 0 to 3. As a result, CA persistence does not appear to have been a large source of error in this data set. Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 207

Lactating Cessation before Collection

Mortality will be biased upward if females with fawns cease lactating before they are collected. Infant ruminants are almost entirely dependent on milk during their ﬁrst month of life and milk production peaks when the fawns are approximately 1 month of age (Robbins et al. 1981). Although by 2 months of age a fawn’s diet consists primarily of vegetation, fawns continue to nurse at reduced levels until about four months of age (Short 1964). Scanlon and Urb- ston (1978) reported a minimum lactation period of 4 months for white-tailed deer, but noted that 55% of mothers were lactating until December. Lactation periods extending 6 months also were recorded by Woolf and Harder (1979). In the NBH, peak fawning occurs around mid-June (Osborn 1994, Benzon 1996, Hippensteel 2000). Assuming a minimum lactation duration of 4 months (Scanlon and Urbston 1978), all females collected during this study should have been lactating if their fawn(s) were still alive. Because all NBH and CBH deer were collected during the last 2 weeks of August, the error introduced by this source of bias should have been inconsequential. However, as collection date increases past 4 months post parturition this will become an increasingly important source of error.

Lactation Persistence

Mortality will be biased downward if females do not stop lactating soon after all fawns die. Information concerning the persistence of lactation after the death of young is limited. Nursing is one of the primary stimuli maintaining lactation and if the mammary gland is not emptied of milk, lactation ceases rapidly (Nalbandov 1964). Similarly, Grosvenor et al. (1970) reported that milk production may start to slow in 12 - 24 hours if nursing does not occur. Bergerud (1964) observed that by 7 days after the death of a calf, udder dimensions of a female caribou (Rangifer tarandus) had decreased by approximately two-thirds. Among cow elk (Cervus elaphus) that lose their calves, lactation ceases within a few days (Bubenik 1982). Unfortunately, because of a lack of information, the potential effects of this source of bias cannot be accurately assessed for white-tailed deer without further research. Its inﬂuence may be minimized, however, by allowing approximately a 2-week buffer between the end of the period of interest and the start of deer collections (i.e., if the period of interest ends in August, then start deer collections the third week in September).

Partial Litter Loss

Utilizing the CAL method, the death of a single fawn in a litter of 2 or more would go undetected, biasing mortality estimates downward. During the 3 years of study, there were 3 cases when 1 fawn of radio-collared twins died. Because a total of 14 collared fawns died, the CAL method could have under- represented the number of fawns that died by 21.4%. For adult female white- tailed deer in good nutritional condition, twinning is common. While there is 208 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) limited information available concerning the frequency with which partial litter loss occurs, it is likely to be a large source of error.

CONCLUSIONS

We identiﬁed ten sources of bias that may inﬂuence the accuracy of using the CAL method to estimate summer fawn mortality. Some are inconsequential or can be avoided by using proper collection procedures. Others are un- avoidable, but their impact can be assessed through the collection of supporting data. Partial litter loss was the bias that introduced the largest amount of error. While the bias associated with partial litter loss cannot be avoided by modifying collection procedures, it may be possible to avoid or minimize this bias by applying the CAL procedure only to ungulates that typically produce only a single offspring per litter. Bison (Bison bison), Dall’s sheep (Ovis dal- li), Stone’s sheep (O. d. stonei), bighorn sheep (O. canadensis), muskoxen (Ovibos moschatus), caribou (Rangifer tarandus), and elk are all examples of North American ungulates for which twinning is uncommon (Demaris and Krausman 2000). At this point in time, estimating summer fawn mortality using CA and lactation status for any ungulate that commonly produces more than one offspring per litter does not seem practical. For ungulates that rarely produce more than one offspring the CAL procedure might prove useful provided additional data is collected to allow the potential impact of biases to be evaluated. Further research is necessary to evaluate the practicality of using the CAL method for other species. If practical, the CAL method of estimating summer fawn mortality could be a valuable tool for collecting initial mortality estimates. Preliminary results derived from the CAL procedure could then be used to help focus more intensive radio telemetry studies.

ACKNOWLEDGMENTS

Funding for this research was provided by Pittman-Robertson funds (Pro- ject W-75-R, Studies No. 7558, 7566, 7567, and 7554, Federal aid to Wildlife Restoration Fund) through the South Dakota Department of Game, Fish and Parks. Thanks are extended to L. Rice, R. Hauk, and S. Grifﬁn, J. Reick, C. De- Perno, D. Flory, D. Gray, D. Knowlton, R. Lasco, J. McCormick, S. Mickelson, M. Muck, K. Wallin, J. Wrede, K. Wintersteen, and many landowners in the Black Hills; without their help, this project would not have been possible.

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LONGITUDINAL PATTERNS IN FISH COMMUNITY COMPOSITION OF UPPER MISSOURI RIVER BACKWATERS

Kipp A. Powell Department of Wildlife and Fisheries Sciences South Dakota State University Brookings, SD 57007

Steven R. Chipps USGS South Dakota Cooperative Research Unit South Dakota State University Brookings, SD 57007

ABSTRACT

The Garrison reach is a 137 km free-flowing section of the Missouri River connecting Lake Sakakawea to Lake Oahe. While fish composition in the main- stem river has been well documented, little is known about fish composition and abundance in associated backwater areas. In September 2001, we sampled 16 backwaters for juvenile fish composition; twelve sites (75%) contained fish. Most sites were relatively close to the river (mean distance = 48 m) and were strongly influenced by water releases from Garrison Dam. A total of 20 fish species were collected from backwater reaches, with most sites containing between 2-10 species. Seventeen of the species captured are native to North Dakota. Fathead minnows (Pimephales promelas) and white suckers (Catosto- mus commersoni) were the most frequently captured species, occurring in 88% of the sites, whereas common carp (Cyprinus carpio), shortnose gar (Lepisos- teus platostomus), white bass (Morone chrysops), golden shiners (Notemigonus crysoleucas), and largemouth bass (Micropterus salmoides), occurred infre- quently (<7%) in backwater areas. Of the 20 species captured, white suckers were most abundant (45% of total catch), while shortnose gar and largemouth bass were least abundant (0.03% of total catch). The Shannon-Wiener Index of diversity (SWI) for each site was positively correlated with distance from Gar- rison Dam. Although impoundments have dramatically altered the natural hy- drograph and floodplain characteristics of the Missouri River, our findings indicate that backwater habitats provide important spawning and rearing areas for fishes, particularly for warmwater fishes.

INTRODUCTION

The distribution and composition of ﬁshes in the upper Missouri River was generally poorly understood prior to mid 1980’s, with few published records. 212 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002)

Hesse et al. (1989) reported 64 fish species found within the North Dakota boundaries of the Missouri River basin. Of these, 53% were considered rare, threatened or endangered by the U.S. Fish and Wildlife Service (USFWS) or the state of North Dakota. In the lower Missouri River, most backwater and floodplain habitats have been lost due to impoundments and channelization (Sandheinrich and Atchi- son 1986). The abundance of off channel habitat (backwaters and tributaries) once provided by the Missouri River may have provided up to 70 percent or more of the icthyoplankton and 90 percent of fish found in the river. As pointed out by Hesse and Sheet (1993), the loss of these important habitats may lead to a significant reduction in fish diversity and biomass in the Missouri River. The objective of this study was to determine species composition and relative abundance of juvenile fishes associated with small (<1 ha) periodically connected backwaters and examine changes in species composition along the Garrison Reach.

STUDY AREA

The Garrison Reach extends from Garrison Dam at river kilometer (rkm) 2234.9 in south central North Dakota to the headwaters of Lake Oahe (rkm 2098.1) near the North Dakota- South Dakota border. For sampling purposes, we stratiﬁed the Garrison Reach into three river segments; tailrace (2234.9- 2204.9 rkm), middle (2184.9-2154.9 rkm) and lower (2134.9-2104.9 rkm).

METHODS

We sampled 16 small backwaters (< 1 ha) for fishes during September 2001; four sites from the tailrace, eight sites from the middle river segment and four sites from the lower segment. Sites were sampled with a 3.2 m by 1.0 m beach seine with 3.0 mm mesh. Seine hauls were standardized to 13 m length hauls pulled perpendicular to shore. Each site was divided into three sections (an upper, middle and lower section) and a single seine haul was made in each section. If a site was not large enough to allow three complete hauls, then one or two standardized seine hauls were made. All fish collected from backwater areas were identified, enumerated and returned to the water. When field identification was not possible, fish were placed in to labeled containers and trans- ported to the laboratory for accurate identification (Pflieger 1997, Eddy 1978 and Lee et al. 1980). We calculated catch per unit effort (CPUE) for individual species from each backwater sampled and expressed relative abundance of fishes as the mean number of fish per seine haul. We characterized fish communities using several measures that included 1) species richness (total number of species per site), 2) frequency of occurrence (percent of sites where a species occurred), 3) percent composition of total Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 213 catch, and 4) the Shannon-Wiener index of diversity (H’) where H’ was calculated as: Σ H’= s (pi) (log2 pi)

where H’ is species diversity, s is the number of species, and pi is the proportion of total sample belonging to the ith species.

RESULTS AND DISCUSSION

We collected a total of 2,951 ﬁsh from backwater areas of the Garrison reach in September 2001. Species composition was comprised of seven orders, 10 families and 20 species (Table 1). All 20 species have been previously en-

Table 1. Percent composition and frequency of occurrence of ﬁshes collected from small seasonally connected backwaters of the Garrison Reach, Missouri River, North Dakota, September 2001. Bold represents non-indigenous species in North Dakota.

Species Total Mean Frequency Number Percent of Collected Composition Occurrence white sucker (Catostomus commersoni) 1334 45.2 56.3 fathead minnow (Pimephales promelas) 782 26.5 56.3 black bullhead (Ameiurus melas) 268 9.1 12.5 Johnny darter (Etheostoma nigrum) 195 6.6 25.0 black crappie (Pomoxis nigromaculatus) 99 3.4 37.5 bluegill (Lepomis macrochirus) 92 3.1 18.8 yellow perch (Perca fluviatilis) 48 1.6 31.3 banded killifish (Fundulus diaphanus) 47 1.6 12.5 emerald shiner (Notropis atherinoides) 30 1.0 18.8 river carpsucker (Carpiodes carpio) 26 0.9 37.5 smallmouth bass (Micropterus dolomieui) 9 0.3 12.5 brook stickleback (Culaea inconstans) 7 0.2 25.0 spottail shiner (Notropis hudsonius) 5 0.2 12.5 white bass (Morone chrysops) 4 0.1 6.3 common carp (Cyprinus carpio) 2 0.1 6.3 northern pike (Esox lucius) 2 0.1 12.5 plains minnow (Hybognathus placitus) 2 0.1 12.5 shortnose gar (Lepisosteus platostomus) 2 0.1 6.3 golden shiner (Notemigonus crysoleucas) 1 0.0 6.3 largemouth bass (Micropterus salmoides) 1 0.0 6.3 214 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) countered within the Garrison reach based on recent historical records (Hen- drickson et al 1993, Mizzi 1994, Welker 2000). Seventeen of the species collected are indigenous, while three are non-native to the upper Missouri River. Native species comprised 91% of the fishes captured by number whereas non- native fishes comprised approximately 9.4% of the total catch. White suckers (Catostomus commersoni), fathead minnows (Pimephales promelas) Johnny darters (Etheostoma nigrum) and black bullhead (Ameiurus melas) comprised over 87% of the total catch by numbers. Moreover, white suckers, fathead minnows, black crappie (Pomoxis nigromaculatus), yellow perch (Perca fluviatilis) and river carpsuckers (Carpiodes carpio) were all found in more than 30% of sites sampled. Common carp (Cyprinus carpio), golden shiner (Notemigonus crysoleucas) shortnose gar (Lepisosteus platostomus) and white bass (Morone chrysops) were collected in less than 7% of the sites sampled (Table 1). Catch per unit effort varied appreciably from site to site and among river segments. Mean CPUE was generally highest for white suckers (46 fish/haul) and fathead minnows (44 fish/haul). Catch per unit effort for other species ranged from 0.02 for largemouth bass (Micropterus salmoides) and shortnose gar to 11.8 for black bullheads. Only four species were collected from backwater areas near the tailrace (Fig. 1). White suckers and Johnny darters were the most abundant fishes collected in the tailwater segment and were over three times more abundant than emerald shiners and fathead minnows (Fig.1). Within the middle river segment, 12 species were captured; eight new species not found in the tailrace river segment (Fig. 2). Fathead minnows were the most abundant (mean CPUE=88 fish/haul) followed closely by white suckers (mean CPUE=84 fish/haul). The least abundant species found within the middle river segment were north- Figure 1. Catch per unit effort (number of fish/sine ern pike (Esox lucius) and haul) for species collected from backwater areas the plains minnow (Hybog- within the tailrace river segment of the Garrison nathus placitus), each with Reach, Missouri River (rkm 2234.9-2204.9). a mean CPUE of 0.04 individuals per haul. The middle river segment is the first portion of the Garrison reach where "game fish" appeared in small backwater areas. The presence of yellow perch and northern pike, while relatively low in abundance, indicates that small backwater areas may provide important habitat for game species (Fig. 2). Fifteen fish species were captured from backwater areas in the lower river segment. Seven species collected from the lower river segment were not Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 215 found in sites upstream (e.g. tailrace and middle river segments). Unlike backwater areas upstream, the most abundant species in the lower river segment is a non-native (i.e. black bullhead; Fig. 3). The least abundant species collected in the lower river segment were largemouth bass, plains minnow, and shortnose gar (Fig. 3). Sport fishes collected from the lower river segment represented 47% of the total Figure 2. Catch per unit effort (number of fish/sine catch and included bluegill, haul) for species collected from backwater areas black crappie, yellow within the middle river segment of the Garrison perch, white bass, small- Reach, Missouri River (rkm 2184.9-2154.9). mouth bass, northern pike, and largemouth bass (Fig. 3). In general, small backwater areas in the Garrison reach are dominated by few species with most species comprising a small portion of the total catch. This is further demonstrated by measures of fish diversity (H’) where mean diversity in backwaters areas was relatively low at H’=0.83. However, species diversity was positively (al- beit weakly) correlated with distance from Garrison Dam (correlation analysis; Figure 3. Catch per unit effort (number of fish/sine r=0.43, P=0.09) suggesting haul) for species collected from backwater areas that fish diversity increases within the middle river segment of the Garrison downstream (Fig. 4). Reach, Missouri River (rkm 2134.9-2104.9). Related studies in the Missouri River indicate that total species richness in the Garrison reach is probably higher that that observed in backwater areas. Mizzi (1994) collected 35 species of fish from a 2.0 km portion of the Garrison reach around Apple Creek, south of Bismarck, ND. Similarly, Welker (2001) documented 21 species through out the length of the 216 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002)

Garrison reach. Because sampling methodologies differed among studies, as did sampling effort, mean- ingful comparisons of relative abundance are difficult to make. Comparative analyses of presence/absence, however, would provide at least a measure of taxa richness in the Garr- sion reach and this information has been summarized Figure 4. Shannon-Wiener Index of diversity based on abundance and composition of species by Powell (M.S. thesis, collected from small backwaters of the Garrison South Dakota State Univer- Reach, Missouri River, plotted against distance in sity, in press). Based on da- river kilometers from Garrison Dam, North Dakota. ta presented here, small floodplain backwaters (<1.0 ha) appear to provide suitable rearing habitat for many species of fishes found in the Missouri River. Moreover, small backwaters in the lower most portion of the Garrison reach provide suitable habitat for a variety of gamefishes including centrarchids and generally support more species than sites located in the tailrace area.

ACKNOWLEDGEMENTS

We would like to thank Dr. Craig Paukert for his help with statistical analysis and friendly review of this manuscript. This project was supported, in part, by the U.S. Geological Survey South Dakota Cooperative Research Unit, the U.S. Environmental Protection Agency, the Department of Wildlife and Fish- eries Sciences at South Dakota State University.

LITERATURE CITED

Eddy, S. and J.C. Underhill. 1978. How to know the freshwater ﬁshes, 3rd edition. McGraw-hill, New York. 215 pp. Hendrickson, J. C., J.D. Lee, and L.C. McGregor. 1993. Aquatic investigation of the Missouri River system in North Dakota. North Dakota Game and Fish Department, Fisheries Investigation Report Number 7, Bismarck. Hesse, L. W., and W. Sheets. 1993. The Missouri River hydrosystem. Fisheries 18(5):5-14. Lee, D.S., C.R. Gilbert, C.H. Jenkins, D.E. McAllister and J.R. Stauffer, Jr. 1980. Atlas of North American freshwater ﬁshes. North Carolina Biological Sur- vey Publication number 1980-12 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 217

Mizzi, J.M. 1994. Zooplankton, macroinvertebrate, herpetile, and ichthyofaunal biodiversity of riverine habitat on the upper Missouri River. Masters Thesis, South Dakota State University. Pflieger, W.L. 1997. The fishes of Missouri. Missouri Department of Conserva- tion, Jefferson City. 372 pp. Sandheinrich, M.B., and G.J. Atchison. 1986. Fish associated with dikes, revet- ments, and abandoned channels in the middle Missouri River. Proceedings of the Iowa Academy of Science 93:188-191. Welker, T. L. 2000. Ecology and structure of fish communities in the Missouri and Yellowstone Rivers. Doctoral Dissertation, University of Idaho.

Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 219

ZOOPLANKTON COMMUNITIES OF BLACK HILLS RESERVOIRS

Benjamin M. Holcomb and Steven R. Chipps USGS South Dakota Cooperative Fish & Wildlife Research Unit Department of Wildlife & Fisheries Sciences South Dakota State University Brookings, SD 57007

ABSTRACT

Black Hills reservoirs exhibit considerable variation in nutrient levels and algal biomass. Water clarity, for example, ranges from 2 m in Stockade Reser- voir to 9 m in Pactola Reservoir. Variation in trophic state and fish composition likely has important implications for secondary productivity, though little is known about the zooplankton communities in Black Hills reservoirs. Here, we report results from a comparative analysis of zooplankton composition and abundance in four Black Hills reservoirs: Pactola, Deerfield, Sheridan, and Stockade. We sampled five sites in each reservoir from October 2000 to Oc- tober 2001. Species richness (9 species) was similar in all reservoirs, although species composition varied across systems. Mean annual zooplankton density was highest in Stockade Reservoir (187.7 No./L) and lowest in Pactola Reser- voir (51.4 No./L). Copepods were a dominant component of the zooplankton assemblage in Pactola and Deerfield reservoirs. Moreover, the cladoceran community in Pactola and Deerfield reservoirs was dominated by Daphnia galeata mendotae, whereas in Sheridan and Stockade, Daphnia pulex was the most abundant Daphnia species collected. Because D. pulex often replaces D. mendotae when fish predation is low, information on zooplankton composition may be useful for monitoring changes in planktivorous fish abundance in Black Hills reservoirs.

Keywords

Black Hills, zooplankton, reservoirs, ﬁsh, phytoplankton, trophic cascade

INTRODUCTION

Zooplankton populations represent an important link between primary productivity (i.e. algae production) and higher trophic levels (i.e. fish) in lakes and reservoirs. Herbivorous zooplankton, such as Daphnia, can also play an important role in regulating algal biomass and water quality. Most often, zooplankton are monitored for their abundance, size-structure, and community composition. By characterizing zooplankton composition and abundance, we gain information on food abundance for planktivorous fish (Mills and Schi- 220 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) avone 1983) and the potential for water quality enhancement via biomanipu- lation (Carpenter et al. 1985). Recent studies have shown that when fish predation is low, larger Daph- nia (e.g., D. pulex) typically out-compete smaller Daphnia (e.g., D. galeata mendotae) (Prazakova 1991; Gliwicz and Lampert 1993; Johnson and Goettel 1999). Larger species can survive under lower food thresholds, thus enabling them to out-compete smaller-bodied species even under oligotrophic conditions. However, fish predation can play a key role in structuring the zooplankton community. When planktivorous fish are abundant, larger zooplankton are often reduced via size-selective predation (Brooks and Dodson 1965; Zaret 1980) and the zooplankton community shifts toward a dominance by smaller bodied taxa. Black Hills reservoirs vary significantly in trophic status, water clarity and fish community composition (German 1997; J. Erickson, SDGFP, pers. comm.). Unfortunately, there is little information regarding zooplankton communities in Black Hills reservoirs. The last recorded observations on zooplankton communities in Black Hills reservoirs were performed in 1976 (A. Repsys, SD DENR, pers. comm.). Because herbivorous zooplankton play an important role in lake ecosystems, knowledge regarding seasonal composition and abundance of zooplankton can have important implications for resource managers. The objectives of this study were to 1) quantify and compare zooplankton composition and abundance among reservoirs, 2) evaluate relationships between zooplankton abundance and algal biomass and 3) compare recent observations with historical data on zooplankton composition in Black Hills reservoirs.

STUDY AREA

Four major reservoirs occur within the Black Hills of South Dakota and include: Pactola, Deerfield, Sheridan, and Stockade. Pactola is the largest reservoir with a surface area of 785 acres and a maximum depth of 150 ft. Stock- ade is the smallest reservoir at 120 acres and a maximum depth of 50 ft. Deer- field and Sheridan reservoirs are intermediate in size (about 400 acres) with maximum depths reaching 70 ft. Pactola is a meso-oligotrophic reservoir with secchi depths often reaching 9 m and summer total phosphorus concentrations of 0.007 mg/L (Fig. 1). Deerfield reservoir is primarily mesotrophic (total phosphorus = 0.011 mg/L), whereas Sheridan (total phosphorus=0.032 mg/L) and Stockade reservoirs (total phosphorus=0.124 mg/L) are characterized as eutrophic systems.

METHODS

We sampled Black Hills reservoirs seasonally from October 2000 to Octo- ber 2001. Five sites were sampled in each reservoir, except in winter when three sites were sampled through the ice at each reservoir. At each site, phy- Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 221 toplankton biomass (i.e., chlorophyll a) and zooplankton were sampled. Phytoplankton were collected at the surface level and at 1 m above the sediment using a 1_L Kemmer- er bottle. Chlorophyll a biomass (µg/L) was measured by filtering at least 250 ml of lake water through a 0.7 µm GF/F filter, wrapping the filter in aluminum foil, then freez- Figure 1. Relationship between mean summer water clarity and mean summer surface algal ing the filters for later anal- biomass in Pactola, Deerfield, Sheridan and Stock- ysis. Chlorophyll a was ade reservoirs during summer (July-August) 2001. measured using a Turner Design TD-700 fluorometer after 24 h extraction in 90% acetone. An 11 cm Wisconsin tow net with 63 µm mesh was used to collect three zooplankton samples at each site from the top 5 m of the water column, rinsed into sample vials, and then preserved in 10% Lugol’s solution. In the laboratory, 10% of each sample was identified, counted and ten of each species per sample were measured for total length to determine density, species composition and size structure of the population (Pennak 1989). Biomass was estimated using length-weight regressions from Dumont et. al. (1975). Differ- ences in zooplankton abundance, species composition, size structure, and biomass were determined using analysis of variance (SAS Institute Inc. 1999).

RESULTS

Zooplankton species richness (n=9) was similar across reservoirs, although species composition differed among reservoirs (Table 1). Additionally, species

Table 1. Zooplankton species composition for Pactola, Deerﬁeld, Sheridan, and Stock- ade reservoirs from October 2000-October 2001. X= presence O= absence

Species Pactola Deerﬁeld Sheridan Stockade

Daphnia mendotae XXXX Daphnia pulex XOXX Ceriodaphnia lacustris XXXX Alona costata XXXX Camtocercus macrurus OXOO Bosmina longirostris XXXX Chydorus sphaericus XXXX Diaptomus siciloides XXXX Cyclops bicuspidatus XXXX 222 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002)

abundance differed appreciably across reservoirs. Mean total zooplankton density was highest in Stockade Reservoir (187.7 No./L) and lowest in Pacto- la Reservoir (51.4 No./L) (Fig. 2). However, total zooplankton abundance was similar in Sheridan and Deerfield reservoirs (Tukey’s HSD, P = 0.97). Figure 2. Mean annual zooplankton abundance Zooplankton biomass ap- for Pactola, Deerfield, Sheridan, and Stockade peared to be a better indi- reservoirs from October 2000- October 2001. cator of reservoir produc- Means with the same letter are not significantly tivity, as mean summer al- different (P > 0.05, Tukey’s HSD). Vertical bars represent 1 S.E. gal biomass was positively correlated with mean summer zooplankton biomass in Black Hills reservoirs (correlation analysis, r=0.97; P<0.05; Fig. 3). Copepods were a dominant component of the zooplankton community in Pactola and Deerfield reservoirs, comprising up to 70 percent of total zooplankton biomass (Fig. 4). The percentage of copepods in these reservoirs was significantly higher than those of Sheridan and Stockade (Tukey’s HSD, P = 0.0004). Figure 3. Relationship between mean zooplankton Moreover, the cladoceran biomass and mean algal biomass for August 2001. community in Pactola and Deerfield reservoirs was dominated by Daphnia galeata mendotae, whereas in Sheridan and Stockade, Daphnia pulex was the dominant grazer (Fig. 5). D. pulex was significantly larger than D. galeata mendotae in all reservoirs where it occurred (Tukey’s HSD, P < 0.05). Furthermore, although not significantly different, D. galeata mendotae was generally larger in Sheridan and Stockade reservoirs than in Pactola and Deerfield reservoirs (Fig. 6). While most species increased in abundance from 1976 to 2001, Daph- nia pulex appeared less frequently in Pactola and Deerfield reservoirs in 2001 samples. In fact, of these two reservoirs, only one specimen was collected from Pactola Reservoir in 2001 (Table 2). Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 223

DISCUSSION

Zooplankton community structure differed appreciably among Black Hills reservoirs. Reservoir productivity, as measured by algal biomass, was positively correlated with zooplankton biomass and abundance. However, composition and size structure of herbivorous cladocerans (i.e. Daphnia spp.) Figure 4. Mean percent of cladocerans and cope- differed among reservoirs, pods in Pactola, Deerfield, Sheridan, and Stockade with smaller species (i.e. reservoirs from October 2000 through October Daphnia g. mendotae) gen- 2001. For mean percent copepod biomass, values with the same letter are not significantly different erally more abundant in (P > 0.05, Tukey’s HSD). Pactola and Deerfield reservoirs and larger species (i.e. Daphnia pulex) more abundant in Sheridan and Stockade reservoirs. Daphnia pulex has never been found to dominate Pactola and Deerfield reservoirs, although its presence has been noted in both. When established, D. pulex will often outcom- pete smaller Daphnia (Prazakova 1991). Howev- er, when planktivorous fish are abundant, larger daph- nids like D. pulex are the Figure 5. Mean annual abundance of D. galeata first to be removed (Brooks mendotae and D. pulex for Pactola, Deerfield, Sheridan, and Stockade reservoirs from October and Dodson 1965; Cerny 2000- October 2001. and Bytel 1991). Both reservoirs contain planktivorous fishes although fish community composition varies appreciably among reservoirs. Warm-water fishes such as yellow perch and several speicies of centrarchids (e.g. black crappie and largemouth bass) are common in Sheridan and Stockade reservoirs, whereas Deerfield and Pactola are primarily put-and-take salmonid fisheries (e.g. cold- water fishes). Trout, which are primarily daytime feeders, are effective zoo- planktivorous fishes and may exert considerable predation pressure on open water zooplankton (e.g. exploit large-sized Daphnia) leaving smaller cladocer- 224 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002)

Table 2. Abundance (No./L) of selected zooplankton from Pactola, Deerﬁeld, Sheridan, and Stockade reservoirs from the Fall of 1976 and mean of Falls 2000 and 2001.

Fall 1976a Fall 2000 & 2001 D. D.g. C. B. D. D.g. C. B. pulex mendotae lacustris longi- pulex mendotae lacustris longi- Reservoir rostris rostris

Pactola 0.6 1.6 6.9 1.0 0.06 13.4 0.5 6.9 Deerﬁeld 0.6 5.0 14.1 5.0 0.0 13.8 5.6 17.1 Sheridan 0.0 5.8 2.1 1.9 9.9 13.5 2.1 24.7 Stockade 10.4 9.2 0.0 7.9 19.4 0.74 6.5 4.6 a South Dakota DENR, 1976

ans and copepods to dominate the community. Cope- pods are also known to prey upon small cladocerans and when abundant, can have a negative influence on cladoceran populations. Gliwicz and Lampert (1993) found that copepods were able to influence the zooplankton community by preying on juvenile clado- Figure 6. Mean lengths of D. galeata mendotae cerans, thereby inhibiting and D. pulex in Pactola, Deerfield, Sheridan, and their abundance. Further Stockade reservoirs from October 2000- October evidence of higher preda- 2001. Means with the same letter are not signifi- tion pressure on Daphnia cantly different (P > 0.05, Fisher’s LSD). Vertical bars represent 1 S.E. in Pactola and Deerfield reservoirs is that the size of D. galeata was generally smaller than those collected from Sheridan and Stockade reservoirs (Fig. 6). Typically, Daphnia will mature at smaller sizes when predation pressure is high (Zaret 1980). Diel vertical migration may also explain, to some extent, differences in Daphnia composition and size structure among Black Hills reservoirs. Because Sheridan and Stock- ade reservoirs generally go anoxic near the bottom (e.g. hypolimnion) every summer, it potentially limits the magnitude of diel vertical migration exhibited by Daphnia. Because our sampling was conducted during the day and did not include the entire water column, we could have underestimated Daphnia abundance in Pactola and Deerfield reservoirs if they were distributed in deeper water strata. Zooplankton composition and abundance can provide important information regarding water quality conditions and trends in planktivorous fish abundance. In Black Hills reservoirs, nutrient levels (e.g. total phosphorus) are gen- Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 225 erally lower in Pactola and Deerfield reservoirs than in Sheridan and Stockade (German 1997). Total algal biomass and zooplankton abundance reflect these patterns and indicate that overall productivity in Black Hills reservoirs is driven by nutrient availability. Changes in species composition and mean size of herbivorous Daphnia, however, suggest that predation pressure varies among reservoirs and may be related to differences in fish community composition. Future efforts that document planktivorous fish abundance would provide useful information for exploring the linkage between fish abundance and zooplankton composition/size structure.

LITERATURE CITED

Brooks, J.L. and S.I. Dodson. 1965. Predation, body size, and composition of zooplankton. Science. 150: 28-35. Carpenter, S.R., J.F. Kitchell, and J.R. Hodgson. 1985. Cascading trophic interactions and lake productivity. Bioscience 35: 634-639. Cerny, M. and J. Bytel. 1991. Density and size distribution of Daphnia populations at different fish predation levels. Hydrobiologia. 225: 199-208. Dumont, H.J., I. Van de Velde, S. Dumont. 1975. The dry weight estimate of biomass in a selection of Cladacera, Copepoda and Rotifera from the plankton, periphyton and benthos of continental waters. Oecologia. 19: 75-97. Johnson, B.M. and J.P. Goettl. 1999. Food web changes over fourteen years following introduction of rainbow smelt into a Colorado reservoir. North American Journal of Fisheries Management. 19: 629-642. German, D.R. 1997. South Dakota Lake Protection Water Quality Report. Wa- ter Resources Institute, South Dakota State University, Brookings, SD Gliwicz, M.Z. and Lampert, W. 1993. Body-size related survival of cladacerans in a trophic gradient: an enclosure study. Arch. Hydrobiol. 129: 1-23. Lampert, W. W. Fleckner, H. Rai, B.E. Taylor. 1986. Phytoplankton control by grazing zooplankton: A study on the spring clear-water phase. Limnolo- gy and Oceanography. 31(3) 478-490. Mills, E.L. and A. Schiavone. 1982. Evaluation of fish communities through assessment of zooplankton populations and measures of lake productivity. North American Journal of Fisheries Management. 2: 14-27. Pennak, R.W. 1989. Freshwater invertebrates of the United States, 3rd edition. John Wiley and Sons, New York, NY. Prazakova, M. 1991. Impact of fishery management on Cladaceran populations. Hydrobiologia 225: 209-216. SAS Institute Inc., SAS® Procedures Guide, Version 8, Cary, NC; SAS Institute Inc., 1999. Zaret, T.M. 1980. Predation and freshwater communities. Yale University Press, New Haven Connecticut.

Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 227

FECAL EGG AND OOCYST COUNTS IN DOGS AND CATS FROM ANIMAL SHELTERS FROM SOUTH DAKOTA

M.B. Hildreth, J.A. Bjordahl and S.R. Duimstra Departments of Biology & Microbiology and Veterinary Sciences South Dakota State University Brookings, SD 57007

ABSTRACT

Dogs and cats serve as hosts to a variety of gastrointestinal parasites, some of which, can also infect humans. Little is known of the current prevalence and intensity of these parasites in dogs and cats from South Dakota. From 1994 to 1996, 580 dog fecal samples and 88 cat samples were collected from seven different animal shelters located in the following cities: Aberdeen (85 samples), Brookings (96 samples), Huron (49 samples), Pierre (28 samples), Sioux Falls (32 samples), Watertown (261 samples) and Yankton (29 samples). Samples were store at 4ºC until they were examined via a standard centrifugal sucrose flotation procedure. Parasites eggs were identified to the genus level when possible (i.e. Toxacara eggs, Strongyloides eggs, and Dipylidium eggs) or to the family level when necessary (Taeniidae or Ancylostomatoidea). Coc- cidian oocysts were not further identified to genus or family. At least one parasite was identified in 35.2% of the cat samples and 27.4% of the dog samples. Over 20% of the cats were shedding coccidian oocysts, and the most commonly identified helminth parasite was Toxacara cati (13.6%). Five cats were shedding Taeniidae eggs and only four cats were shedding hookworm eggs. The most commonly identified gastrointestinal parasite in dogs was Toxacara canis (9.1%). Coccidian oocysts were found in 7.9% of the dog samples, and hookworm (Ancylostomatoidea) eggs were found in 7.4%. Taeniid tapeworm eggs were found in 5.2% of the samples; only one dog was found to be infected with Dipylidium. No Trichuris whipworm eggs were found in any of the samples. Prevalences for several of the parasites in this study were significantly different from those reported in a recent national survey involving dogs from strictly urban sites.

INTRODUCTION

Dogs and cats serve as hosts to a variety of gastrointestinal parasites, some of which, can also infect humans. A recent national survey of dog parasites, based on fecal ﬂotations analysis of samples from 2,322 dogs housed in urban animal shelters, found that almost 36% of the dogs harbored Toxocara canis, Ancylostoma caninum and/or Trichiuris vulpis nematodes (Blagburn et al., 1996). Toxocara canis and A. caninum juveniles are infectious to humans 228 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) causing migrating larvae diseases (visceral larval migrans for T. canis; cutaneous larval migrans for A. caninum). Visceral larval migrans is a potentially lethal disease in children, while cutaneous larval migrans is simply a nuisance condition. Other potentially zoonotic parasites were also found in these dogs. The national survey found that only 0.6% of the dogs were excreting taeniidae tapeworm eggs (i.e. Taenia and Echinococcus). Echinococcus eggs from dogs are directly infectious to humans, but Taenia eggs are not. It is not possible to morphologically differentiate Taenia eggs from Echinococcus eggs, yet, it is very likely that most, if not all, of the dogs excreting Taeniidae eggs in the national survey were infected with a Taenia pisiformis because Taenia pisiformis is a cosmopolitan parasite of dogs, and Echinococcus (i.e. Echinoccus granu- losus and E. multilocularis) has a much more restricted distribution and prevalence. An unusually high percentage of wild canids from South Dakota are infected with E. multilocularis (e.g. approximately 65% of the red foxes), and it is possible that dogs from this area are also becoming infected with this tapeworm (Hildreth et al., 2000). Cats can also become infected with E. multilocularis, however, it’s growth is stunted, and egg production is very reduced (Crellin et al., 1981). Nothing is known of the current prevalence and intensity of E. multilocularis in dogs and cats from the United States Northern Plains (Hildreth et al., 1991). However, if the prevalence of taeniid eggs is as low in dogs throughout South Dakota, then this would suggest that the potential number of dogs excreting E. multilocularis eggs must be extremely rare. The central purpose of this study was to determine the prevalence of intestinal parasites in dogs and cats from South Dakota. This prevalence study was based upon identifying eggs and cysts of these parasites in fecal samples from dogs and cats housed in animal shelters located in eastern South Dakota. Focus was given to trematode, cestode and nematode helminths, and to coccidian proto- zoans. Eggs from A. caninum can be differentiated from Uncinaria stenocephala based upon the size of the egg, however, no attempts were made for this differentiation in this study. These eggs were simply reported as members of the family Ancylostomatoidea (i.e. hookworms). Potential protozoan parasites also found in dogs, such as Giardia and Cryptosporidium, were not included in this study. Special attention was given to determining factors affecting dogs excreting Taeniidae eggs.

MATERIALS AND METHODS

From June 1994 to July 1996, fecal samples were collected from 580 dogs housed at seven different animal shelters located in the following cities throughout South Dakota: Aberdeen (85 samples), Brookings (96 samples), Huron (49 samples), Pierre (28 samples), Sioux Falls (32 samples), Watertown (261 samples) and Yankton (29 samples). Within the 3 years of sampling, 29.2% of these samples were collected during 1994; 34.5% during 1995 and 36.2% during 1996. During 1995 and 1996, fecal samples were also collected from 88 cats. These samples were received from only 4 of the cities: Aberdeen (62 samples), Brookings (15 samples), Huron (10 samples) and Watertown (1 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 229 sample). All samples were shipped and stored at 4ºC until examined via a standard centrifugal sucrose flotation procedure (Bowman, 1995). For this procedure, 1.0 g of feces were added to 10 ml distilled water and stirred to suspend the material. The suspension was then poured through a filter funnel into a centrifuge tube, and centrifuged at 2,000 rpm in a clinical centrifuge for 10 minutes. The fluid was then decanted off of the pellet, and the tube refilled (within a couple cm of the lid) with an aqueous sucrose solution (specific.gravity 1.27). After stirring the pellet into the sucrose solution, the tube was again centrifuged at 1,500 rpm in a clinical centrifuge for 5 minutes. Additional sucrose solution was added to the centrifuge tube to form a meniscus at the top of the tube. A 22X22mm coverslip was added to top of the meniscus, and the eggs and oocysts were allowed to float to the top of the tube and adhere to the coverslip for 30-45 minutes. The coverslip was then removed from the tube and placed on a microscope slide. Parasites eggs were identified to the genus level when possible (i.e. Toxacara eggs, eggs, Strongy- loides eggs, and Dipylidium eggs) or to the family level when necessary (Taeni- idae or Ancylostomatoidea). Coccidian oocysts were not further identified to genus or family. Egg and oocyst numbers were reported according to the following intensity scoring system: #1 RARE = 1-2 Eggs/Oocysts/Slide; #2 FEW = 2-10 Eggs or Oocysts/Slide; #3 MODERATE = 10-20 Eggs or Oocysts/Slide; #4 MANY = 20-100 Eggs or Oocysts/Slide; and #5 NUMEROUS = More than 100 Eggs or Oocysts/Slides. Personnel from each of the shelters also provided the following information on each dog: approximate age and predominant breed of the dog, and the date the fecal sample was taken. The various dog breeds were assigned to the seven dog groups identified by the American Kennel Club (www.AKC.org). These groups include: Sporting Dogs (Group I), Hounds (Group II), Working Dogs (Group III), Terriers (Group IV), The Toys (Group V), Non-Sporting Dogs (Group VI), Hearding Dogs (Group VII), and Miscella- neous Dogs. Cochran-Mantel-Haenszel statistics were performed using SAS 6.12 to measure associations between host information (i.e. age of dog, the year and month samples were collected, city location of the animal shelters, and the American Kennel Club groups represented by each dog) and fecal- evaluation results.

RESULTS

The average age of the cats was 1.4 years, and the average age of the dogs was 2.15 years. Forty-one percent (238) of the dogs included in the study were from Group I (Sporting Dogs), and most of these dogs (165) were Labrador re- trievers or Lab-crosses. The remaining dogs were divided among the remaining groups according to the following: Group II (Hounds) - 4.1 %, Group III (Working Dogs) - 13.1%, Group IV (Terriers) - 8.1%, Group V (The Toys) - 3.5%, Group VI (Non-Sporting Dogs) - 5.4%, Group VII (Herding Dogs) - 21.7% and Miscellaneous Dogs - 3.1%. At least one parasite was identiﬁed in 35.2% of all of the cat samples (Table 1). Over 20% of the cats were shedding coccidian oocysts, and the infected 230 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002)

Table 1. Prevalence and Intensity of Intestinal Parasites in Dogs and Cats from South Dakota

Parasite Prevalence Intensity Prevalence Intensity Name in Dogs in Dogs in Cats in Cats

Parasitized 27.41% - 35.23% Ancylostomadae 7.41% 3.02 (1.32) 4.55% 3.50 (1.91) Toxocara 9.14% 3.61 (1.34) 13.64% 3.47 (1.39) Toxascaris 0.17% 3.00 - Trichuris 0.52% 1.00 (0) - Strongyloides 0.17% 3.00 - Taeniidae 5.17% 3.00 (1.53) 5.68% 3.60 (1.95) Dipylidium 0.17% 3.00 Coccidians 7.93% 3.30 (1.47) 21.59% 4.08 (1.38)

cat was releasing many oocysts (i.e. mean intensity score of 4.08). The most commonly identified helminth parasite was Toxacara cati (13.6%). The average infected cat was releasing a moderate number of these eggs (i.e. mean intensity score of 3.47). Five of the cats were shedding Taeniidae eggs; the average cat was shedding moderate to many eggs (mean intensity score of 3.60). Hookworm eggs (Family Ancylostomadae) were found in the least number of cats (i.e. only 4.6%). At least one parasite was identified in 27.4% of all the dog samples (Table 1). The most commonly identified gastrointestinal parasite was Toxacara canis (9.1%). A moderate number of eggs (i.e. mean intensity score of 3.61) were being excreted by the average infected dog. Almost 40% of the infected dogs were releasing 20-100 eggs (i.e. intensity score of MANY); 26.4% of the infected dogs were releasing numerous eggs (Fig. 1). One dog was infected with Toxascaris nematodes and had released a moderate number of eggs into the fecal sample. Coccidian oocysts were found in 7.9% of the dog samples. Over half of the infected dogs were excreting many or numerous numbers of oocysts, and the average intensity score was 3.30. Hookworm eggs were identified in 7.4% of the samples. The average intensity score was 3.02, but the egg numbers were evenly distributed within the 5 scoring categories (Fig. 1). Three dog fecal samples contained Trichuris eggs, and each sample had an intensity score of 1. A moderate number of Strongyloides eggs were found in one of the dogs. Dipylidium tapeworm eggs were also only identified in one dog; this dog was also excreting a moderate number of eggs (Table 1). A total of 30 dogs (5.2%) were infected with Taeniidae tapeworms. No apparent pattern existed to the distribution of the scoring intensities (Fig. 1), but the average intensity score was 3.00. The presence and intensity of Taeniidae infections did not correlate with the age of the dog or the year and month that the samples were taken (p> 0.1). Taeniidae infections were also not associat- Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 231

Figure 1. Intensity Distribution of Intestinal Parasites in Dogs from South Dakota. Rare or #1 = 1-2 eggs/oocysts per slide; Few or #2 = 2-10 eggs/oocysts per slide; Moderate or #3 = 10-20 eggs/oocysts per slide; Many or #4 = 20-100 eggs/oocysts/slide; and Nu- merous or #5 = more than 100 eggs/oocysts per slides. ed with the city location of the animal shelters. There was a general association between the American Kennel Club groups of dogs and the level of Taeni- idae infection (p<0.01). Dogs grouped as herding dogs (Group VII) represented 46.7% of the 30 infected dogs (prevalence of 11.1% for that group). Sporting dogs (Group 1) possessed 23.3% of the infected dogs, but the prevalence for this group was only 4.0%. None of the 31 non-sporting dogs were infected with Taeniidae tapeworms. The remaining 9 infected dogs were distributed among the remaining groups of dogs.

DISCUSSION

More than one-third of the cat fecal samples tested in the present study contained one or more parasite eggs/oocysts. Fortunately, only a few of these cat parasites are typically infectious to humans. One-fifth of the cats were shedding coccidian oocysts. Lindsay and Blagburn (1991) list 5 genera of coccidian parasites commonly found in domestic cats. Three of these genera contain species that are infectious to humans (i.e.Cryptosporidium parvum, Toxo- plasma gondii and Sarcocystis spp.). It is likely that some of the oocysts found in the cats from this South Dakota study were Toxoplasma and Sarcocystis oocysts, but the special diagnostic tests needed to reliably diagnose Cryp- tosporidium infections was not used in this study. It’s more likely that most of the oocysts were from the genus Isospora. Neither species of Isospora in- 232 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) fecting cats (I. felis and I. rivolta) are zoonotic, but they can cause gastroen- teritis in young and/or immunocompromised cats (Lindsay and Blagburn, 1991). Hammondia hammondi and the species of Besnoitia infecting cats are also not zoonotics, and only rarely cause intestinal problems in their feline host. The zoonotic significance of Toxocara cati is still being debated, but its importance to human heath is minor compared with T. canis (Roberts and Janovy, 2000) even though more that 10% of the cats from South Dakota were infected with this ascarid nematode. Only a few cats were shedding hookworm eggs. It is likely that any species of hookworm can cause cutaneous larval migrans, but Ancylostoma braziliense appears to be the most common agent (Schad, 1994). Most of the hookworm eggs seen in this study were probably Ancylostoma tubaeforme because A. braziliense is more restricted to the Gulf Coast region of the USA (Bowman, 1992). Very few Taeniidae eggs were found in cats from this study. By far, the most common cat taeniid tapeworm found throughout the world is the rat-cat tapeworm, Taenia taeniaeformis. This species is not infectious to humans, and is, very likely, the only species represented by the Taeniidae eggs present in the infected cats from this study. More than one-fourth of the dog fecal samples tested in the present study contained one or more parasite egg/oocysts. The most common and most zoonotically important parasite found in these dogs was T. canis. Toxocaria- sis is thought to be one of the most common human helminth infections, at least in the temperate climates of the world (Schantz, 1983). Roughly 98% of puppies and 20% of adult dogs in the United States are infected with T. canis (Roberts and Janovy, 2000). The recent national survey of dog parasites by Blagburn et al. (1996) found that 14.5% of the dogs were shedding T. canis eggs. Prevalence is obviously dependent on age, but results from the South Dakota study were similar to the national survey. The zoonotic potential of T. canis juveniles has been firmly documented, and this species is the primary agent for visceral larval migrans. Less than 10% of the dogs were shedding hookworm eggs. The majority of these eggs were a combination of A. caninum and U. stenocephala eggs, and both species can cause cutaneous larval migrans in humans. In the national survey, A. caninum was found in 19.2% of all the dogs tested in the survey, but its prevalence was strongly influenced by region, with 36.5% of the southeastern dogs being infected while only 2.6% of the western dogs were infected (Blagburn et al., 1996). Therefore, it is not surprising that the hookworm prevalence results from the present study were midway between that of the western dogs and the mid-western dogs. Only 1.0% of the dogs from the national study were infected with U. stenocephala, and this did not vary significantly with region. Therefore, the vast majority of hookworm eggs found in South Dakota dogs were likely A. caninum. Only 4 genera of coccidians are listed by Lindsay and Blagburn (1991) as parasitizing dogs. Since C. parvum was not included in this study, all of the oocysts found in the South Dakota dogs were members of either Isospora, Hammondia or Sarcocystis. Only the genus Sarcocystis has species that are zoonotic. Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 233

The prevalence of taeniid tapeworms were almost ten times higher in the present study from South Dakota dogs than from the national survey. Taeni- id infections originate in dogs from the consumption of infected mammalian intermediate hosts (e.g. lagamorphs, rodents or ungulates). Because the national survey focused on dogs from large cities (Lindsay and Blagburn, 1991), many of these dogs would likely have limited access to the intermediate hosts of taeniids. The higher prevalence of taeniid infections in the South Dakota dogs may have resulted from the fact that dogs from small cities and surrounding rural areas were used in this survey, and these dogs would likely have greater access to the intermediate hosts. This idea is supported by the predominant types of dogs included in the South Dakota study. Almost 63% of these dogs were classiﬁed either as sporting dogs or herding dogs. These breeds are very capable of catching and killing rabbits and rodents. Another potential explanation for the higher prevalence of taeniid infections in South Dakota dogs relates to the added possibility of E. multilocularis infections in this area but not nationally. This possibility illustrates the need for additional studies speciﬁcally evaluating the prevalence of E. multilocularis in dogs from this region. Recent developments in the creation of a coproantigen diagnostic test for E. multilocularis based upon dog fecal samples should greatly facilitate this type of survey (Deplazes et. al, 1999).

ACKNOWLEDGEMENTS

Special thanks are given to the personnel at the various animal shelters who provided access to the fecal samples. Additional thanks to Keith Mertz and Darlene Buschenfeld who provided technical assistance during this project. This project was supported in parts by Miles, Inc. and by the South Dako- ta Graduate Research Fund.

LITERATURE CITED

Blagburn, B.L., D.S. Lindsay, J.L. Vaughan, N.S. Rippey, J.C. Wright, R.C. Lynn, W.J. Kelch, G.C. Ritchie, and D.I. Hepler. 1996. Prevalence of canine parasites based on fecal ﬂotation. Compend. Contin. Educ. Pract. Vet. 18:483- 510. Bowman, D.D. 1992. Hookworm parasites of dogs and cats. Compend. Contin. Educ. Pract. Vet. 14:585-595. Bowman, D.D. 1995. Georgis’ Parasitology for Veterinarians. W.B. Saunders Company, Philadelphia. 294-295p. Crellin, J.R., A.A. Marchiondo, and F.L. Andersen. 1981. Comparison of suitability of dogs and cats as hosts of Echinococcus multilocularis. Am. J. Vet. Res. 42:1980-1981. Deplazes, P., P. Alther, I. Tanner, R.C.A. Thompson, and J. Eckert. 1999. Echinococcus multilocularis coproantigen detection by enzyme-linked im- munosorbent assay in fox, dog, and cat populations. J. Parasitol.:85(1):115- 121. 234 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002)

Hildreth, M.B., M.D. Johnson, and K.R. Kazacos. 1991. Echinococcus multilocularis: A zoonosis of increasing concern in the United States. Compend. Contin. Educ. Pract. Vet. 13:727-740. Hildreth, M.B., S. Sriram, B. Gottstein, M. Wilson, and P.M. Schantz. 2000. Fail- ure to identify alveolar echinococcosis in trappers from South Dakota in spite of high prevalence of Echinococcus multilocularis in wild canids. J. Parasitol. 86:75-77. Lindsay, D.S. and B.L. Blagburn. 1991. Coccidial parasites of cats and dogs. Compend. Contin. Educ. Pract. Vet. 13:759-765. Roberts, L.S. and J. Janovy 2000. Foundations of Parasitology. McGraw Hill Co. Inc., Boston. 405-431 Schad, G.A. 1994. Hookworms. Pets to humans. Ann. Intern. Med. 120:434-435. Schantz, P.M. Emergent or newly recognized parasitic zoonoses. Compend. Contin. Educ. Pract. Vet. 5:163-172. Abstracts of Senior Research Papers

presented at The 87th Annual Meeting

of the South Dakota Academy of Science

Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 237

GOOD WASPS GONE BAD: INVASIVE SPECIES IN A PARASITOID GUILD ON A NATIVE LEGUME

Paul J. Johnson and Arvid Boe Insect Research Collection Plant Science Department South Dakota State University Brookings, SD 57007

ABSTRACT

Canada milkvetch (Astragalus canadensis L.) is a native legume widespread through the coterminous United States and Canada. It is an important plant for herbivores and granivores in natural ecosystems. Our objectives were to ascertain the insects feeding upon milkvetch seeds, to describe the general life history features of these insects, and to determine frequencies of seed predation and primary parasitism. Two populations of Canada milkvetch were sampled in Brookings County, South Dakota. The first population was field grown at the SDSU agricultural farm in Brookings and the second was a wild population at the Oak Lake Field Station. Seed feeding beetles and their parasites were reared from bulk samples in 1998, and 783 segre- gated seedpods in 1999. All pods were dissected to count damaged pods and seeds, and to recover un-emerged seed predators and parasitoids. The primary seed predators of Canada milkvetch are the bruchid beetle Acanthoscelides perforatus (Horn)(Coleoptera: Bruchidae) and the weevil Ty- chius liljebladi Blatchley (Coleoptera: Curculionidae). These beetles appear to have similar niches. Adults of both species are active on their host plant at the same time. Larvae of both species feed on the developing seeds and occa- sionally within the same seedpod, with no clear evidence of spatial or temporal separation. Larvae of A. perforatus overwinter in the final instar inside the pods, whereas larvae of T. liljebladi exit the pods in late July and overwinter as pupae in soil. The combined frequency of pod occurrence by the beetles was 46% and 71% for Oak Lake and Brookings populations, respectively. Based on location, A. perforatus occurred in 20% of pods at Oak Lake and 54% of pods at Brookings, while T. liljebladi occurred in 26% of pods at Oak Lake and 17% of pods at Brookings. The primary parasitoids of A. perforatus are adventitious chalcid wasps (Hymenoptera), Dinarmus acutus Thomson (Pteromalidae) and Eupelmus vesicularis (Retzius) (Eupelmidae). The primary parasitoids on T. liljebladi are the native wasps Eurytoma tylodermatis Ashmead (Eurytomidae) and an apparently undescribed Mesopolobus sp. (Pteromalidae). Total mean parasitism rates from 783 seedpods were 10% for T. liljebladi and 22.5% for A. perforatus. Based on location, A. perforatus parasitism occurred in 20% of pods at Oak Lake and 25% of pods at Brookings, while T. liljebladi parasitism occurred in 4% of pods at Oak Lake and 16% of pods at Brookings. 238 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002)

Acanthoscelides perforatus is a native bruchid beetle found widely in the U.S. and Canada, and is apparently host specific to Canada milkvetch. No other bruchid beetles are known from Canada milkvetch. Dinarmus acutus was clearly the dominant parasitoid of A. perforatus. Eupelmus vesicularis was the only other wasp reared from this beetle during this study. Dinarmus acutus is a Eurasian species and was first reported from the U.S. in 1936 along the east coast. It was reported from Oregon in 1938, and then purposely distributed in Oregon during 1963 for the biological control of the European vetch bruchid, Bruchus brachialis Fabricius. This wasp is a bruchid specialist and was reared from the native Acanthoscelides aureolus on Glycyrrhiza lepidota and Acan- thoscelides submuticus on Amorpha fruticosa in the northern Great Plains. Na- tive parasitoids and D. acutus were observed from these latter bruchids. How- ever, no native parasitoids were reared from A. perforatus. Eupelmus vesicularis is also a European wasp first reported under a now synonymous name from Minnesota in 1916. This wasp was observed by us as a primary parasitoid of A. perforatus. It is also known as a hyperparasite of D. acutus, but this behavior was not observed by us. The absence of native parasitoids and the high incidence of parasitism by an exotic parasitoid on the native A. perforatus suggest a disruption of seed predator and primary parasitoid guild structure. Native parasitoids are reported for numerous native bruchid beetles, often more than one parasitoid per bruchid species. A prediction for one or more native parasitoids on A. perforatus is reasonable. However, the available data suggest that in the northern Great Plains the native A. perforatus parasitoids were displaced by the exotic D. acutus. Evidently, expanded surveys of A. perforatus and its parasitoids are needed to test our observations and determine the geographic extent of this native parasitoid displacement. Since D. acutus was deliberately released for the biological control of exotic bruchids on crop plants, at least in Oregon, our observations support contentions of others that past biological control programs presented collateral and unintended consequences to native insect communities and insect/plant relationships. Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 239

COMPARATIVE MORPHOLOGY OF SEVERAL SWITCHGRASS CULTIVARS

Martine Zamy and Arvid Boe South Dakota State University Brookings, SD 57007

ABSTRACT

Switchgrass is currently used for forage, revegetation, conservation, wildlife habitat, and beautification in the northern Great Plains. It is also being evaluated for its potential as a biomass/bioenergy crop in that and other regions of the United States and Canada. Breeding new switchgrass cultivars for one or more of the above purposes may involve selection for a particular growth habit or plant form/shape. We studied the morphology of several cultivars in new (3-year-old) and old (16-year-old) stands in eastern and central South Dakota to determine if there was a genetic basis for differences in stand (e.g., tillers/ft.2, ratio of vegetative to reproductive tillers) and individual tiller (e.g., number of leaves/tiller, leaf-to-stem ratio) characteristics that would make one cultivar superior to another for a particular purpose. We found consistent large differences among cultivars for tillers/ft.2, number of leaves/tiller, the percentage of tillers that produced seedheads, and the partitioning of dry matter between leaf and stem tissues. Our research resulted in detailed comparative descriptions of the stand and individual tiller characteristics of the cultivars Blackwell, Cave-In-Rock, Dacotah, Forestburg, Ne- braska 28, Pathfinder, Sunburst, Summer, and Trailblazer in a diversity of environments in the northern Great Plains. In addition, we also determined the effect of environmental variability on the expression of those morphological traits. The information we gathered from this study should be very useful for selecting a particular cultivar for a specific use. For example, if the need is to protect the soil from erosion and provide a dense stand for wildlife habitat, one might select a cultivar that produces many tillers/ft.2 and has a high percentage of tillers that produce seedheads. On the other hand, if the primary need is pasture, a cultivar that produces mostly vegetative tillers that have a high leaf- to-stem ratio may be the most desirable. The data should also be useful for selecting the appropriate germplasm as breeding material for developing new cultivars for specific purposes.

Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 241

POLYMORPHISM IN THE AGOUTI-RELATED PROTEIN (agrp) GENE IN SWINE

Kent J. Donelan, Nels H. Granholm and Carl A. Westby Department of Biology and Microbiology and Donald M. Marshall Department of Animal and Range Sciences South Dakota State University Brookings, SD 57007

ABSTRACT

The agouti-related protein (agrp) gene encodes a gene product that regu- lates components of energy homeostasis – appetite, food intake, body composition, physical activity, basal metabolic rate, and others. The protein encoded by agrp binds to melanocortin receptors (MC4-R) of hypothalamic neurons causing an increased appetite when it overwhelms or out-competes _-MSH, another naturally occurring and appetite-suppressing ligand of MC4-R. The purpose of this project was to isolate and sequence the porcine agrp from diverse breeds of pigs. Using the DNA nucleotide sequences of each breed-specific pig, we characterized different alleles or variations among the various breeds of pigs. Porcine agrp sequences were extended using thermal asymmetric interlaced (TAIL)- polymerase chain reaction (PCR). From this extended transcript, sequence specific primers were designed and standard PCR was used to amplify the agrp sequence. PCR products were purified and sequenced directly. Sixteen samples representing pure or various crossbreeds have been sequenced. The open reading frame (ORF) of porcine agrp consists of 978 nucleotides. Sequence data revealed multiple areas of nucleotide variation that may represent polymorphisms. Information provided by this study may be valuable to pork producers. By optimizing the genetic regulation of weight gain, carcass quality and other aspects of energy balance, these data may help to produce healthier pigs with higher quality meats. This project was supported by funds from the SDSU Agricultural Experiment Station – Project numbers SD186H, SD60H, and SD191H.

Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 243

TRANSGENE DISPERSAL AND EFFECTS IN FEED AND FOOD FROM ROUNDUP READY SOYBEANS

C. Carter, S. Hansen, R. Scott, B. Turnipseed, and L. Wrage Department of Plant Science

P. Krishnan Department of Nutrition and Food Science

R. Thaler Department of Animal and Range Sciences

T. Cheesbrough Department of Biology/Microbiology

South Dakota State University Brookings SD 57007

ABSTRACT

A modified EPSP synthase gene provides tolerance to the herbicide glyphosate in Roundup Ready soybeans. We analyzed the dispersal of this transgene from Roundup Ready plots to nearby non-transgenic soybeans, and examined the effects of the transgene and transgenic protein in food and feed. The transgene was detected by PCR in non-transgenic soybean plots at dis- tances of up to 18.5 ft from the Roundup Ready plots. The pattern of dispersal suggested that most field dispersal appeared to be due to carryover of Roundup Ready seed during planting or harvesting, rather than cross-pollina- tion. The Roundup Ready gene and protein were examined in flour, bread, soymilk and tofu made from Roundup Ready vs. non-transgenic soybeans. The transgene was detected by PCR in soy flour and in the interior of bread loaves. However, Strip Test (ELISA) results were negative for all bread samples, indicating that the transgenic protein was degraded during baking. In soymilk, the transgene and protein were detected before processing, but not after processing. No tofu samples were positive by either PCR or Strip Tests. There were no differences between Roundup Ready and non-Roundup Ready seeds in terms of in protein content, oil content, phenolics or minerals (ash). There were no differences in weight gain, growth or development of swine fed Roundup Ready vs. isogenic non-transgenic soybeans.

Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 245

SPECIES RICHNESS AND NESTING SUCCESS OF NEOTROPICAL MIGRANTS IN NATURAL RIVER CORRIDORS AND ANTHROPOGENIC WOODLANDS IN SOUTHEASTERN SOUTH DAKOTA

Dale Gentry Department of Biology South Dakota School of Mines and Technology Rapid City, SD 57701

David Swanson Department of Biology University of South Dakota Vermillion, SD 57069

ABSTRACT

Recent declines in Neotropical migratory species have been documented at breeding grounds in North America. These declines are attributed to a variety of factors including habitat fragmentation on wintering and breeding grounds, as well as loss of habitat necessary for migratory stopover. In addition to reducing available forested habitat for breeding, forest fragmentation is generally associated with decreased nesting success in woodland nesting birds. Most of the native nesting habitat in the northern Great Plains is found in riparian corridors. Additional woodland nesting habitat has been established within the last century as isolated shelterbelts and woodlots associated with farmsteads. Our study compares relative abundance, density, and nesting success between these two habitats. Nest searching and point counts were done during the summers of 2000 and 2001. One hundred niety-eight nests of 12 species were found in woodlots and 218 nests of 22 species were found in corridors. Comparison of nesting success between the two habitats of all species combined was not signiﬁ- cantly different. However, one species, the American Robin (Turdus migrato- rius) did show a signiﬁcantly higher nesting success in native riparian corridors compared to woodlots. Point counts conducted in each habitat showed that some native species were found only in corridors but all species found in woodlots were also found in corridors. These results suggest that woodlots do serve as suitable nesting habitat for species that use both habitats, but woodlots do not replace native woodland habitat because all native nesting species do not use them.

Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 247

ISOLATION AND PARTIAL CHARACTERIZATION OF THE HAPLIOD STAGE OF USTILAGO MAYDIS FROM SMUTTED EARS AND TASSELS OF FIELD CORN (ZEA MAYS) FROM SOUTHEASTERN SOUTH DAKOTA

Jodie Linneweber and Donna Hazelwood Dakota State University Madison, SD 57042

ABSTRACT

Ustilago maydis, causal agent of common corn smut, is a warm temperature wound pathogen that infects corn at all growth stages. From one field in Minnehaha County South Dakota, an increase in incidence and severity of infection in field corn (Zea mays) was reported growing seasons 1999, 2000, and 2001. The purpose of this study was to isolate the haploid state of U. maydis from field collected teliospores, to partially characterize the haploid isolates, and to determine if compatible mating types had been isolated. Ustilago maydis has two distinct growing phases. The first is a haploid, yeast-like form that reproduces by budding and is nonpathogenic on corn. The second phase, pathogenic on corn, is dikaryotic filamentous. Isolates of U. maydis collected from two fields in Minnehaha County, SD late summer 2001, were utilized to determine conditions for isolation of the haploid state. Sporulation was examined at temperature regimes ranging from 4 C to 34 C. The most rapid rate of sporulation occurred at 34 C. Isolation of the haploid state was accomplished by manipulating germinating teliospores and basidia to ensure that a dikaryon was not produced. Multiple subinoculations were performed. Inoculation of wounded ‘Early Sunglow’ sweet corn seedlings involved application of a liquid suspension of U. maydis. Isolates examined did not exhibit filamentous growth on PDA nor did they produce symptoms on in- oculated corn plants. We concluded that these isolates are haploid, but are not compatible mating types.

Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 249

GENE EXPRESSION PATTERNS IN SEGMENTING RADIALS OF ZEBRAFISH, DANIO RERIO

Patricia L. Crotwell and Paula M. Mabee Department of Biology University of South Dakota Vermillion, SD 57069

ABSTRACT

What is the evolutionary origin of the synovial joint? Although the vertebrate limb is a well-understood model system of comparative developmental genetics, the evolutionary origin of joints has never been studied. A conserved pattern of development involving a sequence of condensation, segmentation and synovial joint formation in regions of segmentation produce the characteristic skeletal branching pattern of vertebrates with limbs (tetrapods). Fishes also exhibit segmentation in skeletal development, in regions including gill arches and median and paired fin skeletons. Fish fin supports, or ‘radials,’ segment, yet do not have the joints that are characteristic of tetrapod limbs. We hypothesize that the genetic mechanisms underlying the formation of joints in tetrapod vertebrates arose from those involved in segmentation of fish fin skeletons. To test this hypothesis, we examined expression patterns of genes known to be critical in synovial joint development using in situ hybridization methods. In this method, labeled antisense RNA strands, or probes, were as- sembled and introduced to fixed zebrafish. A color label allowed us to visu- alize precisely where genes were being expressed in situ, or literally ‘on site.’ Our data indicated that similar to tetrapods, key joint-related genes such as Gdf5, Collagen II, Wnt14, and noggin were expressed in the developing median fin endoskeleton in regions of segmentation.

Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 251

RELATIONSHIPS AMONG IKONOS SATELLITE IMAGERY, AIRBORNE SCANNING LIDAR DATA, AND GROUND-BASED TREE INVENTORY DATA IN A PONDEROSA PINE FOREST VIA SUBPIXEL INTERPRETATION

X. Chen, L. Vierling, D. Dykstra, and W. Capehart South Dakota School of Mines and Technology Institute of Atmospheric Sciences Rapid City, SD 57701

E. Rowell Horizon’s Incorporated Rapid City, SD 57701

T. DeFelice Raytheon Corporation USGS EROS Data Center Sioux Falls, SD

ABSTRACT

Structural and functional analyses of ecosystems require high accuracy information of component fractions within an ecosystem, especially the vegetation components. Spectral unmixing is a technique to estimate the fractions of vegetation and other ground components that occur within remote sensing image pixels, thereby improving spatial mapping capabilities. The purpose of this project is to research the performance of spectral unmixing using IKONOS images, and to compare these results to laser altimetry (LIDAR) and ground measures of forest structure. In the summer of 2001, ground data were collected along 10x150 meter belt transects radiating from the Black Hills Ameriflux tower in the standard eight directions. These data included tree species identification, diameter at breast height, tree height, percent and type of ground cover, and effective leaf area index (LAIe). IKONOS imagery was acquired over the site in July 2000. Air- borne scanning LIDAR data was acquired in October 2001. No thinning or fire activity occurred at the site between data collection dates. The comparison of IKONOS unmixing, LIDAR data, and ground data showed a significant positive correlation between LAIe and the IKONOS-derived tree/shade fraction. LAIe and the open grass endmember fraction were significantly negatively correlated. The Enhanced Vegetation Index (EVI) was negatively correlated with all measures of tree density, including the canopy LIDAR return fraction and tree/shade endmember fraction. Conversely, EVI exhibited a very strong positive correlation with the open grass endmember frac- 252 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) tion. These results indicate that IKONOS data can serve to make the important distinction between tree canopy coverage and exposed understory background, which contains grass or bare soil, near peak summertime greenness. Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 253

SEASONAL VARIATION ON BROWN TROUT DIET IN RAPID CREEK, SOUTH DAKOTA

S. Martin Digler and Kerri T. Vierling South Dakota School Of Mines and Technology Rapid City, SD 57701

ABSTRACT

Brown trout (Salmo trutto) in Rapid Creek, South Dakota have been noted to be in poor condition, and this condition may in part be related to factors affecting food availability. The objective of this study was to examine how seasons (fall vs. summer) affect brown trout diet in Rapid Creek. In each season, adult brown trout were collected in three 100m sites along Rapid Creek: one site was upstream of Rapid City, one site was in the middle of Rapid City, and one site was downstream of Rapid City. In October 2000, 60 fish from each site were captured using electrofishing techniques; their stomachs were pumped and stomach contents were preserved in 70% ethanol. Stomach sample contents were identified to the order level (and to family level if possible), were dried at 45 degree Celsius and weighed. The same procedure was followed in June 2001. A total of 360 stomach samples were analyzed (180 from each season). Our analysis indicates that trout ate more in the summer as compared to the fall (34.2g vs. 20.3g). Samples identified from the stomach of brown trout included Diptera (flies and midges), Ephemeroptera (mayflies), Osteichthyes, Hemiptera, Coleoptera and Gastropoda. Among these, the Ephemeroptera and Diptera were the major diet components during both seasons. The results of this suggest that seasons can influence trout diet, and potentially affect the condition of fish. These data will provide information that will be helpful in analyzing the current poor condition of brown trout in Rapid Creek.

Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 255

URBAN GRADIENT INFLUENCE ON BROWN TROUT DIET IN RAPID CREEK, SOUTH DAKOTA

Irene Cherni and Kerri T. Vierling South Dakota School of Mines and Technology Rapid City, SD 57701

ABSTRACT

Brown trout (Salmo trutta) in Rapid Creek, South Dakota have been noted to be in poor condition. As Rapid Creek travels through Rapid City, it is subjected to a variety of industrial activities that may negatively alter the water quality and fishery habitat. The objective of this study was to examine the influence of the urban gradient on brown trout diet. There were three 100m sites (reaches) designated for the purpose of this study: upstream, midstream, and downstream. Sixty adult brown trout were captured from each reach using electrofishing techniques; their stomachs were pumped and the stomach contents were preserved in 70% ethanol. Stomach sample contents were examined for the presence of invertebrates, which were later identified to the order level (and in some cases to the family level if possible), dried at 45ºC and weighed. The urban gradient influenced the diet of brown trout. However, the results were not as consistent as speculated. The presence of pollution tol- erant species in Ephemeroptera (mayflies) and Diptera (flies and midges) were dominant diet components for fish in all three sites and not just downstream where the water would have been in the poorest condition. No invertebrate indicators of good water quality (i.e. Plecoptera) have been found in any of the three reaches. The results of this study will be helpful in analyzing the brown trout condition as well as the overall status of Rapid Creek.

Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 257

AN EVALUATION OF THE EFFECTS OF FORMALIN AND HYDROGEN PEROXIDE TREATMENTS ON THE EXTERIOR EGG MEMBRANE OF CHINOOK SALMON (ONCHORHYNCUS TSHAWYTSCHA)

Hans Stephenson and Mark Gabel Biology Department Black Hills State University Spearﬁsh, SD 57799

Mike Barnes McNenny State Hatchery Department of Game, Fish and Parks Spearﬁsh, SD 57799

ABSTRACT

The presence of bacteria and fungi in hatchery water stocks can have se- rious effects on the rearing of Chinook salmon (Onchorhyncus tshawytscha) eggs. Antimicrobial chemical treatments evaluated in this test were formalin and hydrogen peroxide. In an attempt to understand the unknown mechanisms by which these chemicals increase egg survival, we have begun analyses using scanning electron microscopy to determine microbe levels and changes in characteristics of the external egg membrane. Initial observations indicate that formalin treated eggs have fewer microbes attached to the external egg membrane compared to peroxide and untreated control eggs. Peroxide treatments produced variable results, but were superior to no treatment. Pore plugs in the external egg membrane exhibited a variable degree of degradation that may be related to chemical treatments.

Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 259

A MORE BIODEGRADABLE QUATERNARY AMMONIUM COMPOUND

Keith Hartman and Gary Earl Augustana College Sioux Falls, SD

ABSTRACT

The purpose of this research is to develop a more biodegradable surfac- tant, through a single pot synthesis. It is also hoped that the reactants used will be safer for the industrial worker. This will be accomplished through increased ether chain content on a polyglycoside. Glucopon 625, an alkylpolyg- lycoside with a 12-carbon fatty chain on carbon-6, was alkoxylated and reacted with butylene oxide. This resulted in long ether chains originating on the secondary alcohols. The product was extremely basic when put into a water solution, indicating that the deprotonated oxygens were still present. In the next step, the alkoxylated product was reacted with epichlorohydrin, producing an intact epoxide. Finally, a tertiary amine was reacted with the epoxide to form the desired quaternary compound. It is anticipated that the greater oxygen concentration increases the biodegradable of the product. Many of the above procedures and ratios were varied in an attempt to ﬁnd the optimum amount of lipid-like groups, ether chain lengths, and amine ratios.

Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 261

INITIAL EXAMINATION OF POSSIBLE NOVEL MACROCYCLES VIA CAChe

Ethan Englund, Arlen Viste, and Jetty Duffy-Matzner Department of Chemistry Augustana College Sioux Falls, SD 57197

ABSTRACT

There are numerous cyclic compounds that are biologically active through the trapping and transport of sodium/potassium ions. Using CAChe, we have examined well known biologically active macrocycles in an attempt to design an ideal novel macromolecule derived from a,b unsaturated carbaldehydes as monomers for such macrocycles. Nonactin and crown ethers were examined and compared to our proposed structures. Using MOPAC (the Molecular Or- bital Package) in CAChe, the ideal geometries and thermodynamic factors were determined via semi empirical and empirical calculations. Variables included size of the macromolecules, cavity dimensions utilized to trap metal ions, and diverse substituted carbaldehyde monomers. (Fujitsu 2002)

LITERATURE CITED

Fujitsu. 2002. CAChe 5.0. http://www.cachesoftware.com/

Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 263

SYNTHESIS OF α,β-UNSATURATED CARBALDEHYDES, KETONES AND 1,4 ADDITION TO α,β-UNSATURATED ALDEHYDES

Jerrid Kruse, Andrea Votroubek, Jetty Duffy-Maztner Department of Chemistry Augustana College Sioux Falls, SD 57197

ABSTRACT

Our group is examining the formation of novel macrocycles using innova- tive dihydrofuran carbaldehydes as building blocks. Various sodium alkoxides underwent Michael Addition to nitroalkenes. The nitroalkene initially used was nitrostyrene made via a Henry Reaction combining benzaldehyde and ni- tromethane in ammonium acetate and acetic acid under reﬂux. The nitrostyrene was added to the sodium alkoxide of propargyl alcohol through a Michael Addition forming 2-[2-propyn-1-oxy]-1-nitro-2-phenylethane. This Michael Adduct underwent an Intramolecular Silyl Oleﬁn Cycloaddition to form the dihydrofuran carbaldehyde 2,5-dihydro-2-phenyl furan-4-caraldehyde. The use of other sodium alkoxides combined with various nitroalkenes to form α,β-unsaturated carbalehydes with different substituents will also be examined. These varied substituents could lead to the synthesis of dihydrofuran ketones as well. These caraldehydes and ketones will potentially be used to form macrocyclic polyesters with possible antibiotic properties.

LITERATURE CITED

Gairaurd, C.B. and Lappin, G.R. J. Org. Chem. 1953, 18,1. Dr. Jetty Duffy-Matzner’s Graduate Thesis. Wade, L.G. Organic Chemistry 3rd Ed. Prentice Hall, New Jersey, 1995.

Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 265

RIDGE DETECTION USING ARTIFICIAL NEURAL NETWORKS

Rishi Kishore and John M. Weiss Department of Mathematics and Computer Science South Dakota School of Mines and Technology Rapid City, SD 57701

ABSTRACT

Ridge detection in digital imagery, unlike edge detection, is an area that has received relatively little attention. Ridges are long, narrow structures that are perceptibly brighter than the background. Examples of ridges include roads in aerial images and vessels in medical images. The ability to rapidly and accurately label ridge pixels in a digital image is an important first step in many computer vision applications. Artificial neural networks (ANNs) have proved useful in various pattern recognition tasks. In this work, we investigated the effectiveness of ANNs for ridge detection. We trained a three-layer feed-forward backpropagation network to detect ridges in digital images. The input and the hidden layers had the same number of perceptrons (roughly equal to the maximum ridge width), while the output layer had a single perceptron. The input training set consist- ed of digital images with strong ridge content (scenes of lightning strikes, containing ridges of varying width and intensity). The input vector was composed of normalized pixel intensity levels, sampled from a small neighborhood about each pixel. The output data set in this supervised training approach was the normalized intensity output of a more traditional ridge detector. The ridge detection results we obtained using ANNs were equal and in some ways superior to the traditional ridge detector. The traditional ridge detector worked well on thinner ridges, but was not as effective in detecting wider ridges. The ANN-based ridge detector appeared to "generalize" well enough to detect ridges of varying width and brightness, giving it a significant advantage over more traditional approaches.

Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 267

THE EFFECTS OF HYPERBARIC ENVIRONMENT-INDUCED pH CHANGES ON THE GENERATION TIMES OF PSEUDOMONAS AERUGINOSA

Contessa E. Creager, Sarah O. Hammar, Mitchell W. Jacobs, John Wayne Kalda, Spring J. Mayer, Brandon J. McCall, Jenny M. Volkert and William J. Soefﬁng Natural Science Area University of Sioux Falls Sioux Falls, SD 57105

Keywords

Carbon dioxide, Generation time, Hyperbaric, pH, Population dynamics, Pseudomonas aeruginosa

ABSTRACT

Contingent to continuing investigations regarding the chemotherapy and characterization of otitis induced by exposures to hyperbaric environments, this study examined the changes in generation times of Pseudomonas aeruginosa subcultured in pH-adjusted Mueller-Hinton broth media under normobaric (101 kPa) and hyperbaric (161 kPa) conditions. Growth media were adjusted to three pH levels, 6.9, 7.0 and 7.1, using a 0.02 N NaOH solution. Standard plate count method was used to evaluate population increases during the exponen- tial growth phase of cultures raised under the experimental conditions. Results were used to calculate generation times for each experimental condition. Mean generation times were subsequently compared in the evaluation of the inﬂu- ence that each microenvironment had on the capacity for the populations to expand. The shortest mean generation time (48.0 minutes) for cultures raised in normobaric environments occurred in media adjusted to pH 7.0. Corre- spondingly, the shortest mean generation time (54.7 minutes) for cultures raised under hyperbaric conditions occurred in media adjusted to pH 7.1. Gas absorption studies, previously and concomitantly conducted with the aforementioned population dynamics study, indicate that the pH of culture medium decreases by 0.1 when exposed to the experimental hyperbaric conditions. The high partial pressure of carbon dioxide (488 Pa) occurring in the experimental hyperbaric environment is believed to be the major contributor to this change in pH. Subsequently, mathematical analysis of the hyperbaric and normobaric generation time trends indicate a reproducible and corresponding relationship that suggests that the hyperbaric microenvironment produces generation time trends similar, but not equivalent, to those observed in normobaric 268 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) environments. The correlation of these trends is most signiﬁcant when the pH shift of the media used in hyperbaric experimental conditions is considered; whereby, normobaric pH values of 6.9, 7.0, and 7.1 are changed to 6.8, 6.9, and 7.0, respectively. Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 269

OBSERVATIONS ON THE COLONIAL MORPHOLOGIES, CELL-CELL ADHESION CHARACTERISTICS AND ANTIBIOTIC SENSITIVITY OF PSEUDOMONAS AERUGINOSA IN A HYPERBARIC ENVIRONMENT

Brandon J. McCall, Sarah O. Hammar, Nicolas L. Strasser and William J. Soefﬁng Natural Science Area University of Sioux Falls Sioux Falls, SD 57105

Keywords

Antibiotic sensitivity, Cell-cell adhesion, Gentamicin sulfate, Hyperbaric, MIC testing, Morphology, Oxygen, Pseudomonas aeruginosa

ABSTRACT

Consequential to efficacy testing of gentamicin sulfate on Pseudomonas aeruginosa conducted in manned-undersea habitats, changes in colonial morphologies, cell-cell adhesion characteristics, and antibiotic sensitivity were observed. Standardized minimum inhibitory concentration (MIC) testing was conducted on P. aeruginosa under hyperbaric (161 kPa) incubation conditions using a concentration gradient of gentamicin ranging from 0.5 µg/ml to 4.0 µg/ml by two-fold concentration intervals. Bacteria were cultivated in 96- well culture plates and incubated at 37ºC in a humidified hyperbaric air environment, where the atmospheric concentration of oxygen varied from 34% to 35% s.e. (≈34 kPa). Positive, negative and contamination controls were conducted with each experimental sample. The sensitivity of standardized MIC testing methods to elevated carbon dioxide concentrations (488 Pa) has already been reported by S.O. Hammar, et al. (Proc. SD Acad. Sci. 80:441-442, 2001). Observations made during the aforementioned study and this study suggest that discoveries made by M.A. Kenward, et al. (Microbios 28:47-60, 1980) and B.P. Berdal, et al. (Acta Path. Microbiol. Immunol. Scand. Sect. B, 93:157-158, 1985) on the changes in colonial morphologies and adhesion characteristics of P. aeruginosa may be occurring at much lower oxygen concentrations and ambient pressures. In the hyperbaric incubation conditions of this study, large, convex, mucoid colonies, termed "giant colonies" by Kenward et al., were observed in stock cultures growing on nutrient agar plates. Additionally, a significant increase in cell clumping at the air-media interface was observed, suggesting an increased cell-cell adhesion characteristic had developed as a result of exposure to the hyperbaric environment. An increased sensitivity of 270 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002)

P. aeruginosa to gentamicin was observed. Bacteriostasis was achieved with a 2 µg/ml and 4 µg/ml gentamicin concentration under the hyperbaric experimental conditions and normobaric control conditions, respectively. When the experimental cultures were transferred to and further incubated at normobaric conditions (101 kPa), bacterial growth rapidly replaced the bacteriostasis previously observed; whereas, experimental samples continuously incubated in hyperbaric incubation conditions maintained the bacteriostasis. The enhancement of the aforementioned characteristics by hyperbaric environments has some relevance to the persistence of and chemotherapy of otitis externa. Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 271

NUTRITIONAL VALUES OF PSORALEA ESCULENTA

Giani Perera and Neil Reese Dept. of Biology and Microbiology South Dakota State University Brookings, SD 57007

ABSTRACT

Psoralea esculenta (Indian breadroot, Tipsin, Scurfpea, Prairie turnip, Timpsula) is a member of the family Fabaceae. It is a perennial herb that flow- ers in May to early July. Indigenous to the Great Plains, it can be found growing on foothills, bluffs, upland and midland prairies and sometimes along roadsides. The prairie turnip has a deep taproot that is swollen in the middle and tapering at the ends. This tuber is used in a manner similar to a potato, but differs in taste and texture. The edible portion is covered with a coarse brown husk. Prairie turnips are valued by Native Americans as an important traditional food, and were and are a staple food source for Native Americans. They are gathered during flowering and the tuber can be eaten raw, dried and used in soups, or ground into a high quality flour. Prairie turnips were obtained from prairies in Brookings Co. and Shannon Co., SD. Turnips were ground using Pulverisette 19. with a 6 mm screen, followed by grinding in a Cyclotec 1093 Sample Mill (1 mm screen). This powder was tested for - proteins, fat, fiber, calcium, copper, iron, magnesium, manganese, phosphorus, potassium, selenium, sodium, starch, zinc, ash, moisture and nitrogen free extracts. The analyses showed the roots contain 53-70% starch and about 7.5% protein on a dry weight basis. Psoralea is also a good source of iron, zinc and magnesium. Available selenium varied by location, with the East River plants having <0.05 ppm Se, but the West River plants having almost 0.1 ppm Se, on a dry weight basis.

Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 273

EXTRACTION, PURIFICATION, AND IDENTIFICATION OF UNIQUE PROTEIN IN ROOT OF (PSORALEA ESCULENTA)

Marcus Braaten and R. Neil Reese South Dakota State University Brookings, SD 57007

ABSTRACT

Plains Indians have been known to harvest the taproot of Psoralea esculenta as a dietary staple. Nutritional analysis has shown protein levels higher than average for common root crops. The edible portion of the root was milled to a ﬁne powder, extracted in a Tris buffer (pH 7.6), and the protein molecular weight was determined by electrophoresis, providing an approximation weight of 28.8 kDa. Isoelectric focusing demonstrated that the major protein component has a pI of approximately 6.0. Ion exchange chromatography and gel permeation were used to purify the protein, and the product identity was conﬁrmed by electrophoresis. Preliminary analyses indicate that this protein contains all of the essential amino acids but, as with most legume proteins, me- thionine content is rather low.

Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 275

EFFECTS OF LETHAL YELLOW AND MAHOGANY MUTATIONS ON REPRODUCTION IN FEMALE MICE

Trynda N. Oberg and Nels H. Granholm Department of Biology and Microbiology South Dakota State University Brookings, SD 57007

Maureen R. Diggins Department of Biology Augustana College Sioux Falls, SD

ABSTRACT

In previous studies, black (a/a) and yellow (Ay/a) mice were found to significantly differ in their reproductive capabilities. The presence of the lethal yellow gene (Ay) increases the production of agouti protein, which has long been associated with obesity and infertility, in the yellow mice. A mutation (mahogany or mg) has been found to suppress the effects of obesity in yellow mice. The objective of this study was to determine the effects of this mutation on fertility in female mice. We wanted to determine if the infertility of the yellow mouse could be reversed by the presence of the mg mutation. Overall, three experiments were conducted (qualitative histological analysis of ovaries, ovulation rates of prepubertal mice, and progesterone assays) using four genotypic combinations of Ay and mg: +/+ a/a (black), +/+ Ay/a (yellow), mg/mg a/a (mahogany black), mg/mg Ay/a (mahogany yellow). Fol- lowing superovulation, histological analysis of the ovaries of mice aged 25-26 days suggested that mg enhances follicular development (ovary function estimate) in control but not superovulated mice. Superovulation of 48 females 25- 26 days old indicated significant difference between black females and the other three genotypes. Analysis of serum progesterone levels (ELISA, ALPCO Di- agnostics) showed that the mg mutation significantly lowered progesterone levels in black and yellow females (P<0.05) compared to +/+ black and yellow females on days five and 12 of placement with a male. Time to copulation, total progeny per litter, progeny weight, and progeny survival (alive on day 30) were also measured as reproductive parameters. Analyses of all four compound genotypes of mice indicate that reproduction in females is affected by the mg mutation as well as the Ay mutation. This project was supported by funds from the SDSU Agricultural Experiment Station – Project numbers SD-60H and SD-191H (NHG/TNO), Biology Department of Augustana College (MRD), and the FOE’s Ehrmann Cancer Fund (NHG).

Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 277

DETERMINATION OF INFECTIVE GASTROINTESTINAL NEMATODE JUVENILES ON SPRING PASTURES

K. J. Mertz, M.B. Hildreth, and W.B. Epperson Departments of Biology/Microbiology and Veterinary Sciences South Dakota State University Brookings, SD 57007

ABSTRACT

To alleviate economic loss associated with gastrointestinal nematodes, spring strategic deworming programs are becoming a popular practice among many cattle producers. The advent of persistent anthelmintics has made it possible for producers to effectively control emerging infective juveniles on pasture. The premise of strategic deworming programs is to synchronize the application of persistent anthelmintics with the parasite’s life cycle to diminish future generations of nematodes. However, recommendations on timing the application of long lasting anthelmintics within a given geographic region have been based primarily on theoretical considerations. To maximize the use of anthelmintics and reduce the number of internal parasites on pasture, strategic deworming programs must be synchronized to coincide not only with the emergence, but also the decline in ﬁrst generation infective juveniles on pasture. The objective of this study was to determine the availability of over-wintering infective gastrointestinal nematode juveniles on pasture, in relation to spring strategic deworming practices. A pasture, divided into 12 paddocks, was "seeded" with gastrointestinal nematode eggs by daily rotating eight year- ling replacement heifers, with known levels of subclinical nematodiasis, through the paddocks. Infective juveniles were allowed to develop from the subsequent eggs and over-winter on paddocks until the following spring. The following spring, four groups of 15 fall born calves were placed on designated paddocks at two-week intervals starting on May 31, and ending on July 27. The calves were removed from designated paddocks after two weeks and placed in a controlled non-grazing environment for 21 days. Measurement of juvenile levels on pasture was performed through analysis of individual fecal samples obtained from the calves 21 days after removal from paddocks. The arithmetic average eggs/gram declined from 22.86 to 8.52 from period one to period two. The average eggs/gram for periods three and four were 3.13 and 0.40 eggs/gram respectively. Geometric transformation, to control for aggre- gation of parasite loads in host populations, revealed a linear decline over the four time periods (p=0.0001, R2=0.8163). The decline in eggs/gram from grazing periods 1 to 2 and throughout the rest of the grazing season is indicative of infective juvenile decline on summer pastures.

Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 279

DETERMINING THE FEASIBILITY OF RESTORING RIVER OTTERS IN SOUTH DAKOTA

Alyssa M. Kiesow and Charles Dieter Department of Biology South Dakota State University Brookings, SD 57007

ABSTRACT

Currently river otters occupy half their historic range in the United States and Canada. The status of the river otter in South Dakota is unknown; it is a state-threatened species. The purpose of this study was to determine the current status of river otters in South Dakota and to evaluate state river systems to ascertain the feasibility of restoring river otters in South Dakota. The sampled rivers were selected by buffering specific features, such as stream ranks 3-7, low gradient, and permanent water flow, using South Dakota GAP stream reach and watershed data. Study sites were selected along each river according to beaver activity, habitat quality (e.g. riparian areas), and accessibility. At each study site a riparian habitat transect was conducted recording floral and faunal species and a water sample was collected analyzing overall water quality. State employees and public conservation/management groups were contacted for further information on river otter sightings, habitat suitability, and restoration effort viewpoints. Rivers were rated (1=poor to 5=excellent) based on stream characteristics, watershed features, water quality, logistical factors, and prey availability. The five highest rated river systems in South Dakota are the Big Sioux R. (91), Little White R. (90), James R. (87), Bad R. (86), and North Fork Whetstone R (86). For a successful river otter restoration in South Dako- ta, recommend releasing river otters in the five highest rated rivers in South Dakota.

Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 281

COMPARATIVE ANALYSIS OF WING BEAT FREQUENCY AND EYE SPAN IN STALK-EYED FLIES (DIPTERA: DIOPSIDAE)

John G. Swallow University of South Dakota Vermillion, SD

Jilian M. Fazio and Gerald S. Wilkinson University of Maryland College Park, MD

ABSTRACT

Sexual selection often results in exaggeration of morphological and behavioral characters when individuals of one sex experience a mating advantage through female choice or male competition. Stalk-eyed flies have eyes placed laterally from the head on elongated peduncles. Distance from eye to eye can be up to 3-fold longer than the total body length. Sexual dimorphism in eyestalk length has arisen several times in the family, leading to dramatic variation between species and sexes. Among sexually-dimorphic species, evolution of increased eye span resulted from both female choice and male competition. Those interested in aerodynamic adaptation of flight have looked to these secondary sexual characters for trade-offs between exaggerated male traits and fit- ness. Costs generated by elongation of eyestalks could create selectively important variation in flight performance limiting further elaboration. Recent analyses have shown that eyestalk elongation resulted in reduced in aerial performance and compensatory changes in body dimensions (Swallow et al. 2000). We undertook a comparative analysis of wing beat frequency and eye span to investigate potential biomechanical trade-offs. The 14 species measured varied in average eye span (range 2.0 – 14.2 mm) and degree of sexual dimorphism (male – female eye span; range = -0.44 – 7.84 mm). Wing beat frequencies were measured using OFIDIS (Qubit Systems). Morphological variables measured were eye span, body length, wing length, and thorax width. Data were analyzed using phylogenetically independent contrasts. Across species, wing length was inversely related to wing beat frequency (r2 = 0.59; P < 0.001). Within species, eye-span dimorphism was inversely related to wing beat frequency dimorphism (r2 = 0.61; P < 0.001); increased eye span, relative to females, resulted in reduced wing beat frequency.

Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 283

LATE CRETACEOUS DINOSAURS, BIG BEND NATIONAL PARK, TEXAS

Julia T. Sankey Department of Geology and Geography Vassar College Poughkeepsie, NY 12604

ABSTRACT

Dinosaurs are not as well known from southern North America compared to more northern areas such as Alberta. Big Bend National Park, Texas has one of the southernmost terrestrial records for the Late Cretaceous in North America. Work in Big Bend provides an important and new geographic perspective on dinosaur diversity during the ﬁnal ten million years of the Creta- ceous from southern North America, leading up to the mass extinctions at the Cretaceous/Tertiary (K/T) boundary. Dinosaur teeth were collected from microfossil sites from the Late Cretaceous upper Aguja and lower Tornillo formations, spanning the late Campanian to late Maastrichtian (~74 to 67 Ma). In addition to detailed descriptions and illustrations of the dinosaur teeth, measurements and bivariate plots quantify their variation, allowing comparison with well-studied collections from similar-aged deposits. Taxa include herbivorous dinosaurs: pachycephalosaurid (dome-head), ceratopsid (horned), ankylosaur (armored), hadrosaurid (duck-billed) and carnivorous dinosaurs: tyran- nosaurid, Saurornitholestes cf. S. langstoni, Saurornitholestes sp., Richar- doestesia cf. R. gilmorei, R. isosceles, and "Paronychodon". Most taxa are represented by teeth from hatchlings or juveniles, demonstrating that these dinosaurs nested in the area. "Paronychodon" and Saurornitholestes sp. are rare and occur only in the Tornillo Formation, initial evidence that the Maastrichtian and Campanian theropod faunas in Big Bend were distinct, a similar pattern in northern assemblages. Absent from both the Campanian and Maastrichtian in Big Bend are Dromaeosaurus and Troodon, common in northern areas. More research is needed to conﬁrm this pattern and to better document the dinosaur diversity in the Big Bend area.

Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 285

VERTEBRATE PALEOECOLOGY FROM THE LATE CRETACEOUS HELL CREEK FORMATION, NORTH DAKOTA: SURFACE COLLECTION VERSUS SCREEN WASHING

Georgia E. Knauss and Andrew A. Farke Museum of Geology and Depat. of Geology and Geological Engineering South Dakota School of Mines and Technology Rapid City, SD 57071

ABSTRACT

Geologists and paleontologists have extensively studied the Late Creta- ceous Hell Creek Formation of southwestern North Dakota for the past ﬁfteen years. This work has produced a high-resolution stratigraphic framework in which numerous fossil sites have been tied to the K/T (Cretaceous/Tertiary) boundary. Microfossil sites consisting of teeth and small bone accumulations have been surface collected by the Pioneer Trails Regional Museum (PTRM). However, sole reliance on this collection method may bias the fossil sample. This study compared fossil collections produced by surface collection and screen washing of bulk sediment. It was hypothesized that each method would collect different elements of the fauna. For this study two sites were evaluated: PTRM V86002 and PTRM V89003. PTRM V86002, produced 1,180 specimens in eight seasons of surface collection. Screen washing of 167 kg of sediment has produced 489 specimens to date. PTRM V89003, produced 715 specimens in eight seasons of surface collection. Screen washing of 140 kg of sediment has produced 1,642 specimens to date. In both sites preliminary analysis showed marked disparity of faunal proportions between the two collecting methods. For instance, the sharks were 17 percent of the screen washed specimens in PTRM V89003, while they represented less than one percent of the surface collected specimens. Dinosaurs were less than one percent of the screen washed specimens in PTRM V89003 and 24 percent of the surface collected specimens. These differences could result from collection bias, erosional bias, or unknown factors. Reliance on one method of collection could severely bias paleoeco- logical interpretations.

Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 287

MORPHOLOGY OF THE CORNUAL SINUSES IN CHASMOSAURINE HORNED DINOSAURS (ORNITHISCHIA: CERATOPSIDAE)

Andrew A. Farke Museum of Geology and Dept. of Geology and Geological Engineering South Dakota School of Mines and Technology Rapid City, SD 57701

ABSTRACT

The ceratopsids, or horned dinosaurs, are well-known for their bizarre cranial ornamentation consisting of horns, frills, and knobs of bone. While the external features of the ceratopsid skull are well-described, many aspects of their internal cranial anatomy are poorly known. Hollow cavities called cornual sinuses occupy the basal portion of the postorbital, or brow, horns in many members of the subfamily Chasmosaurinae. The sinuses are absent in Chas- mosaurus, relatively small in Anchiceratops, and variably developed in Tricer- atops, Pentaceratops, and Torosaurus. Their presence or absence is not known in Arrhinoceratops. The cornual sinus cavities of most chasmosaurines have not been examined, so they were described as a ﬁrst step towards understanding their function and evolution. The cornual sinuses are relatively simple structures in chasmosaurines, consisting of an ovate cavity that extends into the postorbital horncore. These sinuses are domed distally, and small folds of bone may be present on the sinus cavity walls. The bone enclosing the cornual sinus cavity is extensively vascularized in some specimens. The function of the sinuses is problematic, but it is possible that they eliminated structurally un- necessary bone from the horncore, thereby reducing skull mass.

Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 289

MICROVERTEBRATE FAUNA RECOVERED FROM THE HELL CREEK FORMATION (LATE CRETACEOUS), SOUTHWESTERN NORTH DAKOTA

Marcus Ross and Tim Pranger Department of Geology and Geological Engineering and Museum of Geology South Dakota School of Mines and Technology Rapid City, SD 57701

ABSTRACT

The Hell Creek Formation is a Late Cretaceous fluvial system approximately 67 million years old. Three microfossil sites from the Hell Creek For- mation were sampled in conjunction with the Pioneer Trails Regional Museum (PTRM V86002, V88018, and V89003). We conducted this study in order to dis- cover the vertebrate fauna represented by microvertebrate fossils and in order to determine the richness of fossils per site. This was determined by the number of identifiable vertebrate fossil remains per unit weight of sediment. Sed- iment was collected and screen washed with fine-mesh screen to remove the fine-grained particles. Fossils were picked out of the screened material and identified. Vertebrate taxa recovered from the sites include elasmobranchs, ba- toids, teleost fish (including lepisosteids, amiids, and perciforms), salamanders, frogs, lizards, crocodilians, dinosaurs (theropods, hadrosaurs, and ceratop- sians), and mammals (multituberculates, metatherians, and eutherians).

Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 291

POTENTIAL EFFECTS OF GENETICALLY MODIFIED SOYBEANS ON MAMMALIAN FETAL, BREAST-FED, POSTNATAL, PUBERTAL AND ADULT DEVELOPMENT

Denise Brake and Donald P. Evenson Department of Chemistry and Biochemistry South Dakota State University Brookings, SD 57007

ABSTRACT

The health safety of genetically modiﬁed soybeans was studied using the mammalian testis (mouse model) as the sensitive biomonitor of any potential toxic effects. Pregnant mice were fed a GMO-soybean or a Non-GMO diet through gestation and lactation. After weaning, the young male mice were maintained on the respective diets. At 8, 16, 26, 32, 63 and 87 days after birth, three male mice and an adult reference mouse were killed, the testes surgical- ly removed, and the cell populations measured by ﬂow cytometry. The results showed that GMO foodstuffs had no effect on macromolecular synthesis or cell growth and differentiation as evidenced by no differences in the percentages of testicular cell populations (haploid, diploid, and tetraploid) between the GMO-soybean-fed mice and those fed the Non-GMO diet. There were also no differences in litter sizes, gross behavior, and body weights of the two groups. These data strongly support the view that foodstuffs containing Roundup Ready soybean meal do not cause any negative health effects in the sensitive stages of mammalian developing systems and in the adult animal.

Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 293

COMPARISON OF TWO TECHNIQUES FOR THE DETECTION OF DNA FRAGMENTATION IN SPERM CHROMATIN: SPERM CHROMATIN STRUCTURE ASSAY AND COMET

David Tritle, Lorna Jost, Kay Kasperson, and Donald P. Evenson Dept. of Chemistry and Biochemistry South Dakota State University Brookings, SD 57007

The Sperm Chromatin Structure Assay (SCSA) and Comet Assay represent two tests used to determine DNA fragmentation in infertile human sperm ejac- ulates. This study approached the comparison of CVs and correlations between measurements of DNA fragmentation as measured by both assays. Uti- lizing assay of the same sample by both tests as well as direct Comet Assay of cells ﬂow cytometrically sorted from the SCSA determinations were as objective as possible. This presentation covers the preliminary work toward this comparison and will include the CV and correlation measurements but not the ﬂow sorting.

Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 295

DEVELOPMENT OF A GFP CONSTRUCT TO DETERMINE THE LOCALIZATION OF A PORCINE REPRODUCTIVE AND RESPIRATORY SYNDROME VIRUS PROTEIN

William D. Arndt, Susan Ropp, and David Benﬁeld Department of Veterinary Science South Dakota State University Brookings, SD 57007

ABSTRACT

Porcine reproductive and respiratory syndrome virus (PRRSV) is the causative agent of reproductive disorders in breeding stock and severe respiratory disease in neonates in swine herds worldwide. The 15-kb positive-po- larity, single-stranded RNA genome of PRRSV contains 7 ORFs. To date, the role that ORF 4 plays in PRRSV replication is unclear. As a first step in eluci- dating this role, the purpose of this study was to generate a construct that would allow us to investigate the localization of the protein encoded by ORF4 (GP4) in MARC cells. To this end, a construct was generated to produce a GP4 protein fused with a green fluorescent protein (GFP) tag. To create this construct, ORF 4 was amplified from the genome of the PRRSV strain 23983 by polymerase chain reaction (PCR) and tested by gel electrophoresis to confirm that the PCR product had the correct expected molecular weight. The PCR product was then inserted into a GFP Fusion TOPO vector (pcDNA3.1/NT- GFP-TOPO) and the recombinant vector was transformed into TOP 10 E. Coli cells. Colonies containing the recombinant construct were selected and analyzed by DNA sequencing to confirm that the ORF 4 gene was integrated into the vector correctly. The plasmid was then purified and transfected into MARC- 145 cells and tested for expression of the GFP protein by immunofluorescence assay. GFP is normally distributed throughout the cell, showing fluorescence in both the cytoplasmic and nuclear compartments. Cells transfected with the GP4-GFP fusion construct were inspected to see if the pattern distribution was altered by the presence of GP4.

Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 297

LONG-TERM PLANKTON AND NUTRIENT CHANGES IN PRAIRIE LAKES

Lois Haertel

ABSTRACT

Plankton and nutrient chemistry data are presented from six lakes, Picker- el, Enemy Swim, Cochrane, Hendricks, Oak and Bitter, sampled during two time periods, 1970-1979, and 1990-1997. In addition, data from Lakes Roy, East Oakwood, and Tetonkaha, sampled between 1988 and 1994, are compared. All of the data bases presented include samples taken from spring, early summer and late summer seasons of the lakes. The lakes range from saline to fresh and from eutrophic to hypertrophic, as determined by depth and the ability of winds to resuspend nutrient rich sediments back into the water column. The combined data base for phytoplankton genera includes 837 samples from 1970-1979 and 677 samples from 1988-1997. The data base for zooplankton genera includes 851 samples from 1970-1979 and 685 samples from 1988- 1997. The data base for physical and chemical variables varies in number. Physical variables include temperature, water transparency (secchi depth), station depth, and electrical conductivity. Chemical variables include Chloro- phyll α, nitrate-nitrogen, ammonia-nitrogen, total nitrogen, soluble reactive phosphorus, total phosphorus, silica, iron, manganese and major cations and anions. Several of the lakes show decreases in both water transparency, and total phosphorus levels, and two of the lakes show decreases in total nitrogen between the 1970's and the 1990's. Differences in methodology might account for measured changes in the phosphorus levels. Decreases in the larger zoo- plankters between the decades, especially Daphnia spp. may influence the water transparency. Phytoplankton genera show random changes within lakes. In Lake Cochrane, erosional events in the 1970's and and artificial drains both into and out of the lake in the 1990's may have influenced the phytoplankton composition and abundance. Severe winterkill during the 1970's influenced the changes in phytoplankton abundances in Lakes Hendricks and Oak. Ex- treme water level changes influenced Bitter Lake in the 1990's resulting in greatly lowered conductivities and a phytoplankton change from an Anacystis- dominated saline lake assemblage to an Aphanizomenon-dominated assemblage including freshwater taxa not previously recorded from Bitter Lake. Most variables tested show great similarity between decades, indicating current stability of the prairie lake ecosystem. However, as inputs into the lake create sediments and increase shallowness, lakes that are currently eutrophic may come in danger of hypertrophy from sediment resuspension. The complete paper will be published in the 2003 Proceedings.

Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 299

PLANKTON-ENVIRONMENT ASSOCIATIONS IN PRAIRIE LAKES

Lois Haertel

ABSTRACT

Food chains and trophic interactions between microscopic plankton of lake remain a source of controversy as different experiments have produced widely contradictory results. Many interactions have been documented in the laboratory, but the extent of their existance in nature remains unknown. In addition, biotic interactions are influenced by physical and chemical environmental factors which vary widely in freshwater lakes, selecting for organisms which can vary their method of nutrition as needed. This study evaluates the relative importance of interspecific and physical and chemical interactions in Prairie Coteau Lakes Data bases included six prairie lakes sampled between 1970 and 1979 and nine prairie lakes sampled between 1988 and 1997. In addition, all lakes sampled 1970-1979 were tested together as were all lakes sampled 1988-1997. Thus, seventeen separate data bases were analyzed for interactions among plankton taxa and between plankton taxa and physical and chemical data. Not all variables were measured in every lake and not all plankton taxa were present in every lake, so frequency of significant interaction was in most cases determined for less than seventeen separate tests. Data was not combined between the 1970's and 1990's because of differences in methodology. As no mathematical transformation rendered the phytoplankton data normal, the significance of interactions between taxa and the environment was determined by Spearman's Correlation Coefficients. The relative association between variables tested was measured by the per cent frequency of significant correlation between any two variables. Variables analyzed included first, abundance of all other plankton species; second, weather variables (7-day prior rainfall, 7-day prior solar radiation (1970-1979 only) 7-day prior wind stress and depth of snow the preceding winter); and third, physical and chemical factors measured including station depth, water transparency (Secchi depth). temperature, electrical conductivity, major ions, and plant nutrients (nitrogen, phosphorus, silica, iron and manganese). Specific examples were discussed.

Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 301

USE OF STABLE ISOTOPES TO DOCUMENT THE INVASION OF TALLGRASS PRAIRIE BY DECIDUOUS FOREST IN NEWTON HILLS STATE PARK IN SOUTHEASTERN SOUTH DAKOTA

Craig Spencer, Matt Bukrey, Jason Smalley, Erik Wang, Dan Opheim, and Mike Chapman Biology Department Augustana College Sioux Falls, SD 57197

ABSTRACT

Newton Hills State Park contains native prairie relics that appear to be shrinking due to invasion by forests. At present, the park contains small prairie relics on scattered ridgetops interspersed between ravines dominated by Tilia americana (basswood) and Ulmus americana (American elm). We collected soil cores across a transect of vegetation types in an attempt to quantify historical vegetation changes over the last several hundred years through analysis of stable carbon isotopes (δ13C) in soil organic matter. Tallgrass prairie vegetation typically produces a δ13C value around -14‰, while trees average around -25‰. Data from our cores showed carbon isotope values in the deepest and oldest sections averaging -19.57‰ suggesting a mixed grassland forest environment with values near the surface averaging -24.33‰ suggesting a shift towards forest dominance following European settlement. We analyzed a total of 12 cores and each one showed this shift to more negative δ13C values near the surface. Plant survey data provide additional evidence that is consistent with conclusions drawn from the isotope data. At present, the ravines are densely forested with climax species and the hillsides are dominated by intermediate species (Ostrya virginiana (ironwood) and Quercus macrocarpa) (bur oak)). Finally, the ridgetops support pioneer tree species (Zanthoxylum americanum (prickly ash) and Juniperus virginiana (juniper)) interspersed with native grasses (Andropogon gerardi (big bluestem) and Sorghastrum nutans (indian- grass)). Taken together our data provide evidence that the forests are indeed expanding onto the ridgetop prairies due to ﬁre suppression following Euro- pean settlement.

Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 303

THE UTILITY OF AERIAL PHOTOGRAPHS, FIELD OBSERVATIONS, AND INVERTEBRATE FOSSILS IN THE CONSTRUCTION OF GEOLOGIC MAPS

Frank J. Varriale and Alvis L. Lisenbee Department of Geology and Geological Engineering South Dakota School of Mines and Technology Rapid City, SD 57701

ABSTRACT

During the past 26 years several techniques have been used to elucidate the surface and subsurface geology of the Galisteo and Ojo Hedionda quadrangles in Santa Fe County, New Mexico. A simplistic geologic map, published in 1975, was constructed from aerial photographs and supported by minimal field investigation. During the summers of 1998 and 1999 field observations were conducted for the production of entirely new maps at a scale of 2000’ per inch. This investigation greatly enhanced the accuracy of geologic maps of this region. However, the interpretation of surface and subsurface geology in several areas remained a mystery. To resolve these problems several stratigraphic sections were measured and invertebrate fossil taxa (Cephalopods, Bivalves, & Gastropods) were collected. Specimens were identified to the species level and compared to known biostratigraphic and chronologic range charts. This allowed both relative and absolute age dates to be determined for the measured sections. Measured sections were then compared to each other and correlated with other formations in surrounding basins based on paleontologic, chronologic, and lithologic character. As a result, a greater understanding of the surface and subsurface geology of the Galisteo and Ojo Hedionda quadrangles has been achieved. Lithologic units that once evaded classification are now identified, while others have been located in areas were they were previously undetected. Renaming of identified units has also occurred as a result of index fossil discovery, and units once though to be separate are now recognized as the same unit repeated by normal faulting.

Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 305

OBSERVATIONS OF A CHARGED LIGHTNING CHANNEL IN A HIGH PLAINS THUNDERSTORM

Tom Warner, John Helsdon, Andrew Detwiler, Qixu Mo, and Donna Kliche Institute of Atmospheric Sciences South Dakota School of Mines and Technology Rapid City, SD 57701

ABSTRACT

The Severe Electrification and Precipitation Study (STEPS) took place from 17 May - 20 July 2000 in northeastern Colorado and northwestern Kansas. The broad goal of STEPS was to achieve a better understanding of the interactions between kinematics, precipitation production, and electrification in severe thunderstorms on the High Plains. Resources used in the project included three Doppler radars, a three-dimensional lightning mapping array, the Na- tional Lightning Detection Network, mobile thermodynamic and electric field soundings, a mobile mesonet, and the South Dakota School of Mines and Tech- nology T-28 Storm Penetration Aircraft. On 29 June 2000, all STEPS resources sampled a severe thunderstorm that exhibited many of the project’s desired criteria. The T-28 aircraft sampled a charged channel left by an intracloud lightning flash that took place 0.68 seconds travel time in front of the aircraft. Imag- ing probes provided information on hydrometeor concentrations, shapes and electrical charges, while electric field meters sensed the ambient electric field. Analysis of data collected by the T-28, along with data from the radars and lightning mapping array, provides insight into lightning’s role in charge redis- tribution and the electrification processes occurring in thunderstorms.

Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 307

A NEW MEANDERING STREAM COMPLEX FROM THE BIG BADLANDS OF SOUTH DAKOTA

Robert W. Meredith and Jennifer Cavin South Dakota School of Mines and Technology Rapid City, SD 57701

ABSTRACT

The White River Badlands of South Dakota preserve a sequence of Oligocene terrestrial deposits. Stream channels were first reported in the early 1950’s but since then there has been little research. A newly discovered Brule Formation meandering stream channel deposit was studied in the North Unit of Badlands National Park. The channels are from 30 to 33 meters in width and up to 1.5 meters thick. The primary sedimentary structures observed included epsilon, trough, and tabular cross-bedding, all of which dip to the northeast. A vertical trench was made through one of the outcrops in order to examine fine scale features. The channel deposit consists of 5 units distinguished by lithology and primary sedimentary structures. All but unit 1 are composi- tionally and texturally immature feldspathic litharenites. Unit 1, the channel floor, lies above an olive green scoured clay and consists of a medium to coarse matrix supported clay intraformational conglomerate. Units 2 and 3 are thinly laminated, thinly bedded sandstones laid down during upper flow regime representing the lower and upper point bar respectively. Unit 4 is a thinly laminated, very thinly bedded sandstone intercalated with thin mud- stones representing an overbank deposit. Unit 5 is a thinly bedded sandstone deposited during upper flow regime at flood stage. Together, these units suggest that the meandering stream had a low flow velocity and carried a mixed load. Quartz and feldspars comprise the light minerals while actinolite, biotite, magnetite, and hornblende were the most common heavy minerals. This suggests a Northern Black Hills source.

Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 309

USING THE PENTAX TOTAL STATION AND ARCVIEW TO ANALYZE THE FOSSIL ACCUMULATIONS WITHIN THE CONATA PICNIC GROUND EXCAVATION, BADLANDS NATIONAL PARK, SOUTH DAKOTA

Daniel B. Lien Museum of Geology South Dakota School of Mines and Technology Rapid City, SD 57701

ABSTRACT

During the summer of 1993, two visitors stumbled upon a paleontological treasure in the Big Badlands of South Dakota, an Oligocene bone-bed located in the Scenic Member of the Brule Formation now known as the Big Pig Dig. To date, this site has yielded over 6,000 fossils. The faunal list for this excavation consists of the Archaeotherium, Subhyracodon, Mesohippus, Leptomeryx, Merycoidodon, Ischyromys, and a Nimravidae. The bone density at the site can be as high as 100 bones per square meter. This can make interpretation problematic at best. The purpose of this project was to determine if this fossil accumulation was the result of a single event and to determine individual bone association within the high-density accumulation. To aid in the interpretation of this site, precise field maps were created using a surveyor’s transit, the Pentax Total Station, and the GIS program, Arcview. These maps show details not visible in typical field maps. Entering bone coordinates and identifiers directly into a table allowed for rapid assimilation and interpretation of the data with minimum mod- ification. These tables were then converted into shape files and used to develop overlays in Arcview, a spatial modeling program. By isolating and examining the data files for a single taxon, it is now possible to determine the extent of an individual animal within the quarry. This method has proven successful in showing the association between fossils bone elements at the Pig Dig with a high degree of certainty.

Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 311

HYDROLYTIC STABILITY TESTING OF POLYMER MATRIX COMPOSITES REINFORCED WITH FIBERS HAVING NOVEL SURFACE TREATMENTS

H. Du, C. Struckman, D.H. He, W.M. Cross, J.J. Kellar, and D.A. Boyles South Dakota School of Mines and Technology Rapid City, SD 57701

ABSTRACT

A novel layer was grafted to continuous SiO2 fibers. FT-IR spectroscopy was used to characterize the grafted fiber surfaces. The grafted fibers were used as reinforcements in polymer matrix composites that were fabricated by the hand lay-up technique. Three point bending tests were then performed on the composites samples. The composites samples were also evaluated to compare their hydrolytic stability with and without grafted layers, and with traditional surface treatment (silane coupling agent).

Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 313

FOREST CLASSIFICATION OF THE BLACK HILLS, SD USING DECISION TREE ANALYSIS ON TM AND AVIRIS DATA

Sylvio Mannel, Doug Hua, and Maribeth Price Geology Department South Dakota School of Mines & Technology Rapid City, SD 57701

ABSTRACT

In this project we tested different remotely sensed data and various methods to classify forest cover in the Black Hills, SD. We applied decision tree analysis to AVIRIS (Airborne Visible/Infrared Imaging Spectrometer) scenes and compared the results to multi-temporal Landsat TM 5 (Thematic Mapper) scenes over the same area. TM5 is satellite data with 6 solar bands and a pixel size of 30m. AVIRIS measurements, which had a 20m resolution, were taken from an airplane. The hyperspectral AVIRIS measures in 224 solar bands and produces a smooth spectrum of the land cover. Decision tree classification is a machine-learning alternative to other more traditional classification approaches, such as unsupervised and supervised classifications. It emerged within the last decade from business applications. Training and test data came from intensive vegetation surveys completed in 2000 and 2001 and from digital ortho- quads. Our analysis successfully discriminated between the dominant species, ponderosa pine, and smaller stands of white spruce and aspen/birch. In the spatially abundant Ponderosa Pine we were able to distinguish between open, medium and dense stands. The lower number of our training sites in the less abundant aspen/birch and spruce stands did not permit us to separate densities for these two cover types, and in any event, these species usually occur only in medium to dense stands in the Black Hills. Ancillary data, such as elevation, slope and aspect did not considerably impact the classification results. Decision trees produced similar accuracies to those obtained with more con- ventional classification approaches. Multi-temporal TM data was slightly better than single-season AVIRIS.

Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 315

PHARMACEUTICALS, BINDING SITES, AND PHARMACOLOGICAL EFFECT

Rosina Barbara Flasch Hendrickson and Arlen Viste Department of Chemistry Augustana College Sioux Falls, SD 57197

ABSTRACT

Cardiovascular pharmaceuticals have been classified in many different ways. Each class has specific desired pharmacological actions and pharmaceuticals within each class may have similar structures. In addition, pharmaceuticals with similar structures might have different pharmacological actions. Algorithms, such as those developed by the American Heart Association (AHA) for its Advanced Cardiac Life Support (ACLS) courses, recommend certain pharmaceuticals and certain classes of pharmaceuticals as primary selections. In the AHA’s 2000 revision of its ACLS Algorithms, amiodarone was added as a pharmaceutical of choice for treatment of Ventricular Fibrillation/Pulseless Ventric- ular Tachycardia. (ACLS.net 2002) This pharmaceutical replaced lidocaine as the first choice pharmaceutical following epinephrine. (Bledsoe 1997) Through molecular modeling, an attempt was made to understand how amiodarone acts as a stronger pharmaceutical in treatment of Ventricular Fibrilla- tion/Pulseless Ventricular Tachycardia than lidocaine does. Amiodarone has become known for its competitive inhibition of thyroid hormone to the thyroid hormone receptor subtype beta 1. (Drvota 1995) Protein Databases were searched for the structure of thyroid hormone receptors containing a binding site for amiodarone or the thyroid hormones that it inhibits. The Protein Data Bank (Protein Data Bank 2002) yielded two proteins that might provide a binding site; however, it was not clear whether either of these proteins were the ones mentioned by Drvota as being the most prevalent in the human heart which might yield cardiovascular results. (Rastinejad 1995) The first avenue, therefore, was to examine the suggestion that amiodarone’s antiarrhythmic properties are due to its interaction with the thyroid hormone. (Drvota 1995) Two thyroid hormones, Levothyroxine Sodium and Liothyronine Sodium, were chosen for structural comparison with amiodarone and lidocaine. After examining the chemical structure of the four pharmaceuticals, molecular calculations were conducted through the use of HyperChem. From these calculations, a graphic representation of the HOMO and LUMO states of the pharmaceuticals was obtained. The comparison of chemical structure and HOMO and LUMO states indicated that amiodarone is similar to the thyroid hormones, while lidocaine differs significantly from the other three pharmaceuticals. Based upon these findings, it was concluded that the chemical structure and HOMO and LUMO states of pharmaceuticals is not enough to determine the effects that binding would have on pharmacological effect. However, if cardio- 316 Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) vascular function is enhanced by interaction with thyroid hormones (Drvota 1995), these calculations indicate a possible way in which amiodarone is more likely to interact with these hormones than lidocaine is.

LITERATURE CITED

ACLS.net. 2002. ACLS.net Ventricular Fibrillation (VF)/Pulseless Ventricular Tachycardia (PVT) Algorithm. http://www.acls.net.newalgo/vfpvt.htm. Accessed: 12 February 2002. Bledsoe, Brian E, Robert S. Porter, and Bruce R. Shade. 1997. Paramedic Emergency Care. 3rd Edition. Upper Saddle River, NJ: Brady Prentice Hall. Drvota, V., B. Carlsson, J.Hagglad, and C. Sylven. 1995. Amiodarone is a dose- dependent noncompetitive and competitive inhibitor of T3 binding to thyroid hormone receptor subtype beta 1, whereas disopyramide, lignocaine, propafenone, metoprolol, dl-sotalol and verapamil have no inhibitory effect. Journal of Cardiovascular Pharmacology. 26: 222-226. Protein Data Bank. 2002. http://www.rcsb.org/pdb/ Rastinejad, Fraydoon, Thomas Perlmann, Ronald M. Evans, and Paul B. Sigler. 1995. Structural determinants of nuclear receptor assembly on DNA direct repeats. Nature. 375: 203-211. Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 317

NEST SITE CHARACTERISTICS OF EASTERN WILD TURKEY FEMALES IN AGRICULTURAL LANDSCAPES OF NORTHEASTERN SOUTH DAKOTA

Roger D. Shields and Lester D. Flake Wildlife and Fisheries Sciences South Dakota State University Brookings, SD 57007

ABSTRACT

We used radio transmitters to monitor nest site selection in 65 eastern wild turkey (Meleagris gallopavo silvestris) females in a mosaic of deciduous forest (<10%), cropland, and grassland in northeastern South Dakota during 1999 and 2000. We marked the area near incubating females without approaching and ﬂushing them and returned after the nesting attempt (n=99) to locate the actual nest and obtain information on nest site characteristics, primarily microhabitat. Wild turkeys primarily nested in shrubland (3% of study area and 31% of nests), idle grassland (13% of study area and 44% of nests), woodland (4% of study area and 10% of nests), and pastureland (29% of study area and 8% of nests) vegetation types. To determine characteristics important for selection of nest sites by female wild turkeys in shrubland, grassland (including idle grassland and lightly grazed pastureland), and woodland, we compared characteristics at nest sites with randomly selected paired sites within respective vegetation types. Nest success was signiﬁcantly lower (P = 0.051) for woodland sites (20%) than either shrubland (63%) or grassland (58%) sites. Grassland and shrubland, the latter primarily interspersed with grassland, provided safe nesting habitat compared to woodland sites. In general, females sought nest sites that increased the opportunity of producing a successful hatch. Vegetation at nest sites offered a high degree of horizontal visual obstruction of the nest bowl (91.7% and 20.6% cover from 0-60 cm and 60-180 cm, respectively) as well as visual obstruction throughout the surrounding 10-30 m area (69.3% and 11.5% cover at 0-60 cm and 60-180 cm, respectively). Dense ground cover vegetation around the nest bowl further provided substantial overhead obstruction of the nest bowl (53.5% cover).

Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 319

CORRELATION OF BIOCHEMICAL AND MOLECULAR PROFILES FOR 35 RELATED ISOLATES OF METHICILLIN RESISTANT STAPHYLOCOCCUS AUREUS

Amy J. Stephenson Basic Biomedical Health Sciences University of South Dakota School of Medicine Vermillion, SD 57069

Lona Haas and Patricia M. Tille Division of Natural Sciences Mount Marty College Yankton, SD 57078

ABSTRACT

Staphylococcus aureus are spherical gram positive cocci typically isolated from human skin and mucous membranes. Approximately 2.4% of clinically isolated S .aureus are methicillin resistant. These organisms also produce a wide array of virulence factors which ensure their survival and increase the severity of the infectious process. This research was conducted to determine the optimal growth conditions and biochemical proﬁle for 35 isolates of methicillin resistant S. aureus. Vari- ous growth conditions were examined to determine optimal temperature, pH and NaCl concentration. Isolates were also examined for the presence of hem- loysins and antibiotic resistance patterns. This data was compiled and compared to previous molecular studies in order to identify any correlations among the biochemical proﬁles and the molecular analysis.

Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 321

BIODIVERSITY ASSESSMENT OF THE JAMES RIVER BASIN

Chad Thury, Christy DeRoos, John Gullikson, Krisma Dewitt, James Sorenson and Patricia M. Tille Department of Natural Science Mount Marty College Yankton, SD 57078

ABSTRACT

The purpose of this study was to examine the biodiversity of the James Riv- er Basin located in a small region of South Eastern South Dakota. The James River ﬂows primarily through agricultural areas and is subject to deterioration as a result of feedlot run-off, herbicide and/or pesticide use. Soil depth throughout this region is very shallow, which allows for surface water to move rapidly into the aquifer creating widespread sources for water contamination. This study was subdivided into four areas: 1.) physical and chemical properties of water; 2.) physical and chemical properties of soil samples; 3.) water and soil microbiota; and 4.) water and soil macroinvertebrates. Random sites along the James River were sampled upstream as well as downstream of Yankton, South Dakota. This study was used to develop baseline data and protocols for the implementation of a 2-year study examining the biodiversity along the James River Basin as an indicator of environmental contamination.

Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 323

DIVERSITY, DENSITY AND BREEDING STATUS OF THE AVIFAUNA IN THE SCHRAMM ADDITION AT PONCA STATE PARK, NEBRASKA

Eric T. Liknes Department of Biology University of South Dakota Vermillion, SD 57069

ABSTRACT

Ponca State Park, Dixon Co., is located primarily on the bluffs along the Missouri River in northeastern Nebraska. Recently, the park acquired an area of adjacent floodplain and backwater with plans of restoring this area to native vegetation. The objective of this study was to assess the avifauna within the biotic communities of the Schramm addition and to establish a baseline for further studies after the restorations are completed. Sampling effort was focused on the two principal habitats adjacent to a backwater area. A mosaic of relatively mature patches of trees (approximate canopy height 10-15 m) predominated by cottonwood (Populus deltoides) and open fields was characteristic of the upland area, while a mosaic of early successional scrubby trees (approximate canopy height 3-5 m), predominated by cottonwood and willow (Salix spp.) was characteristic of the habitat on the peninsula created by the backwater. The two habitat types surveyed in this study are increasingly rare and fragmented along the Middle Missouri River as upstream dams heavily regulate river flow. Variable circular-plot point counts were conducted from 29 May through 07 August 2001 along two roughly parallel 1 km transects separated by at least 100 m. Point-count censuses were conducted in conjunction with mist-netting. A total of 56 species was detected. Of those 56 species, 50% were Neotropical migrants, 37.5% were temperate resident species, and 10.7% were permanent residents. The two habitats were significantly different in terms of diversity and shared a Morisita’s Index of community similarity of 0.769. The two habitats were also different in terms of density, and relative abundance, but not capture rate.

Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 325

SYNTHESES AND APPLICATIONS OF NEW ORGANOBORON REAGENTS FOR SUZUKI CROSSING-COUPLING REACTION

Jiangtao Hao and David A. Boyles Department of Chemistry and Chemical Engineering South Dakota School of Mines and Technology Rapid City, SD 57701

ABSTRACT

Arylboronic acids have been used extensively in Suzuki reactions. Diaryl- borinic acid and triarylborane should be more reactive in Suzuki reactions according to their molecular structure. However, diarylborinic acids have been ig- nored due to their difﬁculty of synthesis and isolation, and the utility of triarylborane is compromised by sensitivity to oxidation. A practical procedure has been developed for the preparation of bis(4-benzyloxyphenyl)borinic acid in high purity and yield. Furthermore, a simple but powerful strategy for ad- dressing the problem of triarylborane has been examined: convert air-sensitive triarylborane into an air-stable potassium ﬂuoride salt. These diarylborinic acid and triarylborane salts serve as reactive Suzuki reaction partners. The details of the syntheses and their preliminary applications to the Suzuki reactions are described in this presentation.

Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 327

USE OF ARYLDIAZONIUM TETRAFLUOROBORATES FOR SUZUKI-TYPE CROSS-COUPLING REACTIONS WITH DIARYLBORINIC ACIDS

Joseph Marshall, Jiangtao Hao, and David A. Boyles Department of Chemistry and Chemical Engineering South Dakota School of Mines and Technology Rapid City, SD 57701

ABSTRACT

Both aryl tetrafluoroborates and arylboronic acids are common substrates for the popular Suzuki cross-coupling reaction which has found recent, extensive use in the synthesis of bisaryl compounds. No literature reports any men- tion of the reaction of an aryl tetrafluoroborate with a bisarylborinic acid, however. Our previous investigations of the greater reactivity of the later bisarylborinic acids in the Suzuki reaction has prompted us to react them with diazonium tetrafluoroborates in order to overcome solubility problems and to further enhance the rate of reaction of the former compounds. The diazonium tetrafluoroborates of three substrates have been synthesized. Each was reacted with the bis(4-benzyloxyphenyl)borinic acid to compare their feasibility of the proposed reaction. The results of these reactions will be described.

Proceedings of the South Dakota Academy of Science,Vol. 81 (2002) 329

MODIFIED SUZUKI REACTION USING BISARYL TOSYLATES WITH DIARYLBORINIC ACIDS

Celeste Mercado, Jiangtao Hao, and David A. Boyles Department of Chemistry and Chemical Engineering South Dakota School of Mines and Technology Rapid City, SD 57701

ABSTRACT

The Suzuki cross-coupling reaction is a recent reaction which has largely supplanted the classic coupling of aryl bromides at extreme conditions using a copper catalyst. Although many variations of the Suzuki reaction are currently in development, phenolic substrates typically have required conversion to the corresponding trifluoromethanesulfonate (as the aryl triflate) with subsequent reaction with arylboronic acids. Recently, we have undertaken to modify this reaction by use of the less expensive and more robust aryl toluenesulfonates (aryl tosylates). These compounds should, in principle, react with diarylborinic acids which have found extensive use in our laboratory. Accordingly, the tosylate of bisphenol A was prepared as an inexpensive and more stable alternative to the bistriflate previously utilized. This compound was reacted with the electron-rich bis(4-benzyloxyphenyl)borinic acid under catalytic conditions in an attempt to transfer both aryl groups. The results of this reaction are described.