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PROCEEDINGS
of the
Indiana Academy
of Science
Centennial Year
Founded December 29, 1885
Volume 94
1984
DONALD R. WINSLOW, Editor
Indiana University
Bloomington, Indiana
Spring Meeting
April 27, 28, 1984
Brookville, Indiana
Fall Meeting
November 1, 2, 3, 1984
Butler University
Indianapolis, Indiana
Published at Indianapolis, Indiana
1985
1. The permanent address of the Academy is the Indiana Academy of Science, 140 North Senate Avenue,
Indianapolis, Indiana 46204.
2. Instructions for Contributors appear at the end of this volume.
3. Exchanges. Items sent in exchange for the Proceedings and correspondence concerning exchange arrangements
should be addressed:
John Shepard Wright Memorial Library of the Indiana
Academy of Science
140 North Senate Avenue
Indianapolis, Indiana 46204
4. Proceedings may be purchased through the Library at $12.00 for each volume.
5. Reprints of technical papers often can be secured from the authors. They cannot be supplied by the Library
nor by the officers of the Academy.
6. The Constitution and By-Laws reprinted from Vol. 74 are available to members upon application to the
Secretary. Necrologies reprinted from the various volumes can be supplied to relatives and friends of deceased members
by the Secretary.
7. Officers whose names and addresses are not known to correspondents may be addressed in care of the Library.
Editorial Board
1984
Donald R. Winslow Benjamin Moulton
Chairman and Editor John Pelton
Hans Andersen Carl Sartain
Rita Barr Alfred Schmidt
Ernest Campaigne J. Dan Webster
Robert F. Dale John O. Whitaker
James Gammon Bernard S. Wostmann
James Kellar Frank Young
Gene Kritsky
PUBLICATIONS AVAILABLE FROM THE ACADEMY
HISTORY OF THE INDIANA ACADEMY OF SCIENCE,
Daily, W. A. and Daily, F.K. $9.00 ppd
ECTOPARASITES OF MAMMALS OF INDIANA,
John O. Whitaker, Jr. $8.95 ppd
DISTRIBUTION OF THE MAMMALS OF INDIANA,
Russell E. Munford $3.00 ppd
VEGETATION OF THE LIFE ZONES IN COSTA RICA,
John O. Sawyer & Alton A. Lindsey $4.00 ppd
AMPHIBIANS & REPTILES OF INDIANA,
Sherman A. Minton, Jr. $6.00 ppd
Order from and make check payable to: Indiana Academy of Science. Send to Dr. Benjamin
Moulton, Dept. of Geography and Geology, Indiana State University, Terre Haute, IN
47809.
TABLE OF CONTENTS
Page
Preface to the Centennial Volume
Theodore J. Crovello, President 3
Officers and Committees for 1984 4
HIGHLIGHTS OF THE SPRING MEETING
Brookville Historical Tour, John Newman 13
Our Brookville Bond, Fay Kenoyer Daily 18
The Making of David Starr Jordan, Gary A. Sojka 22
Reports from field trip leaders:
Geology, Curtis H. Ault and John R. Hill 29
Ornithology, William H. Buskirk 30
Zoology, Sherman A. Minton 30
Minutes of the Spring Meeting
(Executive Committee) 31
PICTORIAL HIGHLIGHTS OF THE FALL MEETING
Welcome to Butler University, John G. Johnson
President, Butler University 35
Welcome to the Fall Meeting, Theodore J. Crovello,
President, Indiana Academy of Science 36
Centennial Address 37
Dinner for Senior Academy Officers 38
Executive Committee Meetings 39
Noon Luncheon 41
Poster Sessions 42
Indiana Junior Academy of Science 43
Special Acknowledgment 44
Minutes of the Fall Meeting (Executive Committee) 48
Minutes of the Fall Meeting (General Session) 54
Minutes of the Budget Committee Meeting 57
Annual Financial Report 59
Annual Report, Indiana Junior Academy of Science 64
Necrology, Fay Kenoyer Daily, Necrologist 69
New Members for 1984 76
ADDRESSES AND CONTRIBUTED PAPERS
Presidential Address
"Computers, Education, and Artificial Intelligence,"
Theodore J. Crovello 80
"Speaker of the Year" Address, 1984-85
"The Contributions of the Nightshade Family (Solanaceae)
to Human Welfare," Charles B. Heiser, Jr 88
*Abstracts
iii
iv Indiana Academy of Science Vol. 94 (1985)
Anthropology
C. Michael Anslinger— Debitage Classification Systems* 93
Ruth Brinker— Mann Site Figurines* 93
Frank Burkett and Donald R. Cochran— The Commissary Site (12-Hn-2)
Revisited* 93
Mark Cantin and C. Michael Anslinger— Holland Chert Quarries/ Workshops
Near Huntingburg, Dubois County, Indiana* 93
Mary Ellen Carpenter and Robert E. Pace— Test Excavations at the Smith Site,
(12-Vi-86), Vigo County, Indiana* 94
Catharine A. Carson — A Description of Kenneth Chert* 94
Della Collins Cook— Three Cranial Tumors from Late Woodland Sites: Diagnosis
and Cultural Implications* 94
Edmond J . Furia— A Useful Morphological Characteristic of Two Toed Sloth Hair* 94
Ronald Hicks— The Year at Drombeg* 95
Misty Jackson and Robert E. Pace — Towards Predicting Loss of Archaeological
Resources from River Channel Migrations* 95
James A. Mohow— A Preliminary Survey of the Maumee River in Allen County,
Indiana* 95
P. Ranel Stephenson— Woodland Sites and Ross Soils: A Correlation in the Upper
White River (West Fork) Drainage* 95
Curtis H. Tomak— Some Late Archaic Manifestations in Indiana* 96
Botany
Blair Brengle and William Stillwell — Effect of Cytokinins on Erythritol
Permeability to Phosphatidylcholine Bilayers* 97
Rita deCassia, G. Borges, William R. Chaney and Phillip E. Pope — Nonspecificity
with Varied Effectivity in Mycorrhizal Associations* 97
Vonda Frantz — Insect Pest Control in the Greenhouse: Alternatives to Commer-
cial Toxins* 98
Ralph J. Green, Jr. and Philip T. Marshall — Oak "Leaf Tatters": A Malady
of Unknown Cause in Indiana* 98
Romesh C. Mehra and E. Boyts — G-banding in Lens culinaris and Vicia faba* 98
H.S. Bhella — Response of Muskmelon to Within-row Plant Spacing 99
H.S. Bhella and G.E. Wilcox — Stem Length as an Estimator of Muskmelon Growth 105
K. Michael Foos and Judith A. Royer — Isolation of the Coprophilous Fungus,
Pilobolus, from Wayne County, Indiana 109
Jonathan Leeds, Lynne Bemis, Rita Barr and Frederick L. Crane — A New
Amine as an Uncoupler of Chloroplast Electron Transport 113
Gayton C. Marks, William W. Bloom and Jeffrey G. Boyle — A Rapid Method
for the Determination of Barley Seed Viability 117
Patricia W. Reed — Population Studies of Threatened and Endangered Plants of
Barker Woods Nature Preserve, LaPorte County, Indiana 121
'Abstracts
Table of Contents v
G.L. Reed and W.R. Stevenson — Bacterial Wilt Resistance in Commercial
Muskmelon Cultivars 131
Rosemary Rodibaugh and Connie Weaver — Improving Efficiency of Iron Up-
take by Soybeans 141
Gail E. Ruhl, Richard X. Latin, Paul C. Pecknold and Donald H. Scott — A
Compilation of Plant Diseases and Disorders in Indiana — 1984 145
Cell Biology
Kathy Burek and Robert J. Stark — Effect of Acetylcholine Stimulation on
Cytosolic Chloride in Parotid Acinar Cells* 151
Edwin M. Goebel and Deborah A. McMahan — Physiological Studies of
Azospirillum amazonense* 151
Ralph A. Jersild, Jr. — A Brief History of the Cell Biology Section, Indiana Academy
of Science* 152
Michael S. Maloney — Concanavalin A Inhibits Oral Regeneration in Stentor
coeruleus by Binding to the Cell Surface* 152
John W. Munford and Thomas Koenig — The Effect of Fasting on Sodium Pump
Activity in Rat Skeletal Muscle* 152
Jeanette M. Schepper and James P. Hughes — Increased Binding of Growth Hor-
mone following Cleavage by Rabbit Liver Plasmalemma* 153
A.C. Snyder, A.R. Coggan and J.J. Uhl — Protein Degradation after Eccentric
Exercise* 153
Martin A. Vaughan, Timothy J. Mulkey and Charles W. Goff — Calmodulin
Stimulation of ATP — Dependent Ca2+ Uptake in Maize Root Microsomes* 154
Henry C. Womack— The Effect of Illumination on the Rat Pineal as Measured
by MSH Activity* 1 54
James P. Holland, Richard Brooks and Erich Weidenbener — Plasma Pro-
gesterone, Blastocyst Steroidogenesis and Blastocyst Survival in Rats with Altered
Thyroid Status 155
Lisa B. Nass, Annette L. Schlueter and Grayson S. Davis — Chick Limb Duplica-
tions Produced by Retinoic Acid Releasing Microimplants 161
Chemistry
Sepehra Akhavan, Kristen Faust and Bruce Storhoff — Ambidentate Phosphine
Ligands: Phosphine-amine and Phosphine-imidate Complexes of Tungsten* 167
Stasia A. Barnell, Beth E. Beeson and Lynn R. Sousa — The Synthesis of a Crown
Ether that May Exhibit Metal Cation Enhanced Fluorescence* 167
Mohammad Behforouz, Joseph L. Bolan and Michael S. Flynt —
2,4-Dinitrophenylhydrazones: A Modified Method for the Preparation of these
Derivatives and an Explanation of Previous Conflicting Results* 167
Mohammad Behforouz and K.E. Mennen — Wittig Reaction: Stable Ylides in the
Preparation of 7, 5-unsaturated-|3-Ketoesters* 168
Mohammad Behforouz and M.E. Ogle — Synthesis of /3-Carbolines Dervied from
2-Amino-3-(3-indolyl)-butyric Acid (0-Methyltryptophan)* 168
'Abstracts
vi Indiana Academy of Science Vol. 94 (1985)
Stanley L. Burden and Phillip W. Schultz — Coulometric Titrations: Low Cost
Alternatives for Computer Controlled Titrations* 169
Mark Cisneros and Joe Kirsch — Temperature Dependent Infrared Studies of the
Hydrogen Bonding in Aliphatic Alcohols* 169
Sally K. Dotterer and Kenneth L. Stevenson — Spectra and Equilibria of the
Thiocyanate Complexes of Copper (I) in Aqueous Solution* 169
Jennifer L. Dyke and John A. Mosbo — Steric and Electronic Effects upon cis:trans
Distributions in W(CO)4(L)(L') Complexes when L and L' are Phosphorus
Ligands* 170
Bernice Ellis, Kevin Cooksy, James M. Anderson and Harry W. Jarrett — A
Simple, Reproducible High Performance Liquid Chromatography Separation
of Amino Acids with Picomole Sensitivity* 170
Maureen L. Hill, Patrick Gallagher, Jeff Macri and F.W. Kleinhans — An
Electron Spin Resonance Method for the Measurement of Liposomal Leakage* 170
J.C. Huffman, R.A.D. Wentworth, W.E. Streib and C.J. Huffman — Hindered
Ligand Systems: Structure of the cis,cis- 1,3, 5-Tris (pyridine-2-carboxaldimine)
cyclohexane Complexes of Fe(II) and Ni(II) Ions* 171
Nathan E. Kastelein, Phillip E. Klunzinger, Edward J. Ciesla, Claudia Rishaw,
Cynthia L. Roth and Stanley L. Burden — Robots in the Chemistry
Laboratory, Part I: A High Speed RS-232C Serial Communications Link for
Controlling a HERO I Robot from an Apple II Plus Microcomputer* 171
Nathan E. Kastelein, Phillip E. Klunzinger, Edward J. Ciesla, Cynthia L.
Roth, Claudia Rishaw and Stanley L. Burden — Robots in the Chemistry
Laboratory, Part II: Software for Controlling a HERO I Robot from an Apple
II Plus Microcomputer via a High Speed RS-232C Communications Link* . 171
Richard A. Kjonaas — Reaction Sequence Alteration in the Acetoacetic Ester Syn-
thesis of Ketones* 172
LeRoy Kroll and Bruce Storhoff — Functionalized Crown Ethers* 172
Steve Newnam and James P. Rybarczyk — A Trace Metal Analysis of Coal and
Acid Rain* 1 72
Laura Pokorney and James R. Rybarczyk — Conclusion of Acid Rain Monitor-
ing in Central Indiana* 173
Eugene Schwartz — Atomic Polarizations of Transition Metal tris-3-
Pentanedionates* 173
John Scircle and Joe Kirsch — Temperature Dependent Infrared Studies of the
Hydrogen Bonding in Aliphatic Alcohols* 173
Joseph R. Siefker and Kenneth R. Kimmerle — A Study of the Coordination Com-
pounds of Some of the Transition Metals Using 2(2-Aminoethoxy)-Ethanol as
a Ligand and l-Methyl-2-Pyrrolidinone as a Solvent* 173
Daniel K. Wunderlich and Myong-Ku Ahn — An Investigation of Aluminum Con-
centrations in Water* 1 74
Christopher L. Bush and Raima M. Larter — Sensitivity Studies of a Computer
Model for the Peroxidase-oxidase Oscillating Reaction 177
Joe Kirsch, Shannon Lieb and Mark Cisneros — A SCC MO Calculation on the
Tetracyanoethylene-benzine Complex 181
*Abstracts
Table of Contents vii
Kristine S. Kurtz and Kenneth L. Stevenson — Spectra and Photochemistry of
the Chloro Complexes of Copper (I) 187
Barth H. Ragatz, Gina Modrak and Ericka Baeske — Evaluation of Sample Pre-
treatments as Potential Methods of Enhancing Phospholipid Extraction from
Human Amniotic Fluid 193
Barth H. Ragatz, Gina Modrak and Patricia S. Conn — Comparison of Two
Simple Methods for Determining Lecithin/Sphingomyelin (L/S) Ratios in Human
Amniotic Fluid Samples 197
Barth H. Ragatz, Gina Modrak and Mike Engle — The Effects of Oligolysines
and Polylysines on Human Platelet Aggregation Induced by Polylysines,
Adenosine Diphosphate, and Epinephrine 203
Ecology
James R. Aldrich — Pipewort Pond, a Unique Wetland with Atlantic Coastal Plain
Elements in Elkhart County, Indiana* 209
James W. Berry — Competition for Ownership of Webs in the Semi-social Spider
Cyrtophora moluccensis of Yap (Caroline Islands, Micronesia)* 209
Alex Burgin and David T. Krohne — Regional Low Density and Extinction in
Populations of Peromyscus leucopus* 209
Spencer A. Cortwright — Predator-determined Structure in Amphibian Pond
Communities* 210
Michael A. Ewert and Craig E. Nelson — The Complex Relationship of Embryonic
Development to Incubation Temperature in Turtles* 210
Scott Ferson and Daniel D. Stockton — A Competitive Ecotone between Hard-
wood and Relict Hemlock Communities* 210
Burnell C. Fischer and John A. Kershaw, Jr. — Development and Analysis of
a CFI Data Base for Indiana* 211
George S. Libey and Gary E. Miller — Biofiltration in Intensive Culture Systems:
Design Considerations* 211
Molly Morris — Sexual Selection and Alternative Mating Strategies in Hyla crucifer
and Hyla chrysoscelis* 212
Craig E. Nelson — Do Tadpoles Die for their Siblings?* 212
George R. Parker and Donald J. Leopold — Tree Species Dynamics in an Old-
growth Deciduous Forest since 1926* 212
Stephen A. Perrill — Male Mating Behavior in Hyla cinerea* 213
Phillip E. Pope, William R. Chaney and William R. Byrnes — Hardwood Tree
and Ground Cover Establishment on Reclaimed Mineland and Unmined Reference
Sites in Indiana* 213
Brad Semel and Douglas C. Andersen — Interactions among Mast, Small Mam-
mals, and Insects, and their Implications* 213
Rod Walton— Density-dependent Mortality on Galls of the Goldenrod Gall Fly,
Eurosta solidaginis* 214
David K. Apsley, Donald J. Leopold and George R. Parker — Tree Species
Response to Release from Domestic Livestock Grazing 215
"Abstracts
vin Indiana Academy of Science Vol. 94 (1985)
Steven E. Backs, Sean T. Kelly, P. Decker Major and Brian K. Miller—
Characteristics of Drumming Habitat of Ruffed Grouse in Indiana 227
Virgil Brack, Jr.— The Foraging Ecology of Some Bats in Indiana 231
John S. Castrale, Robert E. Rolley and William J. Pfingsten— Legal Game
Harvest by Indiana Landowners Hunting without a License 239
Michael A. Homoya, D. Brian Abrell, James A. Aldrich and Thomas W. Post—
The Natural Regions of Indiana 245
Eric S. Menges and Thomas V. Armentano— Successional Relationships of Pine
Stands at Indiana Dunes 269
Edwin R. Squiers— The Roots of Ecology in Indiana 289
Engineering
David D. Chesak — The IAS Engineering Section: A Brief History* 293
S. Dhawale, G. Cragnulino and D.D. Macdonald — Stress Corrosion Cracking
of Sensitized Austenitic Stainless Steels in Basic Acid Solution Containing Sulfur
Oxyanions* 293
Andrew Hollerman — Engineering and Science Education's Dilemma: Inadequate
Science Programs in the Public School System* 293
Scott Oblander and W.W. Bowden — Prediction of the Variation of Azeotropic
Composition Using the Gibbs-Konovalov Theorem* 293
Dennis West and W.W. Bowden — The PVT Behavior of Compressed Liquids* 295
Paul W. Mueller, Roger M. Hoffer and John E. Jacobson — Evaluation of
Landsat Thematic Mapper Data for Classifying Forest Lands 297
Entomology
Jaime E. Araya and John E. Foster — Effect of Barley Yellow Dwarf Virus Infec-
tion of Wheat and Oats on the Life Cycle of Rhopalosiphum padi (L.)* . . . 303
William E. Chaney — Efficiency of Pollen Traps with Various Sized Trap Screens* 303
B.H. Chen, J.E. Foster and H.W. Ohm — Effect of Viruliferous and Non-
viruliferous Rhopalosiphum padi (L.) Aphids on Winter Wheat* 304
C. Kudagamage and J.E. Foster — Mass Rearing the Bird Cherry Oat Aphid,
Rhopalosiphum padi (L.)* 304
G.L. Reed and D.K. Reed — Assessment of Numbers of Striped Cucumber Beetle
Adults and Frequency of Feeding Injury on Muskmelon Cultivars* 304
H.V. Scheller, R.H. Shukle, E.S. Furgason and J.E. Foster — Relationship of
Probing Behavior of Sitobion avenae (Fabricius) to Transmission cf Luteoviruses
Causing Cereal Yellow-dwarf Diseases* 305
R.H. Shukle, H.V. Scheller and J.E. Foster — Identification of a Pectinase in
Larvae of the Hessian Fly, Mayetiola destructor (Say)* 305
V. Thirakhupt and J.E. Foster — Preference of the Bird Cherry Oat Aphid,
Rhopalosiphum padi (L.) on Hessian Fly-infested Wheat and Effects on its
Biology* 305
John J. Favinger — Anecdotal History of Entomology in Indiana 307
Philip T. Marshall and James A. Clark — Indiana Gypsy Moth Survey — A History 3 1 3
* Abstracts
Table of Contents ix
Robert W. Meyer — Insects and Other Arthropods of Economic Importance in
Indiana in 1984 323
Jack R. Munsee — Annual Changes in Flea Populations on Three Domestic Pets,
1978-1984 329
David K. Reed and Gary L. Reed — Control of Vegetable Insects with Neem Seed
Extracts 335
John Richard Schrock — Checklist of Adult Carabid Beetles Known from Indiana 341
Charles E. White, Frank N. Young and N.M. Downee — A Checklist of the Aquatic
Coleoptera of Indiana 357
Environmental Quality
William Beranek, Jr. and Elizabeth DuSold — The Determination of the Removal
Rate of Specific Chemicals by the Indianapolis Wastewater Treatment System* 37 1
William Beranek, Jr., Elizabeth DuSold, John Merrill and Martin St. Clair —
A Superfund Risk Assessment in Indiana: A Case Study of the Columbia City Site* 371
William Beranek, Jr. and David Jordan — The Ratio of PM-10 to TSP in the
Atmosphere* 371
Howard E. Dunn, Benjamin P. Miller, Charles B. Macer and Michael E.
Klansmeier — Evaporation Rates of Organic Liquids at Various Wind Speeds
and Temperatures* 372
Denise Benson, Claude D. Baker, Bill J. Forsyth and John S. Castrale —
Herbicide (Alachlor, Atrazine, Linuron and Paraquat) Residues in Deer Mice
Inhabiting Conventional and Minimum Tillage Row-crop Fields 373
Ronald J. Galloy — Acid Rain: A Synopsis 381
Geology and Geography
Konrad J. Banaszak and Theodore K. Greenman — Landfills in Marion County—
A Revisit* 387
K.C. Kuo and T.R. West— Compression Strength Testing of the Springfield Coal,
Coal V, Pike County, Indiana* 387
Alan C. Samuelson— Interpretation of Glacial Geology and Groundwater Problems
in East-central Indiana Using Improved Compilations of Water Well Driller's
Records* 388
J.R. Sans and CD. Potter— Three-dimensional Patterns of Biotic Composition
within the Cloudy Pass Batholith, Washington* 388
William L. Wilson and Donald W. Ash — Geology and Geomorphic History of
the Garrison Chapel Cave System, Monroe County, Indiana* 388
Will H. Blackwell— Evidence of Algal Source of Micrite in a Saluda Coral Zone
in Southeastern Indiana 391
History of Science
Barbara A. Seeley and Gerald R. Seeley— The Rich and Varied Past of the History
of Science Section 395
Microbiology and Molecular Biology
Nancy C. Behforouz — Effect of Cyclosporine A on Leishmania tropica* .... 401
*Abstracts
x Indiana Academy of Science Vol. 94 (1985)
Richard H. Lambert and J.R. Garcia — The Regulation of S-Adenosylmethionine
Synthetase in Candida albicans* 401
M. Langona — A Case of Tuberculosis in the University Setting* 401
M. Langona, S. Bossung and M. Orr — Scabies: A Nosocomial Outbreak*. . . 402
Steven H. Larsen and Joann Hoskins — Three Plasmid Cloning Vectors for Mam-
malian Cells* 402
Linda Madisen and M.E. Hodes — Banking DNA for Future Diagnosis of Hereditary
Diseases* 402
F.H. Norris and M.E. Hodes — An Examination of 495 Splice Junction Sequences* 403
Tom Pugh and Mary Clancy — Transcriptional Regulation of the Sporulation-
specific Glucoamylase of Saccharomyces cerevisiae* 403
James L. Shellhaas — Development of a Model System for the Study of Murine
Leukocyte Chemiluminescence* 404
M. Skaria, J.E. Foster and R.M. Lister — Relationship between Symptomatic
Resistance and Virus Production in Barley Cultivars Inoculated with Barley Yellow
Dwarf Virus* 404
David L. Snyder and Bernard S. Wostmann — Serum Hormone Levels in Germ-
free and Conventional Rats: Effect of Dietary Restriction* 404
I.L. Sun, J.E. Putnam and F.L. Crane — Control of Cell Growth by Trans-
plasmalemma Redox: Stimulation of HeLa Cell Growth by Impermeable Oxidants 407
Physics and Astronomy
Albert A. Bartlett and Richard L. Conklin — The Dynamics of the Population
of the United States* 417
Marshall P. Cady, Jr. — On the Measurement of Thermal Diffusivities with
Bryngdahl Interferometry* 417
Vincent A. DiNoto, Jr. — The Physics of the Grist-mill* 418
Frank K. Edmondson — The National Optical Astronomy Observatories* 418
L. Dwight Farringer — The Manchester Interface Adapter for Commodore and
Apple Microcomputers* 418
Jodi Hamilton and Thomas H. Robertson — Software for Astronomical
Photometry* 419
Lawrence E. Poorman — Licensing and Certification of Physics Teachers by Ex-
amination: What are the Dangers? 419
Thomas H. Robertson and Jodi Hamilton — A System for Astronomical
Photometry* 419
Gerald J. Shea— The Great Southern U.S. Geologic Uplift Observed in the Early
Months in 1984* 419
F.R. Steldt — Astrophotography Using Celestron Telescopes* 420
Nancy Watson and James Watson, Jr. — Using Toys to Teach Physics to Middle
School Students* 420
Samir I. Sayegh and Joseph D. Lawrence — Integer-valued Equivalent Resistances 421
'Abstracts
Table of Contents xi
Plant Taxonomy
James R. Aldrich, Lee A. Casebere and Helene Starcs — The Discovery of Native
Rare Vascular Plants in Northern Indiana* 425
Roxane A. Dupuis and Richard Jensen — A Preliminary Survey of Phenolic Com-
pounds in Sympatric Populations of Quercus shumardii and Q. rubra in Nor-
thern Indiana* 425
J.F. Hennen, R.M. Lopez-F and M.M. Hennen — Rust Species Diversity in
Temperate and Tropical Regions in the Americas* 425
Michael A. Homoya — Additions to the Flora of Indiana: II* 426
Richard J. Jensen and Roxane A. Dupuis — Assessing Variation in Mixed Oak
Communities: Evaluation of Multivariate Analyses of Morphological Data* 426
R.C. Mehra, D. Fisher, S. Brekrus, S. Alwine, J. Palbykin and M.G. Butler —
Linear Differentiation of Alluim cepa, Lens culinaris and Vicia faba
Chromosomes* 426
Paul E. Rothrock — Vascular Flora of Grant County, Indiana: Additions and
Comments* 427
Rebecca A. Strait and Marion T. Jackson — Pre-burning Floral Inventory of
Little Bluestem Prairie, Vigo County, Indiana* 427
Richard J. Jensen — The Red and Black Oaks of Indiana 429
John W. McCain — A Preliminary Review and Multiple-entry Key to the Rust Fungi
on Cyperaceae and Juncaceae in Indiana 447
Thomas W. Post — Additions to the Flora of Pike and Gibson Counties, Indiana 455
Thomas W. Post, John A. Bacone and James R. Aldrich — Gravel Hill Praries
of Indiana 457
Victor Riemenschneider and Patricia Wiese Reed — Vascular Plants of Barker
Woods Nature Preserve, LaPorte County, Indiana 465
Psychology
A.M. Fullenkamp, Kim Duffy, Robert A. Vance and Robert Fischer — Marking
in Submissive Male Gerbils after Contact with a Dominant Male and His Odors* 47 1
Bonnie Gray, Robert Fischer and Gary Meunier — Heterosexual Social Interac-
tions in the Syrian Hamster* 471
Barbara Kane — The Several Themes of Adolescence* 471
Oliver C.S. Tzeng and Roberta Schlossmann — Psychovector Love Scale and
its Differentiability* 472
John M. Vayhinger— Orwell's 1984, Skinner's Walden II, Marx' Classless Society
and other Utopias: An Exploration of Human Expectation and the Psychological
Factors in a "Perfect Society" 472
Roger Ware and Charles Yokomoto — Personality Types and Perceptual-motor
Performance* 473
Walter Hartmann — Munro's Doctrines: A Forgotten Pioneer in Holism and
Hypnosis 475
*Abstracts
xii Indiana Academy of Science Vol. 94 (1985)
Science Education
Marshall P. Cady, Jr. — Ideas Concerning the Use of Computer Data Acquisition
Systems to Improve Teaching Effectiveness within the Laboratory* 483
Walter Cory — A New and Challenging Science Program from AAAS for Grades
7 and 8* 483
G. Earle Francq and Jerry M. Colglazier — Determining Needs: First Step for
Improving Science and Mathematics Instruction in Rural High Schools in North-
western Indiana* 484
D. Fabian Lozano-Garcia' and Roger M. Hoffer — The Layered Classifier: A
More Effective Method for Studying Seasonal Changes in Forest Cover Types
Using Satellite Data* 484
James George — Synthesis Experiments for High School Chemistry 485
Linda Hamrick and Harold Harty — The International Challenge: A Comparison
of Science Education Models from Four Nations* 485
Susan M. Johnson — A New Approach to Fostering Scientific Literacy among In-
diana's Secondary School Students* 485
Paul B. Kissinger and John A. Ricketts — Science Training for the Industrial En-
vironment (STIE)* 486
Rosalie Kramer — Field Biology: A Blow to Provincialism* 486
John Richard Schrock — Speaking of Sex — A Presentation on Terminology for
Students in Reproductive Biology Classes* 486
Richard E. Schuley and Marshall P. Cady, Jr. — Improving the Results of
Molecular Mass Determination Experiments by Using a Microelectronic Ther-
mistor Device* 486
Katharine Sessions — An Introductory Titration for First Year Chemistry Students:
A Comparison of Antacid Effectiveness* 487
Stanley S. Shimer — Using the Microcomputer to Teach Science in the Elementary
Classroom* 487
James T. Streator — Computer Aided Classroom Presentations in Chemistry* . 487
Albert A. Williams — Color Vision: A Lecture Demonstration of Afterimages* 488
Gary E. Dolph — CLIMATE: A Microcomputer Program Allowing Student Prepara-
tion of Climatic Maps for Indiana 489
L. Dwight Farringer, James T. Streator and Albert A. Williams — A Summer
Institute in Microcomputer Applications for Secondary School Science Teachers 499
K. Michael Foos — Use of a Microcomputer to Enhance the Coin Flip Probability
Exercise in the General Biology Laboratory 503
Lawrence Scharmann and Harold Harty — Two-year College Biology Instruc-
tors' Perceptions about their Role Expectations 509
Soil and Atmospheric Sciences
M.F. Baumgardner, N.N. Chaudhuri and S.J. Kristof — Land Cover Classifica-
tion of Rupgang Thana Dhaka, Bangladesh Using Landsat MSS Data* .... 517
William R. Gommel, Douglas W. Poad and John W. Wicker — Air Temperature
Fluctuation in Alabama During the Annular Solar Eclipse on 30 May 1984* 517
♦Abstracts
Table of Contents xiii
Paul Joseph and C.W. Lovell — Engineering Properties of Indiana Peats and
Mucks* 518
C.W. Lovell — Characterization of Indiana Soils by Porosimetry* 518
C.L. Rhykerd, S.E. Fowler, Alfonso de Almeida, A.M. Ferreira, Nuno
Moreira, C.H. Noller and J.L. Ahlrichs — Survey of the Mineral Composi-
tion of Forage Crops in Portugal* 518
C.R. Valenzuela, T.L. Phillips, M.F. Baumgardner and L.A. Bartolucci —
Soils; An Important Component in a Digital Geographic Information System* 519
J. A. Andresen, W.W. McFee, J.L. Ahlrichs and K.T. PawU— Wet Atmospheric
Deposition in Indiana 521
John T. Curran, Albert P. Shipe and Edward C. Yess — The National Weather
Service Rainfall Data Collection Network in Indiana 529
D.P. Franzmeier, H.M. Galloway and J.E. Yahner — Soil Survey in Indiana:
Past, Present and Future 533
Diane L. Klingle and David R. Smith — Gust Fronts in Doppler Radar Data . 547
T.E. Klingler and D.R. Smith — An Analysis of the 28 March 1984 Tornado Out-
break in the Carolinas 555
Ana L. Pires, J.L. Ahlrichs and C.L. Rhykerd — Response of Forage Crops to
Dolomitic Lime 565
Wayne F. Rostek, Jr. and John T. Snow — A Wind Tunnel Investigation of
Roughness Parameters for Surfaces of Regularly Arrayed Roughness Elements 571
John Richard Schrock and Jack R. Munsee — A Comparison of Soils on
Unreclaimed 1949 Indiana Coal Stripmine Surfaces in 1964 and 1981 579
Zoology
James D. Hengeveld — The Adaptive (?) Significance of Brood Reduction in the
Red-winged Blackbird (Angelaius phoeniceus)* 597
John B. Iverson — Patterns of Relative Fecundity in Snakes* 597
Mohinder S. Jarial — Light Microscopic and Ultrastructural Features of the Gut
of the Balsam Woolly Aphid, Adelges piceae Ratz* 597
Michael D. Johnson — Parental Investment in the Bee Ceratina calcarata Robert-
son (Hymenoptera: Xylocopidae): A Preliminary Study* 598
Michael P. Kowalski — Territorial Behavior in the Prothonotary Warbler, Pro-
tonotaria citrea, Between- and Within-season Territory Relocations* 598
James R. Litton, Jr. — A Record of the Freshwater Nemertean Prostoma graecense
(Bohmig) in Indiana* 599
James R. Litton, Jr. — Seasonal Abundance of the Psammic Rotifers of Spicer Lake,
Indiana* 599
William J. Rowland — Visual Signals in Sticklebacks: A Reexamination and Ex-
tension of Some Classic Experiments* 599
Sherman A. Minton — Venom Antigens in Oral Secretions of Colubrid Snakes* 600
Roderick A. Suthers — Physiology of Vocalization by an Echolocating Bird* . 600
Licia Wolf — An Experimental Study of Biparental Care in the Dark-eyed Junco* 601
'Abstracts
xiv Indiana Academy of Science Vol. 94 (1985)
Claude D. Baker, Bill J. Forsyth, Tom Wiles and D. Brian Abrell — Rediscovery
of the Spotted Darter, Etheostoma maculatum, in Indiana Waters: Blue River;
Crawford, Harrison and Washington Counties; Ohio River Drainage, USA 603
Virgil Brack, Jr., Ted T. Cable and Daniel E. Driscoll — Food Habits of Urban
American Kestrels, Falco sparverius 607
Richard L. Buckner, Melvin W. Denner, Daniel R. Brooks and Shareen C.
Buckner — Parasitic Endohelminths from Fishes of Southern Indiana 615
Wynn W. Cudmore — The Present Distribution and Status of the Eastern Woodrat,
Neotoma floridana, in Indiana 621
David L. Daniell — Occurrence of Swimmers' Itch in Northeast Indiana 629
Bill J. Forsyth, Claude D. Baker, Tom Wiles and Charles Weilbaker —
Cottonmouth, Agkistrodon piscivorus, Records from the Blue River and Potato
Run in Harrison County, Indiana (Ohio River Drainage, USA) 633
Thomas W. French — Dental Anomalies in Three Species of Shrews from Indiana 635
Thomas W. French — Reproduction and Age Structure of Three Indiana Shrews 641
Neil J. Parke and Charles E. Mays — Canine Dirofilariasis in Central Indiana 645
D. David Pascal, Jr. and John O. Whitaker, Jr. — Ectoparasites of Pine Voles,
Microtus pinetorum, from Clark County, Illinois 649
Ronald L. Richards — Quarternary Remains of the Spotted Skunk Spilogale
putorius, in Indiana 657
Ronald L. Richards and William R. Wepler — Extinct Woodland Muskox, Sym-
bos cavifrons, Cranium from Miami County, North Central Indiana 667
David M. Sever and Douglas Duff — Survey of the Fishes of the Kingsbury State
Fish and Wildlife Area, LaPorte County, Indiana 673
Marcus D. Webster — Heat Loss from Avian Integument: Effects of Posture and
the Plumage 68 1
Charles Weilbaker, Claude D. Baker, Bill J. Forsyth, Carl M. Christenson
and Ralph W. Taylor — The Freshwater Naiads, Bivalvia: Unionidae, of the
Blue River, a Southern Indiana Tributary of the Ohio River 687
Instructions for Contributors 693
Index 697
Proceedings
of the
Indiana Academy
of Science
Preface To The Centennial Volume
As President of the Indiana Academy of Science during this Centennial Year I am
honored to introduce this volume of its Proceedings.
During this special year I found it valuable to think back over the last hundred
years and to reconstruct what science must have been like over this period in general,
but also more specifically within Indiana. How much scientific knowledge and insight
do we take for granted today that did not exist even ten or twenty years ago, let alone
in 1885? Moreover, what current methods and support resources simply were unknown
or unavailable during the earlier days of our Academy?
Even more important than how much we have learned and grown is the realization
that today each of us is related to the many scientists who have preceded us in the Academy.
Moreover, all eleven hundred of us are the only link to the Academy's future. We join
our past colleagues in the spirit of science and in the scientific endeavor at a time when
our skills and dedication are needed more than ever throughout society. For science is
an essential part of today's world, and will be in the future. Our knowledge and abilities
can help alleviate many world problems, but they can also amplify them. Today we
desperately need a continuously concerned scientific community as well as a scientifically
literate society.
We stand on the threshold of the next hundred years of the Indiana Academy of
Science. Let us dedicate ourselves even more deeply to advancing science in ways that
also contribute to all the people of Indiana and the world!
Theodore J. Crovello,
President
Indiana Academy of Science
Indiana Academy of Science
Officers for 1984
President
President-Elect
Secretary
Treasurer
Director of Public Relations
Editor of PROCEEDINGS
Theodore J. Crovello
Department of Biology
The University of Notre Dame
Notre Dame, Indiana 46556
PHONE: (219) 239-7496
SUVON: +736 + 7496
Benjamin Moulton, Professor Emeritus
Department of Geography & Geology
Indiana State University
Terre Haute, Indiana 47809
PHONE: (812) 234-3870
SUVON: __ + 749 + 9 + 234-3870
Richard L. Conklin
Department of Physics
Hanover College
Hanover, Indiana 47243
PHONE: (812) 866-2151, ext. 348
SUVON: + 719 + 348
Duvall A. Jones
Department of Biology
St. Joseph's College
Rensselaer, Indiana 47978
PHONE: (219) 886-7111, ext. 214
SUVON: + 741+214
Walter A. Cory, Jr.
Coordinator for School Sciences
W. W. Wright 253
Indiana University
Bloomington, Indiana 47405
PHONE: (812) 335-5090
SUVON: + 703 + 55090
Donald R. Winslow
School of Education 201 A
Indiana University
Bloomington, Indiana 47405
PHONE: (812) 335-8658
SUVON: +703 + 58658
Officers and Committees 5
Committee Chairpersons and Special Appointments
I. Elected Committees
1. Academy Foundation Committee: William A. Daily, Chair (SUVON:
+ 9 + 251-4719)
2. Bonding Committee: Mary Lee Richeson, Chair (SUVON:
+ 710+5546)
3. Research Grants Committee: Uwe Hansen, Chair (SUVON:
+749 + 2429)
II. Standing and Ad Hoc Committees, and Special Appointments
4. Academy Representative to The American Association for the Advance-
ment of Science: Walter A. Cory, Jr. (SUVON + 703 + 55090)
5. Academy Representative to The Indiana Natural Resources Commission:
Damian A. Schmelz (SUVON: + 8970)
6. Auditing Committee: John A. Ricketts, Chair (SUVON: + 706 + 4607)
7. Biological Survey Committee: John A. Bacone, Chair (SUVON:
__ + 9 + 232 + 4052)
8. Centennial Program Committee: John B. Patton, Chair (SUVON:
+703 + 52862)
9. Constitution Committee: William R. Eberly, Chair (SUVON:
+ 729 + 309)
10. Editorial Board for the Proceedings: Donald R. Winslow, Chair (SUVON:
+703 + 58658)
11. Emeritus Member Selection Committee: Robert H. Cooper, Chair
(SUVON: + 732 + 0 + 288-9068)
12. Fellows Committee: William Melhorn, Chair (SUVON:
+ 755 + 43277)
13. Financial Planning: Frank Guthrie, Chair (SUVON: + 739 + 312)
14. High School Teacher Research Fellows Committee: Walter A. Cory, Jr.,
Chair (SUVON: + 703 + 55090)
15. Indiana Science Talent Search Committee: Walter A. Cory, Jr. Chair
(SUVON: + 703 + 55090)
16. Invitations Committee: Donald J. Cook, Chair (SUVON + 706 + 4601)
17. Junior Academy Council, Director: Cheryl Mason (PHONE:
219/924-7400)
18. Library Committee: Lois Burton, Chair (SUVON: __ + 9 + 253 + 7798)
19. Membership Committee: Duvall A. Jones, Chair (SUVON:
+ 741+214)
20. Necrologist: Fay K. Daily (SUVON: + 9 + 251-4719)
21. Newsletter Editor: Walter A. Cory, Jr. (SUVON: + 703-55090)
22. Nominations Committee: J. Dan Webster, Chair (SUVON:
+719 + 310)
23. Parlimentarian: Clarence Dineen (SUVON: + 743 + 4525)
24. Preservation of Natural Areas Committee: Marian T. Jackson, Chair
(SUVON: + 749 + 2489)
25. Program Committee: Philip A. St. John (SUVON: +704 + 9411)
26. Publications Committee: Benjamin Moulton, Chair (SUVON:
+ 749 + 9 + 234-3870)
27. Resolutions Committee: William Davies, Chair (SUVON:
+ 710 + 5535)
Indiana Academy of Science Vol. 94 (1985)
28. Science and Society Committee: Alice S. Bennett, Chair (SUVON:
+ 732 + 6875)
29. Speaker of the Year Selection Committee: Richard J. Jensen, Chair
(SUVON: + 743 + 4674)
30. Youth Activities Committee: Susan M. Johnson, Chair (SUVON:
+ 723 + 4043)
The Council
The Council consists of the officers plus the Chairperson of the Science and Society
Committee.
Council Members for 1984:
Theodore J. Crovello, President
Benjamin Moulton, President-Elect
Richard L. Conklin, Secretary
Duvall A. Jones, Treasurer
Walter A. Cory, Jr., Director of Public Relations
Donald R. Winslow, Editor of Proceedings
Alice S. Bennett, Chair of Science and Society Committee
The Executive Committee
The Executive Committee consists of the past presidents, current officers, chair-
persons of the sections, chairpersons of all committees, director of the Youth Activities
Committee, and representatives of affiliated organizations.
For current Executive Committee membership, see parts of this Directory listing
the above positions.
The Budget Committee
The Council:
Theodore J. Crovello, President (and Chair of Budget Committee)
Benjamin Moulton, President-Elect
Richard L. Conklin, Secretary
Duvall A. Jones, Treasurer
Walter A. Cory, Jr., Director of Public Relations
Donald R. Winslow, Editor of Proceedings
Alice S. Bennett, Chair of Science and Society Committee
Immediate Past President: Alice Bennett
Junior Academy Council Director: Cheryl Mason
Library Committee Chair: Lois Burton
Program Committee Chair: Philip A. St. John
Youth Activities Committee Chair: Susan M. Johnson
1984 Committees and Special Appointments
Elected Committees
1. Academy Foundation William A. Daily, Chair (1985)
John Ricketts (1984)
2. Bonding Committee Mary Lee Richeson, Chair (1985)
Donald Hendricks (1984)
Officers and Committees
Research Grants Committee
II. Standing and Ad Hoc Committees
4. Academy Representative to the
American Association for the Ad-
vancement of Science:
5. Academy Representative on the
Indiana Natural Resources
Commission:
6. Auditing Committee:
7. Biological Survey Committee:
8. Centennial Program Committee:
9. Constitution Committee:
10. Editorial Board for the
Proceedings
Uwe J. Hansen, Chair (1985)
Betty D. Allamong (1984)
John H. Cleveland (1986)
John O. Whitaker, Jr. (1987)
James F. Newman (1988)
Walter A. Cory, Jr.
Damian A. Schmelz.
John A. Rjcketts, Chair (1984)
Andrew G. Mehall (1983)
John A. Bacone, Chair
James Aldrich
Theodore J. Crovello
James R. Gammon
Donald Hendricks
Philip A. Orput
Victor Riemenschneider
Harmon P. Weeks
John Whitaker
John B. Patton, Chair
Alice S. Bennett
Walter A. Cory, Jr.
Fay K. Daily
William A. Daily
John F. Pelton
Philip A. St. John
William R. Eberly, Chair
William A. Daily
Clarence Dineen
Donald R. Winslow, Chair
Hans O. Andersen
Rita Barr
Ernest E. Campaigne
Donald P. Franzmeier
James R. Gammon
James H. Kellar
Benjamin Moulton
John F. Pelton
Carl C. Sartain
Alfred Schmidt
John O. Whitaker, Jr.
Bernard S. Wostmann
Frank N. Young
Indiana Academy of Science
Vol. 94 (1985)
1 1 . Emeritus Member Selection
Committee:
Robert H. Cooper, Chair
Harry G. Day
Howard H. Michaud
Winona H. Welch
12. Fellows Committee:
13. Financial Planning Committee:
William Melhorn, (1985) Chair
Stanley L. Burden (1986)
Richard Conklin (1984)
Della Cook ((1985)
Clarence Dineen (1986)
John J. Favinger (1984)
Robert Henry (1985)
Richard Jensen (1986)
James G. List (1986)
Robert D. Miles (1985)
John F. Pelton (1986)
Russell K. Stivers (1984)
Eugene D. Weinberg (1984)
Frank Guthrie, Chair
William A. Daily
Duvall A. Jones
John A. Ricketts
14. High School Teacher Research
Fellows Committe:
15. Indiana Science Talent Search
Committee:
16. Invitations Committees:
17. Junior Academy Council:
18. Library Committee:
Walter A. Cory, Jr. Chair
Alice S. Bennett
Ernest E. Campaigne
Cheryl Mason
Walter A. Cory, Jr., Chair
Jo Ann Jansing
Richard A. Mayes
Van A. Neie
Alfred Schmidt
Harold Zimmack
Donald J. Cook, Chair
Cheryl Mason, Director
William T. Anderson, Jr.
Michael Kobe
Virginia Rhodes
Carroll Ritter
Leota Skirvin
Jane Tucker
Lois Burton, Chair
James A. Clark
William A. Daily
John F. Pelton
Officers and Committees
19. Membership Committee:
20. Necrologist:
21. Newsletter Editor:
22. Nominations Committee:
23. Parliamentarian:
24. Preservation of Natural Areas
Committee:
25. Program Committee:
26. Publications Committee:
27. Resolutions Committee:
28. Science and Society Committee:
Duvall A. Jones, Chair
Robert H. Cooper
Walter A. Cory, Jr.
Marion Jackson
Susan Johnson
Jackson L. Marr
William Melhorn
Fay K. Daily
Walter A. Cory, Jr.
J. Dan Webster, Chair
Alice S. Bennett
William R. Eberly
Clarence Dineen
Marion T. Jackson (1984), Chair
James Aldrich (1986)
John A. Bacone (1985)
Carl H. Krekeler (1984)
Carrolle Markle (Honorary)
George Parker (1986)
Victor Riemenschneider (1984)
Robert C. Weber (1985)
William Weeks (1986)
Winona H. Welch (Honorary)
Philip A. St. John, Chair
Benjamin Moulton, Chair
Lois Burton
Donald J. Cook
Walter A. Cory, Jr.
William Eberly
Willis H. Johnson
J. Dan Webster
John O. Whitaker
William Davies, Chair
Edward C. Miller
Alice S. Bennett, Chair
William Beranek, Jr.
Stanley Burden
Lois Burton
Walter A. Cory, Jr.
Jon R. Hendrix
Gene Kritsky
Elden Ortmann
John Pelton
James Shuler
Edwin R. Squiers
Philip A. St. John
Charles Wier
Howard R. Youse
10
Indiana Academy of Science
Vol. 94 (1985)
29. "Speaker of the Year" Selection
Committee:
30. Youth Activities Committee:
Richard J. Jensen, Chair
Stanley L. Burden
Robert E. Hale
Thomas R. Mertens
Susan M. Johnson, Chair
Lloyd Anderson
Jerry Colglazier
Walter A. Cory, Jr.
Karl Kaufman
Michael L. Kobe
Cheryl Mason
Virginia Rhodes
John A. Ricketts
Stanley Shimer
Leota Skirvin
Jane Tucker
IAS 1984 Section Chairpersons and Chairpersons-Elect
Chairpersons
Donald Cochran
Department of Anthropology
Ball State University
Muncie, Indiana 47306
(317) 285-4927
Phillip E. Pope
Department of Forestry
Purdue University
West Lafayette, Indiana 47907
(317) 494-3590
Ralph Jersild
Department of Anatomy
Indiana University School of
Medicine
Indianapolis, Indiana 46202
(317) 264-8730
Shannon Lieb
Department of Chemistry
Butler University
Indianapolis, Indiana 46208
(317) 283-9410
Chairpersons-Elect
Anthropology
Diane Beynon
Department of Anthropology
Indiana University — Purdue
University at Fort Wayne
2101 Coliseum Boulevard East
Fort Wayne, Indiana 46805
(219) 482-5391
Botany
Austin E. Brooks
Department of Biology
Wabash College
Crawfordsville, Indiana 47333
(317) 362-1400 ext. 350
Cell Biology
Robert Stark
Department of Zoology
DePauw University
Greencastle, Indiana 46135
(317) 653-4776
Chemistry
Dennis G. Peters
Department of Chemistry
Chemistry Building, Room A 112
Indiana University
Bloomington, Indiana 47405
(812) 335-9671
Officers and Committees
11
Edwin R. Squiers
Department of Biology
Taylor University
Upland, Indiana 46989
(317) 998-2751 ext. 386
David D. Chesak
Box 883
St. Joseph's College
Rensselaer, Indiana 47978
(219) 866-7111
Paul Robert Grimstad
Department of Biology
University of Notre Dame
Notre Dame, Indiana 46556
(219) 239-5493
William Beranek
Indiana Center for Advanced
Research
120 E. 38th Street
P.O. Box 647
Indianapolis, Indiana 46223
(317) 264-2827
Edward Lyon
Department of Geography
Ball State University
Muncie, Indiana 47306
(317) 285-1761
Ecology
Richard W. Miller
Department of Zoology
Butler University
Indianapolis, Indiana 46208
(317) 283-9328
Engineering
William Stanchina
Department of Electrical Engineering
Notre Dame University
Notre Dame, Indiana 46556
(219) 239-5693
Entomology
James Haddock
Department of Biological Sciences
Indiana University-Purdue University
at Fort Wayne
2101 Coliseum Boulevard East
Fort Wayne, Indiana 46805
(219) 482-5254
Environmental Quality
Horst Siewert
Department of Natural Resources
Ball State University
Muncie, Indiana 47306
(317) 285-5790
Geology and Geography
John Cleveland
Department of Geology/Geography
Indiana State University
Terre Haute, Indiana 47809
(812) 749-2833
Gene Kritsky
Department of Biology
College of Mount St. Joseph
Mount St. Joseph, Ohio 45051
(513) 244-4401
History of Science
Gerald Seeley
Department of Civil Engineering
Valparaiso University
Valparaiso, Indiana 46883
(219) 464-5120
J. R. Garcia
Department of Biology
Ball State University
Muncie, Indiana 47306
(317) 284-4045
Microbiology and Molecular Biology
Mary Lee Richeson
Department of Biological Sciences
Indiana University-Purdue University
at Fort Wayne
2101 Coliseum Boulevard East
Fort Wayne, Indiana 46805
(219) 482-5546
12
Indiana Academy of Science
Vol. 94 (1985)
Vincent A. DiNoto, Jr.
Department of Physics
Indiana University Southeast
New Albany, Indiana 47150
(812) 945-2731 ext 390
Marion T. Jackson
Department of Life Sciences
Indiana State University
Terre Haute, Indiana 47809
(812) 232-6311
Physics and Astronomy
Ruth Howes
Department of Physics and
Astronomy
Ball State University
Muncie, Indiana 47306
(317) 285-5494 or 285-6268— Dept.
Plant Taxonomy
Victor Riemenschneider
Department of Biology
Indiana University South Bend
South Bend, Indiana 46615
(219) 272-8262
Psychology
Robert Fischer
Department of Psychological Sciences
Ball State University
Muncie, Indiana 47306
(317) 285-1713
Linda Hamrick
The Canterbury School
5601 Covington Road
Fort Wayne, Indiana 46804
(219 432-7776
David R. Smith
Department of Geosciences
Purdue University
West Lafayette, Indiana 47907
(317) 494-3285
Thomas Fogle
Department of Biology
St. Mary's College
Notre Dame, Indiana 46556
(219) 284-4675
Science Education
Gary Dolph
Department of Botany
Indiana University at Kokomo
Kokomo, Indiana 46902
(317) 453-2000
Soil and Atmospheric Sciences
Charles L. Rhykerd
Department of Agronomy
Purdue University
West Lafayette, Indiana 47907
(317) 494-8101
Zoology
James R. Litton, Jr.
Department of Biology
St. Mary's College
Notre Dame, Indiana 46556
(219) 284-4669 or 4671
HIGHLIGHTS OF THE SPRING MEETING
Photo courtesy of Whitewater Publications, Inc.
Members of the Indiana Academy of Science toured Brookville historical sites. John
Newman, former Brookville High School teacher, (far left) conducted the tour.
Indiana Academy of Science
CENTENNIAL YEAR
HISTORICAL HIKE OF BROOKVILLE
Friday, April 27, 1984
At the forks of the Whitewater river, early in Indiana history, were located
millstones. From these grew a town, strategically located to control and process produce
coming down the forks from interior farms. Thus by August, 1808, Brookville was
platted by Amos Butler and Jesse B. Thomas. Soon John Allen and Amos Butler were
platting additions. Why such rapid growth? The forks of the Whitewater served as
"roads" to hinterlands from Cincinnati in which settlers could venture north past the
National Road, or west, towards Indianapolis. Brookville's location quickly drew Hoosier
leaders and a Federal Land office. One can judge the prosperity of a town by its homes
and public buildings. By 1819 log structures were being replaced by brick made at
a factory located on the site of the high school, instead of being burned for each
individual project. Yet with the removal of the land office to Indianapolis in 1825
depression hit Brookville; much land, and even James Brown Ray's home, was in the
hands of the Bank of the United States at Philadelphia or in the hands of speculators.
With the passage of the Internal Improvements Act in 1836, prosperity returned to
Brookville with the building of the Whitewater Canal — new homes and businesses "o/
permanent improvements.'''' Such development began to wane by 1855 when the exodus
of younger members of families for the "West" offset progress. The Civil War brought
13
14 Indiana Academy of Science Vol. 94 (1985)
industrialization to the United States and several new factories to Brookville, as barrel
and paper making. The days of Scotus Gaul Picti, whose members believed that
"Brookville was the greatest town on earth," caused the last business building boon in
Brookville between 1890 and 1915. With the growth of urbanization, Brookville
has more and more been left with her last greatest resources, her history.
To tour Brookville, then, is to do so on her terms — leisurely and curiously. Not
to wonder why Governor James Brown Ray ever lived in Brookville, but to imagine
his coming out of his home to greet you. Or not to mumble about the lack of parking
places in town, but to wonder at the anger of the president of the Brookville National
Bank for the livery stable customers next door who took all the bank customers' spaces
at the hitching rail. Only then can Brookville' s secrets unlock to reveal her uniqueness,
fame and beauty.
1. BRACKEN HILL, in back of St. Michael's Church, was the site of a cabin built
by John Allen, a co-settler of Brookville. Part of his second home, built about 1808,
is still standing. Brick home was built by the Price family in 1837 and remodeled in
1937 by Al C. Brown. This home was long the residence of the Bracken family, William
being a veteran of the Civil War and a regional politician.
2. BIRTHPLACE OF LEW WALLACE— Lew Wallace, soldier, statesman, and author
was born in a home on this site, April 10, 1827. David Wallace, his father, was Governor
of Indiana from 1837 to 1840. His mother, Esther Test, was the daughter of John
Test, first Circuit Judge and Congressman. Marked by the Brookville Kiwanis Club.
3. ST. MICHAEL'S CATHOLIC CHURCH, First Catholic Church on this site in
1845. Present church dedicated March 25, 1862. St. Michael's School was founded
in 1855. First school building, 1875. Present structure erected in 1913. Remodeled in 1956.
4. SITE OF NOAH NOBLE HOME, 353 High, was the residence of Noah Noble,
governor of Indiana from 1831 to 1837. The original home became the Catholic rectory
in 1863 and was moved across the square when the present rectory was built in 1882-1883.
Later his home was torn down.
5. JAIL AND SITE OF OLD YELLOW TA VERN, 459 Main, in 1808 a blockhouse
was erected which in 1811 was converted into a "tavern" and hotel by James Knight.
After his death his wife, Mary, ran the tavern for many years. This structure was
torn down in 1861 and in 1882-1884, the present jail was built here.
6A. COURTHOUSE, Fourth & Main, is the second brick and fifth courthouse used
in Franklin County. The first was the "Yellow Tavern," on the site of the jail, then
a log courthouse on the public square, then in 1816-17 James Knight built a brick
structure which burned in February, 1852. The Lutheran Methodist Church on Fourth
St. was rented as a courthouse until the present one was completed in 1855. This structure
was extensively remodeled in 1912.
6B. VALLEY HOUSE, 450 Main, is on the site of the "Brookville Hotel," operated
by Andrew Wallace, father of Governor David Wallace. This frame structure was
destroyed by fire in 1852, and the present structure, the Valley House, was opened
in December, 1852. This hotel is reputed to be the oldest continuously operating hotel
in Indiana.
7. COURT STREET, in the rear of the Courthouse, contains buildings dating back
to circa 1819. Many law offices, the office of the Indiana American, and a marble
works, were located here in the 1840s through the 1870s.
8. HISTORY OF LOT 37, a business block burned out in February 1852. The buildings
which burned give an idea of the development of Brookville in 1852.
Highlights of the Spring Meeting 15
9. SITE OF OLD WHITE CORNER, 501 Main, was the site of the store of Nathan
D. Gallion, an early merchant of Brookville and competitor of Tyner.
10. RICHARD TYNER HOME AND SITE OF OLD GRINDSTONE CORNER,
512 Main, was the store and residence of Richard Tyner, an early merchant in Brookville.
Since he continuously advertised grindstones, he acquired this name for this corner.
1 1A. HITT-HOWLAND-FARQUHAR-GOODWIN home - Probably built by John
W. Hitt, father of George Hitt, Indianapolis newspaper editor, prior to 1840. Owned
by John Howland, father of Hewitt Howland, former editor of Harpers; John H.
Farquhar, Congressman and Lincoln elector in 1 860; and three generations of the Good-
win family, John R., Charles F., and John P. Goodwin. Originally it was a story
and a half house.
11B. WILLIAMS-BUTLER HOME, 911 Main, was built about 1820 for M.T.
Williams. It is an excellent example of Federal style structure, with a recently added
circular porch. It has long been the home of W.W. Butler, a leading naturalist and
prison reformer in Indiana.
12. O' BYRNE HOME, 912 Main Street, was built in the late 1850s for Judge Wilson
Morrow. This home is known as the home of three judges; in addition to Morrow,
Judge Ferdinand F. Swift and Judge Roscoe C. O'Bryne have lived here.
13. MATSON HOME, 914 Main, was built in 1842-43 for John A. Matson, a
businessman and Whig candidate for Governor in 1849. Matson lost the election locally
since he described himself as a poor man (oh, those ideals of frontier democracy!)
which contradicted the splendor of this home. His son, Courtland Matson, was a
Democratic candidate for governor in 1888. Note the leaded glass in the doorway.
14. "JAIL HOME," 1032 Main, was built in 1852-55 by Henry H. Remy, of brick
taken from the first brick courthouse, 1816, which burned in 1852, and the stone from
the old stone jail. An unusual feature is a stone with a ring in it, to which prisoners
were chained.
15. RYAN-STOUT HOME, 1038 Main, was built about 1878 in neo-Federal style.
This building excellently preserved, reveals the prosperity of Brookville in the 1870s
and 1880s with a large residence, a stone "cellar," a shed and carriage house stable.
16. WILLHITE TOURIST HOME, 1049 Main, was built in 1909 and is noted for
its "Stained Glass Windows" on the North. There is cherry woodwork and doors which
have hand carved designs on them. The home is also furnished with beautiful antiques.
17. HEASOM-COOKSEY HOME, 1001 Cliff was built for A.J. Heasom in 1881.
Heasom, a Civil War veteran, was a local politician and merchant in Brookville. Note
the footstone at the back door and the carriage house in the rear. A.J. Cooksey, a
noted woodcarver and contractor, lived here from about 1906 to 1955.
18. WILSON HOME, 1023 Cliff Street, this property was originally platted June
23, 1875 by Mrs. Jane McCarty. In 1916 it was purchased by Furman W. Hathway
who was a cigar manufacturer. In 1928 Fred and Ethel Rusterholz bought it and owned
it until August 22, 1975 when it was purchased by the present owners.
19. CHURCH OF CHRIST, Tenth & Franklin. This church was built in 1917. This
denomination was organized in Franklin County by Alexander Campbell in 1866.
20. SITE OF THE BROOKVILLE COLLEGE, Tenth & Franklin, on this site in
1819 was a brick factory. Later this two-acre site was transferred to the regents of
Brookville College who built a three story brick on the site of the present older high
school building between 1852 and 1855. This school was in use by the Methodists from
16 Indiana Academy of Science Vol. 94 (1985)
1855 to 1873, when it was sold to the town of Brookville for a public school building,
and the building was razed in 1912.
21A. JAMES BROWN RAY HOME, Tenth & Franklin, was built probably
in 1821, as Ray sold his home in September 1921 to William Noble: Lot 59: "on which
stands the house in which I now live." Noble sold this property to the Bank of the
United States in 1822, which James B. Lile, a teacher of the Franklin County Seminary,
and later of Centerville, bought, in 1833, "one lot of land with an old frame building
thereon called the Brookville Hotel on the north part of lot numbered thirty seven
. . . and the other lot of land with the old frame house on it called the Ray home
. . . being the same premises formerly occupied by Governor Ray and conveyed by
him by deed dated 1 -September 1821 to William Noble." The aristocratic frills on
the building nearly cost Ray the election in 1825.
2 1 B . OLD BROOKVILLE CHURCH AND CEMETER Y, Tenth Street — was built
in 1820-21 by the Methodists who held the property until 1830, selling it to the
Presbyterians who controlled it until 1848, when the Lutheran congregation bought it.
It was abandoned in 1923 and in the 1950s the Baptists held service in this church. The
bell tower was added in 1873. In the 1960s this edifice was restored by the Franklin County
Historical Society. The cemetery was first used in 1816 and was the only one until 1883
when Maple Grove Cemetery was opened.
22. FRANKLIN COUNTY SEMINARY— Fifth & Mill Streets, built in 1828-1831,
is one of four such county seminaries still standing of sixty-five buildings erected in
Indiana. It has been purchased by the Franklin County Historical Society for a museum.
23. LUTHERAN METHODIST CHURCH, Fourth & Franklin, was built in 1849
and was used as the courthouse from 1852 to 1855.
24. VALLEY CHRISTIAN CHURCH, 173 East Fourth Street, the first Christian
denominations came to Franklin County at the turn of the century. The Valley Chris-
tians were established seventeen years ago and the present structure replaced a struc-
ture which burned in 1981.
25. FIRST CHRISTIAN CHURCH, 123 East Sixth Street. The Christian denomina-
tions came to Franklin County at the turn of the century. The present structure was
built in 1848 by the Methodists and sold to the Presbyterians and since 1960 has been
renovated and used by the Disciples of Christ (First Christians) until the present time.
26. BROOKVILLE UNITED METHODIST, 150 East Eighth Street — The Good-
win family was instrumental in founding the Methodist denomination in Franklin County
The present building was built in 1866 with an addition to 1927.
27. ST THOMAS LUTHERAN CHURCH. Ninth & Franklin, built in 1920 and
dedicated in 1924. The Lutherans first organized their church in Franklin County in 1848.
28. LITTLE CEDAR GROVE BAPTIST CHURCH— Oldest church on original site
in Indiana. First services held August 1, 1812. Tradition says that a blockhouse was
erected for protection while the church was being built. Clay for the bricks kneaded
by oxen on this site. Restored by Franklin County Historical Society, the National
Society of Colonial Dames of Indiana, and Franklin County, 1952-1955.
29. THE HERMITAGE, in back of the ball park, tour of the house and grounds.
Oldest part was built by James Speer in 1817 and the wings were added about 1898.
James Speer had a mill just to the southeast which was torn down in 1905. The Hermitage
was the center of a Franklin County art colony, under the direction of J. Otis Adams
and Theodore Steele at the turn of the century. Home was badly damaged in the 1913
flood.
Highlights of the Spring Meeting
17
& &(D
Our Brookville Bond
Fay Kenoyer Daily
Butler University
Box 169
Indianapolis, Indiana 46208
To start our centennial year, a return to the place where our society was con-
ceived seemed appropriate to honor our founder, Amos Butler (Figure 1), who lived
here, and to enjoy the beauty of this region.
Figure 1. Amos William Butler.
Brookville was settled in 1804 by Amos Butler, Senior. His cabin was built just
north of the Hermitage site, and he soon built a flour mill on the river bank nearby.
In 1908, he platted Brookville with the help of Jesse B. Thomas. Many prominent
men have lived here including Amos W. Butler, grandson of the senior A. Butler.
Amos William Butler was born in Brookville in 1860 and was educated in local
schools. He attended Brookville College, Hanover College and Indiana University where
he received an A.B. in 1894, M.A. in 1900, and L.L.D. in 1922. He received an L.L.D.
from Hanover also in 1915. The Brookville College mentioned above used to be located
on Main Street now the site of the Franklin County High School.
Butler was influenced early in life by the well-known naturalist, Dr. Rufus Hay-
mond. As time passed, Butler found other members of the community interested in
scientific subjects including Edgar Quick, Oscar Meynke, Charles F. Goodwin, Rev.
David R. Moore and Clifford R. Case. The Rev. Moore arranged for lectures, some
of which were scientific, in his Presbyterian Church. The lectures were so successful
that Amos Butler discussed with the Rev. Moore the subject of forming a scientific
society. Several people met at the residence of Rev. Moore in 1881 and formed the
Brookville Natural History Society. The Rev. Moore was chosen president; Charles
Goodwin, a banker, was vice-president; Amos Butler, secretary; Edgar R. Quick, editor
of the Franklin Democrat, was correspondent; and John T. Rehme was treasurer. The
group met over the stove store of Emerson Rockafellar for awhile. Evidently, Butler
was co-owner of a tinning business with Rockafellar and Goodwin was influential in
obtaining the Waltz Mansion for their later use. The Brookville National Bank owned
the mansion and donated the parlor for a museum and meeting room. Many promi-
nent Indiana scientists came to lecture at meetings. Among them were David S. Jordan,
professor of zoology at Indiana University; John M. Coulter, Wabash College pro-
fessor of botany; David Worth Dennis, professor of biology at Earlham College; John
Highlights of the Spring Meeting 19
P.D. John, president of Moore's Hill College; Barton W. Evermann, naturalist fur-
thering his education at Indiana University at the time; and Stanley Coulter, botanist
and conservationist, lawyer and teacher at Coates College.
Amos Butler did not confine himself to these scientific activities, however; he
attended American Association for the Advancement of Science meetings where his
circle of scientific friends expanded. It was at one of these meetings that he called
a conference to discuss forming a state-wide organization. He had been corresponding
with other Indiana scientists to see how they felt about it. In his words (2), "In my
endeavor to obtain information on zoological subjects, I corresponded with a number
of scientific men in the state. The results were often very unsatisfactory. I found the
experience of other men the same; so I began thinking of some way to bring those
interested in science in the state together to get acquainted and exchange experiences.
To that end, correspondence was undertaken with John M. Coulter, Charles R. Barnes,
Daniel Kirkwood, T.A. Wylie, David Starr Jordan, Stanley Coulter, R.B. Warder,
Philip S. Baker, O.P. Jenkins, David W. Dennis, J. P.D. John, Richard Owen and
others. Most of these gave favorable responses."
At an 1885 AAAS meeting, Butler met with a number of scientists who decided
that the Brookville Natural History Society, as the most active society in the state,
should call a state meeting of scientists. A committee should be appointed by the
Brookville society to attend to the details of planning. The Rev. Moore, S.P. Stoddard
and Amos Butler were appointed to the committee. Our Academy came into being
December 29, 1885, at the Marion County Courthouse in Indianapolis, Indiana, with
Dr. J. P.D. John of DePauw University presiding and Amos Butler as secretary. At
the meeting, David Starr Jordan was elected president; J.M. Coulter, J. P.D. John
and Rev. Moore as vice-presidents; Amos Butler, secretary; O.P. Jenkins, treasurer;
and J.N. Hurty, librarian. Amos Butler presented a constitution which he had written.
A faded pencil-written copy of it can be found in the Indiana Academy of Science
files in our library housed at the State Library Building. The constitution was adopted
with a few changes and charter members signed a roster attached finally to a typed
copy of it. It was decided at the meeting that those persons who signed their names,
had their names signed to the constitution and by-laws at this meeting or had requested
that they be considered original members should be considered charter members.
Evidently 46 persons attended the meeting judging by the signatures. Mr. W. de M.
Hooper signed for Dr. Henry Jameson making 47 signatures at the meeting. Seven
signatures were added by the treasurer with a notation, "by order of the executive
committee" making 54 names of charter members. No later changes in this decision
for determining charter members could be found in the minutes. The constitution and
by-laws with the list of signatures were sent to the secretary, Amos Butler, on January
4, 1886, (less than a week after the meeting) by O.P. Jenkins, treasurer, with a letter
signifying that these were the charter members. The letter became separated from the
signature list and constitution. These and the minute books were unavailable for a
period. They were all brought together and the charter members verified during the
recent writing of an Academy history. The materials are all in the State Library Indiana
Section archives now.
The first Indiana Academy of Science field meeting was scheduled the following
May 20-21, 1886. In Butler's words (1), "It was fitting that the first 'Field Meeting'
of the Indiana Academy of Science should be held at Brookville. There the idea of
such an organization originated. There the steps were taken through the Brookville
Society of Natural History by which scientific investigators of the state were brought
together at Indianapolis, December 29, 1885, to adopt articles of association and ef-
fect an organization."
The Brookville field meeting participants assembled in the town hall for evening
20
Indiana Academy of Science
Vol. 94 (1985)
£ji£
Figure 2. Town Hall, Brookville, Indiana.
A. (left) in 1886. B (right) in
1984.
,1
meetings. The hall as it appeared then is shown in Figure 2A. Figure 2B shows its
present appearance without the upper story and steeple. It is located on Main Street
by the courthouse between Fourth and Fifth Streets. It gave me a feeling of awe as
I stepped into the halls to think that some of our charter members assembled there
almost 98 years ago for a meeting. Amos Butler lists 33 people registered at that first
spring meeting of our society. Among them were 15 charter members. With 7 new
members taken in at that meeting, the Academy had 61 members at that time. Of
that meeting Butler (2) wrote, "Some of us recall a few events of the first meeting —
how disappointed we were that John Coulter was unable to be present for one of
the principle addresses; how well his place was filled by David W. Dennis; that Ever-
mann and others of the Indiana University crowd drove through in a carriage and
on the way made a collection of fishes in the western part of Franklin County. We
had no Kiwanis Club to make such splendid arrangements — but the Brookville Society
of Natural History and its friends did the best they could. . . ."
Perhaps the next speaker will tell you more about that first field meeting as much
of it involved our first president, David Starr Jordan. Suffice it to say, the meeting
was a great success as was the second spring meeting in Brookville in 1923.
Some of the men mentioned here, many of whom were very young when the
Academy was organized, became internationally known. Their fellowship and shared
enthusiasm for their work and this organization, undoubtedly, had much to contribute
to this. Amos Butler was not only known for his scientific pursuits, but they were
interwoven into social work on the State Board of Charities. In the Academy, he serv-
ed as secretary for the first eight years and later was president. In Dean Stanley Coulter's
(3) memorial tribute to him, it was stated that, "The Indiana Academy of Science
Highlights of the Spring Meeting
21
came into being and was kept alive during its earlier years because he refused to allow
it to die. It would not have been born, nor would it have lived, without Butler's genius
for organization — ."
Amos Butler was president in a number of other scientific organizations as well
as the Academy. He not only was founder of this organization and the Brookville
Natural History Society, but also eight more including the Indiana Audubon Society.
There is a good biography of him by Dr. Barton W. Evermann (4) in the Indiana
Audubon Society 1932 Yearbook dedicated to Amos Butler. His bibliography is included.
Our society did not return to Brookville for a meeting after 1923. With the growth
of the society and lack of adequate facilities in Brookville, we came down as far as
the Mary Gray bird sanctuary instead. We just had to strengthen our bond of fellowship,
our Brookville bond, if you will, by returning here in our centennial year.
As Amos Butler said so eloquently at the end of his talk to members of the
Academy the first time we returned here for a meeting in 1923, "Here we are back
to the place where it originated — to Brookville — hail all hail." Or if you prefer today's
vernacular, Hello out there, Brookville, we have returned!
3.
4.
Literature Cited
Butler, Amos W. 1892. Field Meetings. Proc. I.A.S. (for 1891) 1: 9-13.
Butler, Amos W. 1924. Early history of the Indiana Academy of Science. Proc.
I.A.S. 33: 14-18.
Coulter, Stanley. 1938. Memorial tribute to Dr. Butler delivered at the general
session of the Academy, November 5, 1937. Proc. I.A.S. 47: 25.
Evermann, Barton Warren. 1932. Amos William Butler. Indiana Audubon
Soc. Yearbook.
Gary A. Sojka and Fay K. Daily
The Making of David Starr Jordan
Gary A. Sojka*
President, Bucknell University
Marts Hall, Lewisburg, Pennsylvania 17837
On May 21, 1886, the field meeting of the Indiana Academy was called to order
at the town hall in Brookville. David Starr Jordan presided. At that meeting, Jordan
presented two papers: one on Darwin and the other on "How To Go Fishing." In a
later address, his presidential speech to the American Association for the Advancement
of Science in 1910, Jordan talked on "the making of Darwin." He listed the elements
that came together to influence Darwin and helped to prepare him for his great work.
Among these were observations on Darwin's genetic makeup or his "inherent self," his
contact with nature, and, finally, the influence of an inspiring teacher.
In discussing "the making of Jordan," I will examine these same elements. His
*At the time of this address, Gary Sojka was Dean, College of Arts and Sciences, Indiana University, Bloom-
ington, Indiana 47405.
22
Highlights of the Spring Meeting 23
genetic and family background, of course, is no longer easy to analyze; consequently,
I will place more emphasis on the events and mentors that shaped and influenced this
pioneer educator, scientist, internationalist and pacifist.
Jordan was born in 1 85 1 in Gainesville, New Yor k , and was christened simply David
Jordan. The Starr came later. He was the fourth of five children born of Puritan stock,
originally from Devon. His grandfather fought in the Revolutionary War and his father,
Hiram Jordan, was born in 1809, the same year as Lincoln and Darwin. He worshipped
his older brother, Rufus Bacon Jordan, who was 13 years his senior. Rufus went off
to Washington to enlist in 1862 and immediately contracted "Army Fever" and died.
Jordan experienced deep and intense feelings of loneliness for years afterwards. He later
stated that it was at the moment of hearing of his brother's death that he realized the
terrible tragedy of war. Those feelings stayed with him throughout his life and he dedicated
his 1907 book, The Human Harvest, which dealt with the biological effects of war, to
his late brother.
In his childhood he invented for himself an imaginary hero that he named David
Emanuel Starr. It was as a young adult that he added the middle name Starr — an act
that gives some insight into the level of Jordan's self-esteem. As a boy growing up in
a rustic environment, he developed interests in natural history. His earliest scientific obser-
vations were of the stars. He mapped the celestial bodies and endeavored to learn their
names. By the age of 13 he had a complete set of handdrawn star charts. He next turned
to terrestrial geography. He was fascinated by maps, and as a result, he longed to travel
at a very early age. Apparently his first interest in zoology grew from his responsibility
around the Jordan farm to tend the flocks. From his experience in caring for diseased
sheep and from his brother's death, he developed a strong interest in the disease process
and in infection. His first scientific notes were published in The Cornell newspaper, fol-
lowed quickly by a botanical paper in the American Naturalist and a paper on hoof rot
in sheep in the Prairie Farmer, all in 1871. In addition to his interest in microbiology
and zoology, botany fascinated the young Jordan. While yet quite young he developed
a scientific as well as an esthetic interest in the flora that surrounded him. He combined
his cartographic and botanical interests and began to observe, record, and map the biota
of different locales on and about the Jordan homestead.
At age 14, in his own words, "being considered a youth of promise and otherwise
apparently harmless," he was admitted to the female seminary of Gainesville run by two
young women recently come from Mount Holyoke. There he learned French, but his primary
literary interest, not surprisingly, was Thoreau. It was during this time that Jordan first
developed his life-long interest in baseball. The boys of the town organized a ball club
called the Gainesville Suaves. They dressed the part with bright red pantaloons and ac-
companying oriental panache. It was in a baseball game that Jordan broke his nose in
a violent collision while chasing a flyball. This accident produced his most distinctive
physical feature as later portraits show. Baseball remained an important part of Jordan's
entire life. He was involved early and seriously in the question of whether a curve ball
actually changed its direction of flight or was an optical illusion. He also took great pride
in being able to hit with what he considered remarkable power.
Jordan originally had planned to go to Yale University. However, he won a Cornell
scholarship and he entered that new school on March, 1869, with $75.00 in his pocket.
Cornell University was seven years old at that time. Significantly, he had the same $75 .00
upon graduation three years later, thus proving that wealth was not needed to complete
an education at Cornell as it would have been at other eastern schools of that era. This
fit, of course, with Ezra Cornell's desire that a young man should be able to work his
way through school, either through manual labor or help to the university.
By 1870, at the age of 19, he already had become a lecturer in botany. He was also
a very clubby person. He lived at the Grove Boarding Club, became an early member
24 Indiana Academy of Science Vol. 94 (1985)
of Delta Espilon, and later joined the AAAS and Sigma Xi, which was founded at Cornell
in 1896 with Jordan as a charter member. In his autobiography Jordan goes into con-
siderable detail discussing the merits of the college fraternity system.
The early days at Cornell were marked by crude facilities and cramped quarters,
but Jordan noted that they were characterized by a kind of enthusiasm and pioneer spirit
that are hard to maintain in days of prosperity. "At the time we were all young together,
freshman students, freshman professors, freshman president, without experience or tradi-
tion to guide or impede, but we had youth and we had truth and not even the gods have
those," he observed in his autobiography. But, whatever were the conditions at Cornell,
Jordan met the two persons who would prove to be his most influential
teachers: Louis Agassiz from Harvard, a non-resident professor and the most famous
naturalist of his time, and Andrew Dixon White, the young and vigorous president of
the institution. Jordan later went to study with Agassiz who molded his views in science.
White, the enthusiastic and energetic administrator, probably had a greater influence
than any other person on Jordan as an educator. White's view on co-education and cur-
riculum design greatly impressed Jordan. He became noted in these areas at both Indiana
and Stanford, where one can still see the influence of White. White also put great em-
phasis on attracting outstanding scholars. Those he could not get on the Cornell payroll,
he brought in as non-resident professors. Among them, in Jordan's time, were Agassiz
and James Russell Lowell. Jordan appreciated the effect that such persons had on the
young, serious scholar, so he endeavored to "bring in the masters" whenever he was
in charge. The concept of developing strong and autonomous academic departments also
derived from his Cornell experience.
Jordan majored in botany and minored in geology and zoology. He became compe-
tent in French, German, Italian and Spanish. He later added Norwegian and, in his own
words, "some Chinese." Again, we see the interest in things international, for this language
preparation would later make him a facile traveler in Europe. He left Cornell in 1872
with a Master of Science. His Master's thesis was on the " Wildflowers of Wyoming County
in New York." In 1886, he also was awarded the Doctor of Laws degree from Cornell,
which he declined because he didn't believe in honorary degrees.
Upon graduation he had to make a career choice between botany or sheep husban-
dry. He chose the former and took a $1,300 per year appointment at Lombard College
in Galesburg, Illinois. This institution was later absorbed by the much larger and better
endowed Knox College. On the way to Galesburg, he attended his first scientific society
meeting. The AAAS was meeting in Dubuque, Iowa. One of America's great scientists,
Asa Gray, was in attendance. Jordan was mightily impressed by the social and intellec-
tual atmosphere. As noted, he later became the President of the AAAS, but it was this
first meeting which clearly indicated to him that scientific meetings were a valuable exer-
cise and ought to be regularly attended.
Jordan did not do well at Lombard, which he felt suffered from small and narrow
thinking. He was probably fired, but, in any case, he left after one year. He next went
to Appleton Collegiate Institute as principal. It was there that he turned to the study
of fishes because of their relative abundance and ease of study. Appelton Collegiate In-
stitute was then absorbed by the much larger Lawrence University.
From Appleton, he went to the high school in Indianapolis to be in charge of natural
science. The following are his first impressions of the city of Indianapolis. "The capital
of Indiana at first sight seemed singularly monotonous, being perfectly level and laid
out in regular squares around a central circle. The streets, moreover, were lined with silver
maple, a second-rate shade tree, which do not appeal to me. The people said I would
learn to love the town; as a matter of fact, I did! Among other reasons because it contained
an unusual number of clear-headed and broad-minded citizens." Among those good citizens
were William Henry Harrison, James Whitcomb Riley, and Dr. William B. Fletcher,
Highlights of the Spring Meeting 25
a pioneer in the humane treatment of mental disorders.
In the summer of his first year at the Indianapolis high school, he was elected without
warning, according to him, to the professorship of biology at the nascent Northwestern
Christian University at Irvington. Before it even opened its doors, it was renamed Butler
University. In 1878-79, a schism occured at Butler between those who wanted a progressive
scholarly insitution tied to the main trends in American scholarship and those who wanted
closer ties to the Christian Church. As a consequence, faculty began to leave. Jordan's
colleague, A.W. Brayton, tried to secure a post at Indiana University. Jordan may have
done a "Miles Standish" on his colleague by going to Bloomington to testify on his behalf.
As a result of his efforts on behalf of his friend, Jordan, rather than Brayton, was offered
the position at Indiana University. Jordan was very critical of what he found in Bloom-
ington. He had little good to say of the clergymen who headed the place. It was rural
and back woodsy and suffered from a fixed curriculum. There were a few things that
appealed to him, however. He had great respect for four of the faculty: Daniel Kirkwood,
Theophilus Wiley, Elisha Ballantine, and Richard Owens. He also appreciated the "Bates
School of Philosophy." It seemed there existed a unique and quaint situation in Bloom-
ington for Henry S. Bates, the local shoemaker, who, though possessing little formal
education, loved literature and philosophy and talking with students. Indiana University
students could be found gathering around his shoeshop at various hours of the day and
early evening discussing natural philosophy and literature among themselves and with
the cobbler, "Professor" Bates. From the time Jordan joined the Indianapolis high school
until the time he left for Stanford, he spent 17 years in Indiana. In that time he visited
each of the 92 counties and claimed friends and acquaintances in each of them.
In the summer between his leaving Galesburg and the beginning of his appointment
in Appleton, he went to Penikese Island to study with Agassiz, who had a wonderful,
revolutionary idea for a summer school of science for teachers of science and natural
history. The summer session was one response to the perception that there was a crisis
in the schools concerning math and science education. This was the first educational ex-
periment of its kind in the United States, and Jordan was privileged to be a part of it.
The Penikese experiment was called the Anderson School of Natural History. The first
class was composed of 35 men and 15 women living together in make-shift quarters and
separated from each other simply by sheets and other cloth materials hung in the middle
of the room. Agassiz felt that this was a missonary work of the highest importance because
he believed he had gathered around him people who had great influence on the young
of the nation.
Agassiz was a gifted observer and naturalist; he was not a Darwinian evolutionist,
but rather a natural theologist. He tried to build a bridge between religion and science.
He felt that his own studies were "just glimpses into divine plans," and that, as he noted
in 1887, "Our task is complete as soon as we have proved His existence." Agassiz died
the first winter after the Penikese summer. The Anderson School opened again the following
summer, but this time under the direction of Louis Agassiz's son, Alexander. Jordan
was again present. Mottos and slogans of Agassiz were printed on bed sheets and hung
on the walls of the crude shelterhouse in an effort to try to make the presence of the
master a little more real. Some of these slogans, such as "study nature, not books,"
"strive to interpret what really exists," "be not afraid to say I don't know," hung around
the room. After the second summer, the Anderson School closed forever. Interestingly,
15 years later Jordan's student, Carl Eigenmann, took those banners from the walls of
the crude shelterhouse to Woods Hole, the natural successor to the Penikese experiment.
It seems strange to us today that Jordan, so prominently remembered among the
early Darwinists as a contributor to the modern evolutionary theory, should have been
so keenly influenced by Agassiz, a believer in theistic evolution. However, at that time,
there was a number of differing views on the formation and mutability of species, Darwin's
26 Indiana Academy of Science Vol. 94 (1985)
natural selection theory being only one of them. Natural selection was interpreted by
the Darwinists of that time as gradual preferential survival of creatures with slight positive
variations. Theistic evolution, as espoused by Agassiz and his followers, stated that variation
occurs; however, it is not random but rather directed toward some purposeful end by
a creator's will. Probably the greatest intellectual challenge to Darwinian natural selection
at that time was Lamarckism. Only one aspect of Lamarck's earlier theory applies, of
course, that being the inheritance of acquired characteristics. Interestingly, Lamarck's
theory was put forth in the year 1809, the same year that Darwin, Lincoln and Jordan's father
were born. In this theory, characteristics acquired in the lifetime of the organism are supposed
to be passed on in some way to the offspring. Another prevalent idea of the day was
orthogenesis — evolution consistently directed along a single path by forces originating
within the organisms themselves. These involuntary trends were thought to unfold without
reference to the demands of the environment. And, finally, there was the so-called mutation
of significantly new forms. The mutations occur at random and are non-adaptive. In
the late 19th century, mutations were thought to create new populations instantaneously
which were separate and distinct from the originals, a rather different interpretation than
we have today.
The following quotation describes the influence that Agassiz's teaching had on Jor-
dan's intellectual development with regard to biological evolution:
Agassiz had no sympathy with the prejudices exploited by weak and foolish
men in opposition with Darwin's views. He believed in the absolute freedom
of science and that no authority whatever can answer beforehand the ques-
tions we endeavored to solve. An attitude strikingly evidenced by the fact that
everyone, especially trained by him, afterward joined the ranks of the evolu-
tionists. He taught us to think for ourselves not merely to follow him. This,
though I accepted his philosophy regarding the origin and permanence of species
when I began serious studies in zoology, as my work went on there imper-
manence impressed me more and more strongly. Gradually, I found it impossible
to believe that the different kinds of animals and plants had been separately
created in their present forms. Nevertheless, while I pay tribute to Darwin's
marvelous insight, I was finally converted to the theory of divergence through
natural selection and other factors, not by his arguments but rather by the
special facts unraveling themselves before my own eyes. The rational meaning
of which he had plainly indicated. I sometimes said that I went over to the evolu-
tionist with the grace of a cat the boy leads by its tail across the carpet.
All of Agassiz's students passed through a similar experience and most of them
came to recognize that in the formation of every species, at least four elements were in-
volved, those being the resident or internal factors of heredity and variation and the ex-
ternal or environmental ones of selection and segregation. Actually, by 1869, or 10 years
after the publication of The Origin of the Species, acceptance of evolution was widespread
and firmly established. What was in doubt, and still is, is the precise mechanism of evolution.
Darwin favored sympatric natural selection: slow, gradual selection of better fit individuals
in the population until a new species finally emerges, formed alongside the original which
may or may not then become extinct. The modern synthesis with Mendelian genetics
gave a mechanism for the insertion of variation into the scheme. However, the fossil record
does not accord easily with his model since it appears more abrupt or punctuated. Darwin
himself decried that fact and tried to explain it away by pointing out that the fossil record
was probably incomplete. It was some of Jordan's work on minnows and darters and
other non-migratory fish that first suggested the importance of geological isolation as
a crucial factor of allowing the buildup of differentiating factors in the speciation process.
This concept, which later became known as "Jordan's Law," has been incorporated in
the modern theory of punctuated equilibrium.
Highlights of the Spring Meeting 27
In his years at Indiana University, Jordan continued to do field work on fishes.
He involved students in his work and considered this as part of his teaching. Significantly,
he never gave up that enterprise during his years as President at Indiana or Stanford.
He always taught and carried on research. His interest in travel continued while at Indiana
University. He spent a good deal of time on the West Coast doing a systematic study
of the fishes of the Pacific Coast for the United States government. He also began to
go regularly to Europe with students. His command of languages and his training as
a naturalist made him an excellent tour guide for students. On one of his trips he and
some of his students climbed the Matterhorn. This adventure gave him a popular subject
for the numerous lectures that he gave around the state of Indiana. These frequent Euro-
pean visits also convinced him of the foolishness of protective tariffs. He tended to believe
in a world community and once wrote a satiricial essay about protective tariffs. He attended
international scientific meetings whenever possible, and ultimately brought great distinc-
tion to the fledgling Indiana University when he won the gold medal at the First International
Fishes Congress, held in London in 1883, for his collective writings on the "Taxonomy
of Fishes."
In 1884, Lemaul Moss resigned and Ballantine was made the acting president of
Indiana University. Jordan succeeded him in 1885. He had to find money for new buildings,
he had to battle the legislature for funds, and he had to convince the population of a
backward state that a college education was worth having. Some things change only slowly.
Much of what he liked best about Cornell he brought about at Indiana. In 1881
as a faculty member, he tried to introduce a new curriculum with electives but he failed.
In 1 886, as President, he did away with a fixed curriculum and introduced a strong depart-
mental system, electives and a major professor system for indepth study in the last two
years. He often pointed with pride to Carl Eigenmann, who took biology instead of Latin
in this new system, as one of the successful examples of his educational initiatives. Jordan
made a major effort to expose students to the masters. He worked hard to assemble a
fine faculty in Bloomington and, later, he hurt Indiana University badly when he took
most of the best with him when he left for Stanford. He also tried to bring impressive
and important personages to campus, not the least of those being the young naturalist,
Theodore Roosevelt.
In the foregoing, I have made a modest attempt to find the elements in Jordan's
background and experience that equipped him for his role as scientist, internationalist,
antiwar advocate, and pioneer educator. There can be little doubt that both Stanford
and Indiana Unviersity benefitted from the influence exerted on the young Jordan by
Andrew Dixon White in his years at Cornell. It is also clear that Agassiz, though holding
opposing views himself, set Jordan on the path to important scientific contributions.
Jordan's early experiences with social and scientific societies made him a willing and
able leader and developer of the important scientific and educational institutions springing
up around him.
Perhaps one of the best indications of the power of these early influences on Jordan's
subsequent career comes from an oft-told Jordan anecdote. It is said that Jordan always
recognized the students at Indiana and called them by their given names. Some years
after Jordan left Bloomington to be President at Stanford, an Indiana University graduate
from the Jordan era was visiting in Palo Alto. As they strolled around the Stanford
campus, Jordan nodded and smiled at the Stanford undergraduates as they passed by
but he did not acknowledge them by name. When the I.U. man inquired about this, Jordan
reportedly replied, "Every time I remember the name of an undergraduate, I forget the
name of a fish." I wonder if Jordan also remembered an Agassiz quotation emblazoned
on one of those bed sheets that Eigenmann later carried to Woods Hole: "The memory
must not be kept too full or it will spill over."
28 Indiana Academy of Science Vol. 94 (1985)
Reference Materials
Days of A Man
The Eclipse of Darwinsim
Archives of Indiana University
Highlights of the Spring Meeting 29
Geology field trip, April 28, 1984
Curtis H. Ault and John R. Hill, Leaders
About 15 geology enthusiasts traveled by car and van through the valley of the
Whitewater River, along the historical Whitewater Canal, and among the wooded hills
from Brookville westward past the old milltown of Metamora to view and examine
the landforms and rocks that form the scenic region near Brookville. Bold topographic
relief, the result of differential erosion of glacial deposits and bedrock, gives the
Brookville area a rugged appearance that reflects the character and self reliance of
both the founders and the present-day population of Franklin County.
Examination of the rocks exposed in the Derbyshire Quarry, about 15 miles west
of Brookville, revealed qualities of the local bedrock that has given the region its
reputation for durable building stone since the early 1800s. The flaggy limestone of
the Laurel Member of the Salamonie Dolomite (Silurian) is now being quarried at
Derbyshire for building stone that is used for veneer in Indiana and Ohio, and for
crushed-stone aggregate for local use. Limestone of the underlying Osgood Member
of the Salamonie, and the Brassfield Limestone (Silurian), which has a distinctive brassy
color and coarse chrystallinity, are also quarried at the Derbyshire operation.
The overlying glacial deposits were evident along the field trip route, particularly
as valley-train terraces flanking the Whitewater valley. About 15 feet of Illinoian or
older till, containing at least one and probably two paleosols, are exposed above the
bedrock in the Derbyshire quarry.
The small Derbyshire Falls, which was recognized in the early literature, is visible
from the quarry, but the valley containing the falls appears to be in imminent danger
of flooding or burial with waste overburden from the quarry. Participants of the field
trip had the opportunity to ponder the alternative of placing the quarry at this locality
to the eventual detriment of the scenic setting of the falls versus the possible greater
cost of importing crushed stone from more distant points.
Mr. Charles Holzhause, a second-generation stone merchant, discussed the value
and the use of a variety of building stone that he sells from his stone yard at Metamora.
His building stone included several varieties of the Laurel limestone that we examined
in the Derbyshire Quarry. Mr. Holzhause emphasized the use of a large number of
rock types for different kinds of construction and to satisfy individual personal tastes.
We expanded our concepts of how veneer and other building stone could be used and
displayed and gained an appreciation for the effort and expense that go into this type
of masonry.
All participants of the trip became avid paleontologists at the last stop, where
a wide variety of fossils had weathered out of a 100- foot section of Dillsboro Formation
(Ordovician). The fossiliferous Dillsboro is exposed at the Brookville Lake dam on
the north side of Brookville and consists of thin blue limestone flags interbedded with
shale. The limestone contains numerous fossils, and the common brachiopods
Rafinesquina and Platystophia, the bryozoan Hallopora, and the cup coral Streptelasma
are present in great abundance. Fragments and whole specimens of trilobites such as
Isotelus are much less common but more exciting to find.
Heavily laden with fossil-bearing rocks, we reached the end of this most-satisfying
centennial Academy field trip just as the rain began to fall. The timing couldn't have
been better.
30 Indiana Academy of Science Vol. 94 (1985)
Ornithology field trip, April 28, 1984
William H. Buskirk, Leader
The Ornithology walk at the Spring Meeting of the Indiana Academy of Science
at Brookville ran from 6:00 to 8:00 a.m. and was attended by nine people. The trip
visited an area along the scenic drive on the southeast side of the Brookville Reservoir.
Numbers of migrant birds were seen or heard. This included Indigo Buntings, Ovenbird,
Black-throated Green Warbler, etc. A highlight was the simultaneous sighting of both
Scarlet and Summer Tanagers!
Zoology field trip, April 28, 1984
Sherman A. Minton, Leader
The zoology field trip made 28 April in conjunction with the Brookville meeting
was chiefly concerned with herpetology. The group assembled about 9 a.m. near Derby-
shire Quarry and proceeded down Silliman's Hollow. Two-lined salamanders (Eurycea
bislineata) and northern dusky salamanders {Desmognathus fuscus) were plentiful along
the small rocky streams. Eggs of the former species were found. Long-tailed salamanders
{Eurycea longicauda) were also noted in this habitat. Redbacked salamanders (Plethodon
cinereus) were numerous in adjacent woodland along with smaller numbers of slimy
salamanders (Plethodon glutinosus). Of special interest was the collection of a specimen
of the ravine salamander {Plethodon richmondi). This species is known from only a
few sites in extreme southeastern and east central Indiana and had not previously been
recorded from Franklin County. Another interesting find was an adult salamander
of the Ambystoma jeffersonianum complex. It has not been determined if it represents
the diploid species, jeffersonianum, or the all-female triploid form, platineum. A large
adult of the small-mouth salamander (Ambystoma texanum) also was taken. A few
frogs, probably the green frog (Rana clamitans), were noted along the main stream
in the hollow. Calls of the American toad (Bufo americanus) and spring peeper (Hyla
crucifer) were heard. Box turtles (Terrapene Carolina) were plentiful, and a pair was
observed courting. Snakes observed included one adult and a juvenile black racer
(Coluber constrictor), two adult ringneck snakes (Diadophis punctatus), an eastern
garter snake (Thamnophis sirtalis), and a small juvenile of the banded watersnake
(Nerodia sipedori). Most of these amphibians and reptiles were merely observed or
released at site of capture after identification or photography. A few specimens were
retained by John Iverson and Sherman Minton for deposition in museum collections.
Highlights of the Spring Meeting 31
Indiana Academy of Science
SPRING MEETING
OF THE EXECUTIVE COMMITTEE
April 27, 1984
MINUTES
President Theodore J. Crovello called the meeting to order at 4:00 PM in the Knights
of Columbus Hall, Brookville, Indiana.
Minutes of the Executive Committee meeting of October 27, 1983, were approved
as distributed.
TREASURER'S REPORT
Treasurer Duvall Jones distributed a report of the Academy's finances as of April
26, 1984:
1984 Income $15,844.24
1984 Expenditures 13,787.23
Balance
Academy Accounts 10,668.86
Administered Accounts 16,007.43
Total $26,676.29
REPORTS OF ELECTED COMMITTEES
Academy Foundation Committee
William A. Daily, chairman, reported that on March 3 1 , 1984, the Invested Income
Account had a total market value of $118,548.84, of which $4,727.25 was spent on April
4 to assist in publication of the Proceedings. On March 3 1 the market value of the Foun-
dation Account was $40,812.38 and the market value of the John S. Wright Fund was
$716,729.99.
Research Grants Committee
Benjamin Moulton presented the report for Uwe Hansen, Chair. Academy Research
Grants in the amount of $10,335 have been awarded. A list of the grants is appended
to these minutes.
REPORTS OF STANDING COMMITTEES
Academy Representative to the A A AS
Walter Cory will attend the annual meeting of the AAAS and requests suggestions
for ways the national organization might assist the Indiana chapter.
Academy Representative to Indiana Natural Resources Commission
Damian Schmelz reported some examples from the 77 items on the agenda of the
April meeting of the Commission.
Constitution Committee
William Eberly, Chair, requested input from all committees and officers on ways
that current practice differs from statements in the Constitution. The Committee plans
to have a proposed revision of the Constitution in the hands of members 30 days before
the Fall meeting of the Academy.
Emeritus Members Selection Committee
Robert Cooper, Chair, recommended that the following members be granted Emeritus
status:
Dr. Francis D. Hole, Madison, WI
Dr. R. Emerson Niswander, North Manchester, IN
The recommendation was approved.
32 Indiana Academy of Science Vol. 94 (1985)
High School Teachers Research Fellows Committee
Walter Cory, Chair, reported that two Fellows have been accepted for Summer 1984.
They are being encouraged to present papers at the Fall meeting.
Indiana Science Talent Search Committee
Walter Cory, Chair, reported that of 42 entrants, 25 were selected as finalists and
13 of those were declared winners. Kappa Kappa Kappa provides funding for the finals
in the competition and for two $1000 scholarships.
Invitations Committee
Walter Cory reported that the 1985 meetings will be hosted by Indiana University,
Bloomington. The Spring meeting will be at Brown County State Park on April 26-27.
The Fall meeting will be on the Bloomington campus on November 15-16.
No meeting sites have been selected for 1986 and beyond.
Junior Academy Council
Susan Johnson reported that the Spring meeting of the Junior Academy is in pro-
gress at Hammond.
Cheryl L. Mason has replaced Leota Skirvin Smith as Director of the Junior Academy.
Library Committee
Lois Burton, Chair, reported that Volume 92 of the Proceedings has been distributed
to 683 Academy members and science clubs. 205 copies have been sent to libraries, in-
stitutions, and scientific societies.
The "Requisition for Printing" for Volume 93 was submitted to the Department
of Administration on March 20. 825 cloth bound and 525 paper bound copies will be
printed. The amount to be paid by the State will be $8900.
Membership Committee
Duvall Jones, Chair, reported 604 paid memberships as of April 26, with 420 on
file from 1983 not paid for 1984. 1 1 1 members and clubs were dropped for nonpayment
of 1983 dues; some of these have now been reinstated.
A member has asked if the $300 dues for Life Membership might be paid in three
$100 annual installments. There was no immediate objection, but a discussion ensued
of that question and the related one concerning the use of Life Membership dues as sources
of income through interest. President Crovello recommended that the matter be referred
to the Financial Planning Committee.
The question of exchanging membership lists with other organizations as a possible
way of promoting membership was raised. Current policy is that this is not done unless
in some specific case it is deemed advantageous.
Newsletter Editor
Walter Cory reported that the Spring Newsletter had not appeared because of ad-
ministrative problems in his office. Absence of the Newsletter has led to confusion among
the Section Chairs concerning the Call for Papers for the Fall meeting.
It was agreed that instructions should be given to a person as soon as he or she
becomes Chair-Elect . A meeting for incoming Chairs and Chairs-Elect should be con-
sidered as part of the agenda for the Fall meeting.
Program Committee and Centennial Committee
Philip St. John, Program Chair, reported that details of speakers and program for
the Fall meeting at Butler University are in progress. Edwin Squiers, Ecology Chair, said
that his section is planning a poster session describing graduate programs offered by univer-
sities in Indiana, and suggested that other sections might consider this. As part of the
Centennial observation, sections are encouraged to solicit papers about the histories of
their disciplines in Indiana.
Minutes of the Executive Committee 33
Publications Committee
Benjamin Moulton, Chair, reported that Academy monograph sales have been
responding to promotional efforts. The Centennial volume is at the publisher, and a special
cover has been designed.
Jones and Burton asked if the price of back issues of monographs should be increased
to cover increased costs of production and mailing.
Resolutions Committee
President Crovello called for suggestions for special resolutions in connection with
the Centennial observance.
Science and Society Committee
Alice Bennett, Chair, presented the report.
The Committee is implementing its responsibility to relate to State government through
the participation of several of its members on the Task Force on Science and Education.
Its goal is to provide recommendations for ways of improving scientific literacy and
capabilities of Indiana citizens.
The second responsibility of the Committee is to provide a means for the dissemina-
tion of information to the people in Indiana. It is planning to arrange for the Fall meeting
a symposium on the topic of Artificial Intelligence. If possible, the proceedings will be
published in cooperation with the Publications Committee.
Gene Kritsky of the History of Science section is preparing an important collection
of portraits of Charles Darwin. It is possible that funds from the Science and Society
budget might be used to assist in the preparation and display of the collection, and that
it be exhibited at the Fall meeting.
Youth Activities Committee
Susan Johnson, Chair, reported on the Junior Academy and the Science Talent
Search, and participation in the International Science Fair. The Committee is pursuing
plans for public recognition of outstanding science teachers and for a workshop for science
teachers on the topic of science clubs and research in the schools.
The meeting was adjourned at 5:15 p.m.
Following a buffet dinner two talks appropriate to the Centennial theme were
presented:
Fay Kenoyer Daily told of the group of scientists who gathered in Brookville one
hundred years ago to found the society that grew into the Indiana Academy.
Dean Gary A. Sojka of Indiana University traced the life of David Starr Jordan,
a giant in the early history of the Academy and the University.
A good number of people joined on Saturday in field trips into the surrounding
countryside.
Respectfully submitted,
Richard L. Conklin, Secretary
34
Indiana Academy of Science
Vol. 94 (1985)
Indiana Academy of Science
Spring, 1984, Grant Applications Funded
Research
No. Principal Investigator/
Institution & Dept.
Grants Committee Actions
Title
Funded
10
11
12
13
14
15
16
17
Mark Binkley
ISU Geography
S. Cortwright
IU Biology
S.W. Dhawale
IU East Chemistry
R. Faflak
ISU Geology
H. Feldman
IU Geology
S. R. Ferson
State U of NY
B. Fuchs
ISU Life Sc.
J. Hengeveld
IU Biology
M. A. Hughes
IU Biology
Ralph Joyner
BSU Chem.
R. Doug Lyng
IUPU Ft. Wayne Bio
Robert Pinger
BSU Bio
A. Roux
Notre Dame Bio
Curtis Tomak
In Dept Highway
Rod Walton
IU Biology
Wm. Wilson
ISU Geology
Licia Wolf
IU Biology
Base Study for the Development of
a Synoptic Climatology ... $ 500.00
The Role of Predation and the Meta-
community in the Community . . . 550.00
Corrosion of Some Copper Alloys
and Metals in Thiosulfate . . . 550,00
Quaternary Stratigraphy and
Chronology of Terraces . . . 470.00
Spatial Distribution of Macrofauna
& Paleoenvironmental . . . 370.00
Ecotones between Eastern Hemlock
Communities and Surrounding . . . 600.00
Beta-Adrenergic Receptors in
Muric Splenic T Lymphocytes 750.00
The Adaptive Significance of
Brood Reduction . . . 700.00
Cold-Temperature Activity in Two
Freshwater Turtles . . . 400.00
Insertion Reactions of Tin and
Germanium Phthalocyanines 450.00
A Preliminary Investigation on
Using Cultured Mouse . . . 570.00
Serology Survey of Selected
Indiana Vertebrates . . . 650.00
Microdistribution of Trichoptera
Populations in Juday Creek 650.00
Archaeological Research at
Alton Site 900.00
Factors Influencing the Distribution
of the Goldenrod . . . 450.00
Lithologic and Base Level Controls
on Cavern Positions . . . 800.00
Biparental Care in the Monogamous
Dark-eyed Junco; . . . 975.00
TOTAL $10,335.00
PICTORIAL HIGHLIGHTS
OF THE FALL MEETING
Welcome to Butler University, Clowes Hall. John G. Johnson, President, Butler University.
35
36
Indiana Academy of Science
Vol. 94 (1985)
Welcome to the Fall Meeting. Theodore J. Crovello, President, Indiana Academy of
Science.
Highlights of the Fall Meeting
37
Centennial Address: Past and Future Roles of Interdisciplinary Societies. Philip H. Abelson,
Editor, Science.
38
Indiana Academy of Science
Vol. 94 (1985)
Dinner for Senior Academy Officers, Officers Elect, Committee and Section Chairs and
their guests. Krannert Room, Clowes Hall.
Highlights of the Fall Meeting
39
* * « m ■ ~
Executive Committee Meetings often generate problems.
40
Indiana Academy of Science
Vol. 94 (1985)
Participants sometime provide solutions.
Highlights of the Fall Meeting
41
Gymnasiums provide space for informal luncheons.
42
Indiana Academy of Science
Vol. 94 (1985)
■si^Y
>-?!
Poster Sessions are becoming increasingly popular.
Highlights of the Fall Meeting
43
The Junior Academy sessions provide opportunities for student interactions.
SPECIAL ACKNOWLEDGMENT
Indianapolis, Indiana
November 2, 1984
During the Centennial Meeting, the Academy membership acknowledged those who
have been members for 50, or more, years. The 1984 membership roster included the
following 49 members who have an accumulated service to the Indiana Academy of Science
of 2,653 years.
Name and Address
Section
Year Joined
Number of Years
Adams, William B.
B
1919
65
703 Anita Street
Bloomington, IN 47401
Aldred, Jacob William H.
C
1929
55
R.R. 5, Box 8
Florence, AL 35630
Baldwin, Ira CBR 1919 65
1806 Van Hise Hall
University of Wisconsin
Madison, WI 53706
Bochstahler, Lester I. CGP 1920 64
422 Davis Street
Evanston, IL 60201
Brubaker, Ralph SGE 1932 52
Box 141
Leesburg, IN 46538
Campbell, Mildred F. AZB 1931 53
29 North Hawthorne Lane
Indianapolis, IN 46219
Caylor, Harold D. ZCR 1931 53
303 South Main Street
Bluffton, In 46714
Cooper, Robert H. RBC 1934 50
R.R. 9, Box 298
Muncie, IN 47302
Decay, M.H. George ACG 1929 55
715 Meridian Street
West Lafayette, IN 47906
Dunham, David H. RBZ 1920 64
230 Connolly Street
West Lafayette, IN 47906
Fidlar, Marion M. CG 1931 53
1040 Vista View Drive
Salt Lake City, UT 84108
Fulford, Margaret B 1929 55
Department of Biological Sciences
University of Cincinnati
Cincinnati, OH 45221
44
Special Acknowledgment
45
Name and Address
Geisler, Florence E.
3717 North Riley Avenue
Indianapolis, IN 46218
Gettlefinger, W.C.
9 Lincoln Road
Indianapolis, IN 40223
Girton, Raymond E.
47 Cordone Drive
San Anselmo, CA 94960
Gould, George E.
848 Kent Avenue
West Lafayette, IN 47906
Gray, Nina E.
Trust Department
First National Bank of Normal
Normal, IL 61761
Guard, Arthur T.
1845 Woodland Avenue
West Lafayette, IN 47506
Haas, Flora A.
1010 Lafayette Road
Crawfordsville, IN 47933
Hazlett, Donald C.
Russellville, IN 46175
Headlee, W. Hugh
762 North Riley Avenue
Indianapolis, IN 46201
Hennion, George F.
141 East Lasalle Avenue
South Bend, IN 46617
Hougham, Naomi M.
300 North Water Street
Franklin, IN 46131
Johnson, Willis H.
Department of Biology
Wabash College
Crawfordsville, IN 47933
Jordan, Esther K.
400 Circle Avenue
Kerrville, TX 78028
Lang, Maud O.
R.R. 2
Richland, IN 47634
Lemon, Lola M.
Box 113
Larwill, IN 46763
Section Year Joined Number of Years
BG 1925 59
BP
BOH
EZ
BZ
BTL
BT
G
BEZ
BZG
ZY
RB
B
1928
1928
1934
1928
1929
1914
1930
1926
1928
1922
1928
1931
1933
1929
56
56
50
56
55
70
54
58
56
62
56
53
51
55
46
Indiana Academy of Science
Vol. 94 (1985)
Name and Address
McCormick, Robert N.
211 Stradling Road
Muncie, IN 47304
Mathias, Harry R.
123 East Evers Avenue
Bowling Green, OH 43402
Mellon, Melvin G.
338 Overlook Drive
West Lafayette, IN 47906
Michaud, Howard H.
301 East Stadium Avenue
West Lafayette, IN 47906
Miner, William B.
710 Normal Road
Dekalb, IL 60115
Murray, Merritt J.
2718 Oakland Drive
Kalamazoo, MI 49008
Payne, Elmer C.
440 River Road
Chatham, NJ 07928
Plasterer, Eiffel G.
R.R. 5, Box 245
Huntington, IN 46750
Richter, Arthur
8872 Westfield Boulevard
Indianapolis, IN 46240
Roehm, John C.
102 Harold Drive
Hot Springs, AR 71901
Rothrock, Henry S.
3 Red Oak Road
Wilmington, DE 19806
Shock, Nathan W.
6505 Maplewood Road
Baltimore, MD 21212
Slusser, Mack W.
611 North Lebanon Street, Apt.
Lebanon, IN 46052
Smithberger, Andrew T.
53085 Oakmont Park East Drive
South Bend, IN 46637
Sperry, Theodore M.
1413 South College
Pittsburgh, KS 66762
Section Year Joined Number of Years
ZRC 1931 53
M 1925
1921
BZH 1929
1928
BZT 1929
1933
PC 1929
1926
GY 1924
1926
OY 1927
1930
1927
BLT 1928
59
63
55
56
55
51
55
58
60
58
57
54
57
56
Special Acknowledgment 47
Name and Address
Section
Year Joined
Number of Years
Tallman, Arthur W.
RC
1928
56
207 Applecore Avenue
Hendersonville, NC 28739
Thompson, Harold B.
P
1934
50
8501 Wicklow
Cincinnati, OH 45236
Webb, Harold D.
P
1930
54
812 West Delaware Street
Urbana, IL 61801
Welch, Winona H.
BR
1924
60
102 West Poplar Street
Greencastle, IN 46135
Welcher, Frank
C
1934
50
7340 Indian Lake Road
Indianapolis, IN 46236
Wick wire, Grant T.
G
1928
56
43 Fenwood Grove Road
Saybrook, CT 06475
Willer, William Arnold
BZZ
1929
55
3890 Hartman Road
Sodus, MI 49126
Witmer, Samuel W.
BZ
1921
63
1325 Greencroft Drive, Apt. 385
Goshen, IN 46826
Wolfe, Harold E.
M
1920
64
2611 East Second Street, Apt. 16
Bloomington, IN 47401
Indiana Academy of Science
FALL MEETING
OF THE EXECUTIVE COMMITTEE
November 1, 1984
MINUTES
President Theodore J. Crovello called the meeting to order at 3:30 p.m. in Gallahue
Hall 108, Butler University, Indianapolis, Indiana.
TREASURER'S REPORT
Treasurer Duvall A. Jones reported the Academy finances as of October 31, 1984:
Current Assets
Checking Account $ 7,822.96
Savings Accounts $19,566.07
$27,389.03
These are assigned to:
Academy Accounts $13,229.80
Administered Accounts $14,159.23
$27,389.03
The total membership as of October 31, 1984, was 1,040.
It was moved, seconded, and voted that the Treasurer's report be accepted.
REPORTS OF ELECTED COMMITTEES
Academy Foundation Committee
William A. Daily, Chair, reported that on October 26, 1984, the Foundation Ac-
count had a market value of $42,424.53 and had earned $2,956.68 during the past year.
The John S. Wright Fund had a market value of $678,504.03; its income was $37,580.08.
The Invested Income Account, with assets of $125, 143.65 had earned income of $8,853. 1 1
and income transfer from the Wright Fund of $33,579.98.
The report was approved.
Bonding Committee
No report.
Research Grants Committee
Benjamin Moulton, reporting for Uwe Hansen, Chair, distributed a list of Academy
Research Grants in the amount of $9,600 and High School Research Grants in the amount
of $1,152. A list of the individual grants is appended to these minutes.
Report of Constitution Committee
The ad hoc committee on the revision of the Constitution and By-laws of the Academy,
William Eberly, Chair, William Daily, and Clarence Dineen was represented by Dr. Eberly.
He presented a four-page report listing suggested revisions. There was some discussion
of the wisdom of discussing revisions without being able to see them in the context of
the entire document. It was agreed that the changes would be discussed and that members
should be able to see the entire revised document before being asked for final approval.
In the ensuing discussion several editorial changes were suggested which will be in-
corporated in the document to be presented to the membership. These minutes will report
only those points where there was substantial discussion and/or issues were unresolved.
In Article 1 , Section 3, the committee recommended empowering the Council, rather
than the Executive Committee to act as an advisory body. There was some objection to this.
The committee recommends deleting Member from the list of membership categories.
Consideration should be given to making it possible for Senior Members to indicate that
they do not wish to receive the Proceedings.
48
Minutes of the Executive Committee 49
The committee recommends that the sponsor of a science club be required to be
a Senior Member of the Academy. It was suggested this might be an imposition because
their commitment to Junior Academy activities precludes participation in Academy events.
By a show of hands, a majority of those present expressed preference for the word
chair to replace chairman wherever it appears.
There was considerable discussion of Article V, Section 1, Part (1) concerning Trustees
of the Academy Foundation. The resulting recommendation was that there be three elected
members with rotating three-year terms, plus the Treasurer of the Academy as an ex-
officio member without vote.
There was no discussion of Article V, Section 2, which describes the standing com-
mittees appointed annually.
Inclusion of the Science and Society chair on the Budget Committee by a revision
of Article VI, Section 4, was questioned.
J. Dan Webster moved that the Executive Committee table the entire report of the
Constitution Committee until the next meeting of the Executive Committee. The motion
was seconded and carried.
REPORTS OF STANDING COMMITTEES
Speaker of the Year
Dr. Charles E. Heiser, Distinguished Professor of Botany, Indiana University, Bloom-
ington, is Speaker of the Year.
Editor
Donald Winslow, Editor, reported that Volume 92 (1982) of the Proceedings was
delivered from Western Newspaper Publishing Co., Inc. The issue was 825 hard bound
and 525 paper copies, at a total cost of $15,332.49. The State of Indiana paid $8,400
and the Academy $6,932.49.
Copy for Volume 93 (1983) was in the hands of the printer on July 2, 1984. Section
chairs are reminded of the December 1 deadline for copy, which has not been honored
by several, resulting in the late submission to the printer. The volume is still in production
and should appear early in 1985.
The Academy offered an honorarium for the best research papers in biological and
physical sciences but the Editorial Board felt that none of the papers submitted for Volume
93 met the criteria.
The biennial budget request was submitted to the State Budget Agency in August.
The request for Fiscal Year 1985-86 is $9,200; for 1986-87 it is $9,400. The 1984-85 ap-
propriation is $8,900.
Committee on Emeritus Members
Dr. John Christian, School of Health Science, Purdue University, was granted
Emeritus Membership.
Nominating Committee
J. Dan Webster, chair, reported the slate of officers who will be nominated for
election at the General Meeting. His motion to nominate these people was approved.
Committee on Fellows
Wilton S. Melhorn, chair, presented the names of seven members to be placed in
nomination at the General Meeting. They were approved for presentation to the member-
ship at the General Meeting.
Youth Activities Committee
Susan M. Johnson, chair, presented the report.
The Committee has identified leading science teachers through a statewide search
process. The two finalists and six semi-finalists will be recognized at the General Meeting.
50 Indiana Academy of Science Vol. 94 (1985)
A grant proposal to the AAAS for the support of research projects by secondary
school students has been funded for $1200.
The committee proposes the establishment of a Science Olympiad, a series of con-
tests by which middle and junior high school students would be encouraged to become
interested in science-related activities.
Dr. Johnson moved that the Academy support the exploration of establishing an
Indiana Science Olympiad. The motion carried.
Invitations Committee
Donald Cook, chair, reported that the 1985 host will be Indiana University at Bloom-
ington, but that no invitations have been received for 1986 and beyond.
Junior Academy of Science
Cheryl Mason, Director, is studying the history of the Junior Academy and hopes
to make its activities parallel those of the Academy. Suggestions can be sent to her at
216 Chemistry Building, Purdue.
Publications Committee
Benjamin Moulton, chair, reported that the inventory of past publications ranges
from 1300 to 2000 copies for each of the past four monographs. Storage areas must be
found.
Progress is continuing on three monographs.
400 copies of The History of the Indiana Academy of Science by William and Fay
Kenoyer Dailey will be available for distribution after the General Meeting. The Academy
owes a debt of gratitude to the Dailys for their completion of this five-year task.
A booklet summarizing the Symposium on Artificial Intelligence has been published
and is being distributed at this meeting and about the state.
Library Committee
Distribution of the Academy Proceedings, Volume 92, has been completed by shipping
304 volumes to foreign exchange agencies. 176 volumes from the library's journals were
bound commercially. 27 volumes and 23 microfiche have been added to the library col-
lection, making the total number of volumes in the library 10,760.
The "Advice of Allotment" ($8900) has been received from the State Budget Agency
to be applied to the cost of printing Volume 93 of the Proceedings. Budget requests for
the next biennium have been delivered to the State Budget Agency: $9200 for 1985/86
and $9400 for 1986-87.
Mrs. Holly Oster of the Indiana State Library has been appointed to succeed Mrs.
Lois Burton as librarian in charge of the Academy library.
Membership Committee
Duvall Jones pointed out that new brochures are necessary but membership categories
are not clear pending revision of the Constitution.
Alice Bennett moved that the Executive Committee authorize publication of a brochure
that does not include the "Member" category. The motion carried.
Representative to the Natural Resources Committee
Damian Schmelz circulated a report that said the Commission met in the field three
times and at the State Museum. Agenda items pertain to oil and gas, water and flood-
ways, forestry and wildlife, coal mining and reclamation.
Resolutions Committee
William Davies, chair, moved approval of a resolution he will present at the General
Meeting, expressing appreciation to Butler University. Approval was granted.
Duvall Jones has suggested seven resolutions concerning education in Indiana. They
were not discussed, but some modification of them may be presented by Jones on the
floor of the General Meeting.
Minutes of the Executive Committee 51
Science and Society Committee
Alice Bennett, chair, reminded members of the Saturday Symposium and called
attention to a Darwin exhibit prepared by Gene Kritsky of the History of Science section .
NEW BUSINESS
Frank Guthrie moved that the initiation and reinstatement fee for members be set
at zero dollars for 1985. The motion was seconded and carried.
Fay Kenoyer Daily, speaking on behalf of Lois Burton, presented Holly Oster, who
is taking care of Academy affairs at the State Library. She has been voted by the State
Library to succeed Mrs. Burton as our representative there. In order to cement this rela-
tionship and because of her devotion and excellent handling of our affairs and excellent
qualifications, Fay Daily moved that Holly Oster be elected an Honorary Member of
the Academy. The motion was seconded and carried.
Adjournment: 5:45 p.m.
Respectfully submitted
Richard L. Conklin, Secretary
52
Indiana Academy of Science
Vol. 94 (1985)
Indiana Academy of Sciences
Fall 1984 Grant Application Funded
Research Grants Committee Actions
Principal Investigator
— Institution
Title
Funded
C. M. Anslinger
ISU
Thermoluminescent Determination of Chert
Tools and Debitage from the Wint Site
$ 800
N. C. Behforouz
Ball State U
The Role of Prostaglandin E2 in the
Immune Response to Leishmania tropica
$1,000
M. B. Berg/R. Hellenthal
Notre Dame
The Role of Chironomids (Diptera:
Chironomidae) in Stream Insect
Productivity
$ 500
A. K. Berndtson/
R. Hellenthal
Notre Dame
Plasma and Follicular Proteolytic En-
zymes in the Ovulation of Brooktrout . . .
$ 650
D. DeManno/K. Tweedell
Notre Dame
The Role of Adenylyl Cyclase in Brook
Trout Oocyte Final Maturation
$ 750
B. B. Dusa/W. Brett
ISU
Murine Natural Resistance to Trypanosoma
Lewisi
$ 850
T. E. Klingler/D. Smith
Purdue
Field Observations and Objective Analysis
of Severe Weather in Indiana
$1,000
0. Kukal/J. Duman
Notre Dame
Control of Cold Hardiness in Arctic Insects
$ 500
K. C. Kuo/T. West
Purdue
Compression Strength Testing of the
Springfield Coal, Pike County
$ 750
C. L. Mason/J. Kahle
Purdue
Renovation and Updating the Biology
Classroom
$1,000
J. Neven/J. Duman
Notre Dame
Freeze Tolerance in Insects
$ 500
G. K. Podila/W. Brett
ISU
In Vitro Translation and Gene Analysis of
Double Stranded RNA Mycovirus
$ 600
C. M. Rogers/
V. Nolan, Jr.
IU-Bloomington
Genetic and Environmental Components of $ 700
Winter Fat Storage in the Dark-Eyed Junco
Total $9,600
Minutes of the Executive Committee
53
FALL 1984
Secondary School Research Grants
Name/Sponsor
Institution
Funded
Loretts Baker
East Noble
$ 150.00
(V. Rhodes)
Kendallville
Sally Bloom
East Noble
100.00
(V. Rhodes)
Kendallville
Eric Bonfield
Marquette
110.00
(D. Christakis)
Michigan City
Julie Goldman
Gage Institute
175.00
(M. Goldberg)
Indianapolis
John Hendricks
Marquette
100.00
(D. Christakis)
Michigan City
Sima Medow
John Adams
57.00
N. Longenecker)
South Bend
Michele Mengel
John Adams
70.00
(N. Longenecker)
South Bend
Jerome W. Naylor
John Adams
70.00
(N. Longenecker)
South Bend
Mark D. Owens
Marquette
50.00
(D. Christakis)
Michigan City
Tammy Sibert
East Noble
170.00
(V. Rhodes)
Kendalville
Ann R. Thorvik
Marquette
100.00
(D. Christakis)
Michigan City
Total:
$1,152.00
Indiana Academy of Science
Minutes of the General Meeting
November 2, 1984
The General Meeting of the Indiana Academy of Science was called to order by
President Theodore J. Crovello at 1:20 p.m. on Friday, November 2, 1984, in Clowes
Hall, Butler University, Indianapolis.
President Crovello opened this one-hundredth meeting of the Academy with reflec-
tions on the value of scientific disciplines working together as they have in Indiana since
1885 and moving forward together into the second hundred years.
President John G. Johnson of Butler University welcomed the Academy on behalf
of the University. Dr. Crovello presented to him and the University a copy of the History
of the Indiana Academy of Science.
Secretary Richard L. Conklin gave a summary of the Executive Committee meeting.
J. Dan Webster, Nominating Committee chair, moved the election of the following
persons as officers and members of elected committees for 1985.
President: Benjamin Moulton
President-Elect: Ernest E. Campaigne
Treasurer: Duvall A. Jones
Director of Public Relations: Alfred Schmidt
Editor: Donald R. Winslow
Academy Foundation Committee member (2 year term): John A. Ricketts
Bonding Committee member (2 year term): Donald Hendricks
Research Grants Committee member: Austin Brooks
The motion was seconded and carried, and those named were declared to be elected.
Resolutions Committee Chair William Davies presented the following resolution:
WHEREAS: The Indiana Academy of Science is deeply grateful to Butler University
for its invitation to hold our 100th annual meeting on their campus; and
WHEREAS: The administration, faculty, and students alike have cooperated in pro-
viding us their facilities for this Centennial Meeting; be it
RESOLVED: That the Academy members here assembled express their sincere ap-
preciation to Dr. John G. Johnson, President of Butler University, for
all the courtesies that have been extended to the Academy during this
meeting. We are especially grateful to Dr. Philip St. John, his staff,
and other participating members of the Butler University faculty, for
the arrangements of the entire program and the comfort and conve-
niences provided the membership. We also express our sincere thanks
to all members who organized and participated in all aspects of the
Centennial Program.
The resolution was adopted unanimously.
Richard Conklin, member of the Committee on Fellows, moved that the following
persons, whose nomination had been approved by the Executive Committee, be elected
to the rank of Fellow:
Ernest M. Agee, Purdue University
Lois Burton, Indiana State Library
Thaddeus J. Godish, Ball State University
William R. Gommel, Indiana Central University
Henry H. Gray, Indiana Geological Survey
John H. Meiser, Ball State University
David M. Sever, St. Mary's College
54
Minutes of the General Session 55
The motion was seconded and carried. Congratulations and certificates of recogni-
tion were extended to those newly-elected Fellows who were present.
Holly Oster of the Indiana State Library, who was elected an Honorary Member
of the Academy by the Executive Committee, was introduced. She acted on behalf of
Lois Burton in accepting a plaque recognizing Mrs. Burton's years of service to the Academy
as Director of the John S. Wright Library.
Recognition copies of the History of the Indiana Academy of Science were presented
to its authors, Fay Kenoyer Daily and William Daily. Benjamin Moulton announced that
copies would be distributed to members after the meeting.
Secretary Richard Conklin moved that the Indiana Academy of Science continue
its affiliate relationship with the American Association for the Advancement of Science.
The motion was seconded and carried.
Duvall Jones presented the following resolutions:
WHEREAS members of the Indiana Academy of Science are seriously concerned
about having adequate numbers of well-qualified science teachers for
the schools of Indiana
BE IT RESOLVED that the Indiana Academy of Science recommends that certifica-
tion through a teaching minor in a science (or mathematics) be limited
to those persons with a major concentration of courses in another natural
science (or mathematics), and
WHEREAS science education at the elementary level is important to the develop-
ment of attitudes toward science,
BE IT RESOLVED that the Indiana Academy of Science favors the increased re-
quirements for science education in the elementary schools, and recom-
mends support for science workshops and science consultants to assist
elementary school teachers at the regional or local level.
After a brief discussion, the resolutions were approved by a majority of those pre-
sent, as indicated by a show of hands.
Fay Kenoyer Daily, Necrologist, reported the deaths of the following Academy members
who deaths had been recorded during the past year:
Bryon G. Bernard
Walter I. Brumbaugh
David H. Dunham
Elmer C. Payne
Edward W. Shrigley
Ruth Wimmer
Susan M. Johnson, Youth Activities Committee Chair, presented awards to the semi-
finalists and finalists in the 1984 Presidential Awards for Excellence in Science Teaching:
Semifinalists:
Gene P. Buzzard, Snider High School, Ft. Wayne
Ronald E. Divelbiss, Leo Junior/Senior High School
Gladysmae Good, Arlington High School, Indianapolis
Carole Goshorn, East High School, Columbus
Michael Kobe, Clay High School, South Bend
John J. Portle, North High School, Bloomington
Finalists:
Nevin Longenecker, John Adams High School, South Bend
Virginia Rhodes, East Noble High School, Kendalville
Mr. Longenecker spoke briefly about his reaction to the White House presentation
of the Presidential Awards.
56 Indiana Academy of Science Vol. 94 (1985)
The Speaker of the Year, Dr. Charles B. Heiser, Jr., Distinguished Professor of
Botany, Indiana University, Bloomington, gave an abbreviated version of the lecture
he will be presenting in that capacity, "The Contributions of the Nightshade Family
(Solanaceae) to Human Welfare."
The meeting adjourned at 2:40 p.m.
Respectfully submitted,
Richard L. Conklin, Secretary
Indiana Academy of Science
Budget Committee Meeting
December 1, 1984
Members Present: Alice Bennett, Richard Conklin, E. E. Campaigne, Walter Cory,
William Daily, Frank Guthrie, Uwe Hansen, Duvall Jones, Susan Johnson, Cheryl Mason,
Benjamin Moulton (President), Holly Oster, Donald Winslow.
Called to Order: 10:10 a.m., in Room 159A, Indiana State Library, Indianapolis.
President Moulton opened the meeting with a brief statement of his hope to work
within the budget being set for 1985, guided by the findings of the Finance Committee
which should be reported at the Spring Meeting. He plans to write letters to the past
twenty presidents of the Academy asking their opinions on the role and operation of
the Academy in its second century.
There was some discussion of financial decisions that must be made by the Publica-
tions Committee: Shall the price of the Proceedings be increased, given the fact that only
$7.00 from dues are available to pay the $1 1 .37 cost per copy. The History of the Academy
cost about $5.50 per copy. It is being distributed free to members but a price needs to
be set for copies sold to non-members. Guthrie suggested $7.50 for additional copies
to members, $10.00 for non-members. Jones wondered if the History could be used as
an incentive in the forthcoming membership campaign. Several combinations of years
of membership and copies of the History with package rates were suggested. Other
monographs might also be used.
Jones moved that publication cost of the brochure describing the Fall, 1984 Sym-
posium on Artifical Intelligence be taken from the Income Trust Fund. The motion was
seconded, and carried after some discussion about whether such publications could be
more appropriately charged to Science and Society or Publications. The motion carried.
The Treasurer presented the proposed budget for 1985, which is attached herewith.
After it had been discussed line by line, Hansen moved approval of the operating budget.
The motion carried.
The following amounts were proposed for items paid from the Trust Fund:
Research grants $23,200
Research fellowships 3,000
Publications 20,000 (Butterflies of Indiana)
15,000 (Climate of Indiana)
1,500 (Symposium Brochures)
15,000 (History of the Academy)
Cory moved approval of the budget for Trust Fund and Administered Accounts.
The motion carried.
The Bonding Committee will be instructed to discuss the necessity of bonding the
Treasurer and the Trustees.
Bennett moved that the Academy designate Chase Manhattan Bank as a depository
of Academy funds. The motion was seconded by Mason and approved.
Guthrie presented a suggested revision of membership categories and dues. It was
discussed briefly but no action was taken.
Mason recommended that a liaison person from the host institution at the Fall meeting
be appointed to work with the Junior Academy.
Winslow raised the question of late abstracts delaying publication of the Proceedings.
He was encouraged to send a letter to persons presenting papers asking them to press
their section chairs to submit the abstracts on time.
57
58 Indiana Academy of Science Vol. 94 (1985)
The meeting was adjourned at 12:30 p.m.
Respectfully submitted,
Richard L. Conklin
Secretary
Indiana Academy of Science
1985 Budget
Budgeted
Budgeted
Academy Accounts
Income
Expenses
Dues
9,250
Reprints: Vol. 92 & 93
2,750
2,500
Interest
2,700
Transfer from Administered Account
Reserve funds for Centennial
2,000
Meetings
Program, Printing, and Mailing
Registration Fees & Hospitality
Meals
2,000
President's Contingency Fund
250
Secretary
400
Treasurer
750
Editor's Expenses
400
General Office Expenses
/
450
Officer Travel
150
AAAS Representative
400
Biological Survey Committee
1,000
Centennial Committee
2,000
Finance Committee
250
Junior Academy of Science
1,000
Membership Committee
500
Newsletter
900
Public Relations
150
Program Committee
2,000
Speaker of the Year
700
Youth Activities Committee
1,250
Section Chairmen's Expenses
50
CPA Fees for Tax Returns
550
Miscellaneous
100
Transfers to Administered Accounts
Library Binding
1,500
Proceedings; Mailing
700
Science and Society
750
TOTALS $18,700 $18,700
Indiana Academy of Science
Financial Report
1 January — 31 December 1984
I. ACADEMY ACCOUNTS
Dues
Reprints: Vol. 92 & 93
Interest
Transfer from Administered Accounts
Reserve funds for Centennial
President's Contingency Fund
Secretary
Treasurer
Editor's Expenses
General Office Expenses
Officer Travel
AAAS Representative
Biological Survey Committee
Centennial Committee
Finance Committee
Junior Academy of Science
Membership Committee
Newsletter/Public Relations
Program Printing & Mailing
Speaker of the Year
Youth Activities Committee
Section Chair's Expenses
CPA Fees for Tax Returns
Miscellaneous
Transfers to Administered Accounts
Library Binding
Proceedings: Mailing
TOTALS for Academy Accounts
Administered Account (for budgetary
purposes only)
Meeting Fees and Hospitality
Income
Budgeted
Expenditures
Budgeted
$ 9,192.00
$ 8,000.00
1,773.20
3,250.00
$ 1,943.18
$ 2,850.00
2,859.75
2,000.00
434.58
434.58
3,200.00
21.76
1,000.00
276.55
600.00
626.84
750.00
262.06
700.00
210.21
250.00
150.00
150.00
299.50
300.00
0.00
1,000.00
1,053.90
2,600.00
0.00
450.00
998.49
1,000.00
40.00
800.00
1,050.00
1,050.00
1,554.24
2,000.00
700.00
700.00
700.75
1,650.00
50.00
50.00
500.00
500.00
0.00
100.00
1,500.00
1,500.00
775.00
775.00
$14,259.53
$16,884.58
$12,712.48
$20,775.00
1,983.00
1,500.00
1,172.97
1,500.00
TOTALS for Budgetary Purposes
$16,242.53
$18,384.58
$13,885.45
$22,275.00
II.
ADMINISTERED ACCOUNTS
1 January
1984 Transfers
1984 Transfers &
31 December
Balance
& Income*
Expenditures
Balance
Junior Academy
434.58
$ 0.00
$ 434.58 (T3)
0.00
J.S. Wright Library Fund
134.28
0.00
0.00
134.28
Lilly III Library Fund
2,619.76
0.00
0.00
2,619.76
Lilly V Library Fund
4,500.20
0.00
0.00
4,500.20
Library Binding
3,000.65
1,500.00 (T,)
2,088.45
2,412.20
Proceedings'. Printing
1,952.89
4,727.25 (T2)
4,727.25**
1,952.89
Proceedings: Mailing
550.09
775.00 (T.)
577.51
747.58
Publications: Printing &
3,435.35
901.26 (I)
14,235.90
4,282.61
Sale
14,181.90 (T2)
Research Fellowships
149.25
2,520.00 (T2)
2,520.00
149.25
Research Grants & Awards
-5,210.76
20,435.00 (T2)
20,660.70
-3,936.46
AAAS
1,200.00 (I)
Life Membership
300.00(1)
Science & Society
1,295.09
0.00
1,035.42
259.67
Science Talent Search
948.24
2,525.74 (I)
2,209.37
1,264.61
Meeting Fund
0.00
3,821.25 (1)
2,946.07
875.18
TOTALS
$13,809.62
$52,887.40
$51,435.25
$15,261.77
*I: Income from external sources.
T2:
Transfer from Academy Trust Funds
T.: Transfer from Academy Accounts.
T3:
Transfer to Academy Accounts
** The State of Indiana paid an additional $8,400.00 toward printing of the Proceedings
59
60
Indiana Academy of Science
Vol. 94 (1985)
in.
Balance: 1 January 1984
1984 Income
1984 Expenditures
Balance: 31 December 1984
SUMMARY
Academy
Accounts
$10,809.66
14,259.53
12,712.48
12,356.71
Administered
Accounts
$13,809.62
52,887.40
51,435.25
15,261.77
TOTAL
$24,619.28
67,146.93
64,147.73
27,618.48
IV. BANK BALANCES (as of 31 December 1984)
Super NOW Account
Northwest National Bank, Rensselaer, IN
Money Market Checking Account
Chase Manhattan Bank, Acct #581-1-703239
Savings Accounts
Farmers National Bank, Remington, IN-CD #2408862
#302641
State Bank of Rensselear, IN-CD #S 11766
TOTAL
V. SUMMARY OF TRUST FUNDS
Foundation Account (0043-00-0)
1. Income Account
Income Cash Balance (1/1/84)
Dividends and interest for 1984
Investments sold
Disbursements for 1984
Investments purchased
Research grants
Transfer to principal cash
Income cash balance (12/31/84)
2,000.00
300.00
1,216.64
$ 3,516.64
0.00
3,016.64
500.00
$-3,516.64
$4,810.68
9,063.58
2,000.00
2,000.00
9,744.22
$27,618.48
0.00
0.00
Principal Account
Principal cash balance (1/1/84)
Total receipts for 1984
Transfer from Income Account
Total disbursements for 1984
Investments purchased
Principal cash balance (12/31/84)
Market value of investments (12/31/84)
Total value of account (12/31/84)
$ 7,700.00
475.81
6,200.00
1,216.64
-7,700.00
192.45
192.45
40,979.33
41,171.78
'Carrying value of investments (12/31/84) is $34,828.68.
B. John S. Wright Fund (00430-01-9)
1. Income Account
Income cash balance (1/1/84)
Dividends and interest for 1984
Disbursements for 1984
Commission and fees
Transfer to 00430-02-8
Investments purchased
Income cash balance (12/31/84)
2. Principal Account
Principal cash balance (1/1/84)
Total receipts for 1984
Total disbursements for 1984
Principal cash balance
0.00
38,751.07
$ 4,000.10
33,113.63
$ 4,600.00
$ 41,713.73 $41,713.73
$- 2,962.66
$ 2.19
189,088.65
-189,085.89
$ 4.95
Market value of investments (12/31/84)
Total value of account (12/31/84)
* Negative balance due to computer error; corrected 1/2/85.
"♦Carrying value of investments (12/31/84) is $367,882.23.
$ -2,962.66
$ 4.95
722,499.43
$719,541.70
Financial Report
61
C. J.S. Wright Invested Income Account (00430-02-8)
1. Income Account
Income cash balance (1/1/84)
Total interest for 1984
Investments sold
Disbursements for 1984
Investments purchased
Distributions (grants)
Transfer to Principal account
Income cash balance (12/31/84)
2. Principal Account
Principal cash balance (1/1/84)
Funds transferred from Account 00430-01-9
Investments sold
Disbursements for 1984
Investments purchased
Distributions (Grants and Proceedings)
Transfer from Income account
Principal cash balance (12/31/84)
Carrying value of investments (Income account) (12/31/84)
Carrying value of investments (Principal account) (12/31/84)
Total value of account (12/31/84)
$ 0.00
12,212.01
11,500.00
-11,100.00
-2,520.00
-6,877.34
$ 3,214.67
$ 5,475.21
33,113.63
108,200.00
-109,319.16
-39,044.15
6,877.34
$ 5,302.87
$ 3,214.67
i 5,302.87
4,100.00
101,404.39
$114,021.93
D. Total Assets of Trust Accounts (12/31/84)
Market Value
Income Cash
Principal Cash
of Investments
Total
Account 00430-00-0
$ 0.00
$ 192.45
$ 40,979.33
$ 41,171.78
Account 00430-01-9
-2,962.66
4.95
722,499.43
719,541.72
Account 00430-02-8
3,214.67
5,302.87
105,504.39*
114,021.93
Totals (12/31/84)
$ 252.01
$ 5,500.27
$868,983.15
$874,735.43
""Carrying value of investments
VI. NOTES
Membership as of 31 January 1985: The Treasurer's records show that the Academy has 1112 paid memberships
for 1984: 51 sustaining, 2 sustaining family, 508 senior, 33 senior family, 255 regular, 13 regular family, 1 10 student,
106 emeritus, 3 honorary, 4 life, and 27 club memberships.
6 members deceased (included in totals above)
101 members on file from 1983, but not paid for 1984.
143 new members for 1984 (included in totals above).
13 previous members reinstated in 1984 (included in totals above).
7 persons resigned.
105 individuals dropped for nonpayment of 1983 dues.
Dues structure for 1984: $ 2.00 for student memberships
5.00 for memberships and club memberships
10.00 for senior memberships
25.00 for sustaining memberships
2.00 additional for family memberships
300.00 for life memberships
150.00-500.00 corporate memberships
50.00-100.00 institutional memberships
Reprints: All authors of papers in Volume 92 of the Proceedings have paid for the reprints which they ordered.
Cost of the reprints to the Academy was $1 ,943. 18. Authors paid the Academy $2,044.25 for reprints.
Publications: Sales of reprints, monographs and Proceedings in 1984 totaled $2,618.46.
Research Grants: Funds totaling $21,087.00 have been awarded to: M. Binkley (Indiana State), H-W
Chang (Indiana U.), S. Cortwright (Indiana U.), S.W. Dhawale (Indiana U.), R. Faflak (Indiana State
U.), H. Feldman (Indiana U.), S.R. Ferson (State U of NY), B. Fuchs (Indiana State U.), J. Hengeveld
(Indiana U.), M.A. Hughes (Indiana U.), R. Joyner (Ball State), C. Kirkner (Notre Dame), J.M. Kwolek
(Indiana U.), R. Lyng (IUPU— Ft. Wayne), M. Mandracchia (Notre Dame), R. Pinger (Ball State),
S. Ropski (1SU Life Sc), A. Roux (Notre Dame), C. Tomak (IN Dept. Highw.), Vierma & Feldman
(Indiana U.), R. Walton (Indiana U.), Wm. Wilson (Indiana State U.), L. Wolf (Indiana U.), CM.
Anslinger (Indiana State U.), N.C. Behforouz (Ball State U.), M.B. Berg/R. Hellenthal (Notre Dame),
A.K. Berndtson (Notre Dame), D. DeManno (Notre Dame), B.B. Dusai (Indiana State U.), T.E. Klinger
(Purdue), O. Kukal (Notre Dame), K.D. Kuo (Purdue), C.L. Mason (Purdue), L. Neven (Notre Dame),
62
Indiana Academy of Science
Vol. 94 (1985)
G.K. Podila (Indiana State U.), CM. Rogers (Indiana U.), L. Baker (East Noble H.S.), T. Barker
(East Noble H.S.), S. Bloom (East Noble H.S.), E. Bonfield (Marquette H.S.), J. Goldman (Gage In-
stitute), J. Hendricks (Marquette H.S.), S. Medow(John Adams), M. Mengel (John Adams), J. Naylor
(John Adams), M.D. Owens (Marquette H.S.), T. Sibert (East Noble H.S.), A.R. Thorvik (Marquette H.S.).
Grants Received: Kappa Kappa Kappa Sorority made $2,500 available for awards and expenses for the Science Talent
Search for Indiana high school students.
The American Association for the Advancement of Science granted $1,200 to be used by high school
students for research.
VII. BUDGET FOR 1985
The following budget was approved by the Budget Committee in the meeting of
1 December 1984.
Anticipated Income
Academy Accounts
Dues
Interest
Reprint Charges to Authors: Vol. 93 & 94
Centennial expenses (from reserves)
Administered Accounts
Meeting Fees
Total
$ 9,250.00
2,700.00
2,750.00
2,000.00
2,000.00
$ 18,700.00 $ 18,700.00
Budgeted Expenditures
Academy Accounts
Reprints
President's Contingency
Secretary
Treasurer
Editor's Expenses
General Office Expenses
Officer Travel
AAAS Representative
Biological Survey Committee
Centennial Committee
Finance Committee
Junior Academy of Science
Membership Committee
Newsletter/Public Relations
Program Printing & Mailing
Speaker of the Year
Youth Activities Committee
Section Chairmen's Expenses
CPA Fees for Tax Returns
Miscellaneous
Transfers to Administered Accounts
Library Binding
Proceedings: Mailing
Science and Society
Total for Academy Account
Administered Account
Meeting expenses
Budgetary Deficit
Total
$ 2,500.00
250.00
400.00
750.00
400.00
450.00
150.00
400.00
1,000.00
2,000.00
250.00
1,000.00
500.00
1,050.00
2,000.00
700.00
1,250.00
50.00
550.00
100.00
1,500.00
700.00
750.00
$ 18,700.00
$ 1,500.00
$ 20,200.00 $
20,200.00
-1,500.00
Trust Funds
Anticipated Income and Expendable Funds
IAS Foundation (00430-00-0)
J. S. Wright Fund (00430-01-9)
Invested Income Account (00430-02-8)
TOTAL
$ 2,500.00
35,000.00
114,000.00
$151,500.00
Financial Report 63
Approved Expenditures
Fiduciary Fees 4,000.00
Research Grants for Senior Academy Members 22,500.00
Research Fellowships for Secondary School Teachers 3,000.00
Publications
Proceedings— Volume 93 5,000.00
Climate of Indiana 1 5 ,000.00
Butterflies of Indiana 20,000.00
Symposium booklet (Science and Society) 1,500.00
Brochure on research grants 700.00
Awards for outstanding research papers 300.00
TOTAL $ 72,000.00
****************************************************************************************************
Restricted Accounts (accounted for elsewhere)
Anticipated Income
AAAS Funds for High School Student Research Grants $ 1,200.00
Tri-Kappa funds for Science Talent Search 2,500.00
Meeting fees 2,000.00
Sale of Publications 1,000.00
Income from Foundation Account 300.00
TOTAL 7,000.00
Anticipated Expenditures
Research Grants Committee — Junior Academy Grants $ 1,200.00
Science Talent Search 2,500.00
Meeting expenses (Hospitality) 1,500.00
Publications 300.00
Awards for outstanding research papers 300.00
(Funds from Foundation Account)
TOTAL $ 5,800.00
Respectfully submitted,
Duvall A. Jones, Treasurer
We, the undersigned, have audited the Treasurer's records for the Indiana Academy of Science for the year
of 1984 and have found them to be accurate and in order.
Andrew G. Mehall John Ricketts
1985 1985
INDIANA JUNIOR ACADEMY OF SCIENCE
Senior Division Presentation Schedule
November 2, 1984
9:30 Student:
School:
Title:
9:45 Student:
School:
Title:
10:00
10:15
10:30
10:45
11:00
11:15
9:30
9:45
Student:
School:
Title:
Student:
School:
Title:
Student:
School:
Title:
Student:
School:
Title:
Student:
School:
Title:
Student:
School:
Title:
Alternate
Student
School:
Title:
Student:
School:
Title:
Student:
School:
Title:
Butler University
Life Science
Tim Burgess
Center Grove High School (Greenwood)
"An Examination of the Phytotoxic Effects of Sulfur and
Nitrogen Dioxides on Jack Pine Trees (Pinus banksiana) and
Alfalfa (Medicago sativa)"
Richard Berry
Donald E. Gavit High School (Hammond)
"Determining the Concentration at which 2,4,5 Trichlorophenox-
yacetic Acid Changes from an Herbicide to a Plant Growth
Stimulant on Hydroponically Grown Pisum sativum''''
Sally Bloom
East Noble High School (Kendallville)
"A Study of Histoplasma Culture and Treatment in Vitro"
Ted Couillard
Highland High School (Highland)
"Cortical Bone Thickness in Chirikiv Island Eskimos"
Scott Seay
Center Grove High School (Greenwood)
"The Importance of Vitamin A"
Loretta Baker
East Noble High School (Kendallville)
"Studies on Gossypol: II. Antifertility Effects in Nutritionally
Deprived Male Hamsters, Mesocricetus auratus
Tonette Atkins
Paoli Junior-Senior High School (Paoli)
"Synergism Identification of Antibiotic Combinations Phase II:
Toxicity Determination Utilizing Tissue Culture Techniques"
Valerie Lamos
Canterbury High School (Fort Wayne)
"Wholeist Versus Serialist Learners: A Classroom Applicable
Diagnostic Tool for Learning Preference with Variables of Lobe
Dominance, Scanning Approach and Age"
Robert Beglin
Marquette High School (Michigan City)
"Electrical Cell Hybridization and the Discovery of a New Cell
Line"
Physical Science
Peter Hershberger
Canterbury High School (Fort Wayne)
"Computers in the Workplace: Radiation Dosage in Millirads
among Personnel Involved in Office Cathode Ray Tube Work
as a Function of CRT Year of Manufacture"
Anne Tseng
Highland High School (Highland)
"Manipulation of DNA with Restriction Enzymes"
64
Junior Academy Report
65
10:00 Student: Gene DeFelice
School: Donald E. Gavit High School (Hammond)
Title: "An Inexpensive Method for Drawing Space-Fill Molecules"
10:15 Student: Annie Carson
School: Bishop Chatard High School (Indianapolis)
Title: "Extraction, Separation, and Purification of Anthocyanins for
Use as a Natural Food Colorant"
10:30 Student: Mark Owens
School: Marquette High School (Michigan City)
Title: "Holographic Interferometry: Determination of Pressure
Mediated Nanometric Deformation as Applied to Reinforced
Plastic Configurations"
November 2, 1984
9:35
9:47
10:11
10:35
10:47
9:59
10:23
Student:
School:
Title:
Student:
School:
Title:
Student:
School:
Title:
Student:
School:
Title:
Student:
School:
Title:
Student:
School:
Title:
Student:
School:
Title:
Junior Division Presentation Schedule
Butler University
Biological Science
Jennifer Dawes
Canterbury Middle School (Ft. Wayne)
"A Microphotographic Investigation of Chloroplastid Concen-
tration and Chlorophyll Density as a Function of Water Tur-
bidity of Selected Area Rivers and Streams"
Janette DeFelice
Donald E. Gavit Middle School (Hammond)
"The Effects of Pharmaceutical Vitamins on the Green Bean
Plant"
Laura Mannion
Donald E. Gavit Middle School (Hammond)
"The Effects of an Acid Rain-Like Solution on Ivy and Petunia
Plants"
Brett Bologna
Marquette HS, (Michigan City)
"The Etiology of Cancer: Bacterial Assaying for the Potential
Carcinogenicity of Nicotine and Chewing Tobacco"
Chris Bardol
Marquette HS, (Michigan City)
"The Preventative Effect of Vitamin E on Crown Gall Tumors
in Helianthus annus"
Physical Science
Mark Skoog
Highland High School (Highland)
"Kirlian Photography"
Meghan Cast
The Canterbury School, (Ft. Wayne)
"Precipating Sulfides: A Comparison of Three Metals as Sulfide
Salts in Relation to Physical Differences in Structure"
Place
1st
Senior Division Paper Presentation Winners
Biological
Name Grade
Loretta Baker
12
School
East Noble HS
66
2nd
3rd
1st
2nd
3rd
Indiana Academy of Science
Tonette Atkins
Rich Berry
Anne Tseng
Gene DeFelice
Mark Owens
Physical
Vol. 94 (1985)
12
Paoli HS
12
Gavit HS
12
Highland HS
11
Gavit HS
12
Marquette HS
1st
2nd
3rd
Junior Division Paper Presentation Winners
Biological
Chris Bardol 9
Brett Bologna 9
Jennifer Dawes 8
Marquette HS
Marquette HS
Canterbury MS
Outstanding Jr. Scientist Nominees
Nominee
Mark Owen
Anne Tseng
Chris Moses
Annie Carson
Loretta Baker
Richard Berry
John Satanele
School
City
Marquette HS
Highland HS
Canterbury HS
Bishop Chattard HS
East Noble HS
Gavit HS
E. C. Roosevelt
Michigan City
Highland
Ft. Wayne
Indianapolis
Kendalville
Hammond
East Chicago
Outstanding Jr. Scientist Award
Name
Grade
School
Winner
Alternate
Mark Owen
Anne Tseng
11
12
Marquette HS
Highland HS
A.A.A.S. Winners
Name
Grade
School
Top Female
Top Male
Anne Tseng
Mark Owen
12
11
Highland HS
Marquette HS
Senior Division Polemic Winners
1st Place
Name
Grade
School
Tim Burgess
Rich Prall
Lorie Knobel
Doug Peters
11
11
11
11
Center Grove HS
(B Team)
Lisa Parsons
Colby Parsons
Eric Todd
Ann Thorvik
2nd Place
12
10
10
10
Marquette HS
(B Team)
Junior Academy Report
67
Charlie Wolven
Lynda Chick
Alan Adad
Ted Couillard
3rd Place
11
10
12
12
Highland HS
(B Team)
Junior Division Polemic Winners
Jenny Dawes
Anne Hayhurst
Katie Posther
Brad Keoun
Pat Abram
Charles Welborne
Heidi Bonfield
Gina Palmer
Chris Sessions
Selena Hariharan
Chris Cranz
Elisa Spindler
Place
1st
2nd
3rd
Canterbury MS
1st
2nd
3rd
8
8
8
(A Team)
2nd Place
9
9
9
9
Marquette HS
(A Team)
3rd Place
8
8
8
8
Canterbury MS
(B Team)
Aerodynamic Contest
Senior Division
Name
Distance
David Caseldine
Sean Eviston
Luke Adams
62.0 ft
55.0 ft
47.2 ft
Junior Division
John Puke
Nanelle Digdigan
Mark Skoog
35.0 ft
33.2 ft
32.8 ft
Minutes of the
Indiana Junior Academy of Science
52nd Annual Planning Meeting
On August 23, 1984, there was an Indiana Junior Academy meeting held for council
members, officers, and club representatives. The meeting was held at Purdue University
in West Lafayette, Indiana. The purpose of the meeting was to plan the fall meeting
and to take under consideration constitution revisions suggested by the Gavit Science
Club. It was decided at this meeting that each club would have one delegate for voting
purposes at the annual fall meeting. This would eliminate a lot of the confusion that
has existed in the past during vote tabulation.
Two days later on August 25, 1984, a meeting was held at Mr. Kobe's home in
Munster, Indiana. The club delegates in attendance at this meeting made revisions to
the constitution and submitted these revisions for club approval at the annual fall meeting.
Respectively submitted
Kimberly M. Canady
Indiana Junior Academy Secretary
On November 2nd 1984 the Indiana Junior Academy of Science held its 52nd an-
nual meeting at Butler University in Indianapolis, Ind. The meeting was opened by Presi-
dent Petra Fuerhaupter.
Thorn Barker, our I.J.A.S. representative to the A.J.A.S., gave a report of the
A.J.A.S. meeting that he attended during the summer in New York City, NY.
The Senior Academy recognized Jr. Academy Director Cheryl Mason and Council
Members Michael Kobe and Virginia Rhodes for their contributions toward the Indiana
Junior Academy of Science.
Keith Hunnings, director of the I.J.A.S. for ten years, was presented a plaque for
his outstanding efforts as past director.
Dr. James Paul George of De Pauw University distributed chemistry lab papers
to those who were interested.
The constitution changes proposed by the Gavit HS Science club, and the subse-
quent alteration re-written by committee were unanimously approved by the membership.
Respectively submitted
Kimberly Canady
I.J.A.S. Secretary
68
Necrology
Fay Kenoyer Daily, Butler University
Br yon G. Bernard
Serena, Illinois LaPorte, Indiana
June 21, 1921 February 7, 1984
Mr. Byron G. Bernard, Biology teacher and Audio- Visual Administrator, was a
native of Serena, Illinois, where he was born on June 21 , 1921 . His grade and high school
education was obtained locally. Higher education followed at the University of Illinois
where he obtained a B . S . degree in 1 940 and an M . S . degree in 1 950. He attended summer
school at Indiana University, University of Hawaii and other universities.
Mr. Bernard began teaching Biology at the University of Illinois High School in
1949 where he taught a year. In 1950, he joined the teaching staff at LaPorte High School
where he taught Biology until 1970. He then had an additional assignment in the Audio-
Visual Department until 1975. At the time, Mr. Bernard became Head of the Secondary
School Audio- Visual Department for LaPorte Community Schools which post he held
at death on February 7, 1984.
Mr. Bernard joined the Indiana Academy of Science in 1950, the year he received
an M.S. degree from Illinois and began teaching at LaPorte High School. His interests
were listed as Botany, Zoology, and Science Teaching. He aided in Science Fair projects
and was a student sponsor at national science fairs.
Other activities included review of science articles and films for professional organiza-
tions and he reviewed films for selection by the Northern Educational Service Center
(39 school corporations).
Mr. Bernard was a member of the Rotary Club and Elks and liked photography
and travel. He would have been 63 years of age in June, 1984, had he survived until that time.
69
70 Indiana Academy of Science Vol. 94 (1985)
Walter I. Brumbaugh
Huntington, Indiana Union City, Ohio
May 4, 1908 August 11, 1983
Mr. Walter I. Brumbaugh's chief interests were Chemistry and Physics and his pro-
fession was in secondary school teaching. He was born May 4, 1908, in Huntington,
Indiana, to Isaac and Ida Belle Brumbaugh.
Mr. Brumbaugh began teaching in 1930 in Lancaster, Indiana, and also taught at
Forest, Indiana and Winchester.
In 1940, Mr. Brumbaugh moved to Union City. There he taught in the Union City
Community High School until retirement in 1973.
The next year after moving to Union City, in 1944, Mr. Brumbaugh joined the Indiana
Academy of Science. He was an Emeritus member at death. He was also a member of
the Union City Church of the Brethern.
After a long illness, death came to Walter I. Brumbaugh on August 11, 1983, at
Crotinger Nursing Home in Union City.
Necrology 7 1
D(avid) H(anon) Dunham
Indianapolis, Indiana Lafayette, Indiana
November 23, 1894 June 3, 1983
Dr. David H. Dunham died June 3, 1983, at 88 years of age after a long illness.
He was Professor Emeritus from Purdue University where he had taught Biology for
many years.
Dr. Dunham was a native of Indianapolis, Indiana, and born November 23, 1894.
He was still young when moving to Hillsboro, Ohio, and then Oxford where he graduated
from high school. He moved to West Lafayette, Indiana, in 1914. During World War
I, he served in the Army. He then attended Purdue University earning a B.S. degree in
1920 and master's degree in 1922. He received a Ph.D. degree from the University of
Wisconsin in 1931. He was an assistant and instructor in Biology at Purdue from 1921
to 1927, then went to the Monroe Cheese Company in Ohio to work as a bacteriologist
for a year. In 1929, he was back at Purdue University teaching Biology. He became assis-
tant professor in 1930, associate professor in 1939 and retired in 1963 becoming Pro-
fessor Emeritus.
While at Purdue, David Dunham served well. His classrooms were alive with activity
and his popular field trips will long be remembered by the alumni. He had a willingness
to teach undergraduate non-major students hoping to provide a broader education among
our citizens. His chief hobby was photography. He owned a trailer and took several trips
each year with it over the United States of America and also through Mexico and Guatemala.
He traveled until he was 86 years old. His research followed bacteriological lines in the
physiological relationship between the rhizobia and Leguminosae and other agricultural
bacteriological problems. He was recognized for his work in Who's Who in Indiana and
American Men of Science.
Dr. Dunham joined the Indiana Academy of Science in 1920, the year of receiving
his B.S. degree from Purdue University. It was recorded that he was in attendance in
the early 1920's at spring Indiana Academy of Science meetings. He listed Bacteriology,
Botany and Zoology as interests. He was honored as a Fellow in 1935 and was an Emeritus
Member at death.
Dr. David H. Dunham died June 3, 1983, at Home Hospital in Lafayette, Indiana,
after a two-year illness. We are indebted to Dr. Samuel N. Postlethwait, J. Alfred Chiscon
and Mary L. Stiller for their fine memorial resolution for Dr. Dunham. After discussing
rapid post-war development of the Biological Science Department and the steadfast
endeavors of the earlier faculty which made it possible, the memorial concludes, "Over
time he too has become a part of the 'Spirit of Old Purdue' that is not easily forgotten."
72 Indiana Academy of Science Vol. 94 (1985)
Elmer Curry Payne
West Lebanon, Indiana Summit, New Jersey
July 3, 1900 March 9, 1984
Dr. Elmer Curry Payne was a very versatile chemist who served in many capacities
during his career. He was born in West Lebanon, Indiana, but by high school age was
living in Indianapolis where he graduated from Shortridge High School. He received an
A.B. degree from Butler University in 1922, A.M. degree from Johns Hopkins Univer-
sity in 1928 and a Ph.D. degree from the University of Cincinnati in 1931.
Dr. Payne found employment in 1923 to 1924 at the Indianapolis Water Company.
Then continued his education. Upon completion, he came back to Indiana in 1932 to
teach Chemistry at Butler University until 1935. He was with the United States Depart-
ment of Agriculture, Food and Drug Administration from 1935 to 1943. Developmental
Chemistry had his attention with Sylvania Electrical Products in Indiana from 1943 to
1951 . In 1952, he became Chief of the Inorganic Chemistry Section of the U.S. Atomic
Energy Commission until retirement in 1965. He followed physiochemical research after
retirement. Many of his works have not been published because they are government
classified by the A.E.C.
Dr. Payne joined the Indiana Academy of Science in 1933 while teaching at Butler
University. He was an Emeritus member at death. He had many affiliations: American
Archeological Society, Association of Official Analytical Chemists, Alpha Chi Sigma,
Sigma Chi, American Chemical Society and American Association for the Advancement
of Science.
Dr. Payne had many hobbies which were too numerous to mention in toto. They
covered a wide range of interests such as philosophy, astronomy, languages, engineering,
oil painting, cooking, ancient weaponry, horticulture, politics, government economics
and finance. His philosophy of the universe was that it is purposeful — it has a purpose.
On March 9, 1984, at 84 years of age, Elmer Curry Payne died in Overlook Hospital,
Summit, New Jersey. When his son, Palmer Payne, was asked for information, he was
very kind to supply much that is incorporated here. It was interesting that when asked
to characterize his father, the son replied, "I don't feel qualified to make such judge-
ment. He was too complex and involved in so many areas of thought and activity. I'm
his son and knew him half a century." This account would certainly testify to that. This
author remembers him from student days as a conscientious, effective, rather shy and
reserved professor of chemistry teaching at Butler University. He taught a good course!
Necrology 73
Edward White Shrigley
Lansdowne, Pennsylvania Tucson, Arizona
February 20, 1908 December 24, 1983
Dr. Edward White Shrigley was a physician and university professor born in
Lansdowne, Pennsylvania, February 20, 1908, to Arthur and Rebecca Shrigley. He was
a student at Iowa State University receiving a certificate in vocational agriculture in 1928,
a B.S. degree in genetics in 1932, and an M.S. degree in 1933. An M. A. degree was received
from Harvard in 1934. At the University of Wisconsin, his doctorate in genetics was
taken under the direction of the renowned M.R. Irwin. It was received in 1937. He also
entered the medical school there receiving an M.D. in 1941 and a two-year internship.
As a Fellow of the International Cancer Research Foundation at Yale University, he was
an instructor and later an assistant professor in Bacteriology from 1942 to 1949, and
was serologist-in-chief at Grace New Haven Community Hospital from 1948 to 1949.
It was 1949 when Edward Shrigley came to Indianapolis, Indiana, to be Associate
Professor of Microbiology at Indiana University School of Medicine. He became a pro-
fessor in 1952 and the head of the department in 1953 and was also Director of Graduate
Programs from 1968 to 1973. He retired in 1975 becoming Professor Emeritus and then
moved to Tucson, Arizona. While Head of the Department of Microbiology at Indiana
University Medical School, Dr. Shrigley organized the graduate program and was largely
responsible for the first Public Health Service Training Grant at the Medical Center.
He was also instrumental in establishing the chaplaincy program there, the philosophy
club and recorder society on campus.
Dr. Shrigley carried out many important assignments during his career such as ser-
vice on the test committee in bacteriology for the National Board of Medical Examiners,
1953 to 1957; on the genetics panel, committee growth, National Research Council in
1955; on the selection committee for senior research fellows in the U.S. Public Health
Service, 1959 to 1962; and on the American Board of Microbiology.
Dr. Shrigley joined the Indiana Academy of Science in 1950. In 1957, he was Chair-
man of the Bacteriology Division. He was honored as Fellow in 1960, and was an Emeritus
member at death. His interests were listed as Zoology, Bacteriology and Cell Biology.
He was active in several other societies including: Fellow of the American Association
for the Advancement of Science; Secretary of the Society for Study of Development and
Growth, 1947 to 1949; Society of Microbiology; Genetics Society of America; American
Association of Immunologists; Editor of the Abstracts Section, Association of Cancer
Research, 1946 to 1948; Academy of Microbiology; Fellow of the New York Academy;
Sigma Xi; Phi Kappa Phi; Phi Sigma; Alpha Omega Alpha and Friends meetings.
The narrow specialization and great expansion in many fields of science were of
great concern to him. Therefore, he encouraged lecturers to cover subjects outside the
scope of their specialization. He himself pursued a diversity of interests including genetics
in animals, viruses and bacteriophage. He was licensed to practice medicine in Indiana,
Connecticut and Wisconsin, and was interested in medical programs in Pakistan, Australia,
Burma and Peru. He studied archeology, anthropology and religions throughout the world
traveling extensively.
Dr. Edward White Shrigley was developing new interests after retirement at Tucson,
Arizona. He was deeply involved in water quality and water supply in a part of the land
where these matters are of utmost importance. He was hospitalized only a short time
suffering from heart failure before he died December 24, 1983. It is fitting to end this
account with a memorial by his children, "Today we gather on this hilltop to return
to the earth the ashes of this beloved man. Together we draw strength in remembering
the way he lived and what he taught each of us. He taught us what is important in life
74 Indiana Academy of Science Vol. 94 (1985)
and what is not. He believed that although the answers in life change, the questions,
throughout time, remain the same. His compassion for others, his humor, his honesty,
his love for learning and his personal concern for the future of mankind are deep and
lasting values "
Necrology 75
Ruth M. Wimmer
Hungtington County, Indiana Fort Wayne, Indiana
December 12, 1903 June 2, 1983
Miss Ruth M. Wimmer was a teacher of Chemistry and Dean of Girls for many
years at Elmhurst High School in Fort Wayne, Indiana. According to her executor, Mr.
William Gordan, she was an only child born to a doctor and his wife in Huntington
County, Indiana. Miss Wimmer graduated from Huntington High School in 1921. She
attended Western College for Women from 1921 to 1923, then went on to the College
of Education at Indiana University. In June, 1925, she received an A.B. degree from
that university, and an A.M. degree in June, 1927.
When Elmhurst High School opened in Fort Wayne in 1932, Ruth was employed
to teach chemistry, mathematics and geography. She was Dean of Girls for 35 years,
too, and became Chairman of the Chemistry Department. She retired in 1967.
Elmhurst High School Principal Richard Horstmeyer described Miss Wimmer
(January, 1984, Chemical and Engineering News) as "very rigid, stern, of the old school."
However, he added that students in her classes received an excellent background in
Chemistry. A former student once nominated her for an American Chemical Society
Award because of her excellent teaching motivating many to become doctors, chemists
and scientists in other specialities.
Miss Wimmer also discovered a better way to reduce resistance of wood separators
for storage batteries and sold the idea to manufacturers in Indianapolis in 1926.
Ruth Wimmer joined the Indiana Academy of Science in 1937 after starting her
teaching career at Elmhurst High School. Chemistry was listed as her chief interest and
she served on the Junior Academy Council and the Youth Activities Committee. She
was an Emeritus Member at death. She also belonged to the American Chemical Society.
At 79 years of age, Miss Ruth M. Wimmer died June 2, 1983, at the Towne House
Health Center in Fort Wayne, Indiana, leaving a truly noteworthy legacy, a $600,000
science scholarship fund. It has been established in memory of her parents who left to
her a nice inheritance which she managed well. Graduates of Huntington (North) and
Elmhurst High Schools are to benefit from the income from the trust fund to pursue
careers in science, medicine and nursing. Of course, this was the tangible legacy. The
intangible is her favorable influence perpetuated in the lives of those she touched.
Indiana Academy of Science
NEW MEMBERS 1984
Abrell, D. Brian, Indiana Heritage Program, 612 State Office Bldg., Indianapolis, IN
46204
Anslinger, C. Michael, Anthropology Museum, Indiana State University, Terre Haute,
IN 47809
Aspley, David K., 340 South Grant, Apt. #1, West Lafayette, IN 47906
Araya, Jaime E., Dept. of Entomology, Purdue University, West Lafayette, IN 47907
Arnold, Paul T., Dept. of Botany, Miami University, Oxford, OH 45056
Averbeck, Janet, Dept. of Biological Sciences, Indiana University-Purdue University
at Fort Wayne, Fort Wayne, IN 46805
Barefoot, Steven T., 1604 North Capitol Ave., Methodist Hospital, Indianapolis, IN
46202
Behforouz, Nancy, Dept. of Biology, Ball State University, Muncie, IN 47306
Bhella, Harbans S., Research Horticulturist, USDA-ARS, P.O. Box 944, Vincennes,
IN 47591
Bicha, Wesley J., Chemical Plants Engineer, 1494 New London, Hamilton, OH 45103
Binkley, Mark, Dept. of Geography, Indiana State University, Terre Haute, IN 47809
Blackwell, Will H., Dept. of Botany, Miami University, Oxford, OH 45056
Boyle, Jeffrey G., 2509 Camelback Rd., Salt Lake City, UT 84121
Brinker, Ruth, Glenn A. Black Laboratory, Indiana University, Bloomington, IN 47405
Carson, Catharine Anne, Box #18 Shively Hall, Muncie, IN 47306
Casebere, Lee A., 601 State Office Bldg., Indianapolis, IN 46204
Chatham, Lloyd, 9345 S. St. Rd. 58, Columbus, IN 47201
Chaney, William E., Dept. of Entomology, Purdue University, West Lafayette, IN 47907
Chen, Young C, Dept. of Biological Sciences, Indiana University-Purdue University
at Fort Wayne, 2101 Coliseum Blvd. East, Fort Wayne, IN 46805
Chen, Bing-Huei, Dept. of Entomology, Purdue University, West Lafayette, IN 47907
Clark, Dennis E., 3920 Centennial N. Drive, Indianapolis, IN 46208
Clark, William R., Dept. of Psychological Science, Ball State University, Muncie, IN
47306
Conover, Diana R., Archaeological Resources Management Service, Ball State University,
Muncie, IN 47306
Cook, Charles K., Dept. of Mathematics, Tri State University, Angola, IN 46703
Cooper, David L., La Porte High School, La Porte, IN 46350
Cortwright, Spencer, Dept. of Biology, Indiana University, Bloomington, IN 47405
Goshorn, Carole R. F., 230 S. Marr Rd., Columbus, IN 47201
Costello, Priscilla, Terre Haute South, 3737 South 7th St., Terre Haute, IN 47802
Culver, J. Bart, P.O. Box 294, North Webster, IN 46555
Crawford, Ronald R., Dept. of Biology, Ball State University, Muncie, IN 47304
Deckers, Lambert, Dept. of Psychological Science, 5959 Broadway, Ball State University,
Muncie, IN 47306
Dustman, Nancy J., Andrean High School, 5959 Broadway, Merrillville, IN 46410
Ebstino, Frank, Northrop High School, 7001 Coldwater Rd., Fort Wayne, IN 46825
Eubanks, Mary, Dept. of Biology, Indiana University, Bloomington, IN 47405
Farlow, James O., Indiana University-Purdue University at Fort Wayne, Fort Wayne,
IN 46805
Flohr, Stephen M., 7001 Coldwater Rd., Fort Wayne IN 46825
Fluegeman, Richard H., Dept. of Geology, Ball State University, Muncie, IN 47306
Forsyth, Bill J . , Dept. of Biology, Indiana University Southeast, New Albany, IN 47 1 50
76
New Members— 1984 77
Foster, John E., Dept. of Entomology, Purdue University, Lafayette, In 47907
Francq, G. Earle, Indiana Department of Education, Division of Curriculum, Room
229 State House, Indianapolis, IN 46204
Friedle, Robert E., Dept. of Educational Psychology, Ball State University, Muncie,IN
47306
Goetz, Frederick W., Dept. of Biology, University of Notre Dame, Notre Dame, IN 46556
Goetz, Rebecca J., Dept. of Botany and Plant Pathology, Purdue University, West
Lafayette, IN 47907
Goldmann, B. R. and Julie S., Gage Institute, 6144 N. College, Indianapolis, IN 46220
Good, Gladysmae, Arlington High School, 4825 N. Arlington, Indianapolis, IN 46226
Grow, Brien N., 635 Barnhill Drive, MS 157, Indianapolis, IN 46223
Harshman, Hardwick W., School of Education, Indiana University-Purdue University
at Indianapolis, 902 W. New York St., Indianapolis, IN 46223
Hartmann, Walter, Dept. of Psychology, Purdue University Calumet, Hammond, IN
46323
Harty, Harold, Dept. of Science and Environmental Education, 202-B Education Bldg.,
Indiana University, Bloomington, IN 47405
Hendricks, Nancy, 5805 East Southport Rd., Indianapolis, IN 46227
Hennen, Joe F., and Mary Hennen, Dept. of Botany and Plant Pathology, Purdue
University, West Lafayette, IN 47906
Hollerman, Andrew, Dept. of Physics, Purdue University, West Lafayette, IN 47907
Huber, Melanie B., 373 South 7th St., Terre Haute, IN 47802
Huffman, Henry, 2309'/2 N. Headley Rd., Bloomington, IN 47401
Jarrett, III, Harry W., Dept. of Biology, Indiana University-Purdue University at
Indianapolis, Indianapolis, IN 46223
Johnsen, Torgeir S., Dept. of Biology, Indiana University, Bloomington, IN 47405
Kane, Barbara, Indiana State University, Terre Haute, IN 47809
Karns, Daryl R., Hanover College, Hanover, IN 47243
Kern, Jr., William H., Dept. of Zoology, University of Florida, Gainesville, Fla. 32611
Kesling, Mark D., The Children's Museum, P.O. Box 3000, Indianapolis, IN 46206
Kim, Sunghee K., Dept. of Home Economics, Ball State University, Muncie, IN 47306
Kirsch, Joseph, Dept. of Chemistry, Butler University, Indianapolis, IN 46208
Kjonaas, Richard A., Dept. of Chemistry, Indiana State University, Terre Haute, IN 47809
Kovach, Warren L., Dept. of Biology, Indiana University, Bloomington, IN 47405
Krohne, David T., Dept. of Biology, Wabash College, Crawfordsville, IN 47933
Kuaalen, Ruth, Dept. of Horticulture, Purdue University, West Lafayette, IN 47907
Kupagamage, Chan, Dept. of Entomology, Purdue University, West Lafayette, IN 47906
Larsen, Steven H., School of Medicine, Indiana University, 1 100 West Michigan St.,
Indianapolis, IN 46223
Leech, Curtis K., Dept. of Psychology, Anderson College, Anderson, IN 46011
Lieb, Shannon G., Dept. of Chemistry, Butler University, Indianapolis, IN 46208
Loucks, Orie L., Butler University, 4600 Sunset Ave., Indianapolis, IN 46208
McCune, III, John E., R.R. 2, Box 155 B, Floyds Knobs, IN 47119
McDonald, Dennis L., Dept. of Biology, Hanover College, Hanover, IN 47243
McGowan, Michael J., Senior Entomologist, Lilly Research Labs, Greenfield, IN 46140
McMillen, Jack D., Dept. of Biology, University of Notre Dame, Notre Dame, IN 46556
Maloney, Michael S., Dept. of Zoology, Butler University, Indianapolis, IN 46208
Marshall, Philip T., Vallonia State Nursery, Vallonia, IN 47281
Matyas, Marsha Lakes, Dept. of Chemistry, Purdue University, West Lafayette, IN 47907
Menges, Eric, Holcomb Research Institute, Butler University, Indianapolis, IN 46208
Meunier, Gary F., Dept. of Psychological Science, Ball State University, Muncie, IN 47304
Morse, Mary Ann, Dept. of Education, Indiana University East, Richmond, IN 47374
78 Indiana Academy of Science Vol. 94 (1985)
Neill, Michael J., Director, ICCE, 902 W. New York St., Indianapolis, IN 46223
Nicks, Anthony Ray, R.R. 2, Box 373, Borden, IN 47106
Nolan, Val Jr., Dept. of Biology, Indiana University, Bloomington, IN 47405
Ossom, EkpoM., Faculty of Agriculture, University of Science and Technology, P.M.B.
5080, Port Harcourt, Nigeria
Oster, Holly, Indiana State Library, 140 N. Senate Ave., Indianapolis, IN 46204
Ott, Karen J., Dept. of Biology, University of Evansville, Evansville, IN 47702
Parke, Neil, Dept. of Biological Sciences, DePauw University, Greencastle, IN 46135
Perrill, Stephen A., Dept. of Zoology, Butler University, Indianapolis, IN 46208
Podila, Gopi Krishna, Dept. of Life Sciences, Indiana State University, Terre Haute,
IN 47809
Post, Thomas W., Div. of Nature Preserves, 601 State Office Building, Indianapolis,
IN 46204
Pribush, Robert A., Dept. of Chemistry, Butler Universitv. Indianaoolis, IN 46208
Replogle, Daniel L., R.R. 3, Kendallville, IN 46755
Reynolds, Gordon, Seymour High School, 1350 W. 2nd St., Seymour, IN 47274
Robertson, Thomas H., Dept. of Physics and Astronomy, Ball State University, Mun-
cie, IN 47306
Robison, Floyd E., Research and Assessment Consultant, Indiana Department of Educa-
tion, Indianapolis, IN 46204
Rodia, Jacob S., Dept. of Chemistry, St. Joseph's College, Rensselaer, IN 47978
Scharmann, Lawrence C, School of Education, Indiana University, Bloomington,
IN 47405
Schell, Barbara J., R.R. 4, Box 294, Floyds Knobs, IN 47119
Schepper, Jeanette, Indiana State University, Terre Haute, IN 47809
Schnitzer, Samuel B., Dept. of Psychology, Indiana State University, Terre Haute,
IN 47809
Schroeder, Christopher C, 513 W. Wabash Ave., Crawfordsville, IN 47933
Secora, Elizabeth V., Fish and Wildlife Biologist, U.S. Fish and Wildlife Service, 718
N. Walnut St., Bloomington, IN 47401
Sessions, Katharine J., 12321 Aboite Ctr. R., Fort Wayne, IN 46804
Shannon, Marilyn M., Dept. of Biological Sciences, Indiana University-Purdue University
at Fort Wayne, Fort Wayne, In 46805
Shellhaas, James L., Dept. of Microbiology, Butler University, Indianapolis, IN 46208
Sitler, Martha, Taylor University, Upland, IN 46989
Stefanavage, Tom, Indiana Department of Natural Resources, Columbia City, IN 46725
Stephenson, P. Ranel, Dept. of Anthropology, Ball State University, Muncie, IN 47306
Stevenson, Kenneth L., Dept. of Chemistry, Indiana University-Purdue University at
Fort Wayne, Fort Wayne, IN 46805
Suthers, Roderick A., Medical Sciences Program, Indiana University, Bloomington,
IN 47405
Taylor, Ralph W., Dept. of Biological Sciences, Marshall University, Huntington, WV
25701
Terry, Roger L., Dept. of Psychology, Hanover College, Hanover, IN 47243
Thirakhupt, Vacharobon, Dept. of Entomology, Purdue University, West Lafayette,
IN 47907
Turbowitz, Neal, Dept. of Anthropology, Indiana University-Purdue University at
Indianapolis, Indianapolis, IN 46202
Tuncay, Atilla, Dept. of Chemistry, Indiana University Northwest, Gary, IN 46408
Tzeng, Oliver C. S., Dept. of Psychology, Indiana University-Purdue University at
Indianapolis, Indianapolis, IN 46223
Ware, Roger, Dept. of Psychology, Indiana University-Purdue University at Indianapolis,
Indianapolis, IN 46233
New Members — 1984 79
Wartzoh, Douglas, Dept. of Biological Sciences, Indiana University-Purdue University
at Fort Wayne, Fort Wayne, IN 46805
Weilbaker, Charles N., Dept. of Biology, Indiana University Southeast, New Albany,
IN 47150
White, Arthur J., Dept. of Biology, Ball State University, Muncie, IN 47306
Wiles, Tom, Dept. of Biology, Indiana University Southeast, New Albany, IN 47150
Williford, L., Northwestern High School, 3431 North 400W, Kokomo, IN 46901
Wolfal, Marcus L., 1460 South 650 East, Columbus, IN 47203
Wunderlich, Daniel K., R.R. 31, Box 455, Terre Haute, In 47803
Young, David, Dept. of Psychology, Indiana University-Purdue University at Fort Wayne,
Fort Wayne, IN 46805
Young, Gary N., Coordinator of Information, Ball State University, Muncie, IN 47306
Bishop Chathard High School Science Club, Sponsor: July L. Lines, Indianapolis, IN
46220
Borden High School Science Club, Sponsor: Thomas Lockmund, Borden, IN 47106
Center Grove High School Science Club, Sponsors: Carolyn Hayes and Wilma Griffin,
2717 South Morgantown Road, Greenwood, IN 46142
Highland High School, Milligrams, Sponsor: Kathy Reitz, Highland, IN 46322
Kahler Middle School Science Club, Sponsor: Connie Murray 456 Elm St., Dyer, IN 4631 1
Munster High School Science Club, Sponsor: Donald Ullxman, Munster High School,
8808 Columbia, Munster, IN 46311
Roosevelt High School Science Club, Sponsor: Maria Dalhoumi, Roosevelt High School,
East Chicago, IN 46312
West Side Senior High School Science Club, Shirley S. Moorehead, 9th Ave. and Gerry
St., Gary IN 46406
ADDRESSES AND CONTRIBUTED PAPERS
COMPUTERS, EDUCATION, AND ARTIFICIAL INTELLIGENCE
Theodore J. Crovello
Department of Biology
The University of Notre Dame
Notre Dame, Indiana 46556
Introduction
When large "maxicomputers" began to appear in the 1960s, we appreciated their
value in banking, in airline reservation systems and in many other areas. But for
educators they were far removed from our daily activities and could be easily ignored.
Today more powerful maxicomputers, the appearance and spread of the totally new
microcomputer, and a growing diversity of uses and users are causing a true Computer
Revolution in our society. What the Industrial Revolution did for our physical abilities,
the Computer Revolution is doing for our minds. At times I wonder how, if they
were alive today, outstanding people like Martin Luther King or Gandhi or Darwin
or Leonardo da Vinci would be using computers, and for what!
An important difference between the two revolutions is that the Computer Revolu-
tion is happening faster, taking only years instead of centuries. While it is called the
Computer Revolution, more than just computers in the narrow sense are involved.
It also includes telecommunications, television as a two way communications device,
and much more. Telematics is a term frequently used to include this wide diversity
of machines, data, and sensing devices. Computers also have become a convenient
scapegoat in society. If a power or credit card company makes an error in our bill,
it is the computer's fault. We know that a true computer error is a very unlikely
explanation, but it fills a necessary psychological function.
One way to appreciate how important computers have become in society is to
consider what would happen if suddenly all computers ceased to function and could
not be fixed or replaced. Requests for instant credit could no longer be filled. Airline
reservations and traffic control would have to be done manually, greatly reducing the
total number of flights. Our salary checks would be slower in arriving. Computer based
patient monitoring would cease. Many military weapons would be unusable. The list
is almost infinite.
Closer to our careers, computers in education would disappear, much like biological
extinction. The question of whether this would be a good or bad event would no doubt
bring a mixed response from educators. Regardless of how one feels about educational
computing, it plays a significant part in American education today. Consequently, as
professional educators we are mandated to become familiar enough with the subject
to make sound decisions for ourselves about its role in the education of our particular
students.
In this paper I will examine the current status of computer assisted education
and one of its possible future directions. To do that I first will review some basic
concepts of computers and of education.
Computers
Computers have been called many things, some of them unprintable. I suggest
the following as a simple, optimistic, nonthreatening, operational definition: computers
80
Presidential Address 81
are an extension of our minds and senses! And in education they also are extensions
of the minds and senses of our students. A corollary emerges from such a definition:
computers can never replace good teachers; they can only enhance their value.
Computers are not just the hardware, the physical machines themselves. Rather,
a computer system has three essential components: hardware; software; and people.
Software refers mainly to the programs written to tell the computer hardware what
to do. People are us! But for educomputing the two most important groups are in-
structors and learners. Decisions made about computers in education must consider
the specifics of all three components of the computer system. Failure to do so has
been costly and frustrating. Perhaps the most common example is when a college
administration purchases a series of microcomputers from the company that submitted
the lowest bid to supply the hardware. Joy turns to sorrow when the administration
learns that few if any educational programs exist for that particular machine, and
the people in that particular educational computing system (the students and educators)
are unable to create them.
The fields of computing and of educational computing are in exponential phases
of growth. No characteristic is changing linearly over time, be it the possible number
of additions per second, the number and diversity of users and uses, or any other
property. It is like every day we find that we can jog faster and faster.
Education
Most of us are professional educators, a noble and essential calling. But do we
really educate? Do we educate effectively? How do we know? Do we or our administra-
tion measure it by the popularity poll of Teacher Course Evaluations administered
to students?
Do we consider holistic aspects of education? That is, do we consider not just
the cognitive intellectual backgrounds and goals of our students but also the affective
domain — the emotional attitudes and motivation towards the subject, both of our
students and of ourselves?
What do we try to maximize in our courses? Is it test scores, student excitement
about the topic, valuable reasoning skills, or?
What type of diagram would each of us draw to summarize the pedagogic com-
ponents of one of our courses? Would items like lecture and textbook emerge as the
most important sources of learning? If you asked each of your students to do the
same, would they draw the same diagram? More importantly, would they indicate
the same components as being the most important in their learning? For example,
you might think that your lectures are the most important component, but they might
say it is other students and the laboratory. Only after such a systems diagram has
been created specifically for a particular course can we decide on a sound basis if
computers can enhance education in it. Computers can be considered as just another
component in such a diagram. But they are special since they have the potential to
affect almost all others to a very significant degree.
Computers in Education
Overlapping and redundant terminology is unavoidable in any rapidly developing
field. Educational computing is no exception. Let me define several commonly used
terms, because each relates to a different and important concept. I use "computers
in education" as a neutral, general term to encompass all elements of educomputing.
It has three major components: computer awareness; computer literacy; and computer
assisted education.
"Computer awareness" is an appreciation of how computers affect us in our
82
Indiana Academy of Science
Vol. 94 (1985)
everyday lives, both individually and as members of society. Grocery store checkout
scanners, computerized brain scans, traffic signal controls, are three specific examples.
"Computer literacy" is the ability of a particular person to perform a particular
task via computer. Let me emphasize that this may not require knowledge of a pro-
gramming language such as BASIC. Examples are a professor or student writing a
program to simulate exponential or logistic population growth, use of a word process-
ing package to prepare term papers, or use of a test bank to prepare chemistry
examinations. Some people include computer literacy as part of computer awareness,
but this causes considerable confusion.
Finally, "computer assisted education" is the use of computers to teach or learn
a subject other than computing. So computers in the physics laboratory or in an earth
sciences lecture are examples of computer assisted education. Relationships among the
three components of computers in education can be summarized using a Venn diagram
(Figure 1). Focusing on computer assisted education, it can involve a topic in basic
science and require no computer literacy beyond how to follow instructions given on
the computer screen. Alternatively, students may be asked to carry out a simulation
of possible outcomes from a nuclear power plant accident. This would involve an overlap
with computer awareness (Figure 1, area 1). Another possibility would require students
to create a simple program to evaluate the effect of different growth coefficients on
population size after 25 generations. This task would involve computer literacy (Figure
1, area 3). Finally, certain activities in computer assisted education can require all
COMPUTERS IN EDUCATION
Figure 1 . The three aspects of computers in education.
Presidential Address 83
three components of computers in education (Figure 1, area 4). For example, each
group of students might be asked to construct and analyze a model of endangered
species to suggest good management strategies.
Given an understanding of computers, education and computers in education,
the essential question still remains: can computers enhance education? This is really
too general a question. Important but more specific questions include the following:
can computers make a subject more attractive, allowing students to internalize it; can
they help students learn a topic more quickly or deeply, i.e., to climb Bloom's cognitive
ladder faster and higher.
Let's consider the lecture component of a course. Why are they often boring
to students? The major reason is because the majority of students frequently remain
totally passive throughout the lecture period. No matter how good the lecturer, most
students are still involved in only one way communication. Yet we ourselves know
that we only really learn a topic when we have to teach it. And the reason is that
we are totally active in the process. What this means is that ultimately computers may
allow educators to carry out more effectively the essential roles of facilitator and expert,
doing what we should — taking up where the computer and any other educational devices
leave off.
In the usual lecture only course, students never get the chance to be tutors. In
such cases, even without computers, the following procedure might be valuable. Take
the last ten minutes of every lecture period, lock the doors, have students simply turn
around to arrange themselves in groups of three or four, and let them teach each
other what was said by the professor in the first forty minutes. One important advan-
tage is that such a procedure would correct those instances when students truly believe
they understand what was said in lecture but in fact do not.
Today computers are used in many ways in education; literature retrieval; data
retrieval; data accumulation; online control of experiments; statistical analysis; graphic
summarization; simulation and modeling; decision making; drill and practice; tutorials;
test generation and administration; course management; and word processing. The list
grows every year.
Computers are used in the above ways because educators believe they will enhance
learning. Any other reason is insufficient. More specific reasons for using computers
in education include the following: increased effectiveness of teaching what we already
teach; increased students' interest in the subject matter; an increasingly active role
for students; a decrease of boring tasks associated with learning; increased ability for
students to learn at their own pace (and according to their own particular diurnal
rhythms!); and an increase in the level of individualized instruction. This last reason
reveals a paradox — that the allegedly impersonal computer might be able to provide
a more personal education. A corollary is that computers could help provide a better
education to a heterogeneous group of students.
Artificial Intelligence
Before considering the future of computers in education, let's review some basic
ideas of artificial intelligence. Its use in education promises to be as important as the
computer itself. Artificial intelligence is the ability of a machine to exhibit intelligent
behavior. This begs the question of what constitutes intelligent behavior, and each
of us probably would indicate the boundary of intelligence differently. For example,
does a word processing program exhibit intelligent behavior? Is a graphing program
that includes automatic scaling of a graph's axes intelligent? Does a disease diagnosis
program have intelligence?
Perhaps the most constructive view is not to consider intelligence as a yes/no
84
Indiana Academy of Science
Vol. 94 (1985)
character, i.e., that a person or machine either has or does not have intelligence. It
seems more useful to consider intelligence as a continuum, such that a particular com-
puter program, just as individual people, may exhibit various degrees of intelligence
depending on the particular skill or intelligence criterion being used. Currently the
field of artificial intelligence reserves the term artificial intelligence for programs that
exhibit higher levels of cognitive behavior. Thus, a disease diagnosis program that
just compared a set of a patient's symptoms against the known symptoms of a series
of diseases would not be considered intelligent. On the other hand, one that incor-
porated expert physicians' procedural knowledge in addition to their factual knowledge
would be considered intelligent.
Artificial Intelligence In Education: A Future Direction
Artificial intelligence in education is the use of intelligent computers to educate.
For example, a program that simply asked a grade school student to solve simple sub-
traction examples would not be considered intelligent. But a program that could do
the following would be considered intelligent: keep track of a particular student's mistakes
TEACHER - STUDENT
INTERACTION
&fs^^^&&&&&£&® THE TEACHER ^^s^s^ss^s^s&ski^^.
1
IP
a
s3
SUBJECT
KNOWLEDGE
lf^
TUTORING ]
KNOWLEDGE
(PEDAGOGY)
KNOWLEDGE
ABOUT THE
STUDENT
tf
^a*a*J^*^^*^^
£
^
I
The STUDENT
Figure 2. /I diagram of teacher-student interaction.
Presidential Address 85
over a series of subtraction examples; determine what particular type of error is being
made; and provide customized remediation to help the student discover and correct
the specific procedural mistake.
Let's paraphrase these ideas by considering what a teacher does. Figure 2 sum-
marizes the interaction between a teacher and a student. Assume the topic is the sub-
traction problem described above, or the study of meiosis in organisms, etc. An effec-
tive teacher must have three types of knowledge: knowledge about the subject matter;
knowledge about the particular student; and pedagogic knowledge sophisticated enough
to help each student in the most effective way.
Figure 3 shows what an intelligent computer program must have to be able to
exhibit intelligent behavior in education. Analogous with the teacher, it must have
three types of knowledge: a model of the particular student's understanding of the
topic being learned; expert knowledge of the topic for comparison with the student's
knowledge; and the ability to tutor each student in the best way.
The similarity between Figures 2 and 3 is obvious. Does this indicate even more
emphatically that the role of educators will decrease? No! On the contrary, I firmly
COMPUTER - STUDENT
INTERACTION BASED ON
ARTIFICIAL INTELLIGENCE
g«M^3«3«30!S!KaS!5.S!THE COMPU TER*M*MSSS3CMi^^
§ 9
[ The EXPERT ! 1
I "7 <
1 _ J£™ ..... X p
1 [ The TUTOR U-J The STUDENT | |
$, K f" ^T MODEL i
3
XmtnfllBtiniJgj/iltlSfiiWtGaiJl
3
I The STUDENT |
Figure 3. A diagram of computer-student interaction based on artificial intelligence.
86
Indiana Academy of Science
Vol. 94 (1985)
believe that every advance in educomputing will underscore the value of educators
and increase their roles for several reasons. First, viewing computers as another com-
ponent in the educational system clearly requires a professional to integrate them with
other components in a way that assures maximum learning. Second, computer pro-
grams will never be available for every topic covered in a course, with respect both
to depth of coverage and in the format most appropriate to a particular class of students.
Third, even if appropriate programs were available for all topics, few would exhibit
high levels of pedagogic intelligence; the time and other resources needed to create
such programs would be a serious limitation. Finally, just as with textbooks, many
programs become outdated as soon as they are available. Some PERSON has to fill
that gap, and that person is the professional educator.
Educational Computing Today and Tomorrow
Let's summarize where educational computing is today and where it might be
in the future. Certainly we can expect continued improvements in hardware, due mostly
to microprocessor technologies that allow more computer power to be put on one com-
puter chip. Almost all microcomputers used in education today are based on an 8-bit
chip. One exception is the Macintosh with a 32-bit architecture. That is an increase
ARTIFICIAL INTELLIGENCE
IN EDUCATION (AIE)
Figure 4. Artificial intelligence in education.
Presidential Address 87
of two orders of binary magnitude. When most educational computers have a 32-bit
or higher foundation, expect parallel increases in software.
More important than hardware are the changes in educational software. Figure
4 relates present and future developments in programs for education. Currently many
educators are involved with Computer Assisted Education (CAE). At the same time
artificial intelligence (AI) is a very active area of computer research. Several current
education programs exhibit high levels of intelligence, but they are prototypes, not
used widely, and require larger machines than those found in the classroom. Never-
theless, we can expect that the triple overlap of computers, education, and intelligence,
i.e., Artificial Intelligence in Education (AIE) will become more common and more
important. One other overlap area appears in Figure 4, that between Intelligence and
Education (IE). It asks us to consider just how intelligently we are educating now,
even without the computer. I do not believe or mean to imply that there is no intelligent
education without computers. But Figure 4 might motivate each of us to consider how
we could increase intelligent education in our particular situation, regardless of whether
computers are being used.
Periodically someone asks if computers in education are just a fad, like teaching
machines and simple uses of television. From my perspective of almost twenty years
in education I can say it is not just a fad and will not go away. One reason is that
the programs will be smarter, but another reason is that more and more educators
will accept intelligent computers both as powerful teaching aids and teaching aides!
THE CONTRIBUTIONS OF THE NIGHTSHADE FAMILY (SOLANACEAE)
TO HUMAN WELFARE
Charles B. Heiser, Jr.*
Indiana University
Bloomington, Indiana 47405
Few would deny that the grass family (Poaceae) and the legume family (Fabaceae)
are the two plant families of greatest importance to humankind. The many contribu-
tions of the nightshade family have not been as widely recognized, but certainly the
family ranks, if not third, at least among the top five plant families for its significance
to humankind. The family has contributed food, medicinal and ornamental plants,
and no genus has contributed more than Solarium which is not surprising for it is
the largest genus in the family, containing several hundred species.
The Irish potato (S. tuberosum) ranks after the major cereals as the most impor-
tant source of food for humans. The potato was domesticated in the Andes, where
some of the wild species are still used. It perhaps was the original freeze-dried food.
The potatoes were allowed to freeze at night and as they thawed the next day they
were stamped with the feet. This process which was repeated for a few days resulted
in a dehydrated product called chuno. The reasons for making chuno were twofold:
the bitter and toxic alkaloids were removed from the potato and the keeping property
of the potato was greatly increased. The potato was introduced to Europe by the Spanish
in the sixteenth century but it was to be some time before it amounted to much there.
The potatoes that first reached Europe were not well adapted to the climate and then
too, many people regarded the plant with suspicion, probably because it was associated
with the other members of the family already known in Europe, most of which were
extremely poisonous. However, it was slowly accepted and then spread rapidly, soon
becoming almost the sole food of the Irish. Then when the blight struck in the middle
of the last century, disaster followed as has been told in fascinating detail by Cecil
Woodham-Smith in The Great Hunger. This led to a huge number of the Irish
emigrating, many to the United States, where they were to have a great influence on
the political life.
Although most of the food plants in the family are native to tropical America,
the eggplant (5. melongena) comes from southeastern Asia. It, too, was thought to
be poisonous when it first reached Europe and was called Mala insana, or mad apple,
for the eating of it was thought to cause insanity. This feeling lingered among some
people until recently. They believed it necessary to soak it in vinegar or salt water
to remove the toxic properties before eating it. Why the plant was called eggplant
puzzled me as a child, for I knew of no egg so large. If one were to see the fruits
of the wild and primitive varieties, however, he would understand, for they do resem-
ble a hen's eggs in size and color.
Returning to the Americas, we find several other species of Solatium with edible
berries, most of them very little known in the United States. One of my favorites
"Indiana Academy of Science, "Speaker of the Year," 1984-85. After it was announced that I had been
selected Speaker-of-the-Year for the Indiana Academy of Science, I received a note of congratulations from Harry
G. Day, Professor Emeritus of Chemistry at Indiana University and a former president of the Academy. In it
he pointed out that my selection for this honor and service reminded him of Ralph E. Cleland, another former
president of the Academy, who had done much for science in Indiana, including fostering the Speaker-of-the-Year
Program. It is most appropriate that I acknowledge Dr. Cleland here, for it was he who, as chairman of the
Department of Botany at Indiana University, brought me to Indiana University in 1947. As a native Hoosier I
welcomed the opportunity to return to the state of Indiana.
88
Speaker of the Year 89
is the naranjilla or lulo (5. quitoense) a shrub cultivated mostly at mid-altitudes in
Colombia and Ecuador. Its fruit yields a juice that has few or no equals. Unfortunate-
ly much of the flavor is lost when the juice is canned so the juice is not as widely
appreciated as it deserves to be. Presently improved canning techniques and freeze-
dried methods are being used which preserve much of the original flavor so that we
may eventually be able to enjoy it in this country. Closely related to this species is
the cocona or tupiru, S. sessiliflorum, which is cultivated in much of the Amazon
basin and whose fruit gives a juice and is also used as a vegetable in meat dishes.
Another South American species with an edible fruit is the pepino (5. muricatum)
of western South America. It is now being cultivated in New Zealand, and fruits from
there reach our markets occasionally through Frieda's Finest Produce Specialties, Inc.
I have done research on all three of these plants, and I am tempted to tell you more
about them, but if I were to do so I would have to omit some of the other important
species in the family.
One that is interesting but not terribly important is the garden huckleberry (S.
scabrum), which of course is not a huckleberry. It perhaps is familiar to some of you,
for it is carried by many seed companies and is grown in Indiana. It is related to
the weedy black nightshades of our gardens, but its origin is a mystery. We are not
even certain as to the continent on which it originated. To my way of thinking it is
much inferior as a food plant to the related wonderberry or sunberry (S. burbankii)
of Luther Burbank which was introduced in the early part of the century but which
has now virtually disappeared.
It is now time to leave the genus Solarium but I should point out that it also
includes medicinal and ornamental plants which I shall speak of later. Also I should
mention that it contains a number of harmful plants. In addition to many poisonous
species, it also includes a number of weeds. The horse-nettle (S. carolinense), well
known in Indiana, is one of our worst weeds. Long rhizomes make it most difficult
to eradicate once it becomes established.
Closely related to Solarium, and considered by some as a member of that genus,
is Lycopersicon, which includes the wild and domesticated tomatoes. All of the wild
tomatoes are native to western South America but all the evidence points to Mexico
as being the center of origin of the cultivated tomato (L. esculentum). How is that
to be explained? Somehow seeds of a wild tomato must have been carried to Mexico
by humans or some other animal, perhaps birds, early in the prehistoric period. Our
cherry tomato which has become so popular in salad bars is probably very similar
to the wild tomato that gave rise to the domesticated one. After its introduction to
Europe the tomato suffered the same fate as the potato and eggplant, and as recently
as the last century it was still regarded as poisonous by some people. At one time
it was called love apple, and solely because of the name, was thought to be an
aphrodisiac. One story of the origin of this name is that it is a transformation of
an Italian name pomi d'oro (apple of gold) into poma amoris (apple of love). Some
of the earliest tomatoes to reach Europe were yellow or golden in color. Yellow tomatoes,
of course, are still grown and supposedly are less acid than red tomatoes and hence
are preferred by some people. Certainly few vegetables are more popular than the
tomato. This popularity cannot be explained on the basis of nutritional value, for
several other vegetables, broccoli, for example, are a better source of vitamins and
minerals.
Another kind of tomato, the tree tomato (Cyphomandra crassifolia) is another
contribution of the family from western South America. I fail to see the resemblence
of the fruit to the tomato in shape or flavor and it is used more as a fruit than a
vegetable. Moreover, the plant is a small tree or shrub, quite unlike our tomato. Today
the plant is also grown in New Zealand and fruits from there occasionally reach our
90 Indiana Academy of Science Vol. 94 (1985)
markets under the name tamarillo. The plant is also advertised by a nursery in this
country for growing in the home, with claims of yields up to 60 pounds. I haven't
recently purchased plants from this nursery, but with the ones I have grown from
South American I am lucky to get two or three fruits to a plant and then only if
I hand pollinate the flowers.
With the increasing popularity of Mexican foods, another Latin American con-
tribution, the tomatillo or tomate {Physalis philadelphica) is becoming common in our
markets. The tomate is a ground cherry or husk tomato related to those of Indiana
which are sometimes collected by wild food fanciers. The tomate was domesticated
in Mexico, and mixed with chili pepper or by itself, is used to prepare the green sauces
so widely found in Mexico and Guatemala on enchiladas and other foods. Our word
tomato is derived from tomate, but in Mexico Physalis is tomate and the tomato is
jitomate. The prefix of the latter was dropped by the Spanish when they carried the
plant to new areas.
In the Andes the fruit of another species, uchuva (P. peruviana) is eaten out
of hand, but apparently the plant is not cultivated there. It is, however, sometimes
cultivated in the United States under the name of Cape Gooseberry. Insofar as I have
been able to learn the plant went from South America to South Africa and from the
Cape of Good Hope it was taken to Australia where it was called Cape Gooseberry.
The fruit does resemble the gooseberry in shape and size, but, of course, the true
gooseberry belongs to a completely different family.
After black pepper, a member of another family, the red or chili peppers are
the world's major spice. The red peppers come to us from Latin America where four
or five different species were domesticated. Nearly all of the ones grown in the United
States, including such as chili, pimiento, cayenne, jalapeno and the sweet peppers,
belong to a single species, Capsicum annuum which was originally domesticated in
Mexico. So far I have said little about the changes that occur when a wild species
becomes domesticated. The peppers afford a good opportunity, for they have been
subjected to detailed study by some of my students and me. The ancestral form, one
of the bird peppers, was originally considered a separate species but now that its close
relationship to the domesticated peppers has been demonstrated it is recognized only
as a variety (C. annuum var. glabriusculum). It has very small berries, red in color,
extremely pungent, readily deciduous and borne erect. In the domesticated peppers
we find an increase in size of the berry, various mature fruit colors in addition to
red, pungent and non-pungent, persistent on the stalk, and either erect or pendent.
Several of these changes largely eliminate the dispersal of the fruit by birds so that
the berries are always available to people. At the same time the plants became depen-
dent on people for their perpetuation. In other parts of tropical America other wild
species were brought into domestication, one of which, C. frutescens, is cultivated
in the United States and is the source of the well-known sauce, Tabasco.
There are several other minor food plants in the family, but it is now time to
turn our attention to the drug and medicinal members, most of which are extremely
poisonous and several of which have been used as hallucinogens. Our earliest record
of these comes from the Old World. Although now little used in medicine, the most
notorious is the mandrake (Mandragora officinarum). The root of the plant was thought
to resemble the human figure, and for this reason many superstitions grew up around
the plant. It was a cure all and served as a love potion, an aphrodisiac, and for knock-
out drops; we know that it functioned effectively at least for the last use. The plant
is mentioned in both the Bible and Shakespeare, and Machiavelli's play, La Mandragola,
is still being produced.
Two other Old World plants of which we find early uses, henbane (Hyoscyamus
niger) and the deadly nightshade (Atropa belladonna) are sources of hyoscyamine and
Speaker of the Year 91
atropine, both widely used in medicine. The latter is the only antidote known for a
number of toxic substances.
The genus Datura is perhaps better known to you than the other medicinal plants
in the family, for one species, the Jimson weed or thorn apple, D. stramonium, said
to be native to Asia, is a fairly common weed in Indiana. At one time this species
was fairly widely used in medicine. Another species, the angel's trumpet, Datura inoxia,
is sometimes grown as an ornamental in this state. It was a sacred plant among
southwestern Indians and was used ceremonially. The tree Daturas of Central and
South America, now placed in the genus Brugmansia, were widely used as hallucinogens
by the native people. One of these, B. sanguinea, whose seeds were used as a narcotic,
has recently been brought into cultivation in Ecuador for the production of scopolamine.
Solarium also provides us with drug plants. One of the newer ones is S. marginatum,
a native of Africa, which became well established as a weed in the Andes. A few years
ago an Ecuadorian chemist, Alfredo Paredes, found that it was a rich source of steroids,
and recently it has been brought into cultivation in Ecuador for the production of
solasodine, which is being used to make anti-inflammatory drugs and birth control
pills. It will be interesting to observe the changes in B. sanguinea and S. marginatum
as they become converted from wild to domesticated plants. Most of our domesticated
plants are of very ancient origin, so we do not have exact records of the changes that
have taken place.
There are other medicinal plants in the family including some of those previously
mentioned as food plants. For example, the red peppers at one time were rather exten-
sively so used and still have a minor role. Tobacco is another plant that one time
was thought to have medicinal value. It was difficult for me to know how to work
this plant into a talk on the contributions of the family to human welfare, for there
are few plants that cause more harm — the connection of tobacco with cancer and other
health problems in humans is well documented. It is, in fact, difficult to find anything
good to say about the plant, but it does kill insects as well as humans and has been
employed in pesticides. The plant was used ceremonially by the American Indians.
Columbus himself saw the plant, and it was to spread more rapidly around the world
after the discovery of the Americans than any plant with the possible exception of
corn. Nearly all of the tobacco cultivated around the world today is Nicotiana tabacum.
The Indians, however, domesticated a second species, N. rustica, that was widely
cultivated in Mexico and the eastern United States. In fact, this species was the first
one cultivated in Virginia, and it wasn't until the English obtained seeds of N. tabacum,
apparently smuggled in from the Spanish colonies, that tobacco growing began to thrive
in Virginia, which, I am sorry to say, it still does today.
The final contribution of the family is a great number of ornamentals. Sometimes
these are overlooked in ethnobotanic surveys, but the appreciation of plants for their
aesthetic value apparently goes back to prehistoric times, and, of course, the produc-
tion of ornamentals is a multi-million dollar business today. Many of the genera already
discussed have furnished us a number of ornamentals, some of which are appreciated
for their fruits rather than their flowers — thus we have the Jerusalum cherry (Solanum
pseudo-capsicum), the Chinese lantern plant (Physa/is alkekengi ) and the ornamental
peppers (various cultivars of Capsicum annuum). For their flowers we have the flowering
tobaccos (several species of Nicotiana), Solanum wendlandi, and several species of
Datura and Brugmansia. More widely grown than any of these is the petunia {Petunia
hybrida), certainly one of the most appreciated of our garden ornamentals, not only
for its beauty but for the fact that it is so easily grown. Our petunia is of hybrid
origin, involving two or more species native to southern South America. From here
also have come two other favorites, Salpiglossis sinuata and Schizanthus pinna tus.
The latter is known under the common names of butterfly flower and poor man's
92 Indiana Academy of Science Vol. 94 (1985)
orchid. Like many of the other common names, the latter is rather misleading, but
the flower does have a slight resemblance to an orchid and certainly the plants cost
less than most orchids.
This concludes the survey of the family which, I think you will agree, is of con-
siderable importance. One word of caution perhaps is in order, however. A few years
ago Childers and Russo (1977) brought together a large number of testimonials from
people who claimed that by giving up the eating of solanaceous plants their arthritis
had been improved or eliminated. However, until definite proof is forthcoming I shall
continue to eat potatoes, tomatoes and chili peppers with enjoyment.
Literature Cited
1. Childers, N.F. and G.M. Russo. 1977. The Nightshades and Health. Somerset
Press, Somerset, N.J.
2. D'Arcy, W.G. (ed.). 1984. Biology and Systematics of the Solanaceae. Colum-
bia University Press, New York.
3. Hawkes, J.G., R.N. Lester and A.D. Skelding (eds.). 1979. The Biology and
Taxonomy of the Solanaceae. Academic Press, London.
4. Heiser, C.B. 1969. Nightshades, the Paradoxical Plants. W.H. Freeman, San
Francisco.
5. . 1984. The Ethnobotany of the neotropical Solanaceae, in G. Prance and
J. Kallunki (eds.), Ethnobotany in the Neotropics. New York Botanical Garden,
New York.
6. 1984 (in press). Of Plants and People. University of Oklahoma Press.
Norman OK.
ANTHROPOLOGY
Chairperson: Donald Cochran
Department of Anthropology
Ball State University, Muncie, Indiana 47306 (317)285-4927
Chairperson-Elect: Diane Beynon
Department of Anthropology
Indiana University-Purdue University at Fort Wayne
2101 Coliseum Boulevard East, Fort Wayne, Indiana 46805 (219)482-5391
ABSTRACTS
Debitage Classification Systems. C. Michael Anslinger, Indiana State University,
Terre Haute, Indiana 47809. In recent years archaeologists have found it useful
to place debitage recovered from archaeological sites into discrete groups which
theoretically represent sequential stages of lithic reduction systems. This provides one
line of evidence for reconstructing past site activities and functions which is a primary
goal of archaeology. However, recent studies have shown that some of the flake attribute
lists traditionally used to place flakes in their appropriate reduction stage are not always
meaningful and may, in fact, be ambiguous. This paper discusses some of the debitage
classification systems used by researchers and reports on the application of a par-
ticular method of classification to a Lake Archaic lithic assemblage from Bartholomew
County, Indiana.
Mann Site Figurines. Ruth Brinker, Indiana University, Bloomington, Indiana
47405. Figurines of human forms are found at the Mann Site in great numbers
and in almost all known contexts. A total of 421 fragments of human figures have
been recovered generally from village midden, but also from pits and mounds. The
figurines have many characteristics in common which makes them "recognizable,"
yet some distinctive individual traits are present as well. A general description of
characteristic forms is presented, variables are quantified, and Mann Site figurines
are compared with figurines from other Hopewellian sites in Ohio and Illinois.
The Commissary Site (12-Hn-2) Revisited. Frank Burkett and Donald R. Cochran,
Ball State University, Muncie, Indiana 47306. Monitoring of a small earthmoving
project at the Commissary site revealed the remains of a prehistoric pit containing
a few human and small mammal bones. A 10 gm sample of wood charcoal from the
pit yielded a calibrated radiocarbon date of A.D. 1180 ± 60. This date antedates the
one radiocarbon date previously acquired from the site, A.D. 635 ± 105, by 500 years.
The Commissary site has been considered to be an early Late Woodland site because
of the earlier radiocarbon date and the one cordmarked ceramic vessel recovered. The
more recent date is contemporaneous with regional Late Woodland Oliver phase sites
and suggests that either the site was in use for over 500 years or that its placement
needs to be reassessed.
Holland Chert Quarries/Workshops Near Huntingburg, Dubois County, Indiana. Mark
Cantin and C. Michael Anslinger, Indiana State University, Terre Haute, Indiana
47809. A recent archaeological survey conducted near Huntingburg, Dubois County,
Indiana, yielded several sites interpreted as chert quarries and/or knapping workshops.
93
94 Indiana Academy of Science Vol. 94 (1985)
The raw material utilized is Holland Chert and its variants. This paper will describe
the provenience and physical properties of the Holland Chert, as well as its utilization
via lithic reduction sequence analysis, and relate this within the framework of the larger
survey.
Test Excavations at the Smith Site, (12-Vi-86), Vigo County, Indiana. Mary Ellen
Carpenter and Robert E. Pace, University of Illinois-Chicago Circle and Indiana
State University, Terre Haute, Indiana 47809. Materials diagnostic of Albee and
Vincennes components have been recovered from the surface of the Smith and other
sites in Sullivan, Vigo, Parke and Vermillion counties. Excavations previously reported
have been either too limited or inadequately reported to firmly establish temporal,
spatial and cultural relationships of Albee and Vincennes. Testing at the Smith Site
was specifically undertaken to address these problems. Preliminary results suggest con-
temporaneity of materials, and either a mixing of the two populations or rapid assimila-
tion of Vincennes material culture by Albee peoples.
A Description of Kenneth Chert. Catharine A. Carson, Ball State University, Muncie,
Indiana 47306. The purpose of this study is to identify and describe cherts that
occur within the Kenneth Limestone of the Wabash Formation of north-central In-
diana. Kenneth chert is most commonly brownish-gray in color with a predominance
of lighter grey to white. The most obvious diagnostic characteristic of this chert is
its mottled, speckled, swirled, or splotched appearance due primarily to the differen-
tial silicification of fossil and burrow inclusions and the surrounding matrix. The above
factors result in Kenneth chert possessing a highly varied appearance. The chert, which
occurs as thin lenses as well as small nodules, is known to outcrop principally in Howard,
Cass, and Carroll counties. Kenneth chert is archaeologically significant as a raw material
source for the prehistoric manufacturing of chipped-stone tools.
Three Cranial Tumors from Late Woodland Sites: Diagnosis and Cultural Implica-
tions. Della Collins Cook. Department of Anthropology, Indiana University, Bloom-
ington, Indiana 47405. Tumors may not leave clear evidence on the skeleton, and
they are less common than traumatic and infectious bone pathology. For these reasons
paleopathologists seldom discuss them. An angioma or meningioma in an adult from
the Koster mound group, Green Co., Illinois, a probable melanotic ameloblastoma
in a child from the Schild mound group, Greene Co., Illinois, and an osteogenic tumor
of the cranial base in an adult from the Alt site, LaPorte Co., Indiana, are presented.
It is unlikely that age-specific frequencies of these tumors were very different in
prehistoric times than they are today. The two adults are likely to have exhibited
behavioral alterations that would have required much attention on the part of care-
givers in their communities.
A Useful Morphological Characteristic of Two Toed Sloth Hair. Edmond J. Furia,
Department of Anthropology, Indiana University, Bloomington, Indiana 47405. The
guard hairs or protective hairs from Choloepus hoffmanni and Choloepus didactylus
were observed using a scanning electron microscope. The morphological characteristic
of these hairs is unique among hairs from all living mammals known to the observer.
A single hair form either Choloepus species splits a minimum of three times in a
geometric fashion progressing from the proximal to distal portion. This "splitting"
can produce what appears to be eight differentiated shafts in the distal portion of
any one hair. This unique quality of two toed sloth hair may prove to be useful to
anyone investigating the composition of coprolithic material. This information may
prove useful also to ecology, zoology, and evolutionary biology.
Anthropology
95
The Year at Dromberg. Ronald Hicks, Ball State University, Muncie, Indiana
47306. It has been known for more than two centuries that Stonehenge is oriented
to mark the summer solstice sunrise, and recent research on other stone circles has
produced claims for their use to mark as many as 13 other dates, creating a 16-month
solar calendar. Observations over the course of a year at one stone circle — Dromberg,
in County Cork, Ireland — which has been known for some time to be oriented on
winter solstice sunset have shown it to be designed also to mark both summer and
winter solstice sunrises but no other dates. Indeed, weather conditions between autumnal
and vernal equinox today, which are not likely to be significantly different from those
prevailing at the time the circle was constructed, are such that chances of making the
necessary observations during that half of the year are slim for any of the proposed
dates except the winter solstice.
Towards Predicting Loss of Archaeological Resources from River Channel Migrations.
Misty Jackson and Robert E. Pace, Indiana State University, Terre Haute, Indiana
47809. Data being collected from riverbank survey of the central and lower Wabash,
White and Eel rivers suggest significant patterns in frequency, size and cultural affilia-
tions of exposed archaeological sites. Explanatory hypotheses being developed and tested
include variables relating to past and present stream dynamics, natural features and
resource base and settlement systems of prehistoric peoples. Preliminary results are
reported.
A Preliminary Survey of the Maumee River in Allen County, Indiana. James A. Mohow,
Indiana University-Purdue University at Fort Wayne, Fort Wayne, Indiana 46805 and
David Diaz, 4512 S. Hanna, Fort Wayne, Indiana 46806. A preliminary surface
survey was conducted along a six mile length of the Maumee River in Allen County,
Indiana between fall of 1980 and fall of 1983. The survey concentrated on the river's
floodplain and its adjacent terraces. As this was a preliminary survey, its prime objec-
tive was to identify archaeological sites and certain chronologically sensitive artifacts
within the research area. A total of 55 prehistoric sites was recorded during the survey
and more than 4,000 artifacts were recovered. The artifacts consist of lithics and ceramics.
Preliminary identification of pottery and tool types reveals that most sites were multi-
component in nature, ranging from Paelo-Indian through Late Woodland times. Since
it is little known archaeologically, the primary purpose of this survey was to achieve
an overview of the area as well as to form a basis for future research.
Woodland Sites and Ross Soils: A Correlation in the Upper White River (West Fork)
Drainage. P. Ranel Stephenson, Ball State University, Muncie, Indiana 47306. A
reconnaissance survey of the Upper White River drainage in Randolph, Delaware,
Madison, and Hamilton counties was focused on gathering data on Woodland habita-
tion sites. The survey was carried out predominantly in the floodplain of the White
River to determine whether particular soil types were selected for occupation. During
the survey, 32 new floodplain sites were discovered; of these, 10 contained pottery
and nine of the sites with pottery were located on Ross soils. Ross soils formed under
mixed hardwoods and prairie grasses and constituted the smallest percentage of the
floodplain soil types present in the counties being surveyed. Areas of Ross soil surveyed
always produced Woodland components whereas surveys of other floodplain soils did
not. A comparison of the locations of Ross soils and Delaware villages in the survey
area also showed that Delaware villages were located adjacent to the larger areas of
Ross soils. It, therefore, appears that Ross soils were selected by the Woodland occupants
of the Upper White River drainage for either occupation or cultivation.
96 Indiana Academy of Science Vol. 94 (1985)
Some Late Archaic Manifestations in Indiana. Curtis H. Tomak, Indiana Depart-
ment of Highways, Indianapolis, Indiana 46204. This paper focuses upon pre-
Riverton Late Archaic manifestations in the valley of the West Fork of White River
upstream into Morgan County and in the valley of the East Fork of White River
upstream into Jackson County. Those two areas are discussed in terms of sites, setting,
cultural assemblages, and occupations or phases. Then some other areas of the state,
particularly in southern Indiana, are considered. This is followed by commentary re-
garding the Late Archaic manifestations under review.
BOTANY
Chairperson: Phillip E. Pope
Department of Forestry, Purdue University
West Lafayette, Indiana 47907 (317)494-3590
Chairperson-Elect: Austin E. Brooks
Department of Biology
Wabash College, Crawfordsville, Indiana 47933 (317) 362-1400 ext. 350
ABSTRACTS
Effect of Cytokinins on Erythritol Permeability to Phosphatidylcholine Bilayers. Blair
Brengle and William Stillwell, Department of Biology, Indiana University-Purdue
University at Indianapolis, Indianapolis, Indiana 46223. Previously, we
demonstrated that the plant hormone kinetin enhances water permeability to several
natural and synthetic phosphatidylcholine bilayers. This enhancement was noted only
with bilayers in the liquid crystalline state. Here we report the effect of kinetin,
benzyladenine, c/s-zeatin and trans-zeal'm on the permeability of erythritol to bilayers
composed of natural and synthetic phosphatidylcholines. Mixed isomer zeatin (75%
trans, 25% cis), kinetin and to a much lesser extent benzyladenine are shown to enhance
erythritol permeability at concentrations from 0 to 1.16 mM with egg lecithin liposomes.
Trans-zeatin and adenine have no effect on permeability over the same concentrations.
C/s-zeatin greatly enhances erythritol permeability to synthetic dimyristoylphosphatidyl-
choline and dipalmitoylphosphatidylcholine bilayers only when the lipids are in the
liquid crystalline state (above the phase transition temperature). 7>a/7s-zeatin is totally
ineffective at altering permeability whether the synthetic bilayers are in the liquid
crystalline or gel states. These results clearly demonstrate for the first time a substan-
tial difference between cis and trans-zeatin on affecting membrane permeability.
Nonspecificity with Varied Effectivity in Mycorrhizal Associations. Rita de Cassia,
G. Borges, William R. Chaney and Phillip E. Pope, Department of Forestry and
Natural Resources, Purdue University, West Lafayette, Indiana 47907. A com-
mon and widespread symbiosis in plants is the mycorrhizal association between roots
and colonizing fungi. Mycorrhiza are not restricted to specific groups of plants, but
occur in practically all families of angiosperms, gymnosperms, and many lower plants.
Most commercial fruit, nut, and forest trees as well as agronomic grain and forage
crops normally form mycorrhiza. Mycorrhizal associations are so common under natural
conditions that a nonmycorrhizal plant is the exception. There is little evidence of
host specificity for mycorrhizal formation. The same fungal species or isolate can colonize
numerous host species belonging to several different families, although there is some
evidence that particular fungi are preferentially associated with particular host species.
However, the effectivity or the degree of nutritional or other advantage resulting from
the symbiotic association can vary widely among fungal and host combinations.
Nonspecificity for fungal colonization of five angiosperm tree species (Fraxinus
pennsylvanica Marsh., Liriodendron tulipfera L., Liquidambar styraciflua L., Plan-
tanus occidentalis L., and Acacia scleroxyla Tuss.) and six species of vesicular-arbuscular
mycorrhizal fungi (Glomus mosseae, G. fasciculatum, G. stunicatum, G. macrocar-
pum, G. epigaeum, and Gigaspora margarita) was shown in greenhouse studies.
However, the effectivity of the various fungal host combinations as determined by
growth of the seedlings was different. A review of literature shows nonspecificity with
varied effectivity to be a common occurrence in mycorrhizal associations.
97
98 Indiana Academy of Science Vol. 94 (1985)
Insect Pest Control in the Greenhouse: Alternatives to Commercial Toxins. Vonda
Frantz, Department of Biology, Indiana University-Purdue University at Indianapolis,
Indianapolis, Indiana 46205. Treatments for control of mealybug on Coleus were
tested over a period of several months. Plants were sprayed approximately weekly with
a mixture of garlic, cayenne pepper, mineral oil, and liquid soap, or with Safer's In-
secticidal Soap. Three different concentrations of the first mixture were tested and
two concentrations of Safer's. One set of controls was sprayed with water and another
was given no treatment. During the first half of the study mealybug adults, young,
and egg masses, were counted before each treatment. During the last half, photogenic
evidence was obtained to document the general effectiveness of each treatment.
Photographic evidence demonstrates that these treatments are effective in total greenhouse
control. All treatments result in fewer mealybugs than when water or no treatment
is applied. Safer's Insecticidal Soap is the most effective even when sprayed less fre-
quently than the mixture.
Oak "Leaf Tatters": A Malady of Unknown Cause in Indiana. Ralph J. Green, Jr.,
Department of Biology and Plant Pathology, Purdue University, West Lafayette, Indiana
47907 and Philip T. Marshall, Forest Pest Specialist, Division of Forestry, Indiana
Department of Natural Resources, Indianapolis, Indiana 46204. In 1983, a previous-
ly unreported malady of oaks, primarily white oak, Quercus alba, was found in a
number of counties in northcentral Indiana. The symptoms include a marked reduc-
tion in the interveinal leaf blade tissue followed by a partial or complete necrosis of
affected leaves. If a second growth flush occurs, these leaves are usually normal, but
reduced in size. The name of oak "leaf tatters" has been used to describe the symp-
tom complex. Symptoms begin in the lower part of the crown and are progressive
the following season. More than 50% of the trees marked with total crown involve-
ment in 1983 failed to leaf out in 1984. No trees under observation have recovered,
to date. Although symptoms have been observed primarily on white oak, other oak
species, especially black oak, Q. velutina, are also affected. Attempts to associate a
specific causal agent with the symptom complex through field observations, laboratory
isolations, electron micrographs of affected tissues and grafting have, to date, been
inconclusive. However, the progressive nature of the symptoms, both on affected trees
and within stands, suggests an infectious agent of some type.
G-banding in Lens culinaris and Vicia faba. Romesh C. Mehra and E. Boyts, Depart-
ment of Biology, Indiana University at South Bend, South Bend, Indiana 46634. In
the last decade and a half, several banding techniques for linear differentiation of
chromosomes have been developed. Some of these are C, G, N, Q and R banding
procedures. Since the discovery of these techniques, revolutionary advances have been
made in mammalia cytogenetics. However, such has not been the case in plants. Where,
of 231,413 plant species, only 90 have been studied by banding techniques. The techni-
que that has provided maximum differentiation of mammalian chromosomes is G-
banding. It is generally agreed that G-banding is produced because of the enhance-
ment of chromomeric organization of mammalian chromosomes. In plants on the other
hand, G-banding has had the least amount of success. It has been suggested that this
is because chromomeres in plant mitotic chromosomes are too close together and banding
procedures cannot reveal such bands. We have attempted G-banding on two legumes,
Lens culinaris and Vicia faba, and have obtained some measure of success. Additionally,
we have been able to reveal chromomeric organization of mitotic chromosomes in several
plants and found that indeed there appears to be a relationship of G-bands and
chromomeric organization. Evidence for the same will be presented.
Response of Muskmelon to Within-row Plant Spacing
H.S. Bhella
USDA-ARS, Vincennes University
Vincennes, Indiana 47591
and
Department of Agriculture
Purdue University, West Lafayette, Indiana 47907
Introduction
The yield response of muskmelons to plant spacing has been investigated in Arizona
(3), California (2,6) and Florida (5), but very little research, if any, has been conducted
in other areas of the United States
Davis and Meinhert (2) and Frazier (3) reported that the total yield and number
of marketable fruits were the greatest when Powdery Mildew Resistant (P.M.R.) No.
45 cantaloupe plants were spaced 30 cm apart in rows 1.8 m apart. Lazin and Simonds
(5) found that increasing distance between plants (decreasing plant population) increased
the number of fruits per plant and mean fruit weight but decreased the total number
and weight of muskmelon cvs. Earli-Dew and TAM-Dew Improved. Holliday (4) ex-
plained the relationship between plant population and crop yield for fruiting crops as
a parabolic curve. With this type of curve, a certain plant population gives a maximum
yield, while larger or smaller populations give lower yields (4).
This study evaluated the effects of 25, 50, 75, and 100 cm within-row plant spac-
ings in rows 2.7 m apart on stem length, leaf area, dry matter, soluble solids, marketable
yield, number of culls and marketable fruit, yield per plant, fruits per plant, fruit
weight, and nutrient content of muskmelon cvs. Burpee Hybrid and Classic on a
southwestern Indiana sandy loam, mixed, mesic Typic Hapludalf soil in 1982 and 1983.
Materials and Methods
Field investigations were conducted in 1982 and 1983. The 15 cm of soils had
a pH of 5.7 to 6.5, 155 to 220 kg/ha available P (Bray P-l), and 260 to 335 kg/ha
available K (IN ammonium acetate extractable), as determined by the Purdue soil testing
laboratory (1). The preplant fertilizer application consisted of 112, 25, .and 140 kg/ha
of N, P, and K, respectively. Plots were sidedressed with 50 kg/ha N five weeks after
transplanting. Granular furadan (Carbofuran) and prefar (Bensulide) were applied
preplant at the recommended rates for insect and weed control, respectively. Black
plastic mulch, 120 cm wide by 32 (i m thick, and drip irrigation hose (3.55 1/hr/m
Tri-Wall® 0.15 mm) with orifices 31 cm apart were simultaneously applied.
The experimental plots were established in a complete randomized block design
with 4 replications and each of the 4 treatments, e.g., 25, 50, 75, and 100 cm distance
between plants, was randomly assigned to a 16 x 2.7 m plot. Three-week-old greenhouse
raised plants (three-leaf-stage) of muskmelon cvs. Burpee Hybrid in 1982 and Classic
in 1983 were transplanted on May 13 each year. Guard rows were planted on both
ends of the experimental area.
A 7-10 day spray schedule was followed throughout the growing period for disease
and insect control. Plots were kept weed free by hand hoeing. All plots were trickle ir-
rigated until tensiometer readings at 30 cm depth reached 33 kPa.
Surface soil samples (0 to 15 cm) and petioles of first fully-expanded leaf near
the growing point were sampled 5 weeks after transplanting in 1983 and analyzed (1).
Harvest data on weight and number of marketable muskmelons were collected
99
100
Indiana Academy of Science
Vol. 94 (1985)
daily. The muskmelons were analyzed for total solids using Bausch and Lomb refrac-
tometer. Unmarketable muskmelons were culled and no data on culling were recorded
in 1982. During the 1983 growing season, data on number of culls were recorded.
Results and Discussion
Stem length. Total stem length per vine measured 32 days after transplanting
for 'Classic' muskmelon in 1983 increased from 215 to 368 cm with the increase in
within-row plant spacing from 25 to 100 cm. Stem length response to within-row com-
petition was expressed by the equation Y = 175 + 2.019X (Figure 1A) or quadratic
equation Y = 132 + 3.708X - 0.0136X2, which suggests that muskmelon stem growth
is a function of within-row plant spacing.
It is interesting to note that the quadratic equation predicts a maximum stem
growth of 384 cm at 130 cm within-row plant spacing. Since the maximum was outside
the parameters of this study, the quadratic equation cannot be meaningfully extrapolated.
Leaf area. A highly significant relationship was observed between leaf area and
within-row plant spacing in 1983 (Figure IB). This relationship was expressed by the
equation Y = 118.5 + 0.204X. This positive relationship indicates that muskmelon
leaf area was directly influenced by within-row plant spacing.
400
•
A /
/ *
350
/
300
I
Lkl
250
,
/ R2- 0.87
Y » 175 ♦ 2.019X
t-
/*
206
150
s
2.9
2.8
2
2.6
2.5
25
50
75
100
125
PLANT SPACING (CM)
140
135 .
R*-* -0.99
Y - 3.116 - 6.169
25 50 75 100
PLANT SPACING (CM)
125
£ 130
2
S 125
120
115
12
10
Ik 8
2
^ 6
R « 0.93
Y - 118.5 + 0.204X
25 50 75 100 125
PLANT SPACING (CM)
R'- -0.48
Y - 1278 - 8.404X
25 50 75 100
PLANT SPACING (CM)
125
Figure 1A-D. Relationship between within-row plant spacing and A) stem length;
B) leaf area; C) dry matter per hectare; and D) culls per hectare in muskmelon cv. Classic.
Botany
101
Table 1 . Effect of Within-row Plant Spacing on Muskmelon Petiole Nutrient Content.
Plant
N
P
K
Ca
Mg
Mn
Fe
B
Cu
Zn
Al
Na
spacing
Percent
(cm)
PPM
25
3.3
0.27
2.73
4.93
0.51
683
174
20
10
41
124
212
50
2.6
0.30
3.06
4.72
0.47
614
156
22
9
42
107
242
75
4.5
0.29
3.14
4.90
0.47
637
166
22
10
43
1 17
205
100
3.1
0.26
2.84
5.31
0.48
642
164
19
10
42
130
217
'Values reported are means of two replications.
Dry matter. Total above ground dry matter, excluding fruit, decreased linearly
with increased within-row plant spacing, e.g., 2.5 mt/ha at 100 cm and 3.0 mt/ha
at 25 cm within-row plant spacing. This highly significant linear relationship was
expressed by the equation Y = 3.116 - 6.169"3x (Figure 1C).
Nutrient content. Data on muskmelon petiole and soil nutrient content for the
1983 growing season are reported in Tables 1 and 2.
Yield. Total marketable yields (mt/ha) was not affected by plant spacing in either
year. These results differ from those reported by Lazin and Simonds (5), who reported
a highly significant decrease in yield from 19.9 to 15.1 mt/ha as within-row plant spacing
increased from 30 to 90 cm under Florida conditions. It is interesting to note that
yield of cv. Classic was 23 percent more than that of cv. Burpee Hybrid.
Culls. Number of culls (unmarketable fruits) per hectare for 'Classic' muskmelon
was highest at the closer spacings and lowest at the wider spacings (Figure ID). These
results are in agreement with Zahara (6).
Number of fruits. As plant spacing increased from 25 to 100 cm, the number
of marketable fruits per hectare decreased (Figure 2A). Furthermore, highly signifi-
cant negative coefficients of determination (R2) between plant spacing and number
of marketable fruits per hectare suggest that number of fruits per hectare is closely
related to plant spacing. These results agree with those of Davis and Meinert (2) and
Frazier (3). Zahara (6) found that as plant spacing increased from 25 x 25-cm to 75
x 75-cm, the number of marketable fruits per 15.2 m row increased from 0 to 27.
In his study, Zahara (6) was dealing with much higher plant populations (18,000 to
160,000 plants per hectare) and the increased competition with increased plant popula-
tion resulted in yield decreases (4). My studies, however, dealt with plant populations
of only 3,600 to 14,500 plants per hectare and only within-row competition. Holliday
(4) concluded that a certain plant population gives a maximum yield, while larger or
smaller populations give lower yields. Zahara's study was probably at the "larger"
population according to Holliday's parabolic curve.
Table 2. Effect of Within-row plant spacing on soil nutrient content.*
Plant
P
K
Ca
Mg
Mn
spacing
(cm)
ppm
25
78
58
310
24
72
50
66
80
370
36
45
75
82
80
340
33
69
100
73
85
350
23
40
'Values reported are means of two replications.
102
Indiana Academy of Science
Vol. 94 (1985)
24
22
2 20
X
5
£ 18
2
16
14
10
5 3
035X
-L.
-L.
25 50 75 100 125
PLANT SPACING (CM)
R - 0.97
. Y - 0.15 ♦ 0.101X
0 25 50 75 TOO 125
PLANT SPACING (CM)
3 6
s!
2 .
/
/ 2
r/ Rz« 0.98
(/ Y - -0.139 + 0.087X
" R2.
0.32 0 .,
2.5
. Y »
1.78 + 0.008X ^/*
2.0
^^
»-
u.
V-
3
UJ
1.5
1.0
^''' R2' 0.79
'+ Y « 1.147 + 0.007X
i i i . j
f5 50 75 100 125
PLANT SPACING (CM)
0 25 50 75 100 125
PLANT SPACING (CM)
Figure 2A-D. Relationship between within-row plant spacing and A) fruits per hec-
tare; B) fruits per plant; C) yield per plant; and D) weight per fruit in muskmelon
cvs. Burpee Hybrid (H + ) and Classic (* *).
Number of fruits per plant. Highly significant linear relationships were established
between fruits per plant and plant spacing during both the years, the coefficient of
determination being 0.94 and 0.98 in 1982 and 1983, respectively (Figure 2B). These
results are in agreement with those of others (2,5).
The quadratic equation for number of fruits per hectare (x 1000) in relation to
within-row plant spacing was Y = 26.375 - 0.153X + 0.0006X2 for 'Burpee Hybrid'
and Y = 22.325 - 0.1302X + 0.00052X2 for 'Classic' muskmelon. Based on these
equations maximum fruits were predicted for 125 cm within row plant spacing. Accord-
ing to these equations, competition ceased at 125 cm within-row plant spacing.
Yield per plant. The mean yield (kg) per plant was highly correlated with plant
spacing each year, increasing significantly as spacing between plants increased from
25 to 100 cm (Figure 2C).
Fruit weight. The mean fruit weight increased significantly as plant spacing in-
creased from 25 to 100 cm (Figure 2D). Lazin and Simonds (5) reported that as distance
between plants increase from 30 to 90 cm, mean muskmelon weight increased from
1.36 to 1.53 kg. The highly significant coefficient of determination between plant spacing
and fruit weight (Figure 2D) suggests that fruit weight is a function of plant spacing
and can be manipulated to meet consumer and/or market demand.
Botany
103
Soluble solids. Within-row plant spacing had a significant effect on soluble solids.
The highly significant linear relationship showed that soluble solids increased as the
within-row plant spacing increased from 25 to 100 cm (Figure 3). Davis and Meinert
(2) and Zahara (6) also reported similar results.
12
B
(•)
Q
5
«/>
H 11
C3
=3
10
Y • 9.75 ♦ 0.026X
+
R2- 0.95
Y • 10.05 + 0.008X
0 25 50 75 100 125
PLANT SPACING (CM)
Figure 3. Relationship between within-row plant spacing and soluble solids in
muskmelon cvs. Burpee Hybrid (H h) and Classic (* *).
Summary
The effects of within-row plant spacings of 25, 50, 75, and 100 cm with row
spacing of 2.7 m on 'Burpee Hybrid' and 'Classic' muskmelons were evaluated in field
studies conducted on a southwestern Indiana sandy loam, mixed, mesic Typic Hapludalf
soil in 1982 and 1983. With increased plant spacing from 25 to 100 cm, stem length,
leaf area, yield (kg) per plant, number of fruits per plant, fruit weight, and soluble
solids increased linearly, whereas dry matter, number of culls, and marketable fruits
per hectare decreased linearly. Plant spacings had no significant effect on soil and
petiole nutrient content and total marketable tonnage.
Note
Joint contribution from USDA-ARS, and Department of Horticulture, Purdue
University, West Lafayette, Indiana. Mention of firm names or trade products does not
imply endorsement or recommendation by the USDA or Purdue University over other
firms or similar products not mentioned.
Literature Cited
1. Anonymous. 1980. Recommended chemical soil test procedures for the North
Central Region. North Central Regional Publication No. 221 (Revised). W.C.
Dahnke (ed.). North Dakota Agr. Exp. Stn. Bull. No. 499 (Revised).
2. Davis, G.N. and U.G.H. Meinert. 1965. The effect of plant spacing and fruit
pruning on the fruits of P.M.R. No. 45 cantaloupe. Proc. Amer. Soc. Hort.
Sci. 87:299-302.
3. Frazier, W.A. 1940. Fruiting of the Powdery Mildew Resistant No. 45 cantaloupe
as affected by spacing. Proc. Amer. Soc. Hort. Sci. 37:831-835.
4. Holliday, R. 1960. Plant population and crop yield. Nature 186 (4718):22-24.
5. Lazin, M.B. and S.C. Simonds. 1981. Influence on planting method, fertilizer
104 Indiana Academy of Science Vol. 94 (1985)
rate, and within row plant spacing on production of two cultivars of honeydew
melons. Proc. Fla. State Hort. Soc. 94:180-182.
6. Zahara, M. 1972. Effects of plant density on yield and quality of cantaloupe.
Cal. Agr. 26:(7):15.
Stem Length as an Estimator of Muskmelon Growth
H.S. Bhella
USDA-ARS, Vincennes University
Vincennes, Indiana 47591
and
G.E. Wilcox
Department of Horticulture
Purdue University, West Lafayette, Indiana 47907
Introduction
Measurement of growth parameters is an essential component of plant science
research. In order to determine stem and root growth response to various cultural,
management, and insect-pest control practices, plant scientists sacrifice live plants from
their experimental plots. This direct measurement of top or root growth, a universally
accepted criterion for growth measurement, is costly, laborious, and time consuming.
It inflates the error variance because intact plants are permanently lost from experimental
plots. Furthermore, this permanent loss of plants reduces plant population and does
not allow repeated observations for measurement of plant growth and development
on the same plant.
Because of these implications, the melon working group consisting of USDA-
ARS and Purdue University horticulturists, entomologists, plant pathologists, and plant
physiologists was looking for an alternative method for muskmelon growth measure-
ment. This study was undertaken as part of a team approach for determining and
documenting a rapid, reproducible, and practically acceptable method for muskmelon
growth measurement, which is as reliable as the top weight method and allows repeated
observations on the same plant. The main objective of this greenhouse study was to
obtain the closest fit regression equations between total stem length and stem and root
weight, which could be used to estimate muskmelon growth with minimal error.
Materials and Methods
Muskmelon cv. Burpee Hybrid seeds were sown on February 2, 1983 and cv.
Saticoy on January 4, 1984 in no. 38 growing trays (Growing Systems, Inc., Milwaukee,
WI 53213) containing Jiffy-Mix (composed of shredded sphagnum peat moss and
horticultural-grade vemiculite) growing medium. The seedlings were grown under
greenhouse conditions at day/night temperature of approximately 30/21 ±3°C and 16-hr.
light from 20-watt Luxor, Vita-Lite lamps suspended 27 cm above the plants with one
bulb per 0.28 m2 of bench space.
Fifteen-day old seedlings (3 to 4 true leaf stage) of cvs. Burpee Hybrid and Saticoy
were transplanted into one-liter black plastic pots using 'Flint shot' sand as growing
medium. Because of difficulties in separation of roots from growing medium for
muskmelon cultivars sown in Jiffy-Mix, 'Classic' muskmelon was direct seeded into
one-liter black plastic pots containing 'Flint shot' sand on March 8, 1984. Plants were
fed daily with Hoagland's solution (1) containing various levels of nitrogen (up to
200 ppm) to achieve wide range of variation in stem and root growth rates. Iron chelate
was added to the Hoagland's solution twice a week.
Muskmelon plants were harvested five weeks after seeding or 3 weeks after
transplanting. Data on total stem length and stem fresh/wet weight were recorded during
harvest. Muskmelon cv. Classic roots were separated from the growing medium (sand)
by gently washing with tap water over a screen and blotted. The stem and root samples
were dried in a forced air oven for 48-hr at about 50°C and then weighed.
105
106
Indiana Academy of Science
Vol. 94 (1985)
Results and Discussion
In this study, we found highly significant positive relationships between stem length
and top growth (Figure 1). The coefficients of determination for muskmelon cvs. Saticoy,
Burpee Hybrid and Classic were 0.95, 0.98, and 0.98 for top fresh weight, and 0.96,
0.97 and 0.99 for top dry weight, respectively. Interestingly, highly significant correla-
tions were also obtained between stem length and total (top plus root) dry weight
(R2 =0.99) and root dry weight (R2 =0.71) for muskmelon cv. Classic. Complete separa-
tion of entire root system from Jiffy-Mix for muskmelon cvs. Saticoy and Burpee
Hybrid was not achieved, thus, their results are not reported. These highly significant
cr
40
_ cv
Y=-
Saticoy
-3.36+0. 508X
i—
30
R2=
=0.95
f*
X
C/3
LU
OS
LL
20
i /
Q_
o
f-
10
0
I 1 1
" '
CV. Saticoy
U3 40 -
£ 30-
00 20
u_
o. 10
o
K 0
3 40
fe 30
J2 20 L
u_
Q. 10 -
o
i—
0
o 2.5
2 2.0
>-
g 1.5
8 1-°
1 °"5
20 40 60 80
STEM LENGTH (cm)
CV. Burpee Hybrid
Y=-4.92+0.476X
R =0.
20 40 60 80
STEM LENGTH (CM)
0 20 40 60 80
STEM LENGTH (CM)
CV. Classic
Y=0. 31+2. 57x10
R2=0.99
20 40 60 80
STEM LENGTH (cm)
100
100
100
100
2
5
ID
2
0
h-
2
1
5
>>
a
1
0
O
1—
0
5
n
20 40 60 80
STEM LENGTH (cm)
CV. Burpee Hybrid
Y=-0.02+2.33xlO"2X
R2=0 . 97
0
2.5
2.0U
1.5
1.0
0.5h
0
20 40 60 80 100
STEM LENGTH (cm)
CV. Classic
Y=0.19+2.35*10-2X
R2=0.99
0 20 40 60 80
STEM LENGTH (CM)
20 40 60 30
STEM LENGTH (cm)
100
0.4
CV. Classic
Y=0.12+2.26xlO"3X
CJ
R2=0.71
*
1—
s
0.3
>-
(Y
o
0.2
* ^^^
O
^^^ *
o
oc
0.1
i.i i
' 1 —
100
Figure 1. Relationships between various growth parameters of muskmelon. All co-
efficients of determination significant at 0.01 level of probability.
Botany 107
correlation coefficients and regression equations support the hypothesis that stem length
is as reliable an indicator of growth as is the destructive harvest for dry weight measure-
ment of top growth.
The results of this study establishes a direct relationship between stem length and
top growth and root growth that can be used to evaluate top and root limiting condi-
tions in muskmelon. Furthermore, this study demonstrates that stem and root weights
can be indirectly predicted with high degree of accuracy by entering stem length in
the regression equations, as well as for the inverse problem of estimating stem length
from stem and root weights.
Highly significant correlations between various growth parameters for three
muskmelon cultivars suggest the stem length method is reproducible. In terms of
simplicity, practicality, and rapidity, both methods are useful means of determining
muskmelon growth. However, the stem weight method has a serious drawback because
of permanent loss of plants from experiment. The stem length method has important
advantages because it allows repeated observations on the same plant. This method
should be useful to plant scientists measuring growth of vine crops.
Summary
A rapid, reproducible, and practically acceptable method for measuring muskmelon
growth, without destroying intact plants was developed. We tested the hypothesis that
total stem/vine length measurement of muskmelon provides the same indications of
growth as does stem/top weight, a universally accepted criterion for plant growth
measurement. Total stem length was found to be highly correlated with top fresh
(R2 = 0.95, 0.98 and 0.98) and dry (R2 = 0.96, 0.97, and 0.99) weights for muskmelon
cvs. Saticoy, Burpee Hybrid, and Classic, respectively. Total dry matter (top plus root;
R2 = 0.99) and root dry weight (R2 = 0.71) were also found to be highly correlated with
stem length for cv. Classic. These highly significant coefficients of determination sug-
gest that stem length in the early stages of plant development is a reliable and reproducible
estimator of muskmelon growth.
Note
Joint contribution from the U.S. Department of Agriculture, Agricultural Research
Service and Department of Horticulture, Purdue University, West Lafayette, IN 47907.
Mention of firm names or trade products does not imply endorsement or recommenda-
tion by the USDA or Purdue University over other firms or similar products not mentioned.
Literature Cited
1. Hoagland, D.R. and D.I. Arnon. 1950. The water-culture method for growing
plants without soil. Revised by D.I. Arnon. CA Agric. Exp. Stn. Cir. 347. 32 p.
Isolation of the Corprophilous Fungus, Pilobolus,
from Wayne County, Indiana
K. Michael Foos and Judith A. Royer
Department of Biology, Indiana University East
Richmond, Indiana 47374
Introduction
Pilobolus is a microscopic zygomycete that grows on the dung of herbivores. And,
while it appears to be widely distributed, it has not been widely recorded. Records
of Pilobolus from North America are particularly uncommon.
Pilobolus has been recorded from Ohio (4), Michigan (1), New York and Penn-
sylvania (5), but no records of Pilobolus from Indiana exist. In this study samples
of dung were collected in Wayne County, Indiana and examined for isolates of Pilobolus.
Methods and Materials
Beginning mid-winter, collections of Pilobolus were made from samples of dung
of herbivores collected at 35 locations in Wayne County, Indiana. These samples were
collected from sheep, beef and dairy cattle, horses, ponies, llamas and deer. Only fresh
samples of dung were collected. These collections were made aseptically in plastic baggies
and were transferred within hours to sterile preparation dishes lined with water saturated
filter paper. All cultures were maintained at room temperature under cool white fluores-
cent lights with an intensity of 320 foot candles with a 12 hour photoperiod until growth
with visible sporangiophores appeared.
Upon maturity Pilobolus discharge their sporangia. These sporangia adhered to
the tops and sides of the preparation dishes because of the gelatinous layer around
the sporangium. Isolates were obtained by removing single sporangia from the sides
or tops of the preparation dishes with a sterile inoculating needle. Each sporangium
was transferred to a petri dish containing dung agar (6). After 1 to 1 1/2 weeks growth
would fill the petri dish depleting the media. At such times hyphal tips were trans-
ferred to fresh media. Active hyphal growth could be seen within 24 hours after transfer.
Development of Pilobolus sporangia is influenced by light, so sporangia were
collected between 9 am and 1 pm daily (the photoperiod was set between 8 am and
8 pm). Sporangia were collected from the lids of petri dishes or from sporangiophores
with sterile inoculating needles or microforceps, and mounted in lactophenol. Spores
were also observed in the lactophenol mounting preparation. Pressure on the coverslip
broke the sporangial wall releasing spores for observation and measurement. Columella
were observed by removing sporangia with microforceps.
The following characteristics were observed and measured:
1. Spore size, shape, color, and wall thickness
2. Sporangium size, shape, and ornamentation
3. Sporangiophore length
4. Trophocyst size and shape
5. Subsporangial swelling size, shape, and color
6. Columella shape
Measurements of taxonomic structures were made both from the original isolates
on dung and later from the growth on dung agar.
Results
The genus Pilobolus has rarely been seen to reproduce sexually. Its normal method
109
110 Indiana Academy of Science Vol. 94 (1985)
of asexual reproduction is by the production of sporangiospores within a sporangium.
The structure of this asexual reproductive complex, and the component structures are
the primary taxonomic characteristics for the genus.
In nature, Pilobolus grows submerged in dung with its sporangial apparatus rising
above the surface. The sporangial apparatus of Pilobolus is unique. It consists of the
sporangium, containing sporangiospores, the subsporangial swelling, the sporangiophore,
and the trophocyst. The sporangium is covered with a thick cutinized wall that is dark-
ly pigmented and rests at the apex of the sporangiophore. This sporangium contains
thousands of sporangiospores. Unlike many zygomycetes, the spores within the Pilobolus
sporangium remain together and act as a sporangial unit. Upon maturity the sporangium
with all of its spores is forceably discharged from the sporangiophore and travels as
far as 8 feet. This characteristic gives Pilobolus its name "hat thrower" (2).
The subsporangial swelling is the portion of the sporangiophore located just below
the sporangium. It is a widened area of the sporangiophore which is light sensitive
and acts in 'aiming' or directing of the sporangium prior to discharge. Below the
subsporangial swelling is the long, slender sporangiophore. The sporangiophore measures
from 1 mm to several centimeters in length in different species and holds the sporangium
above the surface of the substratum.
The trophocyst, a structure unique to Pilobolus, is located at the lower end of
the sporangiophore. It is embedded in the substratum and anchors the sporangial ap-
paratus. The trophocyst may be elongated, somewhat oval, or turnip shaped.
These structures: the sporangium, sporangiospores, subsporangial swelling,
sporangiophore, and trophocyst are the primary characteristics used in the taxonomy
of Pilobolus.
From twenty-eight isolates of Pilobolus from Wayne County, Indiana, four dif-
ferent species were recovered. These species were: Pilobolus cry stallinus, Pilobolus kleinii,
Pilobolus longipes, and Pilobolus roridus.
Pilobolus crystallinus Tode (7)
Pilobolus crystallinus sporangiophores develop in 3 to 4 days, are 1 to 5 mm long,
and are clear to pale yellow in color. Trophocysts develop submerged in the substratum
and are usually 500 /*m long by 350 ^m wide. Sporangiospores are pale yellow ellipses
which measure 9.83 ± 0.90 /xm in length by 6.05 ± 0.75 /*m in width producing a
length to width ratio of 1.62.
Sporangia are covered with a dark, cutinized wall and range from 100-750 /*m
in diameter, with a mean of 205.7 ± 49.7 fim. About 1/3 of the isolates recovered
have polygonal reticulations as described by van Tieghem (8).
Pilobolus crystallinus was isolated in 15 locations in Wayne County during March
through August from the dung of sheep, cattle, donkey, goat, llama, and pony.
Pilobolus kleinii van Tieghem (8)
Pilobolus kleinii sporangiophores measure 2-3 cm in length and arise from dark yellow
turnip shaped trophocysts measuring 300-500 (im in diameter. The trophocysts are often
partially submerged within the substratum. Sporangia are dark, smooth, and cutiniz-
ed. They measure 100-300 /mi across with a mean of 146.25 ± 98.8 /*m, and are about
2/3 as high as wide. The columella are conical and extend deeply into the sporangia.
Sporangiospores are yellow and elliptical, measuring 12.14 ± 1.16 (im in length by
7.57 ± 0.59 /on in width with a length to width ratio of 1.60.
Pilobolus kleinii was isolated in 5 locations in Wayne County between March
and August from the dung of cows, sheep, and goats.
Pilobolus longipes van Tieghem (8)
Pilobolus longipes sporangiophores range from 5 mm to 3 cm (sometimes longer) and
Botany 1 1 1
develop from large trophocytes often 1 mm or more in length. Sporangiophores grow-
ing from freshly collected dung are much longer than those growing from isolates
transferred to samples of sterilized dung or to dung agar. Sporangia are nearly globose,
smooth, dark, cutinized and vary greatly in size from 100 to more than 400 /mi in
diameter. However, the mean diameter for sporangia is 226.3 ± 53.7 /im.
Sporangiospores are subglobose to globose, dark yellow to orange in color and measure
12.23 ± 1.59 fim by 11.23 ± 1.52 /im with a length to width ration of 1.09.
Philobolus longipes was isolated at 7 locations in Wayne County during May
through July. All isolates were taken from horse dung.
Pilobolus roridus (Bolt.) Pers. (3)
Pilobolus roridus sporangiophores are 1 to 2 mm long. The sporangia are smooth
and hemispherical, and average 260.0 ± 22.4 /im in diameter. The trophocysts are
250-300 /im in diameter, nearly spherical, and bright orange in color. The sporangiospores
are pale yellow to colorless, oval in shape and measure 5.79 ± 0.68 /mi in length
and 3.07 ± 0.35 /im in width. The length to width ratio is 1.89.
Pilobolus roridus was isolated in Wayne County during August from deer dung.
Discussion
Even though Pilobolus has been isolated in many places, this is the first record
of isolates from Indiana. The source of the substratum from which the organism was
isolated has been recorded. Even though there seems to be some relationship between
the species of Pilobolus isolated, and the type of dung on which it was found, there
has not been a direct correlation shown. It is, however, interesting to note that P.
longipes was isolated seven times in this study and in all instances it was isolated from
horse dung. The time of year each collection of dung was made was recorded. During
the winter it was difficult to obtain isolates of Pilobolus. Dung of herbivores not ob-
taining at least part of their food by grazing on pasture included no isolates of Pilobolus.
It is easy to speculate that Pilobolus can be isolated only from the dung of animals
grazing on open pasture. Certainly, this was the case in this study.
Many of the taxonomic characters traditionally used with Pilobolus are of ques-
tionable value in separating the various species. This is because of the wide variation
that occurs within a single isolate. Sporangium size, subsporangial swelling size and
shape, and to some degree length of the sporangiophore have little value. The fluctua-
tion in these characters makes them almost useless.
The sporangiospores seem to be the most constant and thus exhibit the most
valuable taxonomic characteristics. The size, shape, and coloration of the sporangiospore
are reliable taxonomic characteristics. Regardless of the size of the sporangium, the
sporangiospores contained within remain remarkably constant.
Acknowledgments
We would like to thank Lori Westberg for her technical assistance, Dr. Dorothy
Adalis for the use of her photomicroscope, and the Indiana University Research Opera-
tions Committee for its financial support in this project.
Literature Cited
1. Bessey, E.A. 1946. Studies of Pilobolus: P. kleinii and P. longipes. Papers
Michigan Acad. Science 32:15-26.
2. Buller, A.H.R. 1934. Researches on fungi. Volume VI. Longmans, Green, and
Co. London. 513 p.
112 Indiana Academy of Science Vol. 94 (1985)
3. Persoon, C.H. 1801. Synopsis methodica Fungorum. Part 1. p. 117-118.
4. Rakestraw, James B. and K. Michael Foos. 1980. Isolation of the coprophilous
fungus, Pilobolus, from Lake County, Ohio. Ohio J. Science 80:20 (Abstr.)
5. Sumstine, D.R. 1910. The North American Mucorales. Mycologia 2:125-154.
6. Swartz, Delbert. 1934. Pilobolus crystallinus in pure culture. Mycologia
26:192-194.
7. Tode, H.J. 1784. Beschreibung des Hutwerfers. Schrift. Gesell. Naturf. Freunde
Bed. 5:46.
8. Van Tieghem, P. 1876. Troisieme memorie sur les Murocinees. Ann. Sci. Nat.
4:335-349.
A New Amine as an Uncoupler of Chloroplast Electron Transport
Jonathan Leeds, Lynne Bemis, Rita Barr and Frederick L. Crane
Department of Biological Sciences
Purdue University
West Lafayette, Indiana 47907
Abbreviations used: DAD-diaminodurene; DBMIB-2,5,8-dibromo-3-methyl-6-
isopropyl-p-benzoquinone; DCMU-dichlorophenyl-dimethylurea; DMBQ-2,
5-dimethylbenzoquinone; DNP-INT -2, 4-dinitrophenylether of iodonitrothymol; FCCP-
carbonylcyanide-p-trifluoromethoxyphenylhydrazone; MV-methylviologen;
TMPD-N-tetramethyl-p-phenylenediamine.
Introduction
In isolated chloroplasts electron transport is coupled to photophosphorylation
(1,2). To study electron transport rates in Photosystem I and II, certain chloroplast
reactions require an uncoupler to be present. The common uncouplers used for this
purpose are FCCP, ammonia and such ionophores as gramicidin (2).
In this study we describe a new amine-type uncoupler, N-[bis-(3,5-trifluoromethyl)-
phenyl]-2,4-dinitro(3-trifluromethyl)-benzamine (DP A, Figure 1), which appears to work
3 w' 3
Figure 1. The Chemical Composition of the Uncoupler, DPA.
best at coupling site 1, located between the two photosystems in the chloroplast elec-
tron transport chain. We show that low concentrations (1 x 10 ~7) are required to
stimulate electron transport 60% or to inhibit the proton gradients associated with
photophosphorylation.
Materials and Methods
Spinach or lettuce chloroplasts were prepared from commercially available sources
by methods previously reported (3). Briefly, about 20g of leaves were ground in a
Waring blender in 100 ml sucrose-NaCl (0.4 M sucrose, 0.05 M NaCl) with 6 on-and-
off bursts of energy. The resulting green suspension was filtered through 10 layers
of cheesecloth and a single layer of Miracloth into 2 50-ml centrifuge tubes. Heavy
particles, such as the remains of cell walls and nuclei, were pelleted after centrifuga-
tion at 600 x g for 2 min and discarded. The supernatant was filtered through Miracloth
113
114
Indiana Academy of Science
Vol. 94 (1985)
into clean tubes and centrifuged at 1,200 x g for 10 min. to collect chloroplasts, which
were suspended in 5 ml SN. Chlorophyll was determined according to Arnon (4).
Oxygen uptake or evolution were measured with a Clark-type electrode connected
to a Yellow Springs Instrument oxygen monitor. Reaction rates were recorded with
a Sargent-Welch SRG recorder. Chloroplast proton pump was assayed by the methods
of Dilley (5).
DPA was synthesized in the Eli Lilly Laboratories and made available through
the courtesy of Dr. Hollingsworth, Purdue Department of Entomology.
Results and Discussion
An uncoupler should stimulate electron transport reactions in low concentrations
(1 x 10" 6 to 1 x 10~9). As Figure 2 and 3 show, DPA meets this criterion. Partial reac-
tions, which are known to involve coupling site 1, such as H20 — MV (+ azide)
and H20 - FeCN (pH 6 or 8) are stimulated from 30-60%, whereas H20 - FeCN
with DNP-INT or H20 - DDMBQ with DBMIB show little stimulation of electron
- MV, pH 8
AFeCN,pH 8
• FeCNtpH 6
■ F«CN, pH 8 (+DNPINT)
D DMBQ.pH 7 (♦DBMIB)
3 10 30
DPA (n molar)
100
300
KXX>
Figure 2. Uncoupling of Photosystem I and II Reactions by the Uncoupler, DPA.
Reaction mixtures contained chloroplasts (0.05 mg chlorophyll), 25 mM Tris-Mes, pH
6, 7, or 8, as shown and electron acceptors or inhibitors in concentrations indicated
below: DMBQ-lOmM; DCMU-5^M; DNP-INT; 10/xM; and FeCN 250 or 500 fiM.
transport, since they accept electrons before coupling site 1 (Figure 4). Likewise, PS
I reactions, which involve this site also stimulate electron transport rates from 40-60%
(Figure 3).
Botany
115
70
60
50
40
55
z
30
Q
1-
20
m
X
z
10
n
rr
0
0
z
0
-10
b
<
-20
3
2
-30
H
en
-40
-50
-60
-70
- Asc. +DAD-«»MV, pH8
O Asc. + DAD-*-MV, pH8 ( + DBMIB)
• Asc. +DAD-*-MV, pH8 (+DNPINT)
■ Asc. + TMPD-^ MV, pH 8
A Asc. +TMPD-*-MV, pH 8 (+DNPINT)
D Duroquinol -*» MV, pH 6
1
10 30
DPA (n molar)
100
300
1000
Figure 3. Uncoupling of Photosystem I Reactions by the Uncoupler, DPA. Reac-
tion mixtures as in Fig. 1 with additional reaction components in concentrations in-
dicated below: ascorbate, ImM; DAD, 0.5mM; TMPD, 5/xM; and duroquinol, 0.5mM.
PSD
Mn
FeCNn pH 8
Asc. + TMPD
ADP+Pi
MV
e"
FO AND
Reductase
Complex
©
NADP
PS I
H20 02
Figure 4. The Z-scheme of Chloroplast Electron Transport, Showing Coupling Site
I, Uncoupled by DPA.
116 Indiana Academy of Science Vol. 94 (1985)
The second criterion for establishing a compound as an uncoupler is to show
that it inhibits the light-dependent proton pump associated with photophosphoryla-
tion. According to Table I, 100 nanomolar DPA (1 x 10 ~ 7 M) inhibits the chloroplast
proton pump 49%. Higher concentrations of the uncoupler lead to > 90°7o inhibition.
These criteria establish DPA as a potent new uncoupler. Only a few known un-
couples, such as FCCP (6,7) or TTFB (8) uncouple in lower concentrations than DPA
(1 x 10" 8 M versus 1 x 10" 7 M, respectively). This new uncoupler involves coupling site
1, located between the 2 photosystems (Figure 4).
Table I. Inhibition of Chloroplast Proton Pump Associated with ATP Formation
DNP cone. ATP INHIBITION
(nM) OtMOLES/mg CHL'HR) («7o)
0 420 —
10 362 14
30 315 25
60 252 40
100 213 49
300 174 59
600 142 76
1000 20 95
Literature Cited
1. Trebst, A. 1974. Energy conservation in photosynthetic electron transport of
chloroplasts. Ann. Rev. Plant Physiol. 25, 423-458.
2. Good, N.E. 1977. Uncoupling of electron transport from phosphorylation in
chloroplasts. In Encyclopedia of Plant Physiol., new series, vol. 5. (Trebst, H.
and M. Avron, eds.), Springer-Verlag, Berlin, pp. 429-436.
3. Seng, T.W., R. Barr and F.L. Crane, 1983. L-Methionine Sulfoximine as a
new electron acceptor in Photosystem I of spinach chloroplasts. Proceed. Indiana
Acad. Sci. 92, 119-123.
4. Arnon, D.I. 1949. Copper enzymes in isolated chloroplasts. Polyphenoloxidase
in Beta vulgaris. Plant Physiol. 24, 1-15.
5. Dilley, R.A. 1972. Ion transport (H+, K + , Mg2+ exchange phenomena), in
Methods of Enzymol., vol. XXIV, part B (A. San Pietro, ed.), Academic Press,
New York, pp. 68-74.
6. Heytler, P.G. 1979. Uncouplers of oxidative phosphorylation, in Methods in
Enzymol., vol. LV, part F (S. Fleischer and L. Packer, eds.), Academic Press,
New York, pp. 462-472.
7. Heytler, P.G. and W.W. Prichard. 1962. A new class of uncoupling agents-
carbonyl cyanide phenylhydrazones. Biochem. Biophys. Res. Commun. 7, 272-275.
8. Jones, O.T.G. and W.A. Watson. 1965. Activity of 2-trifluoromethyl-
benzimidazoles as uncouplers of oxidative phosphorylation. Nature (London) 208,
1169-1170.
A Rapid Method for the Determination of Barley Seed Viability
Gayton C. Marks, William W. Bloom, Jeffrey G. Boyle
Department of Biology, Valparaiso University
Valparaiso, Indiana 46383
Introduction
Barley farmers and shipment processors currently need a more rapid method for
the determination of barley caryopses viability, especially for those grains used for
malting. Not only is the time required for the test significant, but the methods used
must also produce accurate results to ensure an adequate evaluation of the barley sample
for processing. Grain dealers must know the quality of the incoming barley so that
they can transfer it either to the appropriate purchasing company or to proper storage
for future sale and shipment. Since much of the grain is brought to the elevators directly
from the fields, lengthy testing procedures delay unloading trucks and result in delayed
processing and inefficient shipment control.
The current methods of testing viability involve lengthy determination times as
well as complicated procedures. In the most accurate test, one hundred fruits (hereinafter
identified as seeds) are placed in a flat dish on wet filter paper or tissue and allowed
to germinate for approximately 24 hours. Viability is then determined by deriving a
total percentage of seeds from which the coleorhiza (chit) has emerged. While the results
from this test are accurate and easily assessed, the amount of time involved usually
extends beyond the desired limit.
The Schonfeld test involves placing a filter-funnel with 100 barley seeds into a
cabinet at 18-20°C and subjecting the seeds to a saturated water vapor atmosphere.
The seeds are steeped, drained, and re-wet so that after a certain time, the germin-
ability percentage can be observed. In a similar technique called the Schonjahn or
Coldewe method, the seeds are placed in holes within porcelain plates so that the em-
bryo is pointing down. The plates are housed in a container full of sand and water.
After germinating time has elapsed, viability percentages are obtained by counting the
roots growing through the holes (1).
Waller attempted to determine germinability of Phaseolus seeds through electronic
methods. Beans were soaked in water, split, and the radicle removed and connected
to electrodes. An induction coil provided sufficient stimulus and a galvanometer was
then used to measure shock deflection. Viable seeds were identified by greater relative
deflection and non-viable seeds produced no response (3). Later, Fraser used the same
electronic method to test barley embryos and, in fact, confirmed Waller's results (3).
Although these techniques presumably identified differences in living and dead tissues,
no further investigation has been undertaken.
There are a number of staining methods used involving color changes that in-
dicate the viability or non-viability of a seed. Of the various staining methods used,
one early test by Dimitriewicz employed sulfuric acid in a timed observation where
viable seeds turned a deep rose color in five minutes. Further, respiratory activity of
seeds from soaking in meta-, para-, or orth-dinitrobenzene solutions for twenty hours
followed by ammonia for one hour results in an orange color for viable seeds. Selenium
reduction produces a purple color, both of which are also indicative of seed germinability
(1).
In addition, dead seeds have been known to take up barium chloride which can
then be detected by x-rays. Seeds can also take up rasazuria and indigo stains for
color detection observation (1). Another common method involves the biochemical
activity of living seeds whereby endogenous enzymes and substrates reduce
117
118 Indiana Academy of Science Vol. 94 (1985)
2-(p-iodophenyl)-3-(p-nitrophenyl)-5-phenyl tetrazolium chloride to easily detected in-
soluble red formazanes. This is the most popular method used today among grain
elevators and farmers. A sample of at least one hundred seeds is taken from a truck
or storage compartment and individually counted in a dish on the cutting device. Each
seed is then dropped down a small rack until it rests between a secured cutting ridge
at the center of the device. The seed must then be held in place while a blade is slid
across it, longitudinally splitting the barley into two equal pieces. This process exposes
one side of the embryo and is subsequently stained and counted for viability. Although
this method has proven accurate, it requires a 45-60 minute incubation period and
also involves inefficient and lengthy seed preparation.
Materials and Methods
The tetrazolium (2,3,5-triphenyl-2H tetrazolium chloride monohydrate) test for
the viability of seeds is based on the principle that living tissues release hydrogen as
part of the respiratory process occurring in the mitochondria. Hydrogen combines with
the colorless tetrazolium salt and produces a red pigment (2). Since seeds are largely
dehydrated at maturity, it is necessary to hydrate the embryo for the enzyme systems
to function. With intact seeds, this is a slow process. Even when barley grains are
split in the currently used method described earlier, only a small amount of surface
area is actually exposed to the water in the test solution, requiring one to several hours
for a positive test for viability (4). In addition, the handling of the seeds individually
is time consuming.
It is well known that living plant cells can be separated and continue to function.
A method was sought that would reduce the time required for preparation and to
expose more cells of the embryo directly to the hydrating solution. The most suitable
method devised was to subject the barley grains to considerable pressure, thereby flat-
tening the grain and exposing a relatively large surface area to the test solution. A
9cm Whatman filter paper disc was placed in the inverted lid of a 10cm plastic Petri
dish and saturated with a 0.50% solution of tetrazolium chloride. Twenty-five barley
seeds were arranged on the filter paper and covered with the bottom of the Petri dish.
A metal disc 8cm in diameter and 3cm thick was placed inside the bottom of the Petri
dish. This assembly was placed on a Carver hydraulic press and pressed at 16,000-18,000
lbs. Additional test solution was added after pressing to ensure sufficient saturation
of seeds. This resulted, then, in the exposure of abundant embryonic tissue to rapid
hydration and penetration of the tetrazolium salt into the cells. The water activates
the enzyme systems and effects the release of hydrogen ions which then react with
the tetrazolium chloride. Positive tests were detectable by the presence of a pink or
red pigment in the embryonic tissue in less than ten minutes. In twenty minutes, accurate
evaluation of all the seeds could be made.
As a check on the validity of this method, a standard wet-towel germination test,
which required twenty-four or more hours to complete, was made for each sample
of seeds tested.
Results
The results of the viability test by the two testing methods are shown in Table 1.
Discussion
While the tetrazolium chloride test with pressed barley seeds may not legally
substitute for standard tests now in use, it seems to provide a more rapid and suffi-
ciently accurate method to permit sorting for storage and shipping purposes.
Botany
119
Table 1. A comparison of the results of the two methods for determining barley seed
viability.
Group A
Group B
Group C
Group D
Tetrazolium Wei Towel Tetrazolium Wet Towel Tetrazolium Wet Towel Tetrazolium Wet Towel
Test Test Test Test Test Test Test Test
Trial 1
a) tt seeds
100
100
100
100
100
100
100
100
b) Germinated and
93
76
91
89
100
98
93
95
% Viability
Trial 2
a) tt seeds
100
100
100
100
100
100
100
100
b) Germinated and
95
81
92
93
100
99
96
95
% Viability
Trial 3
a) tt seeds
100
100
100
100
100
100
b) Germinated and
92
86
96
94
99
99
% Viability
Trial 4
a) tt seeds
100
100
100
100
100
100
b) Germinated and
88
96
95
100
100
100
% Viability
Trial 5
a) tt seeds
100
100
100
100
100
100
b) Germinated and
93
93
97
99
99
99
% Viability
Average Viability
92.2%
86.4%
94.2%
95.0%
99.6%
99.0%
94.5%
95.0%
Work is being continued to determine the effect temperature has on viability deter-
mination rate. So far, experimental data suggests that tetrazolium chloride reduction
is temperature dependent within certain limits. Rapid methods for counting the samples
and positioning the seeds for testing also are being explored.
Literature Cited
1.
2.
3.
Briggs, D.E. 1978. Barley, Chapman and Hall, New York. 565 p.
Colbry, Vera L., Thomas Swofford, and Robert P. Moore, 1961. Tests for
Germination in the Laboratory. Seeds, The Yearbook of Agriculture, The U.S.
Gov't Printing Office. 441-443.
Crocker, William, and Lela V. Barton, 1957. Physiology of Seeds. Chronica
Botanica Co., Waltham, Mass. 261 p.
Flemion, Florence, and Harriet Poole, 1948. Seed Viability Tests With 2,
3, 5 -Triphenyltetrazolium Chloride. Contrib. Boyce Townsend Inst.
15:243-258.
Population Studies of Threatened and Endangered Plants of Barker Woods
Nature Preserve, LaPorte County, Indiana
Patricia Wiese Reed
233 Hillcrest Road
Michigan City, Indiana 46360
Barker Woods Nature Preserve covers 12 ha in Michigan City, LaPorte County,
Indiana. The geomorphic features of the preserve reflect their origin as part of an
ancient great lake shoreline (Figure 1). The soils on the site are deep, sandy, acid,
100m
500'
♦
Figure 1. Geomorphic Features of the Barker Woods Nature Preserve. 61 cm con-
tour interval.
121
122 Indiana Academy of Science Vol. 94 (1985)
moderate in organic content and characterized by a seasonally high water table. The
preserve includes pin oak — red maple hydric upland depressional woods, and white
oak — red oak — black oak dry mesic upland forest (Tom Post, personal communica-
tion). Riemenschneider and Reed (11) further describes the property and its history.
The purpose of this study was to investigate the population numbers and viability
of seven state threatened or endangered plant species found on the preserve, and to
describe their general physical habitat preferences. The state endangered species are
Carex arctata Boott (Drooping wood sedge) and C. folliculata L. (Long sedge). The
state threatened species include Betula papyri/era Marsh (Paper birch), Epigaea repens
L. (Trailing arbutus), Melampyrum lineare Desr. (Cow wheat), Pyrola americana Sweet.
(Round-leaved shinleaf) and P. elliptica Nutt. (Shinleaf)-
Methods
Data was collected from field observations during 1983, personal interviews, and
a literature search for each species of concern. A contour map of the general surface
of the water table was made using data from U.S.G.S. 7.5' quadrangle maps. Addi-
tional maps were made over a base map with a 61 cm contour interval. Geomorphic
features were interpreted with the assistance of Dr. Mark Reshkin of Indiana Univer-
sity Northwest. Basic soil data (7) were extrapolated to the 61 cm contour map. Two
soil borings were taken with a hand auger and interpreted by Dr. Victor Riemenschneider
of Indiana University South Bend.
Plant nomenclature was based on Kartesz and Kartesz (8) and endangered and
threatened classifications were based on Bacone and Hedge (1). Data on species habitat
and status were collected from the Natural Heritage Programs of Indiana, Illinois,
Kentucky, Michigan, Ohio and Wisconsin.
Sixty-four random one meter square plots were sampled in the northeast corner
of the preserve. This method was abandoned in July and replaced by a walking traverse
of the property. All populations of endangered and threatened plants were counted
and marked. For Epigaea repens, Pyrola americana and P. elliptica the ground surface
covered was measured. Species locations were mapped with a plane table.
Results and Discussion
Carex arctata is common in Michigan and Wisconsin, is endangered in Ohio,
and does not occur in Illinois. In Indiana it is reported only from two sites in La
Porte County.
A total of 1583 individual clumps were found (68% with seed), over most of the pre-
serve on Saugatuck-Pipestone complex, Brems fine sand, Newton loamy fine sand, Oakville
fine sand and Urban land — Morocco complex soils (Figure 2). No C. arctata were
found in the pin oak (Quercus palustris) openings, or in areas of planted pine.
This sedge was most frequent in middle of an old trail, on spoils mounds beside drainage
ditches, and atop low windthrow mounds. Some preference was shown for slightly
high areas (192.6-193.2 m elevation) and for the north slopes of these higher areas.
C. arctata occurred both in areas with and without evident fire scars.
The correlation of Carex arctata with disturbed areas (trails and ditches) may
be due to several physical and biological factors. The trails and ditches are fairly open.
Either the additional available light or the removal of competing growth may be a
critical factor in the growth of the species. In addition, the mounds and the edges
of the ditches are full of small mammal burrows, with the heaviest seed-producing
C. arctata plants growing over heavily burrowed spots. It appears that small mammals
have been a major factor in seed transportation of this sedge.
Carex folliculata is now extirpated in Illinois, is threatened in Wisconsin and Ohio,
500'
i
Legend
o Individual Carex arctata
^ 2-5 Carex arctata
O 6-10 Carex arctata
O 11-40 Carex arctata
Trail
Drainage ditch
Figure 2. Locations of Carex arctata in Barker Woods Nature Preserve, Showing
the Relationship to Trails and Ditches. 61 cm contour interval.
and is uncommon in Michigan. In Indiana it is found only in northern Porter and
LaPorte Counties. Its habitat is usually swampy woods and bog thickets.
Ninety-seven plants were found in Barker Woods, 27% with seeds. The plants
were found only at 192 to 193.2 m elevation on Saugatuck-Pipestone complex soils
at the edge of the southeast pin oak opening (Figure 3). No fire scars were evident.
Historically, the water table stood at 192.6 m elevation, as evidenced today by
the presence of pin oak openings. Mapping by the author of the current water table
showed water under Barker Woods at 191.4 to 191.7 m elevation. This drop is due
to drainage of the general area begun before the turn of the century and continued
to this day. Carex folliculata, a species of wet woods, persists up to 1.8 m above the
124
Indiana Academy of Science
Vol. 94 (1985)
100m
500'
t
Figure 3. Location of Carex folliculata in Barker Woods Nature Preserve, Showing
the Relationship to Flat, Open Forest Floor. 61 cm contour interval.
water table. A soil auger taken in the area of the C. folliculata in September of 1983,
after a dry summer, showed moist soil within 60 cm of the surface. C. folliculata
may be utilizing rain water suspended above the water table atop cemented layers of
sand called iron pans, which are characteristic of the Saugatuck-Pipestone complex soils.
Betula papyrifera is common in Michigan and Wisconsin, and is found (but is
not common) in northern Illinois. In Indiana it is native only to Lake, Porter and
LaPorte Counties. B. papyrifera is considered an early successional species, lasting
only one generation before being replaced by more shade tolerant species (5). Very
young seedlings are very sensitive and need some shade, but as they grow they need
overhead light (9).
A total of 202 individuals were found in the preserve in two groves (Figure 4).
Grove A consisted of 77 widely scattered trees on the north slope of a Glenwood stage
sandbar on Saugatuck-Pipestone complex, Newton loamy fine sand and Brems fine
sand. Grove A is being over shaded by pin oak, red maple, tulip, and sassafras.
Botany
125
100m
i
500'
Figure 4. Locations of Betula papyrifera in Barker Woods Nature Preserve, Show-
ing Relationship to Fire Scarred Areas. 61 cm contour interval.
Immediately north and east of this grove, on adjacent property, the paper birch trees
were larger and healthier. This land was disturbed more recently and the paper birch
are still the dominant trees.
Grove B is a concentration of 125 trees, including many young trees, on Saugatuck-
Pipestone complex soils. Some overshading of birch is occurring, but the area has
been kept fairly open by regular windthrows.
The most recent recorded fire on the preserve occurred in 1967 (Orphie Loomis,
personal communication). Judging from the fire scars present, Grove B burned to a
greater extent than Grove A, perhaps explaining the vigor of Grove B.
Epigaea repens was once common in Michigan, before being collected almost
to extinction. It has been totally eliminated in Illinois. It is frequent in northern Wisconsin
and eastern Ohio. In Indiana the species is reported from eight counties (Allen, Elkhart,
LaGrange, Lake, LaPorte, Monroe and Washington).
Trailing arbutus occurred only in the northeast corner of Barker Woods (Figure 5).
126
Indiana Academy of Science
Vol. 94 (1985)
North boundary of Barker Woods Nature Preserve
1
\
9ul.
/
Jp
^ffJb ___^--^
aMl-i
QMI-I
Q.MI-1
-'>-« m '(''''f"^;
W'-<J(IM|-I
/
"^ H*-3 aui-i
=-"''' m4 7
qMI-l
//
W&
Bor
Not
Ker Woods
art Preserve
1
Figure 5. Locations of Epigaea repens (Er) and Melampyrum lineare (Ml) in the
Two Northeast Sectors of Barker Woods Nature Preserve. 61 cm contour interval.
Three mats, covering 6.5 m2 of ground surface were found on the north side of the sandbar
on Brems fine sand and Newton loamy fine sand.
Reproduction data was not available for 1983, but flowers were present in 1984.
Ants, important for pollination and seed transportation (3), were active on the plants
in 1984.
Epigaea repens grew in one area of intense sunlight, and two areas of partial
shade. Moderate leaf duff covered all plants. Fire scars were present in the area. The
species tolerates five very well (3).
Melampyrum lineare is common in Michigan and Wisconsin, but is considered
threatened in Illinois and Ohio. It is reported from six counties in Indiana (Lake, La
Porte, LaGrange, Porter, Tippecanoe and White). Its habitat varies from bogs to dry,
coniferous woods.
Cow wheat was found only in the northeast section of the preserve (Figure 5).
It occurred on Newton loamy fine sand, Brems fine sand and Saugatuck-Pipestone
complex soils on the north slope of the sandbar. Fire scars were present in the area.
One hundred and thirty-five plants were found, 93% of which bore seed. This
was an increase from 44 plants in 1982. Even plants which were partially grazed or
dried continued to produce flowers and seeds. Laboratory studies (2) have found that
as many as 264 seeds are produced by one plant.
Melampyrum lineare is a non-obligatory root parasite, attaching itself to host
roots or rhizomes by extremely fine roots bearing minute haustoria. Host plants in-
clude dicots, monocots, conifers, ferns and a bryophyte {Sphagnum). Cow wheat can
also be saphrophytic, attaching to humus or dead plant tissue (10). In Barker Woods
the M. lineare was concentrated in a sunny area opened up by the death of a large
wild black cherry tree. M. lineare may have a saphrophytic relationship with the dead
Botany
127
roots of the cherry tree, or it may be parasitic on live plants such as the thick stand
of bracken fern present.
Pyrola americana is common in northern Michigan and northern and central
Wisconsin. It is frequent in eastern Ohio, but is considered endangered in Illinois.
In Indiana it is reported from six counties (LaGrange, Lake, LaPorte, Porter, Steuben
and St. Joseph).
In Barker Woods this species was found in one large, disjunct population, with
five additional, small locations throughout the preserve (Figure 6). Three of the six
locations showed evidence of fire. This shinleaf occurred on Newton loamy fine sand,
Brems fine sand and Saugatuck-Pipestone complex soils, usually atop windthrow mounds
in moderate shade.
The north fence population of P. americana covered 230.1 m2 of ground surface,
while the five smaller areas covered 31.5 m2.
100m
— i
500'
Figure 6. Locations of Pyrola americana and P. elliptica in Barker Woods Nature
Preserve. 61 cm contour interval.
128 Indiana Academy of Science Vol. 94 (1985)
In 1983 only one flower stalk was produced by P. americana across the entire
preserve, and this solitary stalk did not set seed. The lack of flowering may have been
due to unfavorable environmental conditions, such as the dry summer, or to the general
characteristic of the species to reproduce vegetatively (Dr. Eric Haber, personal
communication).
Pyrola elliptica is common in Michigan, Wisconsin and Ohio. It occurs in northern
Illinois, but is becoming more rare. Deam (4) considered this the most common species
of the genus in Indiana. It is reported from 11 counties (Cass, Elkhart, Grant, Kosciusko,
LaGrange, LaPorte, Parke, Porter, Putnam, Steuben and St. Joseph). The Barker Woods
population is the only population in LaPorte Co.
P. elliptica was found in the preserve in only one 22 m2 area of thinly scattered
individuals (Figure 6). One flower stalk with seeds was produced in 1983. The plants
were found on the north slope of the sandbar at the border of the Brems fine sand
and the Newton loamy fine sand, among fire scars.
Of the seven species of concern, all but Carex folliculata have northern affinities.
Pyrola americana, P. elliptica and Epigaea repens were considered boreal relics by
Friesner (6). In this light, north to south profiles of Barker Woods were made, show-
ing the locations of the seven species in relation to the geomorphic features (Figure 7).
In Profile A- A ' Betula papyri/era, Pyrola americana, Melampyrum lineare, and
Epigaea repens were found on the north slope of the sandbar; B. papyrifera was found
on the north slope of a slight rise, and Carex arctata and C. folliculata were found
at the base of the north slope of the Glenwood dune. Profile B-B ' shows C. arctata,
P. americana, B. papyrifera and P. elliptica occurring on the north slope of the sand-
bar, and C. arctata occurring at the base of the north slope of the Glenwood dune.
In Profile C-C ', C. arctata was found on the north slope of two slight rises in the
central low area. The conclusion is that north slopes are preferred habitat for six of
the seven species of concern.
Summary
Populations of Carex arctata, Melampyrum lineare and Pyrola americana are pre-
sent in large numbers in Barker Woods Nature Preserve and are reproducing sexually
or vegetatively. The population of C. folliculata is small but is reproducing. A species
of wet habitats, it does not seem stressed by the lowered water table and may be utiliz-
ing water suspended above the water table by cemented layers of sand. The Betula
papyrifera population is large but is declining due to succession. Epigaea repens and
P. elliptica are present in small numbers and are reproducing. All species of concern,
except Carex folliculata, appear to prefer north facing slopes, reflecting their northern
affinities.
I thank Dr. Victor Riemenschneider for his assistance, the Barker Woods Preserve
Management Committee for funding the study, and the generous contribution of Miss
Margery Barker which endowed the preserve and made such a study possible.
Literature Cited
1. Bacone, J. A. and C.L. Hedge. 1980. A preliminary list of endangered and
threatened vascular plants in Indiana. Proc. Ind. Acad. Sci. 89:359-371.
2. Cantlon, J.E., E.J.C. Curtis and W.M. Malcom. 1963. Studies of Melampyrum
lineare. Ecol. 44:466-474.
3. Clay, K. and N.C. Ellstrand. 1981. Stylar polymorphism in Epigaea repens, a
dioecious species. Bull. Torrey Bot. Club. 108:305-310.
4. Deam, C.C. 1940. Flora of Indiana. Dept. Conserv., Div. For., Indianapolis,
1,236 p.
Botany
129
SOUTH
-640
630
-640
630
Index map of Preterve
Showing location of profiles
Figure 7. North to South Profiles of Barker Woods Nature Preserve, Showing Pre-
ferred Growth Locations of Threatened and Endangered Species. Vertical exaggera-
tion 15X.
Fowells, H.A., compiler. 1965. Silvics of the Forest Trees of the United States.
U.S. Department of Agriculture Forest Service Agri. Handbook No. 271,
Washington.
Friesner, R.C. 1936. Indiana as a critical botanical area. Proc. Ind. Acad. Sci.
46:28-45.
130 Indiana Academy of Science Vol. 94 (1985)
7. Furr, G.F., Jr. 1982. Soil Survey of La Porte County, Indiana. U.S. Department
of Agriculture. U.S. Government Printing Office, Washington. 162 p. + maps.
8. Kartesz, J.T. and R. Kartesz. 1980. A Synonymized Checklist of the Vascular
Flora of the United States, Canada, and Greenland. VII. The Biota of North
America. University of North Carolina Press, Chapel Hill.
9. Marquis, D.A., J.C. Bjorkbom and G. Yelenosky. 1964. Effect of seedbed con-
dition and light exposure of paper birch regeneration. Jour. For. (1964):876-881.
10. Piehl, M.A. 1962. The parasitic behavior of Melampyrum lineare and a note on
its seed color. Rhodora (1962): 15-23.
11. Riemenschneider, V. and P.W. Reed. 1985. Vascular plants of Barker Woods
Nature Preserve, La Porte County, Indiana. Proc. In. Acad. Sci. 94:(in press).
Bacterial Wilt Resistance in Commercial Muskmelon Cultivars
G.L. Reed
USDA-ARS, Vincennes University
Vincennes, Indiana 47591
and
Department of Entomology
Purdue University, West Lafayette, Indiana 47907
and
W.R. Stevenson
Department of Plant Pathology
University of Wisconsin, Madison, Wisconsin 53706
Introduction
The number of available muskmelon, Cucumis melo L., cultivars with resistance
to bacterial wilt, Erwinia tracheiphilla (Smith) Dye, has declined along with the impor-
tance of Midwestern production. However, current transportation costs have created
a resurgence in demand for production in the area. This demand justifies the selection
of resistant cultivars (both varieties and hybrids) since bacterial wilt continues to be
a major disease of muskmelon, cucumis melo L., in the Midwest (1,3 and 6). Though
transmission of the bacterium was demonstrated to occur by insect vectors prior to
the turn of the century (2), (primarily by the striped cucumber beetle, Acalymma vit-
tatum (F.)); no adequate means of protection from the disease existed before the advent
of modern insecticides (1). Control of the vectors by insecticide application has reduced
incidence of the disease, but significant losses still occur (3). No extensive effort has
been made to develop cultivars with high levels of resistance. The development of
bacterial wilt resistant cultivars and hybrids would reduce those losses and provide an
alternative to the current indirect method of disease control.
This research was initiated to evaluate commercially adapted muskmelon cultivars
for resistance to bacterial wilt. The search was initiated in commercial germplasm to
permit easier selection of disease resistant cultivars with horticulturally acceptable traits.
Seed of 187 cultivars were acquired from the vegetable seed industry, the National
Seed Storage Laboratory, Fort Collins, CO, and public melon breeders. This germ-
plasm was screened for resistance in both the field and greenhouse. No cultivar with
adequate resistance for unprotected commercial production was found, but several
cultivars contained resistant plants in frequences adequate to allow the selection of resis-
tant cultivars.
Materials and Methods
Cultivars were evaluated for bacterial wilt resistance in three separate trials. The
number of cultivars and plants tested per cultivar varied between trials and within
each trial due to availability and germination of seed. In 1976, a field evaluation was
conducted where transmission of the pathogen was dependent upon feeding by field
populations of striped cucumber beetles. In 1977 and 1979, seedlings were inoculated
with the bacterium in greenhouse trials.
Field. The 1976 trial was planted in a commercial field located 3.2 km SE of
Vincennes, IN, in the center of a major melon production area about 4.8 km wide
and 16 km long that contained about 810 hectares of muskmelon and 2025 hectares
of watermelon. Field preparation and routine vegetable production practices were pro-
vided by the grower and were identical to those of his commercial fields except that
no pesticide program was applied. Transplants of 67 cultivars were grown in cold frames
131
132 Indiana Academy of Science Vol. 94 (1985)
using veneer "dirt bands" (8.9 x 8.9 x 10.2 cm) filled with spent mushroom compost
as growing medium (4). Seedlings were transplanted at the 3-4 true leaf stage during
mid-May with five plants/row (1.5 m apart) in rows 1.8 m wide. Plots were separated
by 4.6 m wide alleyways. The experiment was replicated three times with five plants
per cultivar per replication. To estimate beetle populations, counts of striped cucumber
beetle adults were made May 24 and June 7. Symptoms and mortality associated with
bacterial wilt were recorded weekly.
Greenhouse. Greenhouse trials were conducted in 1977 and 1979 to provide a
uniform evaluation of cultivars for bacterial wilt resistance. Greenhouses were operated
at 30 C with 24 h light from 40 watt Luxor Vita-Lite Lamps suspended 27 cm above
the plants with one bulb/0.28 m2 of bench space. Seedlings were grown in Jiffy 64®
trays with Jiffy Plus® potting medium. They were watered daily and received no
additional fertilization. Greenhouse trials were designed as randomized complete block
experiments with four replications and 16-24 seeds planted per cultivar per replication.
Seedlings were inoculated with bacterial wilt at the fully expanded cotyledon stage,
five days after planting. Inoculum for the 1977 trial was prepared from an E. tracheiphila
culture isolated at Vincennes, IN, during 1976. Inoculum culture for the 1979 trial
was from a culture provided by H. M. Munger, Cornell University. Inoculum was
prepared according to Reed and Stevenson (3) from inoculated infected muskmelon
seedlings cv. Perlita which exhibited wilting of both cotyledons. Virulence of inoculum
was established on the response of susceptible check cultivars (Charentais Imp. and
Perlita). In 1977, seedlings were inoculated using a #1 cork through which eight randomly
placed pins protruded a distance of 1 mm (3). In 1977, plants surviving initial inocula-
tion were reinoculated (on 1st and 2nd true leaves) to reduce the chance of escapes.
During the 1979 trial, a single cotyledon inoculation was made using the 15-pin in-
oculation dispenser with reservoir described by Reed and Stevenson (3).
Totals of 100 and 185 cultivars were evaluated in 1977 and 1979 respectively.
To determine whether frequencies of resistant plants might vary between sources of
seed of the same cultivar, lots from several companies were tested in 1979, increasing
the number of treatments evaluated from 185 to 323. Due to the large number of
treatments, only two replications were used in 1979. Seedling mortality was recorded
twice weekly during greenhouse evaluations. Percent survival was computed by dividing
the number of surviving seedlings by the number of inoculated seedlings. Percent sur-
vival data presented in Tables 1-4 are means of the percent survival computed for
each replication. During the 1979 trial, surviving plants were visually rated for symp-
toms of the disease. A 1-5 rating scale was used where: 1 = plants without symptoms,
2 = plants with chlorosis or wilting on cotyledons or lower leaves, 3 = plants with chlorosis
or wilting in upper or terminal growth, 4 = plants with chlorosis or wilting throughout
and dwarfed in size and 5 = plants dead.
Results
Resistance evaluations during 1976, 1977 and 1979 are reported in Tables 1 through
5. All of the cultivars were tested in combined trials, but have been grouped into hybrids
(Table 1), commercially available varieties (Table 2), obsolete cultivars (Table 3), breeding
lines (Table 4), and a list of promising resistance sources (Table 5). Mean percent sur-
vival of 46.3, 9.6 and 1.1 were observed, respectively for the 3 trials. For the 1976
evaluation, striped cucumber beetle adult counts averaged 2.75 beetles per plant and
ranged from 0 per plant on the least attractive cultivar to 12.2 on the most attractive.
Percent survival of the susceptible check cultivar Perlita were 60, 2, and 0 and of the
susceptible check cultivar Charentais Improved were 8, 6, and 0 for the three trials.
Cultivars; Wescan, Burrell's Gem, Hales Best, Harvest Queen, Hearts of Gold,
Botany 133
Table 1 . Summary of response of muskmelon hybrids screened for resistance to bacterial
wilt, Erwinia tracheiphila (Smith) Dye.
1976 1977 1979
Cultivar
Alaska Hy
Ambrosia Hy
Ball 1776 Hy
Burpee Hy
Bushwhopper Hy
Canada Gem Hy
Carnival Hy
Chaca tt\ Hy
Chando Hy
Classic Hy
Crenshaw Hy
Croustillan Hy
Dixie Jumbo Hy
Earlisweet Hy
Early Dawn Hy
Giant Hy
Golden Crispy Hy
Gold Star Hy
Harmony Hy
Harper Hy
Honey Drop Hy
Known-You Hy
Luscious Hy
Mainerock Hy
Market Pride F2 Hy
Midwest Extra Early HY
Minnesota Hy 16
Minnesota Hy 26
Oval Chaca Hy
Roadside Hy
Samson Hy
Saticoy Hy
Scoop Hy
Star Headliner Hy
Star Trek Hy
Summet Hy
Sundae Fl Hy
Super Hy
Super Market Hy
Supreme Delight Hy
Sweetie Hy
#
%
tt
%
tt
°?o
Tested
Survival
Tested
Survival
Tested
Survival
—
—
—
—
49
0
—
—
—
—
48
0
—
—
—
—
41
0
15
20
31
10
71
3
—
—
—
—
68
0
—
—
—
47
0
—
—
—
—
43
0
—
—
—
—
46
0
—
—
—
—
73
0
15
67
23
10
48
0
—
—
—
—
62
0
—
—
—
—
61
0
15
40
—
—
49
0
—
—
—
—
61
0
—
—
—
—
48
0
—
—
—
58
0
—
—
—
—
58
0
—
—
—
—
50
0
—
—
—
—
49
0
15
73
36
10
48
0
—
—
—
—
51
0
—
—
—
—
46
0
—
—
—
—
45
0
—
—
—
—
43
0
15
73
54
10
46
0
—
—
—
—
47
0
—
—
—
—
47
0
—
—
—
—
14
0
—
—
—
—
48
0
—
—
—
—
46
0
—
—
—
43
0
15
47
31
19
45
0
—
—
—
—
49
0
—
—
—
—
46
0
—
—
—
—
47
2
—
—
—
—
46
0
—
—
—
—
48
0
—
—
—
—
29
11
15
47
60
10
44
0
—
—
—
—
44
0
Pride of Wisconsin, and Schoon's Hard Shell had the highest percent survival in 1976.
Cultivars; Persianet, Md 63-53, Rocky Ford Poleock, Emerald Gem, PMR-8, Rocks,
and Burreh"s Gem had the highest percent survival in 1977. Cultivars; Burrell's Gem,
134
Indiana Academy of Science
Vol. 94 (1985)
Table 2. Summary of response of commercial muskmelon cultivars screened for resistance
to bacterial wilt, Erwinia tracheiphila (Smith) Dye.
1976
1977
1979
#
%
ff
%
n
%
Cultivar
Tested
Survival
Tested
Survival
Tested
Survival
Amarelo
—
—
—
—
39
0
Banana
15
47
32
20
97
4
Bender's Surprise
—
—
—
—
107
2
Bush Midget
—
—
—
—
67
0
Casaba, Golden Beauty
13
8
59
0
74
0
Casaba, Sun Gold
—
—
33
0
Cavillon Red-Fleshed
—
—
—
—
38
0
Charentais Improved
12
8
24
6
47
0
Chilton
—
—
—
—
77
0
Crenshaw
14
7
41
0
118
0
Crenshaw Golden
—
—
—
44
0
Cum Laude
12
7
35
12
46
0
Delicious 51
15
60
55
5
508
1
Dr. Jaegar's Mildew Res.
—
—
—
—
35
3
Dwarf
—
—
—
—
37
3
Early Delicious 51
—
29
0
77
0
Early May
10
50
27
5
53
0
Early Sugar Midget
—
—
10
0
26
0
Eden Gem
15
47
5
0
43
0
Edisto
15
40
9
5
132
1
Edisto 47
15
60
43
12
227
2
Far North
—
—
—
—
81
0
Fordhook Gem
15
47
48
10
31
0
Four-Fifty (450)
15
60
34
0
37
0
Giant
15
20
47
7
42
0
Gold Cup
15
40
51
10
26
0
Gold Cup 55
—
—
—
—
45
0
Golden Champlain
—
—
—
—
86
1
Golden Honey
15
53
49
5
46
0
Golden Perfection
15
40
37
8
80
1
Gold Lined Rockyford
—
—
—
—
44
0
Granite State
—
—
—
—
48
0
Green Nutmeg
—
—
—
—
47
0
Gulfcoast
—
—
—
—
72
0
Gulfstream
15
27
64
15
76
3
Gusto 45
48
3
46
0
Hales Best
15
80
33
5
222
.5
Hales Best 36 Improved
15
40
42
8
290
1
Hales Best 936
15
60
29
22
27
0
Hales Best Jumbo
15
53
50
14
378
1
Hales Best Jumbo Improved
54
0
Haogen
—
—
—
—
37
0
Harvest Queen
15
80
36
8
370
1
Hearts of Gold
15
80
57
19
472
2
Honey Rock
14
43
41
13
464
2
Honey Rock Improved
—
—
—
—
40
2
Illinois Hardshell
—
—
—
—
42
2
Botany
135
Table 2. — Continued
1976
1977
1979
ft
%
#
%
#
%
Cultivar
Tested
Survival
Tested
Survival
Tested
Survival
Imperial 5
—
—
—
—
37
0
Imperial 45
—
—
—
—
49
0
Imperial 4-50
15
20
57
9
42
0
Imperial 45-S12
15
53
58
2
31
8
Iroquois
15
73
56
8
551
4
Kangold
15
33
50
19
122
3
King Henry
15
20
46
3
35
3
Knight's Early
—
—
—
—
40
0
Mammoth
—
—
58
19
48
2
Midget
—
—
—
—
47
0
Mildew Resistant 45
—
—
—
—
93
1
Minnesota Honey
—
—
—
—
43
0
Minnesota Honey Mist
—
—
—
—
38
0
Minnesota Midget
—
—
36
15
114
2
New Ideal
—
—
29
22
79
1
No 45-SJ
—
—
39
2
27
0
Ogen
11
0
44
2
23
2
Old Time Tennessee
15
0
57
13
39
0
Osage
—
—
—
—
42
2
Pennsweet
—
—
—
—
64
3
Perfection
15
60
59
7
94
0
Perlita
15
27
67
2
79
0
Persian Small
—
—
43
8
31
0
Planter's Jumbo
15
53
48
2
226
.4
PMR-45
14
29
64
9
122
1
PMR-450
13
31
27
2
45
0
Pride of Wisconsin
15
80
38
4
275
1
Queen Of Colorado
15
67
37
0
110
1
Resistance #45
—
41
7
Rio Gold
15
33
49
3
31
0
Rocky Ford
15
53
43
16
261
.4
Rocky Ford, Earliest
—
—
—
—
.45
0
Rocky Ford, Poleock
—
—
31
29
"65
5
Roi du Nord
42
0
Schoon's Hardshell
15
80
42
7
280
.3
Short 'N' Sweet
—
—
—
—
55
0
Shumway's Giant
—
—
—
—
40
0
Sierra Gold
15
7
17
5
33
0
Smith's Perfect
—
—
34
13
177
0
Spartan Rock
15
53
35
6
174
0
Sugar Salmon
15
53
49
20
47
0
Sulphur Resistant 59
—
—
—
—
56
0
Sulphur Resistant 91
—
-
-
-
39
0
Sweet Granite
32
0
Tarn Uvalde
—
—
—
—
49
0
Texas No. 1
—
—
30
5
86
0
Tip Top
—
—
—
—
32
0
Top Mark
—
-
50
11
48
0
Turkey
15
53
55
5
47
0
Yellow Canary
—
—
—
—
14
7
136
Indiana Academy of Science
Vol. 94 (1985)
Table 3 . Summary of response of obsolete muskmelon cultivars screened for resistance
to bacterial wilt, Erwinia tracheiphila (Smith) Dye.
1976
1977
1979
»
°/o
n
%
ft
%
Cultivar
Tested
Survival
Tested
Survival
Tested
Survival
Arizona 13
—
—
—
—
26
5
Burpee's Fordhook
—
—
55
13
47
0
Burrell's Gem
15
93
43
25
25
25
Burrell's Superfecto
—
—
10
10
12
0
Bush, M.M.
—
—
9
8
7
0
Campo
5
20
—
—
45
0
Daisy
15
53
36
9
42
7
Dulce
10
30
36
2
53
0
Early Mayfair
—
—
59
14
41
0
Early Sunrise
—
—
—
—
37
5
Early Wonder
—
—
—
—
8
16
Emerald Gem
—
—
33
29
29
7
Extra Early Hackensack
—
—
20
5
41
0
Extra Early Sunrise
—
—
55
0
36
0
Healy's Pride
15
73
30
3
15
0
Honey Ball
—
—
12
6
46
0
Jenny Lind
—
—
28
5
48
0
Jewel
10
50
22
6
32
12
Kilgore's Hummer
—
—
26
17
46
0
Milwaukee Market
—
—
23
8
58
0
Perfecto
35
23
48
0
Perfecto, Perfected
—
—
61
27
—
—
Persianet
8
25
26
36
44
0
Pink Queen
—
—
20
8
44
0
Queen Anne's Pocket
—
—
13
0
34
3
Rock "O" Honey
—
29
14
45
2
Seneca Delicious
—
—
19
12
46
0
Sheridan
—
—
27
0
46
0
Ward's Ideal
—
—
13
5
49
2
Woodside Winner
—
—
16
5
62
0
Yate's Surprise
-
-
27
4
43
0
Early Wonder, Jewel, Super Hybrid, Imperial 45-S12, Resistant No. 45, and Yellow
Canary, had the highest percent survival in 1979. Cultivars; Burrell's Gem, Early
Wonder, Jewel, Super Hybrid, Yellow Canary, Imperial 45-S12, Resistant No. 45, Daisy,
and Early Sunrise had the lowest mean disease severity ratings in 1979. Twenty-six
cultivars had disease severity ratings of 1.00 for surviving plants in 1979 (Table 5).
Analysis of variance for all three trials demonstrated significant differences be-
tween cultivars. F values of 3.676 for 66/132 df in 1976 and 2.507 for 99/297 df in
1977 were both significant at .01 probability. The 1979 trial was transformed with
Arcsin because of the large number of cultivars with 0 percent survival. The F value
of 1.334 for 184/184 df in 1979 was significant at .05 probability. A Bayes LSD (BLSD)
was used to separate the large number of means (5). For the 1976 trial BLSD's of
31, 36, and 48 indicated significant differences between cultivars at K values of 50,
100, and 500. For 1977, BLSD's of 17.2, 20.1, and 27.3 indicated significant differences.
Botany
137
Table 4. Summary of response of muskmelon breeding lines screened for resistance
to bacterial wilt, Erwinia tracheiphila (Smith) Dye.
1976
1977
1979
#
%
#
%
n
%
Cultivar
Tested
Survival
Tested
Survival
Tested
Survival
AC 67-59
—
—
—
—
34
3
Cobmelon
4
25
32
5
41
0
Doublon
—
—
—
—
40
0
Earl's Favorite
—
—
—
—
14
0
GA-47
5
60
41
14
81
0
Jacumba
5
20
33
0
32
0
MD 63-53
15
47
54
34
51
0
Ogon 9
—
—
—
—
65
0
PMR-5
3
0
33
2
46
0
PMR-6
15
40
19
0
—
—
PMR-8
10
30
18
28
46
2
PMR-29
—
—
—
—
64
0
Purdue 44
—
—
25
4
40
0
Rocks
10
60
28
27
49
0
Santa Claus
—
—
—
—
37
2
Seminole
23
0
Wescan
5
100
23
5
57
0
Yellow Green
15
40
28
7
45
0
For 1979, BLSD's of 14.1 and 18.8 indicated significant differences at K values of
50 and 100.
Discussion
No muskmelon cultivar tested has ad adequate frequency of bacterial wilt resis-
tant plants to be used in commercial plantings without insecticide protection; however,
a substantial number of lines had sufficient frequencies of resistant plants to be used
as germplasm for developing resistant varieties and hybrids. The 1979 trial provides
the best comparison to select cultivars for four reasons.
First, it was the only trial in which plants of the susceptible check cultivars Perlita
and Charentais Improved were completely killed. The 1976 trial was not simply an
evaluation of bacterial wilt resistance; but, also an evaluation of striped cucumber
beetle feeding preference. It is of interest, however, that those lines with 80 to 100
percent survival in the 1976 trial, with the exception of Wescan, had surviving plants
in the 1979 trial. In 1977 some plants of the check cultivars escaped infection with
the disease, even when inoculated 3 times. Of the 7 most resistant cultivars of the
1977 trial, Persianet, Md 63-53, and Rocks proved to be totally susceptible in the 1979 trial.
Second, of the 54 cultivars with frequencies of resistant plants in 1979, plants
of only 3 of these lines were all susceptible, either in 1976 or 1977. Because all 3 of
these cultivars had relatively low frequencies of resistant plants in 1979; the probability
of a resistant plant occurring in the small number of plants tested in the earlier test
could explain these inconsistencies. For instance, Queen Anne's Pocket had 3 percent
survival in 1979, but none in 1977, when only 13 plants germinated for evaluation.
Third, the 1979 evaluation was much more inclusive of cultivars than the earlier
trials.
138
Indiana Academy of Science
Vol. 94 (1985)
Table 5 . Response of cultivars identified as sources of resistance to bacterial wilt Erwinia
tracheiphila (Smith) Dye.
1979
1976
1977
Percent
Survival
Disease Rat
ing
Percent
Survival
Percent
Survival
Cultivar
Overall
Survivors
Burrell's Gem
93
25
25
3.75
1.00
Early Wonder
—
—
16
4.35
1.00
Jewel
50
6
12
4.45
1.00
Super Hy
—
—
11
4.50
1.30
Imperial 45-S12
53
2
8
4.70
1.30
Resistant No. 45
—
—
7
4.70
1.70
Yellow Canary
—
—
7
4.65
1.00
Daisy
53
9
7
4.70
1.70
Emerald Gem
—
29
7
4.88
1.00
Early Sunrise
—
—
5
4.70
1.00
Arizona 13
—
5
4.75
1.00
Rocky Ford Poleock
—
29
5
4.80
1.00
Iroquois
73
8
4
4.81
1.30
Banana
47
20
4
4.85
1.00
Kangold
33
19
3
4.88
2.00
Pennsweet
—
3
4.93
2.00
AC 67-59
—
—
3
4.85
1.00
Queen Anne's Pocket
—
0
3
4.85
1.00
King Henry
20
3
3
4.90
2.00
Burpee Hy
20
10
3
4.85
1.00
Dr. Jaeper's Mildew Resistant
—
—
3
4.85
1.00
Dwarf
—
—
3
4.95
3.00
Gulfstream
27
15
3
4.93
2.50
Santa Claus
—
—
2
4.90
1.00
Ogen
0
2
2
4.93
2.00
Honey Rock Improved
—
—
2
4.90
1.00
Illinois Hardshell
—
—
2
4.90
1.00
Osage
—
—
2
4.90
1.00
PMR-8
30
28
2
4.95
2.00
Rock "O" Honey
—
14
2
4.95
3.00
Star Trek Hy
—
2
4.85
3.00
Honey Rock
43
13
2
4.92
1.60
Mammoth
—
19
2
4.90
1.00
Ward's Ideal
—
5
2
4.95
3.00
Edisto 47
60
12
2
4.93
2.50
Hearts of Gold
80
19
2
4.92
1.20
Bender's Surprise
—
—
2
4.95
1.50
Minnesota Midget
—
15
2
4.97
3.50
New Ideal
—
22
1
4.95
1.00
Harvest Queen
80
8
1
4.91
1.00
Golden Perfection
40
8
1
4.93
1.00
Golden Champlain
—
—
1
4.85
1.00
Hales Best 36 Improved
33
8
1
4.95
2.70
Mildew Resistant 45
—
—
1
4.98
2.00
Pride of Wisconsin
80
4
1
4.95
1.50
Botany 139
Table 5. — Continued
1979
1976
1977
ing
Percent
Percent
Percent
Cultivar
Survival
Survival
Survival
Overall
Survivors
PMR-45
29
9
1
4.99
4.00
Queen of Colorado
67
0
1
4.99
4.00
Hales Best Jumbo
53
14
1
4.97
1.30
Edisto
40
5
1
4.95
1.00
Delicious 51
60
5
1
4.99
1.80
Hales Best
80
5
.5
4.98
1.00
Planter's Jumbo
53
2
.4
4.98
1.00
Rocky Ford
53
16
.4
4.99
1.00
Schoon's Hardshell
80
7
.3
4.99
4.00
1 = plants without symptoms, 2 = plants with chlorosis or wilting on cotyledons or lower leaves, 3 = plants with chlorosis
or wilting in upper or terminal growth but remaining vigorus, 4 = plants with wilting or chlorosis throughout and
dwarfed in size, 5= plants that died.
Fourth, the 1979 trial included an evaluation of disease severity which allows
comparison based on effect of the disease on surviving plants as well as frequency
of surviving plants.
On this basis, muskmelon cultivars; Burrell's Gem, Early Wonder, Jewel, Super
Hybrid, Imperial 45-S12, Resistant No. 45, Yellow Canary, Daisy, and Emerald Gem
are the most useful cultivars for developing bacterial wilt resistance; however, testing
of further lots of Super Hybrid indicated much lower levels of resistant plants. Of
those cultivars, Burrell's Gem, Early Wonder, Jewel, Yellow Canary, and Emerald
Gem had surviving plants without visible symptoms of the disease (Table 5). Of this
group, Burrell's Gem is probably the best source of resistance. It performed well in
all trials, had the lowest overall disease rating in 1979, and had no symptom develop-
ment in surviving plants.
The three cultivars with highest percent survival and six of the 10 with highest
survival in the 1979 trial were obsolete cultivars, pointing out the need for preserving
these irreplaceable resources. Finding the largest number of cultivars with frequencies
of resistant plants to be from obsolete germplasm, indicates that recent trends in com-
mercial muskmelon breeding have decreased numbers of cultivars with resistance to
bacterial wilt. Comparisons of seed lots from several companies showed that though
normal variance was observed, indications of significantly different frequencies of resis-
tant plant between lots from different companies did not occur.
In summary, 54 cultivars were identified which had frequencies of bacterial wilt
resistant plants; but no cultivar tested was adequately resistant for commercial usage
without insecticide protection. Among surviving plants, response to the disease varied;
possibly indicating that different mechanisms for resistance might be present. Con-
sidering the relatively low frequencies of resistant plants and the potential for more
than one resistance mechanism, recurrent selection procedures should be effective in
development of resistant germplasm. This research provides plant breeders with a list
of cultivars which contain frequencies of resistant plants which should assist in the
development of bacterial wilt resistant cultivars.
Acknowledgments
The authors would like to thank Sonja Myers, Bryan Robling, Grace Barrick,
140 Indiana Academy of Science Vol. 94 (1985)
and Jerry Powell for their assistance in screening these cultivars; Dr. Louis Bass of
the National Seed Storage Laboratory, USDA, ARS, for providing most of the obsolete
cultivars and the many commercial seed companies and public muskmelon researchers
which provided seed utilized in the research.
Summary
A decline in Midwestern muskmelon production during the past 30 years has
resulted in fewer cultivars with resistance to bacterial wilt, Erwinia tracheiphila. Levels
of resistance in 187 cultivars were assessed in field and greenhouse experiments. Resistance
was most common in obsolete muskmelon cultivars and least common in current com-
mercial hybrids. Commercially available cultivars which had resistant plants tended
to be older cultivars. Of the 12 cultivars with highest frequency of resistant plants,
half are obsolete. No cultivar tested was sufficiently resistant for commercial produc-
tion without insecticide protection, but resistant plants were observed in many cultivars.
Burrell's Gem, Early Wonder, and Jewel had the highest frequency of resistant plants.
Literature Cited
1. Gould, G.E. 1936. Studies on cucumber beetle control in 1935. J. Econ. Entomol.
29(4):731.
2. Rand, F.V. and Enlows, E.M. 1916. Transmission and control of bacterial wilt
of cucurbits. J. Agr. Res. 6:417-434.
3. Reed, G.L. and Stevenson, W.R. 1982. Methods for inoculating muskmelon with
Erwinia tracheiphila. Plant Dis. 66:778-780.
4. Romanowski, R.R. and Sims, C.E. 1976. Cantaloupe mulching studies in
Southwestern Indiana. Purdue Univ. Hort. Dept., Veg., Crops Mimeo. 76-83.
5. Smith, C.W. 1978. Bayes least significant differences: A review and comparison.
Agron. J. 70:123-127.
6. Watterson, J.C., Williams, P.H. and Durbin, R.D. 1971. Response of cucurbits
to Erwinia tracheiphila bacterial wilt. Plant Dis. Rep. 55:816-819.
Improving Efficiency of Iron Uptake by Soybeans
Rosemary Rodibaugh and Connie Weaver
Department of Foods and Nutrition
Purdue University, West Lafayette, Indiana 47907
Introduction
In order to conveniently study the distribution, chemical form, and bioavailability
of iron in plant foods, plants are intrinsically labeled with isotopes of the mineral.
Usually this involves growing the plants hydroponically and introducing the label via
the nutrient solution.
Iron is the most difficult nutrient to keep in solution in hydroponic systems. The
solubility of iron is highly pH dependent. It is more soluble in acid solutions and
precipitates with phosphates in alkaline solutions (7). Chelating agents combine with
micronutrients such as iron to form soluble complexes or chelates. By increasing the
solubility of iron, these chelates play an important role in transporting iron to the
plant roots (5). Certain chelating agents are more effective than others. EDDHA
(ethylenediamine di (o-hydroxyphenylacetate)) has been shown to be effective over a
wide pH range (pH 4-9) (3) and promote iron uptake-translocation in iron-stressed
soybeans (1). DTPA (diethylenetriaminepentaacetate) is effective over the pH range
4-7.8 (3).
Soybeans typically absorb < 10% of available iron. Accumulation of iron by soy-
beans is less efficient than other trace minerals such as zinc. In an effort to discover
optimal conditions for intrinsically labelling soybeans with 59Fe, this study investigated
the efficiency of incorporation of a single does of 59FeCl3 into hydroponically grown
soybeans under three different conditions: 1) nutrient solution containing DTPA, 2)
nutrient solution containing EDDHA, and 3) root iron stripped with ferrozine prior
to dosing.
Materials and Methods
Soybeans seeds [Glycine Max. (L) Merr. 'Century'] were germinated in vermiculite.
After two weeks seedlings were transferred to 2 liter plastic pots containing a modified
Hoagland-Arnon nutrient solution (4), measured pH 6.4. Iron was added as FeDTPA,
sodium ferric diethylenetriaminepentaacetate (Sequestrene, Ciba Geigy Corp.,
Greensboro, N.C.). The nutrient solution was aerated continuously and replenished
daily. There were six plants per pot and twelve plants per treatment. Plants were grown
outdoors to the flowering stage at which time a single dose of 0.15 /tCi of 59FeCl3 was
added to each pot. After two weeks of exposure to the 59FeCl3, the roots were
removed and discarded. The six plants from each pot were weighed together and assayed
for 59Fe in a whole body gamma counter.
Three treatments were initiated at flowering. Group I plants remained in the
modified Hoagland-Arnon nutrient solution (Table I) throughout the entire study. This
has been the usual procedure for our lab (7). The chelating agent was DTPA. Four
days prior to receiving the 59FeCl3 dose, group II plants were transferred to Chaney's
nutrient solution (Table 1) which contained an excess of EDDHA and had a pH of
7.2. The plants remained in Chaney's nutrient solution for the duration of the study.
Group III plant roots were placed in a solution of 250mM sodium dithionite, a reduc-
ing agent, and 1.5mM ferrozine, a strong Fe2+ chelator, one day prior to dosing to
strip iron from the roots. Nitrogen gas was bubbled through the solution to keep it
oxygen-free, thus preventing reoxidation of Fe2 + . After this treatment, the plants were
transferred to Chaney's nutrient solution for the remainder of the study.
141
142
Indiana Academy of Science
Vol. 94 (1985)
Table 1. Concentration of Individual Elements in Nutrient Solutions
Element
Modified Hoagland-
Chaney's
Arnon Nutrient Solutions'
Nutrient Solution
mM
mM
15
15
8.6
1.12
4
5
1
0.02
2
2
2
5
liM
MM
9
2
46
10
0.77
1
0.32
0.4
—
0.2
0.5
0.2
45
4
45
40
6.4
7.2
N
K
Ca
P
S
Mg
Mn
B
Zn
Cu
Co
Mo
Fe
DPTA
EDDHA
PH
'Ref. 4
2Ref. 2
Results and Discussion
Accumulation of 59Fe by soybeans in the three treatment groups is shown in Table
II. Plants in group I, exposed to DTPA, incorporated 4. 6% of the 59Fe dose into
plant shoots. Plants in group II, exposed to a molar excess of EDDHA, incorporated
10.3% of the dose. This is more than twice as much as for group I. Stripping iron from
the roots of the plants in Group III increased the uptake of 59Fe to 13.0% of the
dose when EDDHA was the chelating agent. The stripping process caused the plants
to wilt and growth was stunted when compared to plants in groups I and II.
Until recently, the best model for iron uptake by soybean plants was proposed
by Chaney et al (1) who found that iron must be in the reduced form (Fe2 + ) to be
Table 2. Efficiency of S9Fe Accumulation by Soybean Plants
Treatment
cpmi9Fe
Total weight of
plants (g)
%5'Fe dose
in plant shoot
Group I
12,1 12 ± 575
DTPA
Group II
27,175 + 4465
EDDHA
Group 111
34,402 ±11,744
EDDHA,
stripped
roots
681
735
459
4.6±0.2C
10.3±1.7l
13.0±4.4^
Different Superscripts denote significant difference at P^0.05
Botany
143
absorbed. It has been known for several years that soybean roots are capable of reduc-
ing Fe3 + -chelates in their immediate vicinity. This reducing ability is greatly enhanced
in iron deficiency.
According to Chaney et al (1), the Fe3 + -chelate is reduced to the Fe2 + -chelate
at the root. The Fe2 + -chelate can be oxidized or can dissociate to free Fe2 + and
free chelating agent. The free Fe2 + can be absorbed by the root, complex with the
chelating agent, or complex with any competing chelating substance in the nutrient
solution.
Evidence is accumulating that the reduction of Fe3 + -chelate is an enzymatic pro-
cess that takes place at the plasmalemma of the epidermal cells on the root surface
(6). This enzyme would be embedded in the plasmalemma and capable of transporting
electrons across the membrane. Recently Sijmons and Bienfait (6) determined that
cytosolic NADPH is the electron donor for extracellular Fe3 + reduction in iron defi-
cient bean roots. They found that the supply of reduced pyridine nucleotides in lateral
roots of iron deficient beans was greatly enhanced, and that the level of cytosolic
NADPH was strongly lowered when iron deficient roots were exposed to extracellular
iron salts. This indicates that electrons are transported from the cytosolic NADPH to
the Fe3 + outside the cell via a transmembrance electron carrier (Figure 1). Sijmons and
Bienfait concluded that one of the functions of trans-plasma transport systems in
plant roots is the reduction of extracellular Fe3 + -chelates which is a necessary step
in the uptake of iron by the roots.
There was an excess of EDDHA in the nutrient solution used in groups II and
III. This excess chelating agent bound both the radioactive and non-radioactive forms
of iron so that there was an equilibrium between the S9Fe3 + -chelate and Fe3 + -chelate
complexes. Therefore, all iron-chelate in the nutrient solution was equally available
R
Pentose -P- pathway
Glycolysis"
Others
RH
NAD(P)H
NAD(P) +
ROOT EPIDERMIS CELL
OUT
Fe3-chel
Fe2!.chel
V
Fe2+
Figure 1 . Diagram for Fe3 + reduction mechanism by root epidermis cells adapted from
Sijmons and Bienfait (6).
144 Indiana Academy of Science Vol. 94 (1985)
for absorption. EDDHA is a more efficient chelating agent than DTP A because it
binds Fe3 + very tightly which keeps it from precipitating out of solution and makes
it more available for absorption. EDDHA has a greater affinity for Fe3 + and a lower
affinity for Fe2 + than does DTP A. When the Fe3 + -chelate is reduced to Fe2 + -chelate
at the root, EDDHA releases the iron very readily. Thus EDDHA delivers iron to
the roots more efficiently than DTPA. Also, soybeans have been shown to adapt to
excess chelating agent in nutrient solutions by increasing their ability to reduce and
absorb iron (1). EDDHA was present in excess but DTPA was not. A third reason
for increased efficiency of accumulation of Fe3 + by group II plants over group I plants
could be the lower iron concentration of the nutrient solution.
The further increase in 59Fe uptake by the plants in group III was due to the
removal of non-radioactive iron from the immediate vicinity of the roots. There was
less competition for absorption between the non-radioactive iron and the 59Fe than
in group II. Therefore, more of the 59Fe was available for absorption. There was an
increase in the uptake of 59Fe, but not necessarily total iron. The drawback to this
increased 59Fe uptake is that the procedure used to strip iron from the roots caused
the plants to wilt and their growth to be stunted.
The data presented here indicate that nutrient solutions containing EDDHA result
in more efficient uptake of an iron label than those containing DTPA. Prior stripping
of iron from roots is not recommended.
Literature Cited
1. Chaney, R.L., J.C. Brown, and L.O. Tiffin, 1972. Obligatory reduction of ferric
chelates in iron uptake by soybeans. Plant Physiol. 50:208-213.
2. Chaney, R.L., 1984. Personal communication.
3. Halvorson, A.D. and W.L. Lindsay, 1972. Equilibrium relationships of metal
chelates in hydroponic solutions. Soil Sci. Soc. Amer. Proc, 36:755-761.
4. Hoagland, P.R. and Arnon, D.I., 1950. The water culture method for growing
plants without soil. Calif. Agric. Exp. Sta. Circular 347.
5. Lindsay, W.L., 1974. Role of chelation in micronutrient availability, in The Plant
Root and its Environment, E.W. Carson, ed. University Press of Virginia,
Charlottesville, VA.
6. Sijmons, P.C., and H.J. Bienfait, 1983. Source of electrons for extracellular Fe(III)
reduction in iron deficient bean roots. Physiol. Plant. 59:409-415.
7. Weaver, CM., H.A. Schmitt, M.A. Stuart, A.C. Mason, N.R. Meyer, and J.G.
Elliot, 1984. Radiolabeled iron in soybeans: intrinsic labeling and bioavailability
of iron to rats from defatted flour. J. Nutr., 114:1035-1041.
A Compilation of Plant Diseases and Disorders in Indiana — 1984
Gail E. Ruhl, Richard X. Latin, Paul C. Pecknold
and Donald H. Scott
Department of Botany and Plant Pathology
Purdue University
West Lafayette, Indiana 47907
Introduction
The Plant Diagnostic Clinic in the Department of Botany and Plant Pathology
at Purdue University is a service of the Cooperative Extension Service, Purdue
Agricultural Experiment Station. The clinic provides a free service to interested per-
sons through the county extension system for accurate identification of weeds, plant
diseases and plant disorders. This paper is a summary of the major plant diseases
and disorders which were diagnosed in the clinic and observed throughout the state
in 1984.
Methods
Plant specimens are submitted to the Plant Diagnostic Clinic from county exten-
sion agents, homeowners, growers, nursery operators, consultants, and others. Specimens
are diagnosed visually or by culturing the pathogen on selected media. Some virus
diseases are diagnosed by the leaf dip (negative stain) technique utilizing the electron
microscope. Once a disease or disorder is diagnosed, appropriate control measures
are suggested. A summary of the samples diagnosed from January 1 through Nov.
26, 1984 is given in Table 1.
Results
The incidence and severity of infectious diseases were greatly influenced by extremes
in environmental conditions in 1984. Weather and site-related problems were com-
monplace. The severe cold temperatures of December, 1983, in conjunction with a
lack of snow cover caused widespread death or injury to both agronomic and ornamental
crops.
Shade and Ornamental Trees
Diseases: Ash anthracnose was exceptionally severe in the southern areas of the state,
resulting in heavy defoliation during May and early June. Anthracnose on sycamore,
white oak and maple was also severe. Sycamores especially showed extensive dieback
and twig infection. Apple scab was the most common disease on crabapples, causing
extensive leap drop throughout the summer. Rust diseases in general were severe,
especially cedar quince rust on hawthorn.
Disorders: The severe cold in December in conjunction with a lack of snow cover caused
widespread death or injury to both landscape trees and nursery seedlings. The sudden
freezing caused extensive damage at the Vallonia State Nursery. Especially hard hit
were black walnut, white, black, English and cherrybark oaks, tulip, ash, and Chinese
chestnut seedlings. Established landscape trees most severely damaged were sweetgum,
ornamental cherry, and purple leaf plum. The northern third of Indiana experienced
severe winter burn to conifers and broadleaved evergreens. White oak "tatters," a
newly discovered disorder of white oak, was very prevalent in the northern half of the
state. The exact cause of this disorder is not yet known. Tree decline and leaf scorch
were the most predominant problems during the summer.
145
146 Indiana Academy of Science Vol. 94 (1985)
Table 1 . Plant samples received in the Purdue Plant Diagnostic Clinic Jan. 1 through
Nov. 26, 1984.
Number of
Plant Speciman
Samples
Diseases
Disorders
Chemical0
Nutritional
AGRONOMIC
Corn
95
36
3!
13
9
Soybeans
101
81
9
5
Small Grain
28
19
6
2
4
Forage Grasses
and Legumes
31
25
4
1
2
ORNAMENTAL
Trees-Shade and
Ornamental
332
114
181
14
5
Shrubs and
i
Groundcover
79
12
54
5
1
Flowers
50
30
9
3
4
House plants
13
7
3
0
0
FRUIT
Tree Fruit
62
26
27
1
2
Small Fruit
41
13
21
4
0
VEGETABLE
100
47
22
13
5
TURFGRASS
45
25
20
1
1
PLANT IDENTIFICATION
167
FORWARDED TO
ENTOMOLOGY
64
—
—
—
—
TOTAL
1208
435
387
79
38
Problems caused by an infectious disease causing agent, e.g. fungus, bacterium, virus, mycoplasma, nematode.
Problem caused by noninfectious environmental stress, e.g. wind, drought, heat, soil compaction.
c Problem caused by herbicide/pesticide misuse.
Problem caused by a nutrient imbalance.
Ornamentals
Diseases: Powdery mildew was the most frequently recorded disease of shrubs and
flowers. Those plants most frequently recorded with powdery mildew infections were
lilac, rose, euonymus, and zinnia. Crown gall on euonymus was very noticeable during
the early spring period.
Disorders: Injury from the severe December cold was most noticeable on cotoneaster,
euonymus, pyracantha, holly, rhododendron and barberry. However, many other
ornamentals also showed cold damage. The extent of cold injury varied, depending
on plant age, location and vigor. Symptoms associated with cold injury were complete
plant death, delayed leafing out, sudden wilt and dieback of new growth, as well as
severe cracking of young exposed tissue.
Tree Fruits
Diseases: Heavy rainfall in late April and May resulted in outbreaks of apple scab
in a number of commercial apple orchards. Cedar apple rust was also prevalent. Of
interest was the very light amount of fire blight. This usually widespread disease was
only noted in a few orchards in the northern part of Indiana during late June. The
Botany 147
most noticeable disease on peaches and nectarines was bacterial leafspot, which caused
mild to moderate leaf injury in isolated orchards. Peach leaf curl and plum pockets
on both peach and nectarine were common during mid and late spring.
Disorders: The extreme cold killed fruit buds of many stone fruits. Peach and nec-
tarine were especially damaged. Only the southernmost part of the state yielded a peach
or nectarine crop. In addition to cold injury on fruit buds, there was extensive cold
damage to stem tissue of all tree fruits, most noticeably stone fruits. Many peach trees
were killed or showed extensive limb death. Cold injury to the roots of various apple
root stocks, especially E.M. 7, was noted in a number of orchards in the southern
portion of the state. Damage was most severe on exposed sites which had no snow
cover during December.
Small Fruits
Diseases: Strawberries were the most frequently submitted of the small fruit specimens.
However, no major infectious diseases were recorded on strawberries during the grow-
ing season. Various leaf spots, Botrytis fruit rot and black root rot were common
diseases on samples submitted to the clinic. Raspberry anthracnose and other cane
infecting diseases were frequently observed on brambles.
Disorders: Cold injury to roots and the root-crown area was the most prevalent disorder
of strawberries and raspberries. Such injury resulted in extensive losses to many com-
mercial growers. Entire fields were killed in certain areas of the state.
Turf grass
Diseases: In general, weather conditions were good for turfgrass growth and develop-
ment during 1984. Disease problems were relatively minor and scattered except for
early spring when wet, cool weather was favorable for development of the Helmin-
thosporium leaf blight and melting out complex.
Disorders: Excessive thatch accumulation continues to be a major cause of turfgrass
problems in home laws.
Vegetables
Hot, dry weather in early June may be responsible for the relatively low levels
of foliage diseases throughout the weeks of summer. Moderate or severe epidemics
of foliar vegetable diseases did not occur until late August and mid September, when
cool nights were accompanied by heavy dews. Significant disease problems were observed
on vegetable seedlings, cucurbits, tomatoes, and crucifers.
Seedling diseases: Damping-off, caused by Pythium spp., was diagnosed in muskmelon
and watermelon seedbeds. Most seedbeds showed less than 1% damping-off. However,
at two locations farmers lost more than 30% of their seedlings to damping-off. Pepper
seedbeds were again plagued by Rhizoctonia spp., which caused a wirestem symptom
and death of young seedlings. The problem occurred mostly in outdoor seedbeds, but
occasional problems were observed among greenhouse grown seedlings.
Cucurbit diseases: The usual melon foliar blights, powdery mildew and Alternaria leaf
blight, were established late in the season and, therefore, resulted in little or no yield
loss. An epidemic of downy mildew developed in the melon crop in southwestern Indiana
during mid September. The disease was established too late to cause significant economic
loss.
A malady associated with environmental stress (nutrient imbalance, acid soil, and
air pollution) occurred on a significant number of melon farms in 1984. The problem
148 Indiana Academy of Science Vol. 94 (1985)
was diagnosed in south-central Indiana (Jackson County) for the first time.
Incidence of bacterial wilt was reduced from levels observed in previous years.
Presumably registration and widespread application of a soil-incorporated insecticide
is responsible for reduced levels of bacterial wilt.
Fusarium wilt was severe in southwestern Indiana wherever growers planted wilt
susceptible cultivars. 'Superstar,' a Fusarium wilt resistant muskmelon cultivar that
accounted for less than 5% of the acreage in 1982, was estimated to occupy more
than 60% of the land planted to muskmelons. Fusarium wilt remained a mild problem
on watermelons because growers have been using cultivars that are more resistant.
Tomato disease: Widespread, serious epidemics of major tomato fruit and foliage diseases
did not develop in 1984. Low incidences of anthracnose, bacterial speck, bacterial spot,
early blight, gray leaf spot, and Septoria leaf spot were observed in many fields. Bacterial
canker caused severe or near total losses of fresh market and processing tomatoes at
a variety of locations throughout the state. Until the seed sources can be accurately
assayed and indexed for presence of the bacterial canker organism, this disease will
continue to be a significant threat to tomato production.
Sclerotinia stem rot was responsible for the near total loss of a field in central
Indiana. The distribution of Sclerotinia infected plants normally is very clustered and
incidence usually is less than 0.01%. Patterns in the field and field history suggested
that the organism was introduced by transplants obtained from other states. The presence
of this disease may present long-term problems because the pathogen will remain in-
definitely in northern soils and also may depress soybean yields.
Crucifer diseases: Black rot of cabbage was observed on the most susceptible varieties
in northwestern Indiana. Downy mildew of cabbage, cauliflower, and broccoli was
observed in commercial fields in mid-September.
Agronomic Crops
Disease - Wheat: Extremely cold temperatures in December, 1983, coupled with no
snow cover resulted in considerable winter kill in the southern half of Indiana. Wheat
in the northern half of the state was protected by adequate snow cover, and only minor
winter kill was observed. Rhizoctonia spring blight was prevalent, primarily in southern
Indiana, and this disease coupled with winter kill resulted in poor stands in many fields.
The cool, wet spring was favorable for the development of Septoria leaf blotch and
some powdery mildew. Dry June conditions, however, kept these diseases from develop-
ing to major yield-reducing proportions. Leaf rust developed throughout the state and
to severe levels in some fields. However, the disease developed late in the growing
season and yield losses were estimated to be small. Take-all was severe in some scat-
tered fields but was not a significant problem in most fields. While a few fields were
sparsely affected with either wheat spindle streak mosaic or barley yellow dwarf virus,
both of these diseases were minor and considerably less prevalent than during the 1983
growing season. A few fields were affected with bunt. Bunt appeared to be primarily
in individual fields in the north-central and north-eastern part of the state.
Diseases - Corn: Cool, wet weather delayed corn planting in many fields. Those fields
that were planted in late April and early May were subjected to heavy rainfall. As
a result, portions of many of these fields were flooded for brief periods of time, and
crazy top (Sclerophthora macrospora), developed in small scattered areas of many of
these fields throughout the state. Overall, however, this disease caused only minor
yield losses. Foliar diseases were at low levels throughout the growing season. Minor
field infections by the organisms that cause the leaf blight phase of Stewart's disease,
southern corn leaf blight, holcus spot, northern corn leaf blight and northern corn
Botany 149
leaf spot were observed. Common corn smut was noted throughout the state, but yield
losses were minor. Stalk rots were prevalent in most fields, with some fields having
50% or more of the plants affected. Gibberella and Fusarium stalk rots were most
common with moderate amounts of anthracnose stalk rot. Fusarium ear rot was the
most common corn disease, however not severe enough to cause significant yield loss.
Only rare, light occurrences of Gibberella ear rot were observed.
Disorders - Corn: Hot, dry conditions coupled with several days of continuous high
winds dried out the upper soil surfaces of many corn fields when plants were starting
to develop the crown root system. These environmental conditions were incompatible
with the proper development of the crown root system, and a condition called floppy
corn developed in widespread areas of the state. Cultivation and/or timely rainfall
alleviated the condition in most fields.
A condition of unknown etiology occurred for the first known time in several
southern Indiana fields. A wide range of symptoms were associated with this disorder.
Abnormal plant growth was first observed when corn plants were in the 4th to 5th
leaf stage. The symptoms were stunted, chlorotic plants with portions of new leaves
emerging from the whorl that were translucent and dead or dying. This tissue death
gave plant leaves a "shot-hole" or "cut-leaf" appearance. The initial symptoms were
rapidly followed by leaf trapping and twisting which produced a downward curvature
of plant tops. Multiple suckers developed in many affected plants. In severely affected
fields, varying numbers of these plants died (as high as 50%). Later symptoms were
stunted to spindly single to multiple plants or highly deformed, severely stunted plants
with extremely shortened internodes. Split stalks and deformed leaves were commonly
found on the shorter plants. As affected plants reached the reproductive stage, tassels
were either absent, did not emerge because of leaf trapping, or, in some cases, emerged
normally. Ear shoot development was variable from none to a single ear shoot at each
of several nodes. Also in many plants, ear shoot development occurred at the top
of the plants where the tassel normally develops. Sometimes only ear shoots appeared
at the tassel's location, while in other cases a combination of ear shoots and tassels
developed. When ears developed, regardless of location on the plant, they were small,
poorly pollinated and had a definite curvature. One of the most striking symptoms
was the development of the ear at the top of the plant. Other symptoms noted on
some plants were abnormally long silks and abnormally long, multiple husks that gave
ears a feather duster appearance.
The disorder was at first thought to be associated only with no-till corn in PIK
ground with a heavy sweet clover residue. However, the condition was later found
in other tillage systems and with different plant residues. However, the condition was
more severe and more prevalent in no-till systems. One severely affected field was
no-till corn into wheat stubble. The condition was observed across several hybrids and
herbicide treatments. Purdue entomologists could find no consistent evidence of insect
injury in the affected fields. Purdue, Kentucky, and Illinois plant pathologists could
find no evidence of a plant pathogen in or on the affected plants that is known to
cause similar abnormalities. Some of the symptoms exhibited by affected plants were
similar to symptoms of the downy mildew disease known as crazy top, but oospores
of the causal agent could not be found. With crazy top, these oospores are readily
found in diseased tissue.
The only consistent factors found in all affected fields were that the planter opening
did not close in no-till, or planting was very shallow with some exposed seed in other
tillage systems. Also, planting dates between May 15 and June 1 and wet soil condi-
tions seemed to be consistent. In nearly every instance (with only 1 known exception),
heavy rainfall occurred within a day to a few days after planting.
150 Indiana Academy of Science Vol. 94 (1985)
It is possible that the abnormal development was due to a hormone imbalance
within the plant, but no one has yet secured evidence as to what caused the imbalance*
The general consensus of opinion at Purdue is that the cause was probably a combina-
tion of factors rather than a single causal agent. The causal agent or agents is or are
probably exceedingly rare, as this was the first time anyone recalls seeing the problem.
Further laboratory and greenhouse experiments are being conducted.
Diseases - Soybean: Pythium and Phytophthora seedling blights were common in fields
planted before mid-May. Rhizoctonia root rot was common and occasionally severe
in many fields. Phytophthora occurred in some fields, but it was not severe. Bacterial
blight, downy mildew, and brown spot were common foliar diseases, but their severity
was not sufficient to cause significant yield reductions. The most damaging soybean
diseases were caused by soil-borne pathogens and did not become evident until mid-
season or later. Brown stem rot was more prevalent and damaging than in recent years.
Charcoal root rot was prevalent in southern Indiana and damaging in several fields.
The soybean cyst nematode was identified in additional fields, especially in the north-
western part of the state, and Sclerotinia stem rot caused yield losses in some central
Indiana fields.
Diseases - Alfalfa: Foliar diseases were prevalent throughout the state before the first
cutting. Sclerotinia crown and stem rot was observed in several fields. This disease
was especially damaging in a few fall seeded fields. The crown, root rot complex was
responsible for killing patches of plants in some fields. Rust developed severely in
a few stressed fields late in the season.
CELL BIOLOGY
Chairperson: Ralph Jersild
Department of Anatomy
Indiana University School of Medicine
Indianapolis, Indiana 46202 (317)264-8730
Chairperson-Elect: Robert Stark
Department of Zoology, DePauw University
Greencastle, Indiana 46135 (317)653-4776
ABSTRACTS
Effect of Acetylcholine Stimulation on Cytosolic Chloride in Parotid Acinar Cells.
Kathy Burek and Robert J. Stark, DePauw University, Greencastle, Indiana
46135. In parotid salivary glands, acetylcholine stimulates fluid, electrolyte, and
protein secretion and hyperpolarizes the basolateral membrane. To examine the ionic
mechanisms involved in this process, we used ion-selective and conventional microelec-
trodes to measure the cytosolic chloride activity (ac,) and basolateral membrane poten-
tial (Em) during acetylcholine stimulation of mouse parotid glands. In unstimulated
cells, aci was 45.2 ± 1.1 mM (n=25) and Em was -33.8 ± 1.6 mV (n = 66).
Acetylcholine at concentrations of 2X 10-9, lx 10-8, IX 10-7, lx 10-6 and lx 10-5 M
produced a decrease in ac, of 3.5 ± 0.3, 4.4 ± 0.7, 8.0 ± 0.5, 9.3 ± 0.7, and 9.5 ±
1.8 mM and hyperpolarized Em by 0.6 ± 0.1, 1.4 ± 0.2, 4.9 ± 0.2, 8.4 ± 0.3 and
8.4 ± 0.5 mV respectively. The inverse relationship observed between Em and log
ac, suggests that the membrane hyperpolarizations occurring in response to acetylcholine
stimulation may be related to the corresponding changes in cytosolic chloride. (Sup-
ported by a Research Grant from the Indiana Academy of Science)
Physiological Studies of Azospirillum amazonense. Edwin M. Goebel and Deborah
A. McMahan, Department of Biological Sciences, Indiana University-Purdue Univer-
sity at Fort Wayne, Fort Wayne, Indiana 47805. Members of the genus Azospirillum
have been shown to fix nitrogen under microaerophilic conditions in both tropical
and temperate regions. The microbe will fix nitrogen either in association with the
roots of non-legume plants or free-living in the soil. Two species within the genus
have been extensively studied. Neither of these species were able to utilize disaccharides
for catabolism. A newly described species, A. amazonense, has been shown to utilize
certain disaccharides, especially sucrose. This species shares the ability to use various
five and six carbon sugars and organic acids with the other two members of the genus.
This species has been shown by others to belong to the genus by means of comparing
G + C ratio and morphological characteristics. The study reported here sought to deter-
mine which compounds could be used by the microbe grown under either nitrogen-
fixing or fixed nitrogen conditions. Growth occurred in all conditions tested; however,
the best growth occurred with glucose, sucrose, citrate, succinate and malate. Growth
also occurred with galactose, rhamnose, xylose, fructose, ribose, and both D- and L-
arabinose. Growth studies in a defined medium containing ammonium sulfate showed
the doubling time to be shortest (1.5-2.5 hours) when either glucose or sucrose was
provided. Growth with fructose or galactose was considerably slower. Attempts have
also been made to isolate Azospirillum species from the soil in the midwestern area
of the U.S. A semi-solid nitrogen-free malate medium was utilized for primary isola-
151
152 Indiana Academy of Science Vol. 94 (1985)
tion. Secondary isolation was accomplished by selecting characteristic colonies grow-
ing on a complex agar medium containing congo red.
A Brief History of the Cell Biology Section, Indiana Academy of Science. Ralph
A. Jersild, Jr., Indiana University School of Medicine, Indianapolis, Indiana
46223. The first meeting of the Cell Biology Section was held during the fall meeting
of the Academy, October 21, 1967, at Indiana University, Bloomington. For two years
prior to this, a number of scientists and technicians from throughout the state of In-
diana and with common interests in electron microscopy had been meeting as a separate
group. By 1967 this group was well-established, and it became clear that a more for-
mal organization was needed. Informal discussions were initiated with Dr. A. A. Lind-
sey, then President of the Academy, for organization as a Section within the Academy.
At the time, the formation of a Cell Biology Section had been under consideration
by the Academy. At its spring 1967 meeting, therefore, the Academy offered to tem-
porarily establish a Cell Biology Section through which our group could present and
determine the extent of interest statewide. The idea was accepted enthusiastically. The
divisional meeting in the fall of 1967 was considered a success, with 12 papers and
3 exhibits presented. The Executive Committee of the Academy subsequently voted
at their spring 1968 meeting to make the Cell Biology Section permanent. It was an
honor for me to serve as the Section's first chairperson. Others from the original group
that were instrumental in organizing the Section include Dr. D. James Morre and Dr.
Edward J. Hinsman, Purdue University; and Dr. James E. Carter, Indiana University
School of Medicine. From 1967 through 1984, 13 different persons have chaired this
Section, representing 11 institutional locations around the state. An average of 12 papers
have been presented yearly during this period by persons with interests in Cell Biology.
Concanavalin A Inhibits Oral Regeneration in Stentor coeruleus by Binding to the
Cell Surface. Michael S. Maloney, Department of Zoology, Butler University,
Indianapolis, Indiana 46208. Loss of the oral feeding apparatus of the ciliate Stentor
coeruleus results in the regeneration of a new one in 8-10 hrs, a process known as
oral regeneration. Cell surface glycoproteins seem to be involved in oral regeneration
as Concanavalin A (Con A), which binds to such proteins, delays oral regeneration.
Binding of Con A to the cell surface of Stentor is indicated by the fact that a-methyl
mannoside completely reverses the effect of Con A on oral regeneration. Crosslinking
of membrane bound Con A receptor molecules may also be involved as succinyl Con
A, which does not crosslink these receptors in other cells, has no effect on oral regenera-
tion. To provide a direct demonstration of Con A binding to the cell surface, cells
were exposed to fluorescein isothiocyanate Con A (FITC-Con A) for 30 min, fixed,
and then examined by fluorescence microscopy. Upon exposure to FITC-Con A, the
Con A is localized on the cell surface as accumulations of fluorescent granules on
the posterior one half of the cell. These granules are always localized in the pigmented
stripes between the rows of body cilia. Smaller fluorescent granules were also found
in a linear array at the base of the membranellar cilia in the gullet area. Quite often
the entire membranellar band was diffusely stained. Fixed cells without FITC-Con A
exposure show none of these features. When cells are treated simultaneously with FITC-
Con A and a-methyl mannoside, there is no binding of Con A.
Supported by a Holcomb Research Fellowship from Butler University.
The Effect of Fasting on Sodium Pump Activity in Rat Skeletal Muscle. John W.
Munford and Thomas Koenig, Department of Biology, Wabash College, Crawfords-
ville, Indiana 47933. It has recently been reported that decreased circulating in-
Cell Biology 153
sulin levels, resulting from either diabetes or fasting, are associated with a significant
increase in intracellular sodium levels in rat skeletal muscle. It has been suggested that
this increase in intracellular sodium results from decreased sodium pump activity. To
test this hypothesis, the effect of fasting-induced hypoinsulinemia on the rate of 22Na
efflux from rat soleus muscle was investigated. In soleus muscles isolated from rats
fasted for 72 hrs, the rates of both total 22Na efflux and ouabain-sensitive 22Na efflux
were decreased by approximately 20% compared to the rates of 22Na efflux of muscles
from fed rats. However, it appears that soleus muscles from fasted rats retain their
sensitivity to insulin since the in vitro treatment of soleus muscles from rats fasted
for 72 hrs with insulin increased the rate of 22Na efflux to the same level as in muscles
from fed rats. The decreased rate of 22Na efflux in muscles from fasted rats may be
the result of a decreased number of sodium pump sites since preliminary data suggests
that soleus muscles from rats fasted for 72 hrs have a decreased number of 3H-ouabain
binding sites compared to muscles from fed rats.
Increased Binding of Growth Hormone Following Cleavage by Rabbit Liver
Plasmalemma. Jeanette M. Schepper and James P. Hughes, Department of Life
Sciences, Indiana State University, Terre Haute, Indiana 47809. Several studies
have shown that proteolytic cleavage can enhance the biological activity of the growth
hormone (GH) molecule. It seemed possible therefore, that proteolytic modification
of GH structure might be a normal function of GH-target tissues. Plasmalemma-enriched
fractions isolated from rabbit liver were found to contain a proteinase(s) which cleaved
the large disulfide loop of human (h) and rat (r) GH. The proteolytic activity was
specific to plasmalemma-enriched fractions in that much lower activities were observed
in microsomal-enriched fractions prepared from the same livers. The plasmalemmal
proteinase(s) may be a trypsin-like enzyme because proteolytic activity was decreased
by the two serine proteinase inhibitors. Inhibition by unlabeled hGH of [125 I] GH
binding to receptors did not prevent cleavage of the tracer; therefore, hormone-receptor
interaction was not required for cleavage of the GH molecule. In binding studies, cleaved
GH associated more readily than did intact hormone with rabbit liver receptors. These
studies suggest that plasmalemma-enriched fractions prepared from rabbit liver con-
tain a proteinase which cleaves the GH molecule in a highly specific manner. Moreover,
it is unlikely that inactivation of GH is the function of this limited proteolysis because
cleaved hormone is bound preferentially by at least a subset of receptors in rabbit liver.
Protein Degradation after Eccentric Exercise. A.C. Snyder, A.R. Coggan and J.J.
Uhl, Human Performance Laboratory, Ball State University, Muncie, Indiana
47306. Net degradation of proteins in skeletal muscle and liver occurs after
exhaustive exercise. Similarly, increases in muscle protein degradation and structural
alternations occur following nonexhaustive eccentric muscular contractions (force pro-
duced in lengthening muscles). The purposes of this study were to determine: 1) if
increasing muscle protein, but not liver protein degradation, occurred following a single
bout of nonexhaustive eccentric exercise, and 2) the association between this muscle
protein breakdown and the activity of the calcium activated factor (CAF), a muscle
protease. METHODS: Male rats were randomly assigned to one of two groups: 1)
sedentary or 2) exercised for 90 minutes down a 16° decline on a treadmill at 16 m/min.
Animals were sacrificed 24 hours following the exercise bout and the appropriate tissues
were removed. RESULTS: Following the exercise, muscle protein degradation was
significantly increased; however, no change in liver protein content was observed. The
activity of the CAF enzyme was not increased in any of the muscles examined follow-
ing the exercise bout. CONCLUSIONS: 1) Muscle protein but not liver protein degrada-
154 Indiana Academy of Science Vol. 94 (1985)
tion increases following a single nonexhaustive eccentric exercise. 2) As the CAF enzyme
is thought to be the initiating enzyme of protein degradation, the exact mechanism
causing the increased degradation following nonexhaustive eccentric exercise remains
unknown.
Calmodulin Stimulation of ATP-Dependent Ca2 + Uptake in Maize Root
Microsomes. Martin A. Vaughan, Timothy J. Mulkey and Charles W. Goff,
Department of Life Sciences, Indiana State University, Terre Haute, Indiana
47809. The ATP-dependent uptake of Ca2 + by microsomal membrane fractions
prepared from 1 cm segments of maize root tips was assayed in the presence of added
bovine calmodulin and calmodulin antagonists. Increased concentrations of bovine
calmodulin resulted in increased ATP-dependent Ca2 uptake by the microsomal
vesicles. The magnitude of calmodulin stimulation over calmodulin depleted controls
ranged from 200-400%. The very specific calmodulin antagonist R24571 inhibited the
ATP-dependent Ca2+ uptake by 90% at a concentration of 10~4M. A concentration
of 0.5mM chlorpromazine, a phenothiozine drug, was required to affect a similar level
of inhibition. Contrary to previous reports, these data strongly suggest that the ATP-
dependent Ca2+ uptake of maize root microsomes is a calmodulin mediated process.
The Effect of Illumination on the Rat Pineal as Measured by MSH Activity. Henry
C. Womack, Ball State University, Muncie, Indiana 47306. Albino rats were kept
in constant light or constant darkness for a period of 24 hours. The animals were
then decapitated and their pituitary glands removed, weighed, and homogenized. The
melanocyte-stimulating hormone (MSH) activity of these glands was assayed by in-
jecting the test material into the dorsal lymph sacs of hypophysectomized frogs. Pinealec-
tomized and sham-pinealectomized animals were subjected to these same experimental
procedures. MSH levels were higher in the pituitaries of those rats kept in constant
light regardless of the age or sex of the animal. The pituitary MSH content of rats
kept in constant darkness elevated significantly about eight hours after the animals
were exposed to light; about twice this amount of time was required for significant
decreases in MSH levels when light-adapted animals were placed in the dark. When
pinealectomized rats were placed in darkness there was no subsequent fall in MSH
levels as in the controls. It is felt that the pineal hormone melatonin may influence
pituitary MSH release by blocking the action of a MSH-release inhibiting factor (MIF)
known to be produced by the rat hypothalamus; the release of melatonin itself is sup-
pressed by illumination.
Plasma Progesterone, Blastocyst Steriodogenesis and Blastocyst
Survival in Rats with Altered Thyroid Status
James P. Holland, Richard Brooks and Erich Weidenbener
Department of Biology
Indiana University
Bloomington, Indiana 47405
Introduction
Studies in our laboratory continue to investigate the mechanism by means of which
thyroid hormone influences reproductive physiology in the female rat. Thyroid hor-
mone has been reported to exert effects upon the reproductive system and pregnancy
in many types of animals, including human beings. These findings have been reviewed
by Leathern (11). However, the mechanisms by means of which thyroid hormone ex-
erts these effects have not been elucidated. In our laboratory, earlier investigations
of thyroidal influences upon reproduction have utilized the technique of experimental-
ly delayed implantation of blastocyst in rats (4), which allows the investigator to con-
trol some of the variables which one encounters in studies of reproduction. For exam-
ple, this technique allows the investigator to control the levels of sex steroids available
during early pregnancy, allows control of the time of implantation, and allows the
separation of progesterone-dependent effects from estrogen-dependent effects. Our earlier
investigations (8, 9) using rats demonstrated that thyroxine, in dosages as low as 8
ug per day, can compensate for progesterone deficiency during the progesterone-
dependent maintenance period of experimentally induced delayed blastocytes. The op-
posite effect was caused by surgical thyroidectomy which further intensifies the detrimen-
tal effects of progesterone deficiency upon the survival of blastocysts. Our studies have
further demonstrated that the thyroidal effect upon progesterone-dependent blastocyst
survival is exerted by means of direct effects upon the blastocyst as well as by means
of effects upon the uterus which indirectly effect the blastocyst. For example, thyroid
hormone was demonstrated to stimulate RNA and protein synthesis in the blastocyst
(1) and was demonstrated to stimulate the activity of a uterine enzyme associated with
blastocyst survival (9). In the present investigations there is a continued examination
of a direct effect and an indirect effect of thyroid hormone upon the rat blastocyst.
For a direct effect the influence of thyroid hormone upon blastocyst steroidogenesis
was examined, and for an indirect effect the influence of thyroid hormone upon plasma
levels of progesterone was examined.
Materials and Methods
Sprague-Dawley-derived female albino rats (Harlan Industries, Cumberland, Ind.)
between 60 and 120 days old were maintained on Wayne Laboratory Chow and tap
water ad libitum. All rats were housed in an animal room at 24 °C with a daily il-
lumination schedule of 14 hours of light and 10 hours of darkness. Hyperthyrodism
was induced by daily injection of 48 /xg L-thyroxine (Sigma Chemical Co.) beginning
at least ten prior to the experiment. Surgical thyroidectomies were performed through
a mid-ventral incision in the neck at least four weeks prior to the experiment.
Blastocyst Cytochemistry
Female rats showing a proestrus or estrus vaginal smear were placed overnight
in cages with adult male rats. Insemination was confixmed on the following morning
by the presence of spermatozoa in the vaginal smear and this was designated as Day
1 of pregnancy. Experimental delay of implantation was accomplished by ovariectomy
155
156 Indiana Academy of Science Vol. 94 (1985)
on Day 3 of pregnancy and daily injections of 0.4 mg of progestrone (the deficiency
dosage as determined in earlier studies; 8, 9). Blastocysts were flushed from the uteri
excised from control, hyperthyroid, and hypothroid rats on either Day 5 of pregnancy
(normal, non-delayed blastocysts) or on Day 8 of pregnancy (the final day of the
progesterone-dependent delay period during delay of implantation). Using a one-milliliter
syringe filled with 0.1 M phosphate buffer (pH 7.4) and fitted with a 25 gauge needle,
the blastocysts were washed and flushed into depression slides. Blastocyst cytochemistry
for the determination of 3/3 hydroxysteriod dehydrogenase was conducted according
to the procedure of Dey and Dickman (5). For each blastocyst cytochemistry experi-
ment, the incubation medium was freshly prepared. This medium was prepared by
adding the following to 9.6 ml of 0.1 M phosphate buffer (pH 7.4): 1.8 mg dehydroe-
piandrosterone, 4.5 mg nicotinomide adenine dinucleotide (NAD), and 2 mg Nitro
Blue tetrazolium dissolved in a minimal amount of dimethyl formamide (all obtained
from Sigma Chem. Co., St. Louis, MO.). An aliquot of 0.5 ml of the incubation
medium was placed in each depression of depression slides. Three to four blastocysts
were placed into the medium of each depression; each depression slide was then placed
in a Petri dish containing moistened filter paper and these dishes were incubated at
37 °C for three hours. As controls, some depressions did not contain the substrate
dehydroepiandrosterone. After three hours the blastocysts were removed from the depres-
sion slides, placed on microscope slides, and at 100X and 430X magnification they
were analyzed for intensity of the formazan reaction and were photographed.
Progesterone Radioimmunoassay
Control, hyperthyroid, and hypothyroid rats were ovariectomized and injected
daily with 0.4 mg progesterone for five days in order to simulate the progesterone
maintenance period in the delayed implantation experiments. On the sixth day (com-
parable to Day 8, the final day of blastocyst delay in the delayed implantation
experiments) the rats were anesthesized with ether and blood samples were removed
by means of cardiac puncture. Heparin dissolved in physiological saline was used as
anticoagulant.
Radioimmunoassay of plasma was accomplished using Coat-A-Count, solid phase
,25I radioimmunoassay kits prepared by Diagnostic Products Corp., Los Angeles, Calif.
(6). Duplicate aliquots of 100 ul of each plasma sample were used for the determina-
tions, incubation time was three hours at room temperature, and the tubes were counted
for one minute in a Beckman Gamma 4000 gamma counter. Corrections were made
for non-specific binding, a seven-point standard curve was established, and the pro-
gesterone levels in the plasma samples were expressed as ng per milliliter. The assay
is sensitive to a minimum of 0.05 ng per milliliter.
Results
Table I summarizes the evaluations of the cytochemical reaction for 3 (3-
hydroxysteriod dehydrogenase in non-delayed blastocysts which were flushed from the
uteri of control, hyperthyroid, and hypothyroid rats on Day 5 of pregnancy. High
amounts of the enzymatic reaction were present in all of these blastocysts, and there
were no differences between blastocysts which were obtained from rats of different
thyroid states. As a control for the reaction, blastocysts which were incubated in medium
without the dehydroepiandrosterone substrate did not show the darkening indicative
of the enzymatic reaction.
Table 2 summarizes the evaluations of the cytochemical reaction for 3 /3-
hydroxysteroid dehydrogenase in delayed blastocysts (maintained on a deficiency dosage
of progesterone) which were flushed from the uteri of control, hyperthyroid, and
Cell Biology 157
Table 1 . Summary of Histochemical Determinations of 3 /3-OH Steroid Dehydrogenase
in Five-day Blastocysts Obtained from Rats of Different Thyroid States (Intact Ovaries — No
Exogenous Progesterone)
No. Blastocysts
Examined
Enzyme
Reaction in
Treatment
Trophoblast
Inner Cell Mass
Euthyroid
Hyperthyroid
Hypothyroid
26
24
17
+ + +
+ + +
+ + +
+ + +
+ + +
+ + +
Blastocysts were incubated for three hours at 37 °C in 0.1 M phosphate buffer containing dehydroepiandrosterone,
NAD, and Nitro Blue tetrazolium.
hypothyroid rats on Day 8 of pregnancy. The blastocysts from control and hyper-
thyroid rats showed approximately the same levels of enzyme activity, except for possibly
higher activity in the inner cell mass area of those from hyperthyroid rats. The blastocysts
Table 2. Summary of Histochemical Determinations of 3 (3-OH Steroid Dehyrogenase
in Eight-day Blastocysts Obtained from Rats of Different Thyroid States (Experimentally
Delayed Blastocysts)
No. Blastocysts Enzyme Reaction in
Treatment Examined Trophoblast Inner Cell Mass
Euthyroid 28 + + + +
+ 0.4 mg Prog.
Hyperthyroid 55 + + + + + +
+ 0.4 mg Prog.
Hypothyroid 16 +
+ 0.4 mg Prog.
All animals were ovariectomized on Day 3 of pregnancy and maintained with progesterone until autopsy on Day 8.
Blastocysts were incubated for three hours at 37 °C in 0.1 M phosphate buffer containing dehydroepiandrosterone,
NAD, and Nitro Blue tetrazolium.
from the hypothyroid rats, however, showed much lower levels of the enzyme activity
and some of these blastocysts were entirely devoid of the enzyme activity. Again, the
blastocysts which were incubated in medium without the dehydroepiandrosterone did
not show darkening.
Table 3 contains the data from the l25I radioimmunoassay determinations of plasma
Table 3 . Effect of Altered Thyroid States Upon Plasma Progesterone Levels in Ovariec-
tomized Rats Injected Daily with 0.4 mg Progesterone for Five Days
Plasma Progesterone +
Treatment No. Rats (ng/ml)
Euthyroid 8 5.27
Hyperthyroid 8 4.55
Hypothyroid 8 9.32*
+ Corrected to uniform body weights.
Progesterone determinations by means of radioimmunoassay (Diagnostic Products Corporation).
*Significantly different from euthyroid and hyperthyroid (P < 0.05) as determined by Tukey and Scheffe analysis.
158 Indiana Academy of Science Vol. 94 (1985)
progesterone in rats of different thyroid states. It can be seen that there was no dif-
ference in plasma progesterone level in control and hyperthyroid rats. On the other
hand, the hypothyroid rats had a significantly higher plasma progesterone level than
controls (9.32 ng/ml and 5.27 ng/ml, respectively).
Discussion
The survival of blastocysts in rats which are ovariectomized on the third day
of pregnancy is progesterone-dependent. Implantation and further development of these
blastocysts are "delayed" since ovariectomy removes the source of estrogen which is
essential for the implantation process (13). The normal daily maintenance dose of pro-
gesterone during delay is 2.0 mg; a deficiency dose of 0.4 mg/day causes a significant
decrease in the number of surviving blastocysts (10). Thyroid hormone has been
demonstrated to exert important effects upon delayed blastocysts during a deficiency
of progesterone. Our earlier studies have shown that these thyroidal effects may be
exerted directly upon the blastocyst or indirectly by means of altered uterine physiology.
The full extent and mechanisms of the direct and indirect effects of thyroid hormone
have not been elucidated. The present studies were conducted to determine whether
maternal thyroid activity can alter the activity of an enzyme in the blastocyst which
is important for progesterone synthesis and whether thyroid hormone influences the
plasma levels of progesterone which may influence uterine physiology.
Using cytochemical determinations of 3 /3-hydroxysteroid dehydrogenase, a key
enzyme in steroidogenesis, investigators (5) have demonstrated that the pre-implantation
blastocysts of the rat and the rabbit synthesize steroid hormones. It has also been
suggested that blastocysts of the rabbit accumulate steroids from the uterine fluid (2).
In the present studies the five-day blastocysts from normal, intact rats all showed high
levels of the 3 /3-hydroxysteriod dehydrogenase regardless of the thyroid status of the
mother. This is not surprising since our data from earlier experiments have all in-
dicated that thyroidal effects only become important during progesterone deficiency.
No such deficiency existed in these intact rats. On the other hand, the delayed" blastocysts
in hypothyroid rats which had been maintained on a deficiency dosage of progesterone
showed lower intensity of the enzyme reaction. This finding correlates well with our
earlier studies. The hypothyroid, progesterone-deficient rats comprise the group which
had the lowest survival of blastocysts (8, 10), and these blastocysts showed the lowest
amount of protein synthesis (1). However, the ability of hyperthyroidism to overcome
the detrimental effects of progesterone deficiency cannot be explained by means of
the 3 /?-hydroxysteroid dehydrogenase studies since blastocysts from control and hyper-
thyroid rats showed approximately the same levels of the enzymatic reaction. In order
to more clearly determine whether thyroid hormone influences steroidogenesis in the
rat blastocyst, we are conducting in vitro steroidogenesis experiments with long-term
(four days) incubated blastocysts using NCTC-135 (GIBCO) nutrient medium which
is changed daily and analyzed for progesterone by means of radioimmunoassay. Recently,
McCormack (12) reported in vitro studies of rat blastocyst steroidogenesis using this
procedure. They found that the blastocyst production of progesterone was low and
variable.
An indirect route by means of which thyroid activity may influence blastocyst
survival is by altering the metabolism of progesterone and its action upon the uterus.
Bradlow et al. (3) reported that thyroid hormone alters the activity of enzymes that
transform progesterone in vivo in hyperthyroid human beings, resulting in a shift toward
the production of 5 a-reduced metabolites of progesterone. Our preliminary studies
reported here show that in rats which were treated with progesterone in a manner
to simulate the delayed implantation studies, hypothyroid rats have a significantly higher
Cell Biology 159
plasma progesterone level than do control or hyperthyroid rats. Since the hypothyroid
rats are the ones with the lowest survival of blastocysts, the higher plasma progesterone
level in these rats may reflect an altered metabolism (utilization, degradation, excre-
tion, etc.) of progesterone which has an overall negative effect upon uterine physiology
and blastocyst survival. Again, the beneficial effects of thyroid hormone cannot be
explained by the present data. Gas chromatographic studies of progesterone metabolism
in rats of different thyroid states are underway to further examine the conversion of
progesterone to related compounds and the excretion patterns of progesterone
metabolites. Also, uterine progesterone receptor binding studies should be helpful in
approaching firm conclusions concerning the effects of thyroid hormone which are
exerted by way of the uterus. Receptors for thyroid hormone were recently identified
in the rat uterus (7); therefore, thyroidal effects upon the uterus are expected to be
significant.
Collectively the present studies concerning blastocyst cytochemistry and plasma
progesterone levels are generally supportive of our earlier findings that thyroid hormone
can influence blastocyst survival by means of direct effects upon the blastocyst and
by methods which influence uterine physiology. Further studies are in progress in order
to more clearly determine the mechanisms which are involved.
Literature Cited
1. Archer, V.G. and J. P. Holland. (1980). Effect of maternal thyroid activity upon
in vitro protein synthesis in the rat blastocyst. Proc. Ind. Acad. Sci. 90:136-142.
2. Borland, R.M. G.F. Erichson, and T. Ducibella. (1977). Accumulation of steroids
in rabbit pre-implantation blastocysts. J. Reprod. Fert. 49:219-224.
3. Bradlow, H.L. D.K. Fukushima, B. Zumoff, L. Hellman, and T.F. Gallagher.
(1966). Influence on thyroid hormone on progesterone transformation in man.
J. Clin. Endocrinol. Metab. 26:831-834.
4. Cochrane, R.G. and R.K. Meyer. (1957). Delayed nidation in the rat induced
by progesterone. Proc. Soc. Exp. Biol. Med. 96:155-159.
5. Dey, S.K. and Z. Dickman. (1974). DeIta-5, 3-beta hydroxysteroid dehydrogenase
activity in rat embryos on days 1 through 7 of pregnancy. Endocrinology
95:321-322.
6. Diagnostic Products Corp. (1982). Progesterone, Coat-A-Count protocol.
7. Evans, R.W., A. P. Farwell, and L.E. Braverman. (1983). Nuclear thyroid hor-
mone receptor in the rat uterus. Endocrinology 113:1459-1463.
8. Holland, J. P., J.M. Dorsey, N.N. Harris, and F.L. Johnson (1967). Effect of
thyroid activity upon delayed implantation of blastocysts in the rat. J. Reprod.
Fert. 14:81-85.
9. Holland, J. P., F.L. Calhoun, N.N. Harris, and N.W. Walton. (1968). Uterine
alkaline phosphatase and blastocyst implantation during altered thyroid activity.
Acta Endocr., Copenh. 59:335-343.
10. Holland, J. P., J.M. Finley, R.D. Kazwell, and F.L. Meshberger. (1970).
Progesterone-dependent blastocyst survival during altered thyroid activity in the
rat. J. Reprod. Fert. 23:143-146.
11. Leathern, J.H. (1972). Role of the thyroid. In Reproductive Biology, H. Balin
and S. Glasser (eds.), pp. 857-876. Excerpta Medica, Amsterdam.
12. McCormack, S.A. and S.R. Glasser. (1981). Hormone production by rat blastocysts
and midpregnancy trophoblasts in vitro. In Cellular and Molecular Aspects of
Implantation, S. Glasser and D.W. Bullock (eds.), pp. 461-463. Plenum Press,
New York.
13. Prasad, M.R.N., S. Mohla, and M. Rajalakshmi. (1969). Hormonal environment
160 Indiana Academy of Science Vol. 94 (1985)
and blastocyst development. In Progress in Endocrinology, C. Gaul and F.J.B.
Ebling (eds.), pp. 939-943. Excerpta Medica, Amsterdam.
Chick Limb Duplications Produced by Retinoic Acid Releasing
Microimplants
Lisa B. Nass, Annette J. Schlueter and Grayson S. Davis
Department of Biology
Valparaiso University
Valparaiso, Indiana 46383
Introduction
Several recent studies have suggested that the limb bud vasculature may act in
determining the skeletal pattern of the limb (3) by establishing metabolic gradients
which would control the local differentiation of muscle or cartilage (1). Furthermore,
systemic application of vitamin A, or its acid, retinoic acid, to developing embryos
has been shown to produce both skeletal malformation and abnormal vascularization
of the limb (7, 4). When microimplants of filter paper containing retinoic acid are
implanted into developing limb buds, they can induce duplications of the limb skeleton
(11), apparently mimicking the action of the polarizing region. Our initial interest was
to observe the effect upon the vasculature produced by retinoic acid implants. However,
we were unable to reliably produce duplications with filter paper implants. An alter-
native implant, an ion exchange bead, was suggested by Bruce M. Alberts, Depart-
ment of Biochemistry and Biophysics, University of California, San Francisco (2). A
comparison of these two carriers revealed that the ion exchange implanting method
was far more reliable, less toxic, and less likely to induce other malformations than
the filter paper implanting method.
Methods
For the paper implants, Rhode Island Red chick eggs were incubated under standard
conditions for three days to stages 18 to 20 (6) while being turned twice daily. The
eggs were then windowed as described by Hamburger (5) except that the windows were
broken into the shell with forceps rather than sawn with a hacksaw blade. This method
is reliable for early stage chicks and much faster than sawing. The amniotic fold directly
over the right wing bud was pulled back using an electrolytically sharpened tungsten
wire probe and a slit was then made into the anterior portion of this bud using the
same probe.
A small piece (0.5mm x 0.5mm) of Whatman diethylaminoethyl cellulose (DEAE)
filter paper was prepared for implanation by being soaked for one minute in one
of a series of concentrations of all trans-retinoic acid (Sigma, type XX) dissolved in
dimethyl sulfoxide (DMSO; Sigma grade 1). The solutions for each experiment were
made from a freshly opened ampoule of retinoic acid and kept in darkness to minimize
decomposition of the retinoic acid. Paper soaked longer than one minute tended to
disintegrate when implantation into the slit was attempted. This paper was then im-
planted into the slit such that the paper extended through the apical ectodermal ridge
and into the limb mesoderm adjacent to somites 15, 16 and 17. A free edge of the
paper remained outside the limb bud. The window was then sealed with cellophane
tape and the egg reincubated.
After seven days, the embryo (now at stage 35 to 37) was removed from the
egg, rinsed in physiological saline and fixed in Bouin's fixative. The fixed embryo
was stained with 1.2% solution of Victoria Blue B dye (Sigma) to stain the cartilages,
dehydrated in a graded ethanol series (50%, 70% and 95%) and transferred to methyl
salicylate to clear the flesh so that the skeletal elements could be examined for
duplications.
161
162
Indiana Academy of Science
Vol. 94 (1985)
For the bead implants, the same method was used up to and including slitting
the wing bud with the tungsten probe. However, instead of introducing the retinoic
acid in filter paper carriers, AG1-X2 ion exchange beads (Formate form, 100-200 mesh,
Bio-Rad Laboratories) were used. These beads, made of a styrenedivinylbenzene
crosslinked lattice with attached quaternary ammonium groups, exchanged electro-
statically bound formate ions for retinoic acid ions when soaked for 20 minutes in
one of a series of concentrations of retinoic acid in DMSO (2). The beads loaded
with retinoic acid were then rinsed twice with ten minute changes of Hank's balanced
salt solution and implanted. The implanted bead was completely surrounded by the
100
90
80
70
O 60
Si
o
§J 50
Q
40
30
20
10
O
or
■A FILTER PAPER (N = 35)
■0 ION EXCHANGE BEAD (N = 117)
-©
f
4
0.1
23456789
RETINOIC ACID CONCENTRATION (MG/ML)
Figure 1. Dose response curves for filter paper and ion exchange bead implants:
percent of surviving embryos with limb duplications versus retinoic acid concentration.
Cell Biology
163
mesoderm just beneath the apical ectodermal ridge. The egg was next sealed with
cellophane tape and reincubated. The embryo was later fixed, stained and cleared as
before. Controls were run for both implanting processes using either filter paper or
beads soaked in DMSO alone. Several bead-implanted and several paper-implanted
embryos were fixed after only one or two days of incubation and examined to see
if the implants had slipped out of the limb.
Results
All control embryos examined one or two days after implantation of either filter
paper or beads still retained their implants. Furthermore, once 10 day embryos were
cleared, it was often possible to find the implant still in the limb. When loaded with
retinoic acid, both types of implant were capable of producing duplications in the
cartilages of the autopod. Bead implants produced duplications when loaded with lower
concentrations of retinoic acid than did the paper implants (Figure 1). Trying to pro-
duce more duplications by increasing the retinoic acid concentration loaded into the
paper produced an increase in the death rate to a value much greater than that obtained
with the bead implants (Figure 2). Moreover, nearly 100°7o of the embryos surviving
100
0 FILTER PAPER ( N = 131)
90
" ION EXCHANGE BEAD (N - 117J
_- -®
80
_ -—
70
^ J&— "
en
60
— "
I
©- "
h-
<
UJ
50
Q
IL
O
40
3
30
20-
10
0
»—
»
A — ■ i
RETINOIC ACID CONCENTRATION (MG/ML)
Figure 2. Dose response curves for filter paper and ion exchange bead implants:
percent of deaths unattributable to contamination or injury at the time of implant
versus retinoic acid concentration.
the paper implant technique were malformed. These embryos developed brain defor-
mations, beak deformations, ectopia cordis (heart exterior to the chest cavity), or
extensive abdominal herniation of the gut. Such abnormalities were common but not
universal in control embryos implanted with filter paper soaked in DMSO alone. Only
one of the 117 surviving bead-implanted embryos showed a detectable malformation.
When the results of the 5mg/ml bead-implanted embryos were examined, substantial
differences in the length of the duplicated digits were noted. Embryos implanted at
earlier stages had longer duplications (Figure 3).
164
Indiana Academy of Science
Vol. 94 (1985)
ai
5
O
m
<r>
a
ui
i-
<
_j
Q.
3
a
u.
o
<T
UJ
CD
5
(N = 35 AT 5 MG/ML)
18
19 20
STAGE OF EMBRYO AT TIME OF BEAD
IMPLANT
Figure 3. Average number of duplicated segments versus the stage of the embryo
at the time the bead was implanted.
Discussion
The data from the two types of carriers showed significant differences. A much
higher rate of duplication was obtained with lower retinoic acid concentrations when
using ion exchange beads as opposed to filter paper. Specifically, an 83% duplication
rate was obtained at a concentration of 5mg/ml of retinoic acid using ion exchange
beads, significantly better than a 17% duplication rate at a concentration of 8mg/ml
using the filter paper carrier (see Figure 1). This may be attributable to the fact that
the bead releases retinoic acid in lower concentrations and over a longer period of
time than does the paper (2). Since neither paper nor bead shows an inclination to
slip out of the limb after proper implantation, we cannot attribute the lower percent
duplication obtained with paper to the failure of paper to remain implanted.
At the same time, the death rate (unattributable to contamination or embryonic
injury at the time of implantation) was much lower for bead-implanted embryos than
for filter paper-implanted embryos. At a 5 mg/ml concentration of retinoic acid using
the ion exchange bead method, a death rate of only 7% was observed, whereas at
a 4 mg/ml concentration of retinoic acid using the filter paper method a 56% death
rate resulted (Figure 2). Because the malformations in the filter paper-implanted em-
bryos occurred in the controls as well as the experimental chicks, it may be that this
effect was due to the filter paper itself, or, more likely, to the relatively large amount
of DMSO each filter paper implant carried. Furthermore, the embryos treated with
retinoic acid-containing paper implants almost universally developed with head defor-
mations, heart exterior to the chest cavity or gut exterior to the abdominal cavity.
In contrast, the ion exchange bead treated embryos showed only one case of deforma-
tion (at a concentration of 5 mg/ml).
The fact that implanting young embryos produced longer (proximal to distal)
duplications is not surprising given the popular model of sequential proximal-distal
specification of limb pattern (8). According to this model, increasingly shorter and
more distal regions of the limb would be labile to alterations, including duplications,
Cell Biology 165
of pattern at later stages (9). We were surprised that a stage 18 implant would produce
duplications in the autopod alone. Experiments in which limb development is inter-
rupted by removal of the apical ectodermal ridge indicate that at stage 18 pattern
specification is not yet effectively complete for the zeugopod or even the most distal
portion of the stylopod (8, 10).
We are convinced that retinoic acid implants are an effective tool for producing
duplications in the pattern of limb cartilages. Of the two carriers we compared, the
ion exchange beads are by far the more reliable and less damaging to the embryo.
We are now examining the early effects of retinoic acid implants upon the vasculature
of the limb bud.
Literature Cited
1. Caplan, A. I., and S. Koutroupas, 1973. The control of muscle and cartilage
development in the chick limb: the role of differential vascularization. J . Embryol.
exp. Morph. 29:571-583.
2. Eichele, G., C. Tickle, and B.M. Alberts. Micro-controlled release of biologically
active compounds in chick embryos: beads of 200 um diameter for the local release
of retinoids. In preparation.
3. Feinberg, R.N. and J.W. Saunders, Jr., 1982. Effects of excising the apical
ectodermal ridge on the development of the marginal vasculature of the wing
bud in the chick embryo. J. exp. Zool. 219:345-354.
4. Fraser, B.A., and A. A. Travill, 1978. The relation of aberrant vasculogenesis
to skeletal malformation in the hamster fetus. Anat. Embryol. 154:111-120.
5. Hamburger, V., 1942. A Manual of Experimental Embryology. University of
Chicago Press, Chicago.
6. Hamburger, V., and H. Hamilton, 1951. A series of normal stages in the develop-
ment of the chick embryo. J. Morphol. 88:49-92.
7. Kochhar, D.M., 1977. Cellular basis of congenital limb deformity induced in mice
by vitamin A. Proceedings of the Second International Conference on
Morphogenesis and Malformation. Birth Defects: Original Article Series 13:111-154.
8. Saunders, J.W. Jr., 1948. The proximo-distal sequence of the origin of the parts
of the chick wing and the role of the ectoderm. J. exp. Zool. 108:363-404.
9. , M.T. Gesseling, and J. Errick, 1976. Inductive activity and enduring cellular
constitution of a supernumerary apical ectodermal ridge grafted to the limb bud
of the chick embryo. Devi. Biol. 50:16-25.
10. Summerbell, D., 1974. A quantitative analysis of the effect of the excision of
the AER from the chick limb-bud. J. Embryol. exp. Morph. 32:651-660.
11. Tickle, C, B. Alberts, L. Wolpert, and J. Lee, 1982. Local application of retinoic
acid to the limb bud mimics the action of the polarising region. Nature, Lond.
296:564-565.
CHEMISTRY
Chairperson: Shannon Lieb
Department of Chemistry, Butler University
Indianapolis, Indiana 46208 (317)283-9410
Chairperson-Elect: Dennis G. Peters
Department of Chemistry, Chemistry Building Room A112, Indiana University
Bloomington, Indiana 47405 (812)335-9671
ABSTRACTS
Ambidentate Phosphine Ligands: Phosphine-amine and Phosphine-imidate Complexes
of Tungsten. Sepehra Akhavan, Kristen Faust and Bruce Storhoff, Department
of Chemistry, Ball State University, Muncie, Indiana 47306. The reaction of
Ph2PCH2CH(R)CN (R = CH3,H) (L) with W(CO)6 with an excess of NaBH4 in dry
ethanol provides excellent yields of as-coordinated (CO)4W[Ph2PCH2CH(R)CH2NH2]
and (CO)4W[Ph2PCH2CH(R)C(OC2H5)NH. The latter are converted to the correspond-
ing phosphine-amine complexes upon reaction with additional NaBH4. These results
are rationalized in terms of a reaction scheme involving a side-on coordinated nitrile
group which is susceptible to nucleophilic attack by ethanol. The coordinated amine
and imidate groups are replaced by PMe2Ph providing mixtures of cis and trans
complexes.
The Synthesis of a Crown Ether that May Exhibit Metal Cation Enhanced Fluorescence.
Stasia A. Barnell, Beth E. Beeson and Lynn R. Sousa, Department of Chemistry,
Ball State University, Muncie, Indiana 47306. A crown ether molecule that con-
tains both a fluorescent chromophore and a potential quencher of that chromophore's
fluorescence is being synthesized. The synthesis is convergent and involves several steps.
Based on literature data concerning the steric requirements for the quenching of
fluorescent singlet states and our understanding of metal cation complexation by crown
ethers, it is probable that some metal cations (Na + , K + , Ca+\ etc.) will enhance
the fluorescence of our crown ether compound. Such a "fluorogenic" crown ether
could prove useful for the quantitative analysis of selected metal cations.
2,4-Dinitrophenylhydrazones: A Modified Method for the Preparation of these
Derivatives and an Explanation of Previous Conflicting Results. Mohammad Behforouz,
Joseph L. Bolan and Michael S. Flynt, Department of Chemistry, Ball State Univer-
sity, Muncie, Indiana 47306. We have found that the conventional methods for
forming 2,4-dinitrophenylhydrazones (2,4-DNPs) usually leave traces of acids com-
plexed with the derivatives and that this has been the major cause for the melting
point discrepancies and controversy throughout the 50 year history of their applica-
tion. A simple modification of the original method, a bicarbonate wash of the 2,4-DNP
crystals, removes the acid and reproducibly gives derivatives with previously reported
or higher melting ranges. A series of aldehydes and ketones was selected and the
2,4-DNPs were prepared by both the conventional and the modified methods. In nearly
all cases the modified method gave products with higher melting ranges. 2,4-DNPs
of several hydroxy ketones previously unattainable by the standard method were also
prepared. Careful studies of the 2,4-DNPs of acetaldehyde and 3-hydroxy-
3-methyl-2-butanone by NMR spectroscopy and differential scanning calorimetric analysis
167
168
Indiana Academy of Science
Vol. 94 (1985)
showed that traces of acids incorporated in the crystals catalyze the interconversion
of the syn and anti forms of the 2,4-DNPs and promote the dehydration of the hydroxy-
carbonyl derivatives thus lowering or changing the melting behaviors of the products.
Wittig Reaction: Stable Ylides in the Preparation of 7,<5-unsaturated-j3-Ketoesters.
Mohammad Behforouz and K.E. Mennen, Department of Chemistry, Ball State
University, Muncie, Indiana 47306. Stable ylides of /3-ketoesters are prepared and
their condensations with aromatic and aliphatic aldehydes to give 7,6-unsaturated-jS-
ketoesters are discussed. These Wittig reactions are sterospecific and give mainly the
E-isomers. ~
0
R = H, Me
X = CI, Br
PPh3
RAr>
PhoP
C02Me
PPh3X'
0
, I'
R-C-H
Base
PPho
0 0
,/
R "R
OMe
Base
CO (OMe)
Synthesis of /3-CarboIines Derived from 2-Amino-3-(3-indoIyl)-butyric Acid (/3-
Methyltryptophan). Mohammad Behforouz and M.E. Ogle, Department of Chemistry,
Ball State University, Muncie, Indiana 47306. Although /3-carbolines are com-
mon structural units and their chemistry has been well documented, their preparations
from 0-methyltryptophan (1) have never been reported. Four aldehydes were reacted
with /3-methyltryptophan via the Pictet-Spengler reaction to form the corresponding
salts of tetrahydro-/3-carbolines (2a-d). The formation and subsequent esterification
and dehydrogenation of these compounds were studied.
An example of such a synthetic route involves the reaction of acetaldehyde with
j8-methyltryptophan in aqueous sulfuric acid at 25° to yield 2a-d. The resulting salt
(2a) was filtered, dried, and dissolved in saturated methanolic HC1 and refluxed under
nitrogen to yield the salt 3a-CHl. The salt was converted to the free base on treatment
with 14% NH4OH to give 3a. The resulting free base was refluxed over 10% Pd/C
in dioxane to yield 4a. Reactions involving the other aldehydes were conducted in an
identical fashion.
+ RCH
CH3CH
HOCH-
CO2CH3
'HC1
Pd/C_
(3a-d)
Chemistry 169
Coulometric Titrations: Low Cost Alternatives for Computer Controlled Titrations.
Stanley L. Burden and Phillip W. Schultz, Department of Chemistry, Taylor Univer-
sity, Upland, Indiana 46989. Software and hardware for several different system
configurations to carry out coulometric titrations under control of an Apple II or He
computer will be presented. One of the main advantages of the coulometric approach
is the elimination of costly titrant delivery systems. The systems described will accom-
modate a variety of budget and equipment limitations. The simplest systems use, in
addition to the Apple and pH meter with BCD readout, only a battery, electrodes
and simple interfacing costing less than $50 to construct. Such a system is useful for
classroom demonstrations or experiments in which shapes of titration curves are the
primary data of interest as opposed to highly accurate end point determinations. The
software presented will plot the data collected in real time and will compute and display
first and second deratives as well as Gran plots in different colors using the high resolu-
resolution graphics. A somewhat more expensive system uses operational amplifiers
for the constant current source and a relay and timer, controlled by the computer,
to stop the titration at a user selected endpoint potential or pH. The software for
this system also permits specifying a pH or potential at which current will begin to
be delivered in short pulses, with user selected intervals between pulses, to permit the
endpoint to be approached slowly. Applications of the use of these systems and typical
data will be presented.
Temperature Dependent Infrared Studies of the Hydrogen Bonding in Aliphatic Alcohols.
Mark Cisneros and Joe Kirsch, Department of Chemistry, Butler University,
Indianapolis, Indiana 46208. The extent of the hydrogen bonding of aliphatic
alcohols in dilute solutions has been studied through the use of temperature dependent
infrared spectroscopy. The equilibrium constants for the hydrogen bonding process
have been determined as a function of temperature. The enthalpy and entropy for
the hydrogen bonding process has been calculated from the temperature dependent
equilibrium constants. The equilibrium constants, enthalpies, and entropies for the
hydrogen bonding process are related to the molecular structure of the alcohols and
the steric hindrance at the hydrogen bonding site.
Spectra and Equilibria of the Thiocyanato Complexes of Copper (I) in Aqueous Solu-
tion. Sally K. Dotterer and Kenneth L. Stevenson, Department of Chemistry,
Indiana University-Purdue University at Fort Wayne, Fort Wayne, Indiana
46805. The objective of this research was to determine the spectra of the copper
(I)-thiocyanato complexes and recheck the published value of the equilibrium constant
for the reaction:
Cu(SCN),2- + SCN" = Cu(SCN)43-
According to Ahrland and Tageson, there are normally three complexes present in
aqueous solutions of CuSCN and SCN" : Cu2(SCN)64 - , Cu(SCN)32~ , and Cu(SCN)43" .
Of these, based on the reported values for equilibrium constants, only Cu(SCN)32_
and Cu(SCN)43~ are present in significant concentrations. To determine the spectrum
of each complex in the equilibrated, anaerobic solution, spectra were run on solutions
of copper (I) thiocyanate, at a constant ionic strength of 5.0M (NaC104 medium) while
varying the thiocyanate concentration between 0.2M and 5.0M. The most significant
data appeared at thiocyanate concentrations lower than 1.0M, but the very low solubility
of CuSCN at these thiocyanate concentrations caused difficulty in obtaining these data.
This was done by filtering saturated CuSCN solutions, taking the spectra, and deter-
mining the copper concentration with atomic absorption spectrophotometry. At thio-
cyanate concentrations above 2.0M, the copper concentration was maintained at 0.005M.
170 Indiana Academy of Science Vol. 94 (1985)
From the spectra, the molar extinction coefficient was calculated at a specific wavelength
for each solution, and, using a computer technique, an attempt was made at recalculating
the value for the equilibrium constant. From this value, the spectra of each of the
two complexes can be determined.
Steric and Electronic Effects upon cis:trans Distributions in W(CO)4(L)(L ' ) Complexes
when L and L' are Phosphorus Ligands. Jennifer L. Dyke and John A. Mosbo,
Department of Chemistry, Ball State University, Muncie, Indiana 47306. The
tungsten complexes W(CO)4(L)(Py) (Py = pyridine and L = PPh2Et or PPhMe2)
have been reacted with two series of electronically and sterically divergent ligands,
PPhx (OMe)3_x and PPHX (NMe2)3.x (where x = 0, 1 or 2). Cis:trans ratios of the
W(CO)4(L)(L') products decrease in the order x = 0>1>2 for the OMe-containing
series, but increase with the number of phenyl groups for the MNe2-containing ligands.
These results are consistent with increasing trans preference as the size of L' increases,
but are also consistent with increasing trans preference as the Tolman electronic
parameter (v) decreases.
A Simple, Reproducible High Performance Liquid Chromatography Separation of
Amino Acids with Picomole Sensitivity. Bernice Ellis, Kevin Cooksy, James M.
Anderson, and Harry W. Jarrett, Department of Biology, Indiana University-Purdue
University at Indianapolis, Indianapolis, Indiana 46223 and Alltech Associates/Ap-
plied Science Labs, Deerfield, Illinois 60015. A method for separating and quan-
titating the amino acids commonly found in protein acid hydrolysates has been developed.
The amino acids are derivatized using o-phthaldehyde reagent and separated using reverse
phase by a method similar to published reports. Our method differs from these in
that it uses a C8 reverse phase column, 0.1% triethylamine in the buffers to improve
the peak shape of HIS, and achieves baseline separation of all amino acids in 16 min
total analysis time. The column (4 x 15 cm) used is inexpensive and various lots of
the column have been shown to behave identically. The limits of detection are in the
10 ~ 12 to 10 ~ '4 mole range. The same column has been used for over 100 analyses without
a guard column or any special precautions with no degradation of performance noted.
The detailed method and factors which influence resolution will be discussed.
An Electron Spin Resonance Method for the Measurement of Liposomal Leakage.
Maureen L. Hill, Patrick Gallagher and Jeff Macri, Department of Medical
Research, Methodist Hospital of Indiana, Inc., 46202, F.W. Kleinhans, Department
of Medical Research, Methodist Hospital of Indiana Inc., and Department of Physics,
Indiana University-Purdue University at Indianapolis, Indianapolis, Indiana
46223. The effectiveness of different compositions of liposomes as drug delivery
systems is dependent on their leakage properties. Diffusion into multilamellar liposomes
was examined by recording the increase in ESR signal amplitude as the spin label,
3-carboxy-Proxyl, diffused into the lipid vesicle. The external spin label signal was
quenched with chromium oxalate. Values for leakage into vesicles obtained via ESR
were compared with studies of leakage out of vesicles by the conventional radioactive
tracer method using 99WTc04. Both radioactive tracer and spin label have a negative one
charge and comparable molecular weights of 163 and 169, respectively. They yielded
qualitatively similar results; however, the ESR data exhibited significantly less scatter
of ± 2% vs. ± 5% with the radioactive tracer method. The data exhibit a fast initial
rate of leakage followed by a slow long-term component. This does not fit a simple
Fick's Law process. It suggests a distribution of liposomal diffusion rates or sizes,
thus a nonhomogeneous system. Membrane compositions considered included distearoyl,
Chemistry 171
dipalmitoyl, and dimystearoyl phosphatidylcholines with varying ratios of cholesterol.
ESR is superior to the radioactive method as it decreases scatter, eliminates the hazards
of working with radioactive materials, and also eliminates the difficulties associated
with disposal of the radioactive wastes.
Hindered Ligand Systems: Structure of the c/s,ra-l,3,5-Tris(pyridine-2-carboxaldimine)
cyclohexane Complexes of Fe(II) and Ni(II) Ions. J.C. Huffman, R.A.D. Wentworth,
W.E. Streib and C.J. Huffman, Molecular Structure Center, Department of Chemistry,
Indiana University, Bloomington, Indiana 47405. Structures of the perchlorate
salts of the title compounds have been determined by single crystal X-ray crystallography.
The Fe(II) complex is monoclinic, space group Cc with a = 17.230(10), b = 9.729(4),
c = 16.061(7)A, beta - 104.79(2)° (at -130 C); Dcak = 1.662 gm/cm3 for Z = 4.
The Ni(II) complex is cubic, space group P2,3 with a = 14.337(6)A (at 20 C.) and
D ! = 1 .474 gm/cm3 for Z = 4. The Fe(II) complex has approximate octahedral coor-
dination while the Ni(II) complex lies intermediate between the trigonal prismatic coor-
dination previously found for the Zn(II) and Co(II) complexes and that of the Fe(II)
ion. A detailed comparison is made of the inter- and intramolecular distortions pre-
sent in this unusual system.
Robots in the Chemistry Laboratory, Part I: A High Speed RS-232C Serial Communica-
tions Link for Controlling a HERO I Robot from an Apple II Plus Microcomputer.
Nathan E. Kastelein, Phillip E. Klunzinger, Edward J. Ciesla, Claudia Rishaw,
Cynthia L. Roth and Stanley L. Burden. Digital Equipment Corporation, St. Louis,
Missouri and the Department of Information Science, Physics and Chemistry, Taylor
University, Upland, Indiana 46989. Small table-top robots are beginning to be
used in the analytical chemistry laboratory to minimize routine and mundane sample
preparation tasks. Although its capabilities are not well-suited for many laboratory
tasks, the low price and variety of sensors on the HERO I robot make it attractive
for introducing laboratory robotics to undergraduate students. As the first phase of
developing an instructional laboratory robotics system, a method of minimizing the
constraints of limited memory and inconvenient programming associated with the HERO
I was needed. To overcome these limitations, an RS-232C communications interface
which runs at 9600 baud has been designed and installed in a HERO I robot. The
interface allows the robot to be controlled via a single RS-232C cable from any com-
puter with RS-232C output capability. The interface and associated cable can be con-
structed for approximately $60. In our application, an Apple II Plus microcomputer
was used to write programs and then download them, one command at a time, to
the robot over the serial communications line. This presentation will focus on the hard-
ware of the communications link. The initial phase of a robotized solution preparation
system was implemented as the first application of this system.
Robots in the Chemistry Laboratory, Part II: Software for Controlling a HERO I
Robot from an Apple II Plus Microcomputer via a High Speed RS-232C Communica-
tions Link. Nathan E. Kastelein, Phillip E. Klunzlnger, Edward J. Ciesla, Cynthia
L. Roth, Claudia Rishaw and Stanley L. Burden. Digital Equipment Corporation,
St. Louis, Missouri, and the Department of Information Science, Physics and Chemistry,
Taylor University, Upland, Indiana 46989. Software has been written for both
the HERO I robot and the Apple II Plus microcomputer which enables programs to
be written on the Apple and downloaded to the robot over an RS-232C line operating
at 9600 baud. A short assembly language routine which is loaded into the robot memory
from a cassette tape enables the more lengthy command receiving and interpreting
172 Indiana Academy of Science Vol. 94 (1985)
routines to be downloaded frcm the Apple. To activate the robot, commands are sent
from the Apple to the robot as a sequence of escape characters which indicate the
appropriate motor to activate as well as the direction, speed and extent of movement.
Commands are sent and executed by the robot individually. Typical command formats
and capabilities will be discussed. Software was also written for the HERO I robot
which will accept these escape sequences, acknowledge to the Apple the reception of
the command and activate the appropriate motor. Since robot movement is much slower
than transmission time plus interpretation time, the next interpreted command is always
available before the robot needs it and the robot motion occurs just as if all of the
commands were residing in the on-board robot memory. This presentation will focus
on the software involved in this system.
Reaction Sequence Alteration in the Acetoacetic Ester Synthesis of Ketones. Richard
A. Kjonaas, Department of Chemistry, Indiana State University, Terre Haute, Indiana
47809. The acetoacetic ester synthesis of ketones is a very important method not
only of making ketones, but also of making new carbon-carbon bonds. This classical
method and its well known modifications, such as the use of j8-ketosulfoxides and
/3-ketosulfones, require hydrolysis, saponification, aluminum amalgam reduction, or
other such treatment to remove the stabilizing group after alkylation has been achieved.
Thus, these methods cannot be used with substrates that are sensitive to these post-
alkylation treatments. We have found at ISU, however, that deprotonation of acetoacetic
acid with two equivalents of base, followed by alkylation, gives an adduct which readily
decarboxylates in situ to give good yields of methyl ketones. This method, which is
essentially an alteration of the reaction sequence of the acetoacetic ester synthesis of
ketones, provides a way of achieving the same goals as that synthesis but without carrying
the organo halide moiety through harsh reaction conditions.
Functionalized Crown Ethers. LeRoy Kroll and Bruce Storhoff, Ball State Univer-
sity, Muncie, Indiana 47306. The previously reported 15-crown-5 ethers functionaliz-
ed with -CH2OH or -CH20-CH2CH =CH2 have been studied as potential starting
materials for phosphine- and phosphinite-crown ethers. In accord, the reaction of
15-crown-5-CH2OPPh2 which has been identified by spectroscopic measurements. In
contrast, the corresponding phosphine derivative, 15-crown-5-CH2PPh2 has yet to be
identified from the reaction between 15-crown-5-CH2OTs and Ph2P~. The reaction
of 15-crown-5-CH2OCH2CH = CH2 with 9-BBN has also been investigated. This yields
a surprisingly water soluble compound which has been tentatively identified as the
corresponding alcohol, 15-crown-5-CH2OCH2CH2CH2OH.
A Trace Metal Analysis of Coal and Acid Rain. Steve Newnam and James P.
Rybarczyk, Department of Chemistry, Ball State University, Muncie, Indiana
47306. In an attempt to determine the origins of acid precipitation collected in
Central Indiana, a thorough trace metal analysis was performed on the samples pro-
vided by volunteer members of the Central Indiana District 656 of the International
Rotary Club. The 2000 samples from this study were analyzed for such metal ions
as Ca, Mg, Na, K, Fe, V, Mn, etc., on a graphite furnace atomic absorption system
and an inductively coupled plasma. The ratios between the amounts of these various
trace metals found in the samples were found to be related to seasonal, geographical,
and pH variations and thus provide an indirect means of "tracing" the source of the
metals. With the volunteer assistance of numerous electric power companies, coal samples
were obtained from various geographic Midwest locations. These coal samples were
thoroughly digested in a modified procedure and then analyzed for their trace metal
Chemistry 173
content. Utilizing the variable geographical and meteorological data for the acid rain
samples, the metal ratios from the coal samples were compared to those measured
from the acid rain.
Conclusion of Acid Rain Monitoring in Central Indiana. Laura Pokorney and James
P. Rybarczyk, Department of Chemistry, Ball State University, Muncie, Indiana
47306. With the volunteer assistance of the Central Indiana District 656 of the
International Rotary Club, thirty precipitation collection stations have been in opera-
tion for the past two years. This collection process has just been concluded with over
2000 samples being analyzed for pH, conductivity, depth, trace metal ion and anion
concentrations. These data have been related to the various meteorological, geographical,
and seasonal conditions at the time of collection. The resulting statistical data base
has revealed definite trends in acid precipitation within Central Indiana, with an overall
volume-weighted average pH of 3.9 for the two-year precipitation study. Several in-
dividual events have been recorded in the highly acidic pH = 2.8 to 2.0 region.
One of the most obvious effects of acid rain in Central Indiana is structural damage.
In conjunction with the above monitoring study, carefully-controlled laboratory weather-
ing studies of Indiana limestone have been conducted. Various types of Indiana limestone
were subjected to pH = 4.0 and 3.0 simulated rain, and the results quantitatively
monitored.
Atomic Polarizations of Transition Metal /ra-3-Pentanedionates. Eugene Schwartz,
Department of Chemistry, DePauw University, Greencastle, Indiana 46135. Results
are presented for the radiofrequency and electronic polarizations of the
^m-3-pentanedionates of vanadium(III), manganese(III), and ruthenium(III) in benzene
solution. The atomic polarizations (the difference between the radiofrequency and the
visible frequency or electronic polarization) for these compounds are, for vanadium(III),
manganese(III), and ruthenium(III), 39 cc, 81 cc, and 35 cc, respectively. The atomic
polarizations for the //7S-3-pentanedionates of the series vanadium(III) through cobalt(III)
peak at manganese(III). The second transition series compound /n'5-3-pentanedionato-
ruthenium(III) has an atomic polarization considerably smaller than its iron(III) analogue.
These results are discussed in terms of contributions to the atomic polarization arising
from absorptions in the high frequency microwave region and in the far-infrared spec-
tral region.
Temperature Dependent Infrared Studies of the Hydrogen Bonding in Aliphatic Alcohols.
John Scircle and Joe Kirsch, Department of Chemistry, Butler University,
Indianapolis, Indiana 46208. The extent of the hydrogen bonding of aliphatic
alcohols in dilute solutions has been studied through the use of temperature dependent
infrared spectroscopy. The equilibrium constants for the hydrogen bonding process
have been determined as a function of temperature. The enthalpy and entropy for
the hydrogen bonding process has been calculated from the temperature dependent
equilibrium constants. The equilibrium constants, enthalpies, and entropies for the
hydrogen bonding process are related to the molecular structure of the alcohols and
the steric hindrance at the hydrogen bonding site.
A study of the Coordination Compounds of Some of the Transition Metals Using
2(2-Aminoethoxy)-Ethanol as a Ligand and l-Methyl-2-Pyrrolidinone as a Solvent.
Joseph R. Siefker and Kenneth R. Kimmerle, Department of Chemistry, Indiana
State University, Terre Haute, Indiana 47809. The purpose for this study was
to investigate the coordination compounds of some of the transition metals using
174 Indiana Academy of Science Vol. 94 (1985)
2(2-aminoethoxy)-ethanol as a ligand and l-methyl-2-pyrrolidinone as a solvent. The
formation of a coordination compound in solution is an equilibrium process. For metal
complexes the central elements of coordination, the transition metal ions, are surrounded
by the coordinating groups or ligands, which at first are the solvent molecules but
later are replaced by the ligand molecules. The formation of the complex may be
represented by the equation:
M + nL r ML
n
and the corresponding formation constant is:
Kf = [MLn]
[M] [L]n
Two independent spectrophotometric methods were used to measure the variation of
the metal ion and metal complex concentrations during the formation of the complex.
Job's method of continuous variations as modified by Vosburgh and Cooper was the
major method used to determine the coordination formula and the formation constant
for the complex. The second method, called the mole ratio or fixed metal method,
was tested for its applicability to this case and was used to give a check for the results
of Job's method. The spectra were recorded with a Cary Model 14 Spectrophotometer.
The coordination formulas and average formation constants from Job's method are
presented below: Nickel (II) Perchlorate and 2(2-Aminoethoxy)-Ethanol Complex
Formula = [Ni(2(2-Aminoethoxy)-Ethanol)2](C104):
Kf = 6.5 104
Copper (II) Nitrate and 2(2-Aminoethoxy)-Ethanol Complex
Formula = [Cu(2(-Aminoethoxy)-Ethanol)2](N03)2
Kf = 2.2 x 105
Cobalt (II) Perchlorate and 2(2-Aminoethoxy)-Ethanol Complex
Formula = [Co(2(2-Aminoethoxy)-Ethanol)3](C104)2
Kf = 1.9 x 1010
Manganese (II) Perchlorate and 2(2-Aminoethoxy)-Ethanol Complex
Formula = [Mn(2(2-Aminoethoxy)-Ethanol)3](C104)2
Kf = 1.5 x 10"
An Investigation of Aluminum Concentrations in Water. Daniel K. Wunderlich,
Department of Science, Terre Haute South High School, Terre Haute, Indiana 47802
and Myong-Ku Ahn, Department of Chemistry, Indiana State University, Terre Haute,
Indiana 47809. In this project we investigated the amount of aluminum obtainable
from water solutions under conditions similar to those used in cooking with aluminum
Chemistry 175
utensils. The aqueous aluminum concentrations were examined as a function of exposed
surface area and pH. The aluminum from sample runs was complexed with
8-hydroxyquinoline and concentrated by extracting with chloroform. The concentra-
tions of aluminum were determined by the molecular fluorescence of the complex,
tris(8-hydroxyquinolato)aluminum(III), at 509 nm. The total aluminum concentrations
ranged from 0.38 to 1.9 ppm for the aluminum surface area between 125 cm2 and
625 cm2, respectively, in 200 ml of distilled water. Aluminum levels in the diet have
recently been suspected of being connected with health hazards including senile dementia
and osteomalacia.
Sensitivity Studies of a Computer Model for the Peroxidase-oxidase
Oscillating Reaction
Christopher L. Bush and Raima M. Larter
Department of Chemistry
Indiana University-Purdue University at Indianapolis
Indianapolis, Indiana 46223
Introduction
The peroxidase-oxidase enzyme catalyzed reaction is considered to have the general
form
02 + 2 YH2 - 2 H20 + 2 Y
where YH2 is a hydrogen donor such as nicotinamide adenine dinucletide (NADH).
In an experiment performed by Olsen and Degn (3) a continuous flow of NADH was
pumped into a reaction mixture containing the peroxidase-oxidase enzyme. A continuous
flow of 02 was supplied by blowing a mixture of nitrogen and oxygen over the sur-
face. When the reaction mixture was stirred, oscillations in the concentrations of NAD
(the oxidized product) and 02 were observed.
Method
A model for the peroxidase-oxidase reaction has been proposed by Olsen and
Degn (3). The essential steps in the mechanism are:
K,
A + B + X - 2X
K2
2 X - 2 Y
K3
A + B + Y-2X
K4
X - P
K5
Y - Q
K6
X_ -* X
K7
A - A
o —
K-7
K8
B0 - B [1]
177
178 Indiana Academy of Science Vol. 94 (1985)
where A is [02], B is [NADH], and X and Y are intermediate free radicals. The rate
constants K,-K8 and initial concentrations A0 and B0 were chosen by comparing com-
puter simulations to experiment. The computer simulations involve the numerical solution
of
dA
dt
dB
dt
dX
dt
dY_
dt
-K,ABX - K3ABY + K7A0 - K_7A
-K,ABX - K3ABY + K8B0
-K,ABX - 2 K2X2 + 2 K3ABY - K4X + K6
2 K2X2 - K3ABY - K5Y [2]
When the constants K,-K8, A0 and B0 are chosen appropriately,1 the solutions to [2]
are found to be oscillating functions of time, t.
The validity of this model was tested using sensitivity analysis. Sensitivity analysis
provides several different kinds of information about the changes in the solution of
a set of differential equations due to changes in the values of its parameters. For this
model, it will allow us to determine which rate parameters affect the oscillation's
characteristics, such as its period. The sensitivity analysis yields sensitivity coefficients
which are gradients of the limit cycle in parameter space. The general form of equa-
tion [2] is:
dC/dt = RjtC,..., CN, a,,..., aj i = 1,..., N [3]
where C is the concentration of the species i, R is an algebraic function describing the rate
of change of C due to chemical reactions, and a ,,..., a are parameters such as rate con-
stants. The solution to equation [3] for an oscillatory mechanism may be written as
00
Cj(t) = ^ [ a!n cos 7ml + b!n sin 2nZl
T T
n = 0
,1 ^„A kl
[4]
where t is the period of the oscillation, and r, an and b^ are all functions of the system
parameters.
Differentiating equation [4] with respect to a paramater ex. gives the following ex-
pression for the sensitivity coefficients:
dci <t\ .. 2n7rt dr V^ L0i cJr1 2nxt
(t) = 2n^t p_ y [^ sin 2mrt . nbi 2mrt
T^ da; W L n 7 n r
n = 0
dcc-}
00
+
n = 0 dai T da]
[5]
Larter, Rabitz and Kramer (2) have shown that [5] reduces to:
dC\ -t dr dC'\
daj " t daj dt
Chemistry 179
(§-),«
where the subscript r on the second term indicates that the period, r, is constant for
that term. The first tern consists of a linear function of time [t/ridr/da )] multiplied
by a periodic function (dC /dt). This equation has also been derived by Tomovic and
Vukobratovic (5). Unless dr/da] = 0, dC'/da. will grow as an undamped oscillation as
t get large. This means that dC'/da- gives us physically meaningful result since it becomes
infinitely large as t progresses.
In constrast, the modified sensitivity coefficient, (dC /dojMt), does give physically
meaningful results since it is well-defined and periodic for a differential equation system
that is structurally stable to perturbation of aj. Since the second term is purely periodic,
the values will be consistent as t progresses. In order to solve for this second term, the
period sensitivity, dr/daj must be determined. First, the sensitivity coefficients in equa-
tion [6] are integrated from t to t + r and evaluated at two different times, t, i.e., t
= t, and t = t2, and then these results are subtracted giving,
dr/da: -
rU + T
I "
t2
dC: p ti . t
t,
aq
daj
J
q (t,) - q (t2)
[7]
Numerical instabilities encountered by Edelson and Thomas ( 1 ) with a similar calculation
were avoided by eliminating choices where the concentration at t, and t2 were approx-
imately equivalent. Now the modified sensitivity coefficient can be found because dC dt
have already been calculated, and t/r can be found by measuring the period, r, with
t already known.
Discussion
The peroxidase-oxidase reaction is an oscillatory reaction with a limit cycle solu-
tion, so we used the above equations to obtain a modified sensitivity coefficient. In
order to oscillate, a reaction must operate far from equilibrium; this system satisfies
this requirement because substrates, A and B, are being continually added. Also, some
product of at step in the reaction sequence must exert an influence on its own rate
of formation; the intermediate free radicals, X and Y, are produced autocatalytically
and thus do influence their own formation.
These two facts seem to indicate the importance of the rate constants for the
addition of the substrates and the autocatalytic steps. We have found this to be true
for the rate constant, K8, which is the rate constant for the addition of NADH. For
most values of K8, the solution is not a true periodic function, and the sensitivity
coefficients do not show unbounded growth with time. These quasi-periodic regions
give some unusual results which appear to include some chaotic regions; Olsen (4)
has also noted these chaotic regions. We intend to investigate this further at a later
date. The solution is truly periodic when K8 is equal to 0.4939; for the periodic solu-
tion, the sensitivity coefficients had the expected time behavior given by equation [6].
Note
1. The values used were K, = 8.5 x 10" 2, K2 = 1.25 x 10\ K, = 4.8675 x 10~2,
K4 = 20.0, K, = 2.0, K6 = 1.0 x 10~\ K7 = .94, K-7 = .1175, K8 = .4939,
A0 = 3.84, Bo = 33.73, X0 = 1.1 x 10~4, Y0 = 3.62 x 10~6.
180 Indiana Academy of Science Vol. 94 (1985)
Literature Cited
1. Edelson, D. and Thomas, V.M. 1981. Sensitivity Analysis of Oscillating Reac-
tions. 1. The Period of the Oregonator. J. Phys. Chem. 85:1555-1558.
2. Larter, R., Rabitz, H. and Kramer, M. 1984. Sensitivity Analysis of Limit Cycles
with Application to the Bursselator. J. Chem. Phys. 80:4120-4128.
3. Olsen, L.F. an Degn, H. 1978. Oscillatory Kinetics of the Peroxidase-Oxidase
Reaction in an Open System; Experimental and Theoretical Studies. Biochim.
et Biophys. Acta 523:321-334.
4. Olsen, L.F. 1983. An Enzyme Reaction with a Strange Attractor. Phys. Lett.
94 A: 454-45 7.
5. Tomovic, Rajko and Vukobratovic, Miomir. General Sensitivity Theory, 1972.
Elsevier, New York.
A SCC MO Calculation on the Tetracyanoethylene-benzene Complex
Joe Kirsch, Shannon Lieb and Mark Cisneros
Department of Chemistry
Butler University
Indianapolis, Indiana 46208
Introduction
The tetracyanoethylene-benzene charge transfer complex is formed through the
interaction of the pi electron density of the benzene and the pi antibonding orbitals
of the tetracyanoethylene. Two sandwich type structures of the complex have been
proposed in the literature (2) and are shown in Figure 1 . Molecular orbital calculations
STRUCTURE
STRUCTURE
^
:.xx
%
H
H. C
C M
H C
C. H
Figure 1. Proposed Structures of Benzene-Tetracyanoethylene Complex.
can be used to determine the nature and energy of the absorption which results from
complex formation. If the calculations are carried out as a function of the benzene-
tetracyanoethylene intermolecular distance, the intermolecular distance that yields the
best agreement with the observed charge transfer energy can be obtained.
Calculations
The MO calculations were carried out on a VAX 11/780 computer. The necessary
input data for the SCC MO program are given in Table 1. The atomic coordinates
are calculated from standard bond lengths and bond angles. These molecular parameters
are also listed in Table 1.
The SCC MO calculations require the evaluation of overlap integrals, coulomb
integrals, and resonance integrals. These integrals are the elements of the secular deter-
minant. Calculation of the overlap integrals, S-, using Slater type atomic orbitals, the
effective nuclear charge, and the atomic coordinates have been described in the literature
(1). Valence orbital ionization energies, VOIE, are used to approximate the coulomb
181
182 Indiana Academy of Science Vol. 94 (1985)
Table 1. Input Data for the SCC MO Program
1 . Total Number of Atoms in the Complex (22)
2. Atomic Number of Each Atom
3. Valence Shell Atomic Orbitals (Carbon and Nitrogen — 2s, 2p , 2p , 2p ; Hydrogen - Is)
4. Total Number of Valence Electrons (74)
5. Charge on Each Atom
6. Electron Population in Each Atomic Orbital
7. Coordinates for Each Atom*
* Bond Type Length (angstroms) Angle (Deg.)
Benzene C-C 1.390
Benzene C-H 1.085
TCNE C = C 1.336
TCNE C = C 1.157
TCNE C-C 1 .450
Benzene C-C-C 120
Benzene C-C-H 120
TCNE C = C-C 120
TCNE C-C = N 180
integrals, H (1). The Wolfsberg-Helmholtz approximation is used to determine the
resonance integrals, H (1). The secular determinant is then solved from these values
of S , H.., H to obtain two sets of MO mixing coefficients (Eigenvectors) and MO
energies (eigenvalues).
The eigenvectors resulting from the calculation are used in a Mulliken population analysis
to calculate new atomic charges for each atom in the complex. These new atomic charges,
output atomic charges, are compared to the input atomic charges. If the input and
the output charges are different, the difference times 0.1 is used as a new input charge;
and the calculation is recycled until the input and output atomic charges converge
and self consistent charges on the atoms are obtained.
Results and Discussion
The benzene-tetracyanoethylene complex has 74 valence electrons and 70 valence
atomic orbitals in its basis set. This basis set and collection of valence electrons will
yield 70 molecular orbitals with the first 37 molecular orbitals being doubly populated
with electrons. The 38th molecular orbital is the lowest unoccupied molecular orbital.
The lowest energy electronic transition, the charge transfer band, will then occur bet-
ween the 37th MO and the 38th MO.
Examination of the eigenvectors for molecular orbitals 37 and 38 show that all
of the atomic orbital coefficients are near zero except those for the pi 2p type atomic
orbitals. This indicates that molecular orbitals 37 and 38 are primarily pi type molecular
orbitals. The values of the eigenvectors of the pi 2p atomic orbitals for molecular
orbitals 37 and 38 are listed in Table 2. The atom numbering system is given in Figure
2 and 3 for each proposed structure. Further examination of the eigenvectors show
that MO 37 is bonding for benzene (C2-C3 and C5-C6), bonding for tetracyanoethylene
(CN groups), and bonding for the complex for both structures. Molecular orbital 38,
however, is antibonding for the tetracyanoethylene part of the complex for both struc-
tures. Finally, it can be noted that the eigenvectors indicate more electron density,
larger values for the eigenvectors, on the tetracyanoethylene for MO 38 than for MO
37 in both structures. In summary, the analysis of the eigenvectors for MO 37, highest
occupied, and MO 38, lowest empty, supports the notion of an electronic transition,
charge transfer, from a benzene pi bonding MO to a tetracyanoethylene pi antibon-
ding MO as a description of the charge transfer band.
Chemistry
183
Table 2. Pi AO Eigenvectors for the Benzene — Tetracyanoethylene Complex
Atom Number*
Structure
A
Structure
B
M037
M038
M037
M038
Benzene
C-l
0
0
0
0
Benzene
C-2
-.36
.38
-.32
-.25
Benzene
C-3
-.36
-.38
-.32
-.25
Benzene
C-4
0
0
0
0
Benzene
C-5
.36
-.38
.32
.25
Benzene
C-6
.36
.38
.32
.25
TCNE
C-7
0
0
.117
.47
TCNE
C-8
0
0
-.07
-.47
TCNE
C-9
.13
-.21
.14
.22
TCNE
C-10
-.13
.21
.14
.22
TCNE
C-l 1
-.13
-.21
-.14
-.22
TCNE
C-12
.13
.21
-.14
-.22
TCNE
N-13
-.29
-.35
32
-.30
TCNE
N-14
.29
.35
.32
-.30
TCNE
N-15
-.29
.35
-.32
.30
TCNE
N-16
.29
-.35
-.32
.30
*Refers to the numbering system in figures 2 and 3
Table 3 shows the energy difference of MO 38 and MO 37, energy of the charge
transfer band, as a function of the benzene-tetracyanoethylene intermolecular distance.
Figures 4 and 5 are plots of this data for each proposed structure. The plots for both
structures show minimum near the observed charge transfer absorption energy for an
intermolecular distance of 2 angstroms, structure A— 386 nm, 2.04 A; structure B— 389
nm, 2.09 A.
STRUCTURE
STRUCTURE
B
N
16
H'
114
'C
9
H
C7 C
C8
C
H
C
-N
15
H Cj^C^^Cii^h
10
H
■N
13
Figure 2. Atom Numbering System for Figure 3. Atom Numbering System for
Eigenvector Analysis in Table 2.
Eigenvector Analysis in Table 2.
184
Indiana Academy of Science
Vol. 94 (1985)
Table 3. E(MO 38) - E(MO 37) versus the Benzene-Tetracyanoethylene Intermolecular
Distance
Structure A
r (angstroms)
1.97
1.98
1.99
2.00
2.04
2.05
2.10
E(MO-38) - E(MO-37) [nm]
413
409
405
401
386
388
403
Structure B
r (angstroms)
2.03
2.04
2.05
2.06
2.07
2.08
2.09
2.10
2.12
E(MO-38) - E(MO-37) [nm]
409
406
402
398
395
391
389
392
398
no
I
O
UJ
00
00
I
a
LxJ
STRUCTURE A
+
410
+
+
+
400
+
390
380
^7n
i i
1 !
r (Angstroms)
Figure 4. Charge Transfer Band Energy vs Benzene — TCNE Intermolecular Distance.
Chemistry
185
m
i
o
:e
LU
I
/">
00
00
I
a
l_Ll
4«:u
STRUCTURE B
410
+
+
400
+ ■+•
+
390
380
i
i i
r (Angstroms)
Figure 5. Charge Transfer Band Energy vs Benzene— TCNE Intermolecular Distance.
Acknowledgments
The authors wish to thank the Holcomb Research Institute and the Butler University
Academic Grants Committee for their funding of this project.
Literature Cited
1. McGlynn, S.P., L.G. Vanquickenborne, M. Kinoshita, and D.G. Carrol, 1972,
Introduction to Applied Quantum Chemistry, Holt Rinehart Winston, New York,
N.Y., 48, 97-140.
2. Mobley, M.J., K.E. Rieckhoff, and E.M. Voight, 1978, Spectroscopic Studies
on the Conformations of Electron Donor Acceptor Complexes of Tetra-
cyanoethylene, J. Physical Chem., 82, 2005-2012.
Spectra and Photochemistry of the Chloro Complexes of Copper(I)
Kristine S. Kurtz and Kenneth L. Stevenson
Department of Chemistry
Indiana University-Purdue University at Fort Wayne
Fort Wayne, Indiana 46805
Introduction
Previous studies (9,8,7,4,1) have shown that in aqueous chloride media, the two
copper(I) complexes shown in the following equation are in equilibrium:
1) CuCl2~ + CI" - CuCl32~
The purposes of this study were to achieve the following measurements of this system
at 5M ionic strength: 1) verification of the equilibrium constant measured by Ahrland
and Tagesson (1), 2) to resolve the ultraviolet charge-transfer-to-solvent (CTTS) spec-
tra of these two complexes, and 3) to determine the quantum yields of the following
photoredox reaction:
2) Cu(I) + H+ = Cu(II) + l/2H2(g)
for each of the two complexes at several wavelengths in the CTTS absorption region.
Procedure
Seven solutions in which chloride ion concentration varied from 0.2M to 5M at
constant ionic strength of 5M and constant hydrogen ion concentration of 1M were
prepared using analytical grade reagents (perchloric acid, sodium perchlorate, sodium
chloride, and hydrochloric acid) and deionized water. Since Cu(I) solutions are readily
oxidized by air, the 0.01 M cuprous chloride solutions were prepared by inserting test
tubes containing preweighed amounts of cuprous chloride into the flasks containing the
solutions, above, and bubble-degassed with argon through septums in the top of the flasks.
The flasks were then tipped and the solid dissolved.
The absorbance spectra of the equilibrated solutions were measured in the 200-340
nm range where the two complexes exhibit CTTS absorption (4). A Beckman ACTA
M-VI spectrometer interfaced with a HP-86 microcomputer allowed the spectra to be
stored on disk for subsequent spectral computations.
The photochemical setup consisted of a Schoeffel 1000-watt mercury-xenon high
pressure arc lamp, a Jarrell Ash quarter meter monochromator, a thermostated 1-cm
cuvette, and a recording gas volumeter (3) for measuring the evolved hydrogen gas.
Light intensities were measured in the cuvette with the potassium trioxalatoferrate (III)
actinometer (5). Volumeter chart traces were digitized and integrated rates determined
with the computer.
Results and Discussion
Figure 1 shows the molar extinction absorption spectra of the seven solutions
of varying chloride ion concentration. The increase in peak absorption at 274 nm with
increase in [CI-] indicates that the trichloro species has a stronger absorption in this
region. Since there are only two complexes in equilibrium in this system, one ca n
show (4) that the measured extinction coefficient is a function of the extinction coeffi-
187
Indiana Academy of Science
Vol. 94 (1985)
CI
o
H
+^
u
c
X
LU
D
O
5000
4000 -
3000 -
2000
f= 1000 -
200
300
320
340
220 240 260 280
Wavolength ( n m )
Figure 1. Molar extinction spectra of 0.01M CuCl in 5M ionic strength medium in
which 0.2<[C1~]<5.0M, at 25°C.
cients of di- and trichloro complexes, e2 and e3, and the equilibrium constant, K, for
equation 1, as follows:
3) e = e3 + (e2 - e3)/(l + K[Cr])
5000
4000 -
o
U 3000
c
L^ 2000
L
O
o
1000 -
200
220
240
260
280
300
320
340
WavG length (nm)
Figure 2. Resolved molar extinction spectra of CuCl2~ and CuCl32~ at 5M ionic
strength, 25°C, assuming K = 0.72.
Chemistry
189
^r
CM
u
Q)
D
+^
C
O
D
a
o. 6
0.5 --
0. 4
0. 3 -
0.2 --
0. 1 --
0.0
[C1-]
Figure 3. Quantum yield of photooxidation of aqueous CuCl versus chloride ion con-
centration, at 274 nm irradiating wavelength, 25°C, 5M ionic strength.
The computer was used to find the value of K which gave the best linear fit of e
vs. 1/(1 + K[C1"]) in the 271 to 281 nm region where the spectra are most sensitive
to chloride ion concentration. This gave an average value of K of 0.72 ± 0.08, which
compares favorably to the value of 0.76 ± 0.07 determined by Ahrland and Tagesson
from electrochemical measurements (1). Using our value of K the computer then
calculated e2 and e3 from equation 3 at all wavelengths, resulting in the resolved spec-
tra for the two complexes, as shown in Figure 2. It is noteworthy that the trichloro
complex has a strong band at 274 nm compared to a weak shoulder somewhat blue
shifted in the dichloro species, whereas both exhibit transitions more nearly equal in
the 230-235 nm region.
Figure 3 indicates that the quantum efficiency of photolysis into the band at 274
nm is depressed by increasing chloride ion concentration. This would imply that the
trichloro species has a lower quantum efficiency than the dichloro complex. One can
show, using Beer's law, that the net quantum yield, <J> , is related to individual quan-
tum yields of the di- and trichloro complexes, <j>2 and (j)3 as follows:
4) <(> = (j)3 +.
(§2 ~ fyl) e2
(e2 + e3K[Cl-])
A suitable linear plot of this function at 274 nm is shown in Figure 4, resulting in
resolved quantum yields for the two complexes. Equation 4 was used for the curve-fit
in Figure 3 using the results of the linear regression. These quantum yields and those
determined at two other wavelengths are shown in Table 1.
These results show that the quantum yields bear an inverse relationship to molar
extinction coefficients, since the trichloro complex, which has the larger absorbance,
has the smaller quantum yield. This may be rationalized by the fact that lifetime of
190
Indiana Academy of Science
Vol. 94 (1985)
E
C
C\J
QJ
E
D
-P
C
D
D
a
U. D
0. 5 -
0. 4 -
^ -Q
0.3 -
jy
0. 2 -
0 /-
0. 1 -
n. n -
1
1
H 1 1
0.00
0. 10
0.20
0. 30
q2/ Ce2 + e3K [CI ] )
Figure 4. Quantum yield of photooxidation of aqueous CuCl versus the function,
e2/(e2-e3K[Cl~]), at same conditions as Figure 3.
any excited state is inversely proportional to the oscillator strength (2). The dichloro
species, which has the lower absorbance, and hence lower oscillator strength, would
have a longer excited state lifetime, thus resulting in a greater probability for the hydrated
electron produced by the photolysis to be scavenged by hydrogen ion (6). Further
experiments are in progress, with the intent of discerning similarities or contrasts with
spectral and photochemical properties of other halo complexes, and what role
sterochemistry plays in these properties.
Acknowledgment
Acknowledgment is made to the donors of the Petroleum Research Fund,
administered by the American Chemical Society, for support of this research.
Literature Cited
1. Ahrland, A., and Tagesson, B. 1977. Thermodynamics of Metal Complex For-
mation in Aqueous Solution. XII. Equilibrium Measurements on the Copper(I)
Bromide, Iodide and Thiocyanate Systems. Acta. Chem. Scand., A31(8):615.
2. Calvert, J.G. and Pitts, Jr., J. N. Photochemistry. John Wiley & Sons, Inc.,
New York, 1966, 173-174 pp.
Table 1: Quantum Yields
Wavelength (nm)
Quantum Yield
265
274
296
CuCh
CuCl,2
1.30 +-0.12
0.178 + -0.010
1.62 +-0.28
0.144 + -0.010
1.47 + -0.25
0.275 + -0.010
Chemistry 191
3. Davis, D.D., and Stevenson, K.L. 1977. A Recording Gas Microvolumeter. J.
Chem. Educ., 54: 394.
4. Davis, D.D., Stevenson, K.L., and Davis, C.R. 1978. Photooxidation of Dichloro-
and Trichlorocuprate(I) Ions in Acid Solution. J. Amer. Chem. Soc, 100(17): 5344.
5. Hatchard, C.G., and Parker, L.A. 1956. A New Sensitive Chemical Actonometer
II. Potassium Ferrioxalato as a Standard Chemical Actinometer. Proc. Roy. Soc.
London, A235: 518.
6. Stevenson, K.L., Kaehr, D.M., Davis, D.D., and Davis, C.R. 1980. Long-Lived
Intermediates in the Production of Hydrogen from Ultraviolete Photolysis of Acidic
Di- and Trichlorocuprate(I) Ions. Inorg. Chem., 19(3): 782.
7. Sugasaka, K. and Fujii, A. 1976. A Spectrophotometry Study of Copper (I) Chloro-
Complexes lin Aqueous 5M Na(Cl,C104) Solutions. Bull. Chem. Soc. Japan, 49(1):
82.
8. Sukhova, T.G., Temkin, O.N., and Flid, R.M. 1970. Electronic Absorption Spectra
of Chloro-complexes of Univalent Copper in Aqueous Solution. Russ. J. Inorg.
Chem., 15(7): 949.
9. Sukhova, T.G., Temkin, O.N., Flid, R.M., and Kaliya, T.K. 1968. Determina-
tion of the Composition and Stability Constants of Chlorocuprate (I) Complexes
in Concentrated Solutions. Russ. J. Inorg. Chem., 13(8): 1072.
Evaluation of Sample Pre-treatments as Potential Methods of Enhancing
Phospholipid Extraction from Human Amniotic Fluid
Barth H. Ragatz, Gina Modrak and Ericka Baeske
Fort Wayne Center for Medical Education
Indiana University School of Medicine
Indiana University-Purdue University at Fort Wayne
Fort Wayne, Indiana 46805
Introduction
It is well known that phospholipids present in human amniotic fluid have been
transferred from the fetal lung compartment to amniotic fluid. Furthermore, these
phospholipids are components of pulmonary surfactant, a fluid necessary for normal
lung physiology in neonates. These phospholipids necessary in surfactant are synthesized
by the Type II alveolar cells and include dipalmityl phosphatidyl choline (lecithin),
sphingomyelin, and phosphatidyl glycerol. The relative levels of sphingomyelin are known
to remain rather constant throughout gestational development, but the levels of both
lecithin and phosphatidyl glycerol increase dramatically beyond week 28 of intrauterine
life. If the phospholipids are extracted from amniotic fluid and chromatographed, it
is possible to predict that normal fetal lung development is occurring when
lecithin/sphingomyelin ratios are greater than 2.0 and when phosphatidyl glycerol is
also detected among the chromatographically resolved spots (6).
Amniotic fluid is a complex analytical matrix composed of water, dissolved salts,
various proteins, cholesterol and other neutral lipids and several kinds of phospholipids.
We decided to evaluate various pre-treatment methods to determine if we could release
more phospholipid from protein binding sites to enhance the extraction of the three
principal phospholipids into chloroform-methanol, and to avoid the emulsions sometimes
generated when amniotic fluid samples are extracted with chloroform-methanol. Potential
pre-treatments could involve the quantitative destruction or removal of undesired com-
ponents from the mixture (protein, cholesterol, or neutral lipids) or enhancement of
the extractibility of the three principal phospholipids into chloroform-methanol (3).
We have evaluated three pre-treatment procedures for this purpose: addition of am-
monium sulfate to alter the activity coefficient of water and permit quantitative removal
of protein components from the analytical matrix; pre-extraction with various non-
polar organic solvents to quantitatively remove cholesterol and/or neutral lipids from
the analytical matrix; or adjustment of the amniotic fluid pH to either acid or alkaline
extremes to alter the partition coefficient of the principal phospholipids into chloroform-
methanol by modification of ionization states of the principal phospholipids.
Materials and Methods
Frozen human amniotic fluid samples were obtained from Parkview Memorial
Hospital, Fort Wayne, Indiana and from University Hospital, Indianapolis, Indiana.
These samples were thawed, pooled and refrozen in 4 ml. aliquots. All samples were
stored at -20°C. Only those samples stored for periods less than nine months were
used and samples obviously contaminated with blood or meconium or heme pigments
were routinely discarded. Ammonium sulfate (A-5132) was obtained from Sigma
Chemical Company, St. Louis, Missouri and was added as various dry powered in-
crements to 4 ml. amniotic fluid samples which had been thawed to room temperature.
After thorough mixing with the ammonium sulfate, the samples were centrifuged at
1000 RPM for five minutes in a Clay-Adams centrifuge and extracted with chloroform
193
194 Indiana Academy of Science Vol. 94 (1985)
and methanol according to the Helena Fetal Tek 200 Procedure (1). The remaining
steps in determination of L/S ratio and detection of phosphotidyl glycerol were according
to the Helena Fetal Tek 200 method also.
For organic solvent pre-extraction tests, ACS reagent grade Matheson, Coleman
and Bell reagents were used, including hexanes (HX 299), ethyl acetate (EX 240), benzene
(BX 220), and tricholoroacetic acid (TX 1045). For organic solvent pre-extractions,
a 4 ml. sample of thawed amniotic fluid was placed in a 15 ml. liquid scintillation
counting vial. The appropriate organic solvent was added in three separate portions
of 3 ml. each. The vial was shaken after each addition and the top organic reagent
layer was withdrawn by a Pasteur pipet. After the third extraction was completed,
a 2 ml. sample of amniotic fluid was drawn off with a measuring pipet from the bottom
aqueous layer and again submitted to the Helena Fetal Tek 200 Procedure. An un-
treated sample was also used in the Helena method to serve as a control.
For trichloroacetic acid pre-treatment, a 4 ml. amniotic fluid sample was mixed
with 8 ml. of chilled trichloroacetic acid. After precipitation had occurred, the sample
was transferred to centrifuge tubes equipped with Bio Analytical Systems filters. Tubes
loaded with 1.5 ml. samples were centrifuged at 2000 RPM's for ten minutes. Since
filters became clogged with precipitates, it was often necessary to transfer partially
clarified liquid to fresh centrifuge-filter apparatus and repeat the centrifugation step
a second time. The combined filtered solutions were adjusted back to pH 7 using 2
M. sodium hydroxide and glacial acetic acid. The resultant aqueous sample was sub-
mitted to the Helena Fetal Tek 200 Procedure.
Finally, amniotic fluid samples were adjusted to extremes of pH using 1 M. sodium
phosphate, analytical reagent grade, supplied by Mallinckrodt, Inc. The pH altered
samples were prepared as usual by the Helena Fetal Tek 200 method.
Results
The effect of ammonium sulfate precipitation of amniotic fluid samples on the
L/S determination can be seen in Table 1. Four values are reported at each treatment
level on four aliquots of the pooled amniotic fluid. It can be seen when ammonium
sulfate is added in amounts which would bring the saturation of water from 20% to
80% that no alteration of the L/S ratio occurs. It was also noted that huge amounts
of protein were precipitated, even at the lowest level of ammonium sulfate addition.
Although the ammonium sulfate pre-treatment does not negatively influence the
extraction of lecithin and sphingomyelin, there is no enhancement of phosphatidyl
glycerol from the altered analytical matrix. Thus ammonium sulfate pre-treatment would
be of no value in the present case.
Table 1. Effect of Ammonium Sulfate Precipitation of Amniotic Fluid on L/S
Determination
Amount (NH4);S04 added
to 4 ml sample L/S Ratio
untreated — ; 1.2
0.56g 1.1; 0.7
1.13g 1.6; 0.8
1.69g 1.2; 0.9
2.26g — -; 1.1; 1.2
1.2
1.2
1.2
1.1
1.2
Chemistry 195
Table 2. Effect of Organic Solvent Pre-Extraction of Amniotic Fluid on L/S
Determination
Organic Solvent L/S Ratio
None 0.9
Benzene 0.8
Ethyl Acetate 0.7
Hexanes 0.8
1.1
1.0
0.6
0.8
1.1; 0.7
0.9; 0.5
0.7; 0.5
1.0; 0.7
Trichloroacetic Acid ppt. formed
Table 2 shows the effects of various organic solvent pre-extractions of amniotic
fluid upon the L/S determination. Once again, benzene and hexane are without effect
on the determined L/S ratio and no additional phophatidyl glycerol was extracted into
the chloroform-methanol treatment of the Helena Fetal Tex 200 Procedure. No attempt
has been made to examine the extent to which cholesterol or neutral lipids may have
been removed by organic solvent extraction. Table 2 suggests that ethyl acetate pre-
extraction lowers the L/S ratio which is calculated. Examination of the densitometer
scans show clearly that ethyl acetate differentially removes lecithin in the pre-extraction
phase. Trichloroacetic acid treatment yields a copious precipitate which apparently traps
phospholipids quantitatively in the precipitating mixture. We have concluded that the
organic solvent pre-extractions examined at present are of no value in the pre-treatment
of human amniotic fluid for the enhancement of phospholipid removal by the Fetal
Tek 200 Procedure.
Table 3 shows the effect of extreme pH adjustment of amniotic fluid before
extraction on the determined L/S ratios. It was seen that there is no significant altera-
tion of the L/S ratio by adjustment of amniotic fluid pH to either pH2 or pH12.
Once again no enhancement of phosphatidyl glycerol extraction into the chloroform-
methanol occurred.
Discussion
Although undesirable contaminants are often removed from an analytical matrix
by pre-precipitation treatments or by pre-extraction, it is clear that the three approaches
presently reported have not been of value in enhancing the removal of phospholipids
from human amniotic fluid. In the past, Gluck et al. have reported that acetone pre-
treatment was useful in enhancing the extraction of lecithin and sphingomyelin (4).
More recently, other reports suggest that acetone pre-treatment is useless (5). A recent
Table 3. Effect of pH Adjustment of Amniotic Fluid (before extraction) on L/S
Determination
Adjusted pH of Sample L/S Ratio
1. 2M sodium phosphate, pH2 1.4
1.3
1.6
1.7
2. 2M sodium phosphate, pH12 3.1
1.3
1.5
1.4
196 Indiana Academy of Science Vol. 94 (1985)
report by Duck-Chong et al. compared various methods of extracting phospholipids
from human amniotic fluid. These authors noted that various pre-treatment exposure,
such as extraction with chilled solvents or chromatography of single phased mixtures
over Sephadex G25 columns caused decreases from 15 to 40% in the total phospholipid
originally present in the sample (2).
This report dramatizes the fact that whatever procedure one uses in determining
L/S ratios, it is necessary to rigorously standardize all conditions and to develop local
standards for fetal lung maturity by comparing statistically the results obtained on
many patient samples with the general health observed in the neonate during subse-
quent postpartum follow-up. We are continuing to evaluate other pre-treatment methods
in our laboratory with the hope of realizing this desired goal of enhancing the removal
of lecithin, sphingomyelin and phosphatidyl glycerol into chloroform-methanol extraction
mixture.
The authors wish to acknowledge the careful preparation of this manuscript by
Ms. Elaine Wilson.
Literature Cited
1. Anonymous. 1982. Helena Fetal Tek 200 Method. Helena Laboratories, Beau-
mont, Texas, p. 1-6.
2. Duck-Chong, C.G., G.J. Baker, S.R. Murdoch and R.M. Price. 1984. Methods
for Extracting Phospholipids from Human Amniotic Fluid Compared. Clin. Chem.
30:271-274.
3. Giese, R.W. 1983. Technical Considerations in the Use of "High-Performance"
Liquid Chromatography in Therapeutic Drug Monitoring. Clin. Chem.
29:1331-1343.
4. Gluck, L. 1971. Diagnosis of Respiratory Distress Syndrome by Amniocentesis.
Am. J. Obstet. Gynecol. 109:440-445.
5. Hill, E.H. 1979. Comparison of Foam Stability Index and Lecithin/Sphingomyelin
Ratio in Amniotic Fluid and Its Predictive Value for Fetal Lung Maturity. Clin.
Chem. 25:1138.
6. Gluck, L. 1978. Evaluating Functional Fetal Maturation. Clin. Obstet. and Gynecol.
21:547-559.
Comparison of Two Simple Methods for Determining
Lecithin/Sphingomyelin (L/S) Ratios in Human Amniotic Fluid Samples
Barth H. Ragatz, Gin a Modrak and Patricia S. Conn
Fort Wayne Center for Medical Education
Indiana University School of Medicine, and
Department of Mathematical Sciences
Indiana University-Purdue University at Fort Wayne Campus
Fort Wayne, Indiana 46805
Introduction
The ability of a neonate to survive after delivery depends largely on proper develop-
ment of its respiratory system. If the Type II alveolar cells are incapable of synthesiz-
ing proper pulmonary surfactant, there is a high probability that the neonate will have
respiratory distress syndrome. Proper pulmonary surfactant is rich in phospholipids,
especially dipalmityl phosphatidyl choline (lecithin) and phosphatidly glycerol (PG).
The latter phospholipid is present in the pulmonary surfactant in tenfold smaller molar
quantities. Since the maternal amniotic fluid is in direct contact with the fetal lung
compartment during gestation, the lung phospholipids are readily transferred and
reflected in relative abundance in the amniotic fluid (10,2,5).
Analytical studies of phospholipids present in amniotic fluid have shown that
the relative levels of sphingomyelin remain relatively constant throughout gestational
development. However, the amounts of lecithin and phosphatidyl glycerol are relatively
low throughout gestation until approximately week 28 when both of these compounds
begin to increase in relative levels asymptotically. If an amniotic fluid sample is carefully
taken by amniocentesis before delivery, it is possible to remove contaminating cells
and to extract the phospholipids from the amniotic fluid. The phospholipids can be
collected in a cholorform layer, concentrated, and subjected to thin layer chromatography
for resolution. After separation, some detection method can be employed and the relative
amounts of sphingomyelin, lecithin, and phosphatidyl glycerol can be detected either
by visualization of a developed color or by a developed fluorophore (10). The detected
phospholipid spots can be measured visually or with a scanning densitometer. If an
L/S ratio greater than 2.0 and the presence of phosphatidyl glycerol are detected it
can be concluded with some certainty that fetal lung development is normal. If a smaller
L/S ratio or absence of phosphatidyl glycerol in the amniotic fluid sample is detected,
it is possible to give fetal retentive drugs and to enhance the synthesis of pulmonary
lecithin and phosphatidyl glycerol by administration of Cortisol to the mother (5).
Various detection systems have been utilized in the past to reveal phospholipids
resolved by thin layer chromatography. These detection reagents have been ionization
sensitive, unsaturation sensitive, or phosphate sensitive. In recent years the Helena
Laboratories (Beaumont, Texas) copper acetate reagent (8000) has enjoyed much
popularity as an unsaturation sensitive detector. Using 42 human amniotic fluid samples,
we have compared the results obtained for L/S ratios by the copper acetate charring
detection method with the use of anilino -1,8- naphthalene sulfonate (ANS) detec-
tion. We have used this latter reagent because of its reported ability to form fluores-
cent complexes with phospholipids and yield similar fluorescence intensities. In earlier
studies in our laboratories we have also found anilino - 1, 8 - naphthaline sulfonate
to be especially sensitive in detecting six phospholipid standards and we found the
response for spot size to concentration load to be linear for most of the phospholipids
over a 100 fold concentration range (7).
197
198
Indiana Academy of Science
Vol. 94 (1985)
Materials and Methods
Purified phospholipid standards (lecithin, sphingomyelin, phosphatidyl glycerol)
were purchased from Sigma Chemical Company, St. Louis, Missouri, and were dissolved
in chloroform at concentrations of two milligrams per milliliter. Glass plates (20 cen-
timeter X 20 centimeter) were coated with a slurry prepared by dissolving Silica Gel
G (Brinkman 7731) 40 grams in 90 milliliters of demineralized water. The silica gel
was coated with a Brinkman apparatus. Plates were air dried at room temperature
overnight and received no additional activation at higher temperatures. Frozen amniotic
fluid samples were thawed to room temperature and two ml. aliquots were placed in
60 ml. separatory funnels. Two ml. of 100% methanol was added and the mixture
was shaken for 20 seconds. Two ml. of chloroform was added and the mixture was
again shaken for 20 seconds. Each emulsified sample was centrifuged in a Clay Adams
Clinical Centrifuge at 2000 RPM's for ten minutes. A Pasteur pipet was used to carefully
remove a one ml. aliquot from the lower chloroform layer. The organic extract was
evaporated to dryness under a stream of nitrogen and was reconstituted with approx-
imately 40 microliters of chloroform.
This entire extract could be placed on a thin layer plate and chromatographed
along with ten microliter samples of the respective phospholipid standards. The plates
were developed in a solvent system containing 68 ml. of chloroform, 28 ml. of methanol,
and 4 ml. of 30% ammonium hydroxide, in a Sigma thin layer chromatography chamber.
Detection of the resolved phospholipids was accomplished by direct visualization follow-
ing spraying with the Helena Laboratories copper acetate spray reagent and heating
to 120°C for ten minutes. Alternatively, a similar aliquot from a given patient amniotic
fluid extract was detected by spraying a solution of 50 milligrams of ANS dissolved
in 100 ml. of methylene chloride. Detection of the resolved spots was obtained with
a Ultra-Violet Products UVSL-25 lamp. Spots were quickly circled with pencil and
a quantity proportional to spot area was calculated by multiplying horizontal diameter
by vertical diameter of each spot. Finally the ratio of lecithin spot area to sphingomyelin
spot area was calculated. The sub-populations of 42 patient samples detected by these
two methods were statistically compared using the Students' t test.
Results
The statistical data obtained from the Students' t test is shown in Table 1. The
result of comparing the two population means with each other statistically indicates
in a two-tail probability that the two methods give statistically significantly different
results. Of course this statistical parameter does not indicate that one method is better
than the other, merely that the two sample populations are statistically different (4).
Table 1: Comparison of L/S Ratios in Amniotic Fluid Determined by Two Detection
Methods on 42 Patient Samples.
ANS Fluorescence Detection Method
Cupric Acetate Charring Method
Diff. of Means
Std. Error Diff. of Mean
Calc. t Value
Two Tail. Probability
Mean of L/S Ratios Std. Dev,
2.56
1.17
2.07
0.94
0.49
0.14
3.39
0.002 (Significantly Different)
Chemistry 199
Table 2: Individual Patient L/S Ratios Determined by the Two Detection Methods
Patient No. ANS Fluorescence Cupric Acetate Charring
1 2.60 0.72
2 2.00 0.75
3 3.00 0.83
4 1.40 0.98
5 2.80 1.30
6 4.50 1.60
7 0.83 1.40
8 1.50 1.20
9 1.20 0.85
10 0.98 1.40
11 0.48 1.20
12 4.00 3.80
13 2.60 4.40
14 2.40 2.10
15 1.50 1.60
16 2.80 1.50
17 2.60 2.30
18 4.30 2.80
19 1.30 1.40
20 3.30 2.50
21 3.30 3.90
22 3.60 3.00
23 2.50 2.70
24 2.20 2.40
25 2.90 3.00
26 5.30 3.10
27 1.60 1.50
28 1.30 2.00
29 3.40 2.60
30 1.40 0.88
31 1.70 1.80
32 2.80 3.70
33 1.80 1.70
34 3.70 3.00
35 2.20 1.50
36 3.20 2.00
37 3.60 1.90
38 4.20 3.20
39 1.40 1.40
40 3.90 3.00
41 4.30 3.50
42 1.20 1.70
It is evident from Table 2 that in 28 of the 42 L/S ratio determinations, these ratios
are larger when determined by the ANS fluorescence method.
Discussion
Published results from several laboratories suggest that the copper acetate charr-
ing method of detection is loaded with problems. Spillman, et al. reported that when
unsaturation sensitive methods such as copper acetate are compared with unsaturation
insensitive methods such as molybdate detection, that the unsaturation insensitive
methods consistently give higher L/S ratios than those methods that are unsaturation
sensitive (8). Touchstone, et al. completed a study of the reactivity of separated
phospholipids toward various charring reagents. They also noted that saturated lecithins
as are commonly found in mature amniotic fluids are rather insensitive to copper acetate
200 Indiana Academy of Science Vol. 94 (1985)
detection. Those lecithins containing at least one unsaturated fatty acid are responsive
to the reagent and multiple unsaturated lecithins are additionally sensitive. Partially
unsaturated samples of phosphatidyl ethanolamine, phosphatidyl serine and phosphatidly
inositol are also responsive to copper aceteate charring reagents (9). Various resear-
chers report that the relative abundance of saturated lecithins (as dipalmityl phosphatidyl
choline) increases dramatically beyond week 30 of gestation (5). Thus, as fetal lung
maturation occurs as reflected by pulmonary surfactant present in amniotic fluid one
can expect the resultant lecithins to become increasingly insensitive to spray reagents
such as copper acetate.
A number of authors have shown that the temperature used to complete the charring
is even critical with copper acetate sprays. Mueller has shown, if sprayed plates are
heated at 120°C that consistently higher L/S ratios are determined, while if plates are
heated to 130°C, consistently lower L/S ratios are determined, using the phospholipid
standards (6). In a classic study by Gluck, et al. comparing other charring methods,
it was noted that differing results are obtained at different temperatures and also results
are dependent on whether calcium sulfate binder is present or absent in the silica gel.
Finally these results were variable with the kind of charring reagent used (3). Brown,
et al. recently reported that as charring times are increased, the resolved sphingomyelin
spot intensified in color while the resolved lecithin spot decreases in color. Thus as
charring time is increased, the L/S ratio appears to decrease (1).
These published results of other researchers certainly indicate that if copper acetate
charring methods are used that all the parameters, such as silica gel source, presence
or absence of calcium sulfate binder, charring temperature, and charring time must
be very carefully and uniformly regulated from one determination to another. In order
to ultimately decide if the sensitive ANS fluorescent reagent is more effective, we will
need to conduct a series of experiments comparing L/S determinations on amniotic
fluid samples for the two detection methods. We will need to have an elaborate neonate
follow-up after the fact to actually determine if the lung development of the newborn
would parallel the estimate yielded by our respective tests.
Summary
We have extracted and chromatographed methanol-chloroform concentrates of
42 patient amniotic fluid samples on air dried silica gel thin layer chromatography
plates. We have detected resolved phospholipids by either direct visualization of cupric
acetate charred spots or by fluorescence of spots revealed after spraying with ANS.
Calculation of spot areas in each case has permitted us to determine
lecithin/sphingomyelin ratios for each sample analyzed by each detection method. We
have shown that the two sample populations are statistically different and that the
saturated lecithin sensitive methods (ANS detection) yields larger L/S ratios for 28
of the 42 patient samples.
The authors wish to acknowledge the typing of this manuscript by Ms. Elaine
Wilson.
Literature Cited
1. Brown, L.M., C.G. Duck-Chong and W.J. Hensley. 1982. Improved Procedure
for Lecithin/Sphingomyellin Ratio in Amniotic Fluid Reduces False Predictions
of Lung Immaturity. Clin. Chem. 28:344-348.
2. Gluck, L. 1978. Evaluating Functional Fetal Maturation. Clin. Obstet. and Gyn.
21:547-559.
3. Gluck, L., M.V. Kulovich and R.C. Borer. 1971. Diagnosis of the Respiratory
Distress Syndrome by Amniocentesis. Am. J. Obstet. Gynecol. 109:440-445.
Chemistry 201
4. Kaplan, L.A. and A.J. Pesce. 1984. Clinical Chemistry: Theory, Analysis and
Correlation. C.V. Mosby Co., St. Louis, p. 287-296.
5. Kikkawa, Y. and F. Smith. 1983. Cellular and Biochemical Aspects of Pulmonary
Surfactant in Health and Disease. Lab. Investig. 49:122-139.
6. Mueller, R.G. 1982. Effect of Charring Temperature on Observed L/S Ratio.
Clin. Chim. Acta. 122:79-83.
7. Ragatz, B.H., B. Otfinoski, G. Modrak and D. Lyng. 1982. Evaluation of Detection
Systems Used to Determine Lecithin/Sphingomyelin Ratios in Amniotic Fluid.
Proc. Ind. Acad. Sci. 91:188-194.
8. Spillman, T., D.B. Colton, S.C. Lynn, Jr. and J. P. Bretandiere. 1983. Influence
of Phospholipid Saturation on Classical Thin-Layer Chromatographic Detection
Methods and Its Effect on Amniotic Fluid Lecithin/Sphingomyelin Ratio Deter-
minations. Clin. Chem. 29:250-255.
9. Touchstone, J.C., S.S. Levin, M.F. Dobbins, L. Matthews, P.C. Beers and S.G.
Gable. 1983. (3-sn-Phosphatidyl) cholines (Lecithins) in Amniotic Fluid. Clin.
Chem. 29:1951-1954.
10. Warren, B.M. 1980. The L/S Ratio—How Does It Relate to Fetal Maturity? Helena
Laboratories. Beaumont, Texas.
The Effects of Oligolysines and Polylysines on Human Platelet Aggregation
Induced by Polylysines, Adenosine Diphosphate, and Epinephrine
Barth H. Ragatz, Gina Modrak and Mike Engle
Fort Wayne Center for Medical Education, Indiana University School of
Medicine
and Department of Biological Sciences, Indiana University-Purdue University at
Fort Wayne
Fort Wayne, Indiana 47805
Introduction
In the past there have been several confusing reports in the literature about the
interaction of polylysine with human platelet-rich plasma (PRP) suspensions. Some
reports indicate that this synthetic polycation can induce platelet aggregation and
stimulate the release reaction when added to PRP suspensions (5). Other investigators
have suggested that at most there is an electrostatic interaction between the positively
charged polylysine and the sialic acid-rich negatively charged platelet surfaces (6). This
polylysine effect has been reported to be independent of polymer molecular weight,
with polymers in the molecular weight range 2500-400,000 Daltons being effective (4,
5, 6, 7, 8, 10). Once again, Metcalf and Lyman report that plasma cofactors may
be required for the polylysine-platelet interaction but Massini et al. report that no
plasma cofactor is required (4, 5). Published reports also indicate that conformational
variations are possible and that extended left-handed polylysine helices effectively in-
teract with platelets and that L, D, and D-L monomers can be present. Metcalf and
Lyman indicate that the beta polylysine conformation interacts with the platelets, while
the random coil conformation is ineffective (4).
It is reported also that the epsilon amino groups of the lysine monomers must
remain intact for the interaction to occur. Succinylation of these groups abolishes ac-
tivity as does deamination, N-acetylation, or N-dinitrophenylation. Various biological
polyanions can also inhibit the polylysine-platelet interaction, presumably by forming
electrostatic complexes with the added polylysine. Included in this category are heparin
and chondroitin sulfates (7).
Since many reagents which induce platelet aggregation or the release reactions
in platelets are dependent on the liberation of arachidonic acid from membrane bound
phospholipids, and the subsequent generation of cyclic endoperoxides from the
arachidonic acid, we decided to use a well known reagent to block the generation of
these derivatives in the arachidonic acid cascade (13). We reasoned that if the polylysine-
platelet interaction is primarily an electrostatic interaction, then impairment of the
biochemical functionality of the platelets probably would not alter it.
It is well known that aspirin (acetyl salicylic acid) is a common pharmacologic
agent which can block the generation of arachidonic acid derivatives (13). Published
studies indicate that aspirin acetylates susceptible protein R groups on at least three
platelet proteins, including the enzyme, platelet cyclo-oxygenase. This particular en-
zyme is involved in the generation of the cyclic endoperoxide intermediates (PGG2,
PGH2) which are precursors to the potent platelet aggregator, thromboxane A2 (2, 4, 9).
In our present study, we have taken platelets from human volunteer subjects who
were either aspirin-free or well aspirinized at the time the platelets were collected, and
we have then studied the interaction of various molecular weight, oligo-and polylysines
with either aspirin-free or well aspirinized platelets in plasma suspension. We have
also studied the interaction of these two kinds of platelet populations by incubating
them with various molecular weight oligo-or polylysines and then adding low doses
203
204 Indiana Academy of Science Vol. 94 (1985)
of adenosine diphosphate to the suspensions 30 seconds later. The dose of adenosine
diphosphate was selected to induce only a mild reversible primary aggregation when
it is added alone. Finally, the two kinds of platelet populations were preincubated
with various oligo-and polylysines and then epinephrine was added in strong aggregating
30 seconds later.
Materials and Methods
Potential platelet donors were recruited and each completed a questionnaire
evaluating disease- free and drug-free state of the donor. Each donor also signed an
informed consent statement developed and approved by the Committees for the Pro-
tection of Human Subjects within IUPUI and IPFW. Approximately 50 ml. of whole
blood was collected into Becton-Dickinson 6419 Vacutainers, specifically designed for
preparation of PRP. These evacuated containers were sterilized, silicone-coated, and
contain 0.5 ml. of buffered 0.129M sodium citrate. The collected whole blood was
centrifuged in a vibration-free Sorvall DuPont T6000 centrifuge at room temperature
for ten minutes at 1000 RPM's. The PRP is carefully pipetted from the top of the
tubes into a plastic container using a plastic Falcon 10 ml. pipet. Platelet poor plasma
(PPP) was prepared by centrifuging the remaining blood components for ten minutes
at 10,000 RPMs in a high speed refrigerated Sorvall centrifuge and the supernatant
resulting was collected. Platelet counts were obtained on an automated Coulter counter
at Veterans Administration Hospital in Fort Wayne, Indiana. All PRP typically had
a platelet count greater than 350,000 platelets/mm3.
Adenosine diphosphate and most of the polylysines were obtained from Sigma
Chemical Company. Some of the polylysines and all oligolysines were obtained from
Vega Chemical Company. Epinephrine was obtained from Bio Data Corporation. Stock
solutions were prepared at appropriate concentrations by diluting the respective reagent
with 0.85% sodium chloride. Dilutions were prepared also using this sodium chloride
solution and all solutions were adjusted to pH7 with an Orion 501 pH meter. The
test reagent solutions and standards were stored in plastic culture tubes at — 20°C in
5 ml. aliquots. Polylysines and oligolysine were dissolved at the appropriate concen-
tration on the day of usage.
Platelet-rich plasma was stored at room temperature and promptly utilized within
4-6 hours after the whole blood was drawn. All aggregation tests were done in a Payton
300 Dual Channel Aggregometer at 37°C. with a constant stirring speed of 900 RPM's.
These conditions are optimal for efficient aggregation and do not cause sufficient shearing
forces to disaggregate platelet clumps. The chart recorder ranges are established on
the two pen recorder system using aliquots of platelet-rich and platelet-poor plasma.
Baseline stability is periodically checked and aggregation standards are added to samples
periodically to insure that platelets are remaining viable. If obvious erythrocyte sediments
or hemolysis is detected in the PRP, it is discarded promptly.
Results
Typical results evaluating the interaction of platelet suspensions with the oligo-
and polylysines at 1 mg./l ml. concentrations are shown in Table 1. It can be seen
that if small oligolysines (lysyl-lysine, pentalysine) or intermediate molecular weight
lysines (molecular 4000-14,000 Daltons) are added to suspensions of normal platelets
or aspirinized platelets, no aggregation effect is observed when monitored for at least
five minute periods. It can be seen, however, when large molecular weight polylysines
(25,000-240,000 Daltons) are added to normal or aspirinized platelet suspensions that
a prompt, complete aggregation occurs. The experimental results suggest that a larger
polylysine (molecular weight = 55,000 Daltons) is required to bring about initial
Chemistry 205
Table 1 . Interaction of Platelet-Rich Plasma Suspensions with Oligo- and Polylysines
(at lmg/ml concentrations)
Compound Added Aspirin Free Platelets Aspirinized Platelets
Lysyl-Lysine — -
Pentalysine - -
4K Polylysine - -
14K Polylysine - -
25K Polylysine +
55K Polylysine + +
90K Polylysine + +
150K Polylysine N.D. +
240K Polylysine N.D. +
( + ) = Complete, Irreversible Aggregation
( - ) = No Effect
N.D. = No Data Collected
aggregation of platelets, but basically it can be concluded that the polylysine-platelet
interaction is independent of the usual functioning arachidonic acid cascade leading
to the production of cyclic endoperoxides and thromboxane A2.
Table 2 shows the effects of pre-incubating normal platelets or aspirinized platelets
with various oligo- or polylysines for 30 seconds before a low dose of adenosine
diphosphate (ADP) is added. This table indicates once again that there is no difference
in responsiveness between the normal platelets and the aspirinized platelets. Further-
more, this series of experiments shows that there is a cooperative interaction between
the larger polylysines (molecular weight greater than 25,000 Daltons) and adenosine
diphosphate. This cooperative interaction between polylysine and ADP can be explained
by a linkage of a discrete polycation receptor with the adenosine diphosphate receptor,
or by the fact that platelets preincubated with polylysines are drawn in closer proximi-
ty to one another and are more readily stimulated by low doses of ADP than is the
case when polylysines are absent. It can certainly be seen that aspirin does not impair
in any way this polylysine and adenosine diphosphate interaction with platelets.
Table 2: Effects of 30 Sec. Pre-Incubation of Oligo- and Polylysines on ADP-Induced
Platelet Aggregation
ADP Added 30" After:
Lysyl-Lysine
Pentalysine
4K Polylysine
14K Polylysine
25K Polylysine
90K Polylysine
150K Polylysine
240K Polylysine
( - ) = No Effect
( + ) = Rapid, Complete, Irreversible Aggregation
N.D. + No Data Collected
Aspirin Free Platelets
Aspirinized Platelets
-
N.D.
-
N.D.
+
+
+
+
+
+
+
+
206 Indiana Academy of Science Vol. 94 (1985)
Table 3. Effects of 30 Sec. Pre-Incubation of Oligo- and Polylysines on Epinephrine-
Induced Platelet Aggregation
Epi Added 30" After: Aspirin Free Platelets Aspirinized Platelets
Lysyl-lysine - N.D.
Pentalysine - —
4K Polylysine - -
25K Polylysine + +
55K Polylysine + +
90K Polylysine + +
150K Polylysine + +
240K Polylysine + +
( - ) = No Effect
( + ) = Enhances Aggregation in Magnitude or Onset
N.D. = No Data Collected
Finally Table 3 shows the effect of pre-incubating normal platelets or aspirinized
platelets with various oligo-or polylysines and then adding a vigorous aggregating dose
of epinephrine 30 seconds after the polycation addition. Once again, the data in Table
3 shows there is no difference in response between the normal and aspirinized platelets.
It can be seen that a minimal sized polylysine (molecular weight = 25,000 Daltons)
is required for this cooperative effect and again, there is a positive interaction between
polylysine pre-incubation and epinephrine addition. As was the case for secondary ADP
induced aggregation, this phenomenon could be explained by the linkage of a discrete
polycation receptor to a discrete membrane epinephrine receptor on the platelets, or
it could be explained alternatively by an electrostatic interaction of the polylysines with
the platelets initially bringing them into spatial proximity to enhance the effect of
epinephrine. Again, it is obvious that the aspirinized, biochemically impaired platelets
yield equally good responses in these polylysine-epinephrine experiments.
Discussion
The present results indicate that polylysine polymers in the molecular weight range
25,000 to 240,000 Daltons are effective in inducing aggregation of platelet rich plasma
when added at concentrations of 1 mg./l ml. In contrast to earlier published studies
with normal platelets, we have observed no aggregation with polylysines of molecular
weight lower than 14,000 Daltons. Many of the earlier studies were done with citrate
addition to whole blood in which the relative concentration of citrate was less care-
fully controlled and not standardized as has been the case in the present study using
the B-D liquid citrate Vacutainers (4, 5, 6, 7, 10). Control of relative citrate concentra-
tion has been shown to be an important parameter in obtaining good platelet aggrega-
tion results in clinical studies (11).
In both normal and aspirinized platelets, we have seen a cooperative interaction
between polylysines and classical aggregating agents, such as adenosine diphosphate
or epinephrine. These positive interactions could be explained by a coupling between
discrete receptor sites on the platelet surface and by an electrostatic interaction of platelets
with polylysines initially. These present results do demonstrate that biochemically im-
paired platelets with an inability to generate the cyclic endoperoxides PGG2 or PGH2,
or the potent aggregating substance, thromboxane A2, give equally good responses
to polylysines or combinations of polylysines with either adenosine diphosphate or
epinephrine. Although the present results alone do not prove conclusively that polylysines
are without biochemical or metabolic effects on the platelets, they are certainly sug-
Chemistry 207
gestive that polylysine-platelet interactions are largely electrostatic in nature. Earlier
published reports by Guccione et al. in which platelets were pretreated with adenosine,
EDTA, or prostaglandin E2 also suggest that platelet functionality is not altered when
polylysines are finally added (3).
In conclusion, there is no difference in response to populations of normal or
aspirinized platelets to any of the tests mentioned above. There is a minimum molecular
weight for polylysines that is required for induction of the aggregation reaction with
B-D Vacutainer prepared platelet-rich plasma. However, since lysine and oligolysines
have been shown to inhibit adenosine diphosphate or thrombin induced aggregation,
it is obviously relevant for us to learn more about this polycation-platelet interaction
(1, 12). The next phase of our research will include transmission electron
photomicrographic studies to determine if the polylysine interaction causes any of the
classic morphologic changes seen in platelets when various conventional aggregating
agents are added.
The authors wish to acknowledge the preparation of this manuscript by Ms. Elaine
Wilson and the platelet counts which were provided by Mr. Zane Smith at the Veterans
Administration Hospital in Fort Wayne.
Literature Cited
1. Agam, G., T.K. Gartner and A. Livne. 1984. Inhibition of Platelet Aggregation
and Endogenous Lectin Activity by Oligoamines. Thromb. Res. 33:245-257.
2. Buchanan, M.R., J. A. Rischke and J. Hirsh. 1982. Aspirin Inhibits Platelet Func-
tion Independent of the Acetylation of Cyclo-Oxygenase, Thromb. Res. 25:363-373.
3. Guccione, M.A., M.A. Packham, R.L. Kimbaugh-Rathbone, D.W. Perry and
J.F. Mustard. 1976. Reactions of Polylysine with Human Platelets in Plasma and
in Suspensions of Washed Platelets. Thrombos. Haemostas. 36:360-375.
4. Hoak, J.C. 1983. Mechanisms of Action: Aspirin. Thromb. Res. Suppl. IV. 47-51.
5. Massini, P., L.C. Metcalf, U. Naf and E.F. Luscher. 1974. Induction of Ag-
gregation and of the Release Reaction in Human Platelets by Polylysine.
Haemostasis. 3:8-19.
6. Mohammed, S.F., H.Y.K. Chuang, P.E. Crowther and R.G. Mason. 1979. In-
teractions of Poly (L-Lysine) with Human Platelets, Correlation of Binding with
Induction of Platelet Aggregation. Thromb. Res. 15:781-791.
7. Mohammed, S.F., H.Y.K. Chuang and R.G. Mason. 1977. Roles of Polymer
Size and 6-Amino Groups in Polylysine-Platelet Interaction. Thromb. Res.
1:193-202.
8. Ragatz, B.H. 1980. Interactions of Various Homopolypeptides with Human
Platelet-Rich Plasma Suspensions. Proc. Ind. Acad. Sci. 90:180-185.
9. Roth, G.J. and P.W. Majerus. 1975. The Mechanism of the Effect of Aspirin
on Human Platelets. I. Acetylation of a Particulate Fraction Protein. J. Clin.
Invest. 56:624-632.
10. Tiffany, M.L. and J. A. Penner. 1976. Polylysine Aggregation of Human Blood
Platelets. Thromb. Res. 8:529-530.
11. Triplett, D.A. (Edit.). 1978. Platelet Function: Laboratory Evaluation and Clinical
Application. Amer. Soc. Clin. Path. (Publisher), p. 64-67.
12. Ts'ao, C, S.J. Hart, D.V. Krajewski and P.G. Sorenson. 1982. Opposite Effect
of Lysine on Platelet Aggregation Induced by Arachidonate and by Other Ag-
gregants. Thromb. Haemostas. 48:78-83.
13. Weiss, H.J. 1982. Platelets: Pathophysiology and Antiplatelet Drug Therapy. Alan
R. Liss, Inc., New York.
ECOLOGY
Chairperson: Edwin R. Squiers
Department of Biology
Taylor University
Upland, Indiana 46989
(317)998-2751 ext. 386
Chairperson-Elect: Richard W. Miller
Department of Zoology
Butler University
Indianapolis, Indiana 46208
(317)283-9328
ABSTRACTS
Pipewort Pond, a Unique Wetland with Atlantic Coastal Plain Elements in Elkhart
County, Indiana. James R. Aldrich, Division of Nature Preserves, Indiana Depart-
ment of Natural Resources, Indianapolis, Indiana 46204. This remarkable wetland
supports a unique assemblage of native vascular plants many of which are commonly
referred to as "Atlantic Coastal Plain disjuncts." Many of the species that occur at
Pipewort Pond such as Fuirena pumila, Psilocarya scirpoides, Rhynchospora
macrostachya, Eriocaulon septangulare, Juncus pelocarpus and Utricularia purpurea
are rare or otherwise noteworthy species for the Indiana flora. The vegetation and
ecology of the wetland is discussed and a species list is presented.
Competition for Ownership of Webs in the Semi-social Spider Cyrtophora moluccen-
sis of Yap (Caroline Islands, Micronesia). James W. Berry, Department of Zoology,
Butler University, Indianapolis, Indiana 46208. In a colonial web each spider builds
its own orb, but wandering individuals frequently challenge the original inhabitant
of the orb. When a spider is introduced into the orb of another spider, one of the
individuals eventually is chased from the orb. Two important factors in deciding posses-
sion of the orb are prior occupancy of the orb and body weight. Disregarding weight
differences, the owner retained possession of the orb about 70% of the time. The
heavier spider, whether the intruder or the owner, was the winner 60% of the time.
In 53 experiments, the time elapsing between the intruder being introduced into the
orb and one of the individuals leaving the orb varied from 30 seconds to more than
four hours.
Regional Low Density and Extinction in Populations of Peromyscus leucopus. Alex
Burgin and David T. Krohne Department of Biology, Wabash College, Crawfords-
ville, Indiana 47933. In the spring and summer of 1984 unusually low densities
of populations of the white-footed deer mouse, Peromyscus leucopus were, encountered
throughout the Sugar Creek drainage in west-central Indiana. Extensive trapping at
seven sites on both sides of the creek indicated that this phenomenon extended for
at least 60 km and included local extinction in at least three sites. Age structures were
biased toward adults in all sites during the low density periods. Sex ratio was heavily
biased toward males on all sites but one during this period. On the one site in which
males did not predominate, recovery from the low density situation began earlier and
continued more rapidly than on other sites. By the end of the summer of 1984, the
209
210 Indiana Academy of Science Vol. 94 (1985)
sites had begun to diverge in density with some recovering at different rates while
others remained extinct.
Predator-determined Structure in Amphibian Pond Communities. Spencer A.
Cortwright, Indiana University, Bloomington, Indiana 47405. Community struc-
ture encompasses the number and relative abundances of interacting species. The
mechanisms producing community structure are interactions among the species and
their relations to the physical environment. Patterns in pond-breeding amphibians sug-
gested that moderate or high densities of a fall-breeding Ambystoma (a salamander
predator on spring-breeders) were associated with low populations of one spring-breeding
salamander and higher populations of a second. A factorial pen experiment was done
using two densities each of the three salamanders (plus constant numbers of other
common amphibians).
The results showed a strong predator effect on the fall-breeding Ambystoma
opacum on two early-hatching species, Ambystoma jeffersonianum and Rana sylvatica,
in both the pen experiment and the pond itself. Two later-hatching species, Ambystoma
maculatum and Notophthalmus viridescens, experienced much higher survivorship in
the presence of predators. Rana clamitans breeds even later, has low palatibility, and
is too large to be consumed the following spring. Thus, timing of prey hatching and
possibly prey behavior may strongly affect prey susceptibility and, hence, community
structure.
In more temporary pools without A. opacum, A. jeffersonianum survives in higher
numbers and appears to depress the survivorship of A. maculatum. Thus A. opacum
appears to cause a reversal in the relative abundances of these two prey species.
The Complex Relationship of Embryonic Development to Incubation Temperature in
Turtles. Michael A. Ewert and Craig E. Nelson, Indiana University, Bloomington,
Indiana 47405. In birds, the incubation period within a species deviates little from
the mean. In turtles, however, the incubation period within a species varies greatly
and healthy turtles hatch following a broad range of durations. Only part of this variation
is attributable to acceleration of development by increased temperature. At a single
temperature, eggs from higher latitudes develop up to 30% faster than conspecifics
from lower latitudes, a difference expressed throughout embryonic differentiation. Other
species have a variably prolonged arrest of development in early stages and, sometimes,
at term, when otherwise they are ready to hatch.
What is the adaptive significance of variable incubation periods? In particular,
what do slower developers gain? In species with environmental sex determination, pro-
longed development may match temperature sensitive phases of development with
seasonal arrays of temperatures more favorable for gonadal development. Alternatively,
some species may be "bet-hedging" to assure that some eggs hatch when environmen-
tal conditions are favorable. We are using calorimetry and respirometry to explore
the energetic implications of these options.
A Competitive Ecotone between Hardwood and Relict Hemlock Communities. Scott
Person, Department of Ecology and Evolution, State University of New York, Stony
Brook, New York 11794 and Daniel D. Stockton, Department of Biology, Wabash
College, Crawfordsville, Indiana 47933. Apparently relict stands of Eastern
Hemlock (Tsuga canadensis) occur along Sugar Creek bluffs at the Allee Memorial
Woods Nature Preserve in Parke County, Indiana, in sites usually occupied by beech-
maple hardwoods. If, despite their complexity and variability, these communities behave
integrally in interaction with each other, the boundary between them is expected to
Ecology 211
be very sharp and display a constant width when measured objectively across perpen-
dicular transects. In this case, local floristic configurations will form a bimodal distribu-
tion in vegetation space lacking intermediate states. All hardwood trees (larger than
7.5 cm dbh) and all hemlock stems (taller than 10 cm) were measured for diameter
and mapped in a 1.8 hectare area which included an extremely sharp ecotone between
communities with and without hemlock. Qualitative estimates of understory and
herbaceous layer composition were also made. Analysis of these data may give weak-
inferential evidence that this ecotone is the result of mutual competitive exclusion by
the two communities along an environmental gradient.
Development and Analysis of a CFI Data Base for Indiana. Burnell C. Fischer and
John A. Kershaw, Jr., Department of Forestry and Natural Resources, Purdue Univer-
sity, West Lafayette, Indiana 47907. Continuous Forest Inventory (CFI) plots were
established throughout Indiana during the late 1940s through the mid 1960s. Many
of these plots were maintained and periodically remeasured. However, few summaries
of the data were attempted and the data, if not lost, was simply put in the file cabinet.
This type of data is essential if forest researchers are to develop models of forest develop-
ment and growth which can be used by forest managers.
The relocation and remeasurement of the Purdue portion of Indiana's CFI plots
is nearing completion. This will result in a data base of over 400 CFI plots (many
originating in the early 1950s) on either Purdue Agricultural Centers and Purdue Depart-
ment of Forestry and Natural Resources woodlands. The initial effort was concen-
trated on these woodlands because the existing data was most accessible and these
plots were considered to be in the "best" condition. Work has begun to assess the
condition of plots and accompanying data bases on State Forests and other woodlands.
Initial analysis of the data has concentrated on the development, and growth and
yield of forest stands and the response of individual trees by species and size class.
Stand growth is summarized by growth component. Gross growth, ingrowth, mortality
and cut for basal area and board foot volume are utilized. Although, individual tree
growth has primarily been an analysis of diameter growth by size class and species,
we are also looking at ingrowth and mortality rates.
Obviously, tree and plot growth rates are quite variable depending on both forest
and site conditions. The summarization of a large data set, such as is available in
Indiana, should allow researchers to test a number of hypotheses on the growth and
management of Indiana forests.
Biofiltration in Intensive Culture Systems: Design Considerations. George S. Libey
and Gary E. Miller, Purdue University, West Lafayette, Indiana 47907. The
growth of the human population is accompanied by a need to increase food produc-
tion. Aquaculture, the cultivation of aquatic organisms, offers the potential for expanding
the human food base. Reconditioning systems for fish culture increase the use of limited
water supplies and maintain necessary water quality parameters. Characteristics or recon-
ditioning systems include:
Removal/detoxification of metabolic wastes
Solid waste removal
Reoxygenation
Temperature control
Disease control
Design constraints are:
Soluable organics concentration
212 Indiana Academy of Science Vol. 94 (1985)
Soluable inorganics concentration
Temperature
Dissolved oxygen
Alkalinity
pH
Devices available include:
Packed tower (trickling filter)
Rotating biological contractor
Fluidized bed-reactor
Tube/plate clarifier
Sexual Selection and Alternative Mating Strategies in Hyla crucifer and Hyla chrysoscelis.
Molly Morris, Department of Biology, Indiana University, Bloomington, Indiana
47405. Observations and field experiments were conducted on a population of
Hyla crucifer (spring peeper) and a population of Hyla chrysoscelis (gray treefrog)
during their respective mating seasons. Data was taken to determine behavioral and/or
morphological characteristics that could influence a male's reproductive success. Close
attention was also given to the location and distinguishing characteristics of the call
sites. In both species, large males did not have a higher probability of mating than
smaller males, nor did I find positive assortative mating of large males with large females
and smaller males with smaller females. Factors that seem to affect mating success
in gray treefrogs include the number of nights spent calling and a male's close associa-
tion with another calling male. Males that spent more evenings calling had a higher
probability of mating. These and other results will be discussed in terms of mating
systems, alternative male mating strategies and game theory.
Do Tadpoles Die for their Siblings? Craig E. Nelson, Department of Biology, Indiana
University, Bloomington, Indiana 47405. When same-age conspecific tadpoles are
grown together, it has frequently been observed that one or a few of the tadpoles
grow well and that the growth of the other tadpoles is severely inhibited. Indeed the
inhibited tadpoles often fail to feed and consequently die. Inhibitability appears to
be selectively disadvantageous and might be expected to evolve out of the population.
Kin-selection could maintain inhibitability if its net effect was an increase in the growth
and/or survivorship of favored siblings. These experiments ask whether the growth
disparity within sibling groups is greater than that within groups of non-siblings. Such
a disparity would strongly implicate kin-selection.
Tree Species Dynamics in an Old-growth Deciduous Forest since 1926. George R.
Parker and Donald J. Leopold, Purdue University, West Lafayette, Indiana
47907. All trees (_> cm dbh) in a 20.5 ha mature deciduous forest on the Tipton
Till Plain of central Indiana were tagged and mapped in 1926. Trees within the central
8.5 ha were remeasured and mapped in 1976. Thirty-two species were recorded in 1976
and 28 in 1926. There was a shift in relative abundance among species due to ingrowth
and mortality. The majority of ingrowth trees within a 5-m radius gap of dead domi-
nant trees were Ulmus americana (30% of total) and Acer saccharum (20%). Low
mortality species (<_25%) included A. saccharum, Aesculus glabra, most Carya spp.,
Celtis occidentalis and most Quercus spp. High mortality species (>75%) included
Fagus grandifolia, Ulmus spp., and Fraxinus nigra. Nearly half (46.9%) of those trees
measured in 1926 were dead by 1976. Stand density and basal area increased 93.9 and
30.8%, respectively, to 320 stems/ha and 31.0 m/ha by 1976. Mortality has average
2.9 stems/ha since 1976 with U. americana accounting for about 30% of those dying.
Ecology 213
Male Mating Behavior in Hyla cinerea. Stephen A. Perrill, Department of Zoology,
Butler University, Indianapolis, Indiana 46208. Hyla cinerea males in two ponds
on Wilmington Island, Georgia, were toe-clipped and freeze-branded for individual
identification. Their activities and rates of mating success were observed over three
breeding seasons, 1979-81. The goal of these observations was to relate their behavior,
location, and physical characteristics to their rates of mating success. Three categories
of behavior were considered (calling, satellite and non-calling) and more than 80%
were found to be either calling or adopting the satellite strategy for most of the obser-
vation period. The least site-specific, least mobile males showed the highest rate of
mating success; the most site-specific, the least successful. In each year, there was a
significant positive correlation between calling activity and mating success. Also, there
were consistent positive relationships between the number of nights the frogs frequented
the study site and their rates of mating success; but mean body size did not appear
to influence the mating success rates.
Hardwood Tree and Ground Cover Establishment on Reclaimed Mineland and Unmined
Reference Sites in Indiana. Phillip E. Pope, William R. Chaney and William R.
Byrnes, Department of Forestry and Natural Resources, Purdue University, West
Lafayette, Indiana 47907. Establishment success, productivity, and compatibility
of ground cover and hardwood tree seedlings planted concurrently and maintained
under the same level of management were evaluated on reclaimed, surface-mined, coal
land and unmined reference sites in southwestern Indiana. Topography, soils, and vegeta-
tion were similar on both sites prior to mining. The mined land was reclaimed for
forest land use under provisions of Public Law 95-87, The Surface Mining Control
and Reclamation Act of 1977. The reference area was cleared of all vegetation and
both sites were disced, limed, and fertilized before planting. Soil physical and chemical
properties were analyzed and compared between sites. One-year-old black walnut (Juglans
nigra L.) and red oak (Quercus rubra L.) seedlings were planted at 2 x 2 meter spacing
concurrently with a mixture of K-31 fescue (Festuca arundinacea Schreb.) and red clover
{Trifolium pratense L.) in spring 1981. Tree rows in one-half of each experimental
unit were treated with amizine (simazine + amitrole) and dalapon to control ground
cover plants and to assess the competitive effects of ground cover on hardwood tree
establishment and growth. After three growing seasons, black walnut and red oak seed-
ling survival was significantly greater on the reference site (88 and 77%, respectively)
than on the reclaimed mineland (50 and 42%, respectively). Chemical control of ground
cover was essential to meet stocking levels of 450 trees/acre specified in Federal and
Indiana reclamation laws. Percent ground cover exceeded 70% of the cover present
on the unmined reference site for three growing seasons, and hence met the initial
requirement of Public Law 95-87. Ground cover biomass was similar on the minesite
and reference areas in 1981, however, it was about twice as great on the reference
site than on the minesite in the 1982 and 1983 growing seasons.
Interactions among Mast, Small Mammals, and Insects, and their Implications for
Oak Management. Brad Semel and Douglas C. Andersen, Purdue University West
Lafayette, Indiana 47907. Interactions among acorn weevils (Circulionidae), short-
tailed shrews (Blarina brevicauda), white-footed mice (Peromyscus), and acorns were
examined to assess the net impact of these animals on acorn germination and survival.
Only 5% of acorns collected in traps within replicate plots at Martell Forest, near
Lafayette, Indiana were found to be undamaged. Sixty-two percent of the 1983 acorn
crop was damaged as a result of Curculionid infestation; arboreal vertebrates damaged
another 29%.
214 Indiana Academy of Science Vol. 94 (1985)
Feeding trials indicated P. leucopus will consume both infested acorns and weevil
larvae; a preference for non-infested acorns over infested acorns was detected in
laboratory food choice experiments. Blarina consumed weevil larvae but did not extract
them from acorns. Peromyscus detected and excavated larvae that had exited host
acorns and entered the soil only from the upper 5 cm of the soil profile in contrast
to Blarina brevicauda, which consumed larvae from within the upper 16 cm of the
soil profile. Field experiments indicated that about 50% of larvae overwinter in the
upper 5 cm of the soil profile; no larvae were noted to overwinter below 21 cm. Other
field experiments indicated that rates of acorn removal by mice decreased as the pro-
portion of weevil infestation increased. An attempt to document a negative impact
by Blarina on Peromyscus populations was inconclusive.
Taken together, our studies suggest that precaution is necessary in designing oak
management programs based largely upon chemical insect control to increase acorn
production; enhancement of the beneficial activities of Blarina may provide an alter-
native strategy.
Density-dependent Mortality on Galls of the Goldenrod Gall Fly, Eurosta
solidaginis. Rod Walton, Department of Biology, Indiana University, Bloomington,
Indiana 47405. The dependence of predator foraging on both local and overall
prey density can have broad implications for the dynamics and stability of a predator-
prey system. Recent discussions of the effects on predator efficiency of predator
aggregation in areas of high prey density, transit times between prey patches and predator
handling times, together with the distribution of prey among patches of a heterogeneous
habitat suggest that different prey distributions should result in different foraging rates
by predators. Prey faced with a strongly aggregating predator, for instance, will benefit
from an underdispersed rather than a clumped distribution. This is especially critical
for sedentary prey. Eurosta solidaginis (Tephritidae) females oviposit in the stems of
Solidago spp. during early spring. Third instar larvae overwinter within spherical stem
galls. During development and over the winter, larvae are vulnerable to several predators:
two species of parasitoid (Eurytoma spp.) a predatory beetle larva (Mordellistena spp.)
and avian predators (e.g. Downy Woodpeckers). This study was undertaken to deter-
mine the distribution pattern of Eurosta galls in a natural habitat, the degree of density-
dependence for each predator, and the theoretical "optimal" gall distribution that would
minimize predation losses under the constraints of a specific suite of predators.
Tree Species Response to Release from Domestic Livestock Grazing
David K. Apsley, Donald J. Leopold and George R. Parker
Department of Forestry and Natural Resources
Purdue University, West Lafayette, Indiana 47907
Introduction
Grazing of domestic livestock has long been a factor which has greatly influenced
forest structure and composition in the Central Hardwood Region, and the quality
and quantity of timber produced. Approximately sixty-six percent of the forests in
the Central Hardwoods Region were subjected to grazing as late as 1947 (4). Prior
to the passage of the Indiana Forest Classification Act in 1921, nearly all of the farm
woodlots in Indiana were grazed (3). Currently, approximately thirty percent of the
forest land in Indiana is grazed (8).
Although many of the farm woodlots in northern Indiana have been protected
from grazing since the passage of the Forest Classification Act in 1921, questions con-
cerning the long-term effect of grazing remain unanswered. The purpose of this report
is to present recent findings of an extensive research project that was initiated in 1930
to monitor the recovery of Central Hardwood forests from grazing by domestic livestock.
Daniel Den Uyl, Department of Forestry and Natural Resources, Purdue University,
established permanent plots in the early 1930s, throughout central and northern Indiana,
and remeasured the plots at approximately five year intervals until the early 1960's.
These plots were established to elucidate recovery processes of these woodlots from
grazing. Den Uyl's initial project and early results are detailed in several publications
(3,4,5,6,7 and 8). The foresight of Den Uyl has provided a unique opportunity to
study the long-term effects of domestic livestock grazing. Research to be reported will
focus on changes in species composition, size-class distributions, and basal area and
density values over the past 50 years on several of Den Uyl's grazed and ungrazed plots.
Study Area
Some of Den Uyl's original plots had been visited from 1970 to 1984; however,
due to recent disturbances in these stands (selective logging) and/or missing data from
past inventories, only a small number of quadrats were suitable for remeasurement.
Four plots were selected and are located in the Deam, Hoffman and Romey (two plots)
woods in the northeastern Indiana counties of Wells, Allen and Adams, respectively
(Figure 1). Each varied in forest type, silvical condition, density of canopy, and grazing
history when they were established in 1931-32 (Table 1). All plots are level to slightly
rolling and have had some selective cutting prior to plot establishment.
Table 1. Characteristics of Plots at time of establishment, 1931-1932 (Diller and Medesy,
unpublished report, Purdue University).
Forest
Silvical
Canopy
Gazing
Last Year
Woods
Type
Condition
Density
Intensity
Grazed
Deam
Upland-Swamp
Fair-poor
70%
Medium-Heavy
1930
Hoffman
Oak-hickory
very good
90%
none
—
Romey
Oak-hickory
good
80%
heavy
1927
See Day and DenUyl (1932) for grazing-intensity criteria.
215
216
Indiana Academy of Science
Vol. 94 (1985)
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Figure 1. County map of Indiana showing location of Deam (1), Hoffman (2), and
Romey (3) woods.
Methods and Materials
In 1931-1932 Den Uyl and associates established 70 plots in 16 northern Indiana
counties. Plot size varied from 0.1 to 1 acre (0.04-0.4 ha), most being 0.5 acre (0.2
Ecology 217
ha) in size (7). Diameter at breast height (dbh; about 1.37 m above ground) was measured
on every tree 0.6 inches (1.5 cm) or larger. Tree species, height and crown class were
also determined for each stem. All stems were numbered with metal tags or paint and
were mapped by location and crown shape. Each plot was remeasured at approximately
five year intervals. Photographs were taken of each plot, and general plot descriptions
were made which included: silvical condition, drainage, topography and density of
crown cover. Grazing history and evidence of disturbance such as fire and cutting
were also recorded when this information was available.
In the fall of 1984 four 0.5 acre (0.2 ha) plots were remeasured. The original
plots were located by maps made in 1931 and 1932. Maps which provided directions
and distances from nearby towns were utilized to locate properties; those which
designated plot locations by distances in chains (1 chain = 20.1 m) and bearings were
used to determine the general plot locations.
Once the plot locations were determined, a staff compass and 100 foot (30.5 m)
tape were employed along with remnant tree tags, crown maps and records of distances
and bearings to relocate quadrat boundaries. At least one metal corner stake from
the original plot was found on three plots, and was used as a reference point. However,
when no corner stakes were found, tagged trees from the original study that were on
or near the boundary in conjunction with crown maps provided adequate information
to estimate boundary positions.
Several of the stems near the western edge of of plot no. 66 in the Romey woods
were cleared for agricultural purposes. In order to eliminate the edge effects, 25 per-
cent of the original plot was not included in the newly established plot, resulting in
a 0.38 acre (0.15 ha) plot. Results for all plots have been expressed in relation to one
hectare for ease of comparison. The data from both Romey plots are presented together
since these plots were very similar.
All of the trees present at the time of the initial inventory were remeasured (dbh),
and the original numbers were recorded. When tree tags were no longer present or
readable, crown maps and dbh measurements from the previous survey were used to
determine original tree numbers. All stems greater than or equal to 1.0 cm dbh that
were not present at the time of the previous survey were classified as ingrowth. Ingrowth
stems were identified as to species, measured (dbh) and recorded.
Diameters were recorded for all stems that forked at or below dbh, and all stems
were measured to the nearest 0.1 cm (dbh). A metric caliper was used to measure
dbh of stems less than 6.0 cm, and a metal diameter tape was used for stems larger
than 6.0 cm.
Data from Den Uyl's research (species, crown class and diameters from each
measurement period) were stored, with the newly acquired data, on magnetic tape for
analysis on the University's computer system. Computer programs were written by
the senior author.
Density (stems ha,~ '), basal area (m2ha~ '), and Importance Values ((relative density
+ relative basal areas)/2) were calculated for each species by plot. Stems were also
separated into 5.0 cm size-classes by species and all species combined.
Results
After grazing, some species which were not present in the initial survey had become
established in the plots. At the Romey woods, two species (Carya tomentosa and Acer
rubrum) colonized the plots with the cessation of grazing (Table 2). Celtis occidentalis,
Fraxinus americana, F. nigra and Prunus serotina were new species to the plot in Deam
woods following grazing (Table 3). Two new species, C. occidentalis and Liriodendron
218
Indiana Academy of Science
Vol. 94 (1985)
Table 2. Changes in density ha '(D), basal area m2ha ' and Importance Values3 (IV)
from 1932 to 1984 on plots #66 and #67, combined, in Romey's woods.
1932
1947
1984
Species
D
BA
IV
D
BA
IV
D
BA
IV
Acer rubrum
0.0
0.0
0.0
5.0
0.1
1.0
3.3
0.0
0.1
Acer saccharum
64.2
2.2
12.2
27.2
1.0
6.6
644.2
3.0
22.3
Carpinus caroliniana
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
Carya cordiformis
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
Carya glabra
74.2
2.4
13.6
66.8
3.2
18.2
32.2
4.2
10.3
Carya ovata
212.5
5.0
34.0
192.8
6.4
44.4
169.6
10.1
24.9
Carya tomentosa
0.0
0.0
0.0
0.0
0.0
0.0
3.3
0.0
0.1
Celt is occidentalis
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
Fagus grandifolia
27.2
1.4
6.7
19.8
0.6
4.4
31.3
1.2
3.3
Fraxinus americana
7.4
0.1
1.2
7.4
0.3
1.8
11.6
0.2
0.7
Fraxinus nigra
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
Juglans nigra
2.4
0.2
1.0
2.4
0.3
1.2
0.0
0.0
0.0
Liriodendron tulipifera
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
Ostrya virginiana
22.2
0.2
2.6
17.3
0.2
2.6
778.4
1.8
21.3
Populus deltoides
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
Prunus serotina
2.4
0.1
0.5
2.4
0.1
0.6
69.2
0.2
1.7
Quercus alba
19.8
1.5
6.0
14.8
1.0
5.2
14.8
1.0
2.5
Quercus bicolor
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
Quercus macrocarpa
5.0
0.4
1.6
5.0
0.6
2.4
0.0
0.0
0.0
Quercus muehlenbergii
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
Quercus rubra
24.7
3.0
10.6
9.9
1.0
4.2
93.9
1.5
4.4
Tilia americana
2.4
0.1
0.4
2.4
0.1
0.6
65.9
0.4
2.0
Ulmus americana
14.8
0.9
3.8
14.8
1.0
5.0
145.8
0.5
4.4
Ulmus rubra
17.3
0.5
3.2
7.4
0.2
1.7
18.2
0.1
0.8
Others'3
27.2
0.1
2.8
0.0
0.0
0.0
56.0
^b
1.4
TOTAL
523.9
18.0
394.4
16.1
2139.1
23.8
IV = (relative density + relative basal area)/2.
bT < 0.05
"may include individuals of Quercus shumardii.
includes Asimina triloba, Cornus spp., Crataegus spp., Lindera benzoin, Staphylea tri/olia and Viburnum prunifolium.
d.
Table 3. Changes in density ha ' (D), basal area M2 MA
values3 (IV) from 1931 to 1984 on plot #49 in Deam's woods.
(BA) and importance
1931
1951
1984
Species
D
BA
IV
D
BA
IV
D
BA
IV
Acer rubrum
4.9
0.7
3.0
4.9
1.0
3.4
14.8
1.6
0.0
Acer saccharum
19.8
1.4
8.4
19.8
1.8
9.0
1,7"4.8
4.1
0.0
Carpinus caroliniana
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
Carya cordiformis
44.5
2.3
16.7
19.8
1.3
7.8
24.7
1.5
0.0
Carya glabra
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
Carya ovata
54.4
3.5
22.4
54.4
4.6
23.9
54.4
6.1
0.0
Carya tomentosa
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
Celtis occidentals
0.0
0.0
0.0
0.0
0.0
0.0
14.8
TR
0.0
Fagus grandifolia
4.9
0.7
2.9
4.9
0.8
3.0
0.0
0.0
0.0
Fraxinus americana
0.0
0.0
0.0
0.0
0.0
0.0
9.9
T
0.0
Fraxinus nigra
0.0
0.0
0.0
0.0
0.0
0.0
93.9
0.3
0.0
Juglans nigra
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
Liriodenron tulipifera
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
Ostrya virginiana
4.9
0.8
1.4
0.0
0.0
0.0
207.6
0.8
0.0
Populus deltoides
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
Prunus serotina
0.0
0.0
0.0
0.0
0.0
0.0
19.8
T
0.0
Ecology 219
Table 3. — Continued
1931
1951
1984
Species
D
BA
IV
D
BA
IV
D
BA
IV
Quercus alba
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
Quercus bicolor
9.9
1.3
5.9
9.9
1.7
6.3
9.9
2.7
0.0
Quercus macrocarpa
9.9
1.6
6.7
9.9
2.4
7.6
9.9
4.1
0.0
Quercus muehlenbergii
4.9
0.6
2.8
4.9
0.9
3.2
4.9
1.6
0.0
Quercus rubra
14.8
3.1
11.7
14.8
4.4
13.2
14.8
8.6
0.0
Tilia americana
9.9
1.7
6.9
29.6
2.1
12.1
158.1
1.8
0.0
Vlmus americana
19.8
2.0
10.0
19.8
2.5
10.5
29.7
0.2
0.0
Ulmus rubra
4.9
-pB
1.2
0.0
0.0
0.0
499.2
1.7
10.0
Others0
0.0
0.0
0.0
0.0
0.0
0.0
207.6
0.2
0.0
TOTAL
207.6
19.0
192.7
23.5
3,098.7
35.4
0.0
IV = (relative density + relative basal area)/2.
BT < 0.05.
r
may include individuals of Quercus shumardii.
includes Asimina triloba, Cornus, spp., Crataegus spp., Lindera benzoin, Staphylea trifolia and Viburnum
prunifolium.
tulipifera, also colonized the Hoffman woods, although this stand supposedly had not
been grazed (Table 4).
Some species disappeared from each plot. For example, Fagus grandifolia no longer
exists in the plot at Deam woods. Juglans nigra and Quercus macrocarpa disappeared
from the Romey plots, undoubtedly in part due to selective cutting of the former species.
Carya glabra, Juglans nigra and Populus deltoides were not tallied during the 1984
inventory in the Hoffman plot. However, most if not all of these species still exist
outside of these plots within the respective woodlots.
Densities have increased from 1931-1932 to 1984 for all species combined on all
plots measured; however, increases are greatest on the plots that had been previously
grazed. Overall density increased from 523.9 to 2139.1 stems ha-1 on the Romey plots,
an increase of 308% (Table 2). Density changes were the greatest on the Deam plot;
stem numbers increased from 207.6 to 3098.7 ha"1, an increase of 1392% (Table 3).
Density values for all species combined on the ungrazed plot increase 86% (from 953.8
to 1774.2 stems ha,-1; Table 4).
On all plots surveyed there was a decrease in density from the first measurement
(1931-1932) to the second (1947-1951). This decrease was greatest on the ungrazed Hoff-
man plot (Table 4); however, the density for this plot was much greater than that
of the grazed Romey and Deam plots at the time of the initial survey (Tables 2 and
3). Records kept by Den Uyl indicate that some selective logging occurred in stands
after 1931-1932, but most of the decrease in density was due to natural mortality.
Since natural regeneration following grazing requires 3 to 15 years to establish (8),
seedlings which colonized previously grazed plots were too small to be enumerated
during the intermediate period.
The two species that contributed to the greatest increase in density from 1931-1932
to 1984 on the combined Romey plots were Acer saccharum and Ostyra virginiana.
Densities increased from 64.2 to 644.2 stems ha" ' and from 22.2 to 778.4 stems ha" '
for Acer saccharum and Ostrya virginiana, respectively. These two species accounted
for nearly 83% of the density on the Romey plots in 1984.
Acer saccharum increased from 19.8 to 1724.8 stems ha" ' on the Deam plot; this
increase accounts for nearly 59% of the total plot increase. Increases in density of
220
Indiana Academy of Science
Vol. 94 (1985)
Table 4. Changes in density ha ' (D), basal area M2 ha ' (BA) and Importance ValuesA
(IV). from 1931 to 1984 on plot #26 in Hoffman's woods.
1931
1951
1984
Species
D
BA
IV
D
BA
IV
D
BA
IV
Acer rubrum
24.7
0.6
2.3
14.8
0.6
2.4
4.9
0.2
0.4
Acer saccharum
118.6
0.7
7.4
93.9
1.2
10.1
207.6
2.2
8.8
Carpinus caroliniana
44.5
0.2
2.6
4.9
T-B
0.5
192.7
0.2
5.7
Carya cordiformis
24.7
0.8
2.7
0.0
0.0
0.0
14.8
T
0.4
Carya glabra
29.7
2.2
5.3
0.0
0.0
0.0
0.0
0.0
0.0
Carya ovata
9.9
0.6
4.9
T
0.4
4.9
T
0.1
Carya tomentosa
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
Celt is occidentalis
0.0
0.0
0.0
0.0
0.0
0.0
4.9
T
0.2
Fagus grandifolia
89.0
0.5
5.5
59.3
0.7
6.3
74.1
1.4
4.0
Fraxinus americana
24.7
0.9
2.8
14.8
1.1
3.0
123.6
2.0
6.2
Fr ax in us nigra
89.0
1.4
7.0
19.8
1.0
3.3
19.8
T
0.6
Juglans nigra
9.9
0.7
1.7
9.9
1.2
2.7
0.0
0.0
0.0
Liriodendron tulipiefera
0.0
0.0
0.0
0.0
0.0
0.0
14.8
0.3
0.8
Ostrya virginiana
123.6
0.4
7.2
74.1
0.4
7.2
232.3
1.4
8.4
Populus deltoides
4.9
0.4
1.0
0.0
0.0
0.0
0.0
0.0
0.0
Prunus serotina
0.0
0.0
0.0
0.0
0.0
0.0
84.0
0.1
2.4
Quercus alba
74.1
2.2
7.5
44.5
2.3
7.6
34.6
3.5
5.7
Quercus bicolor
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
Quercus macrocarpa
14.8
0.8
2.2
9.9
1.3
2.9
9.9
2.4
3.6
Quercus muehlenbergii
Quercus rubra
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
93.9
8.4
18.8
89.0
12.9
23.6
84.0
21.0
31.0
Tilia americana
93.9
6.8
16.2
74.1
6.4
16.8
237.2
1.7
9.0
Ulmus americana
4.9
0.1
0.5
4.9
0.2
0.7
64.2
0.1
1.9
Others0
4.9
T
0.3
4.9
T
0.5
286.6
0.2
8.0
TOTAL
953.8
30.2
568.3
31.1
1,774.2
36.7
IV = (relative density + relative basal area)/2.
T< 0.05.
D
May include individuals of Quercus shumardii.
Includes Asimina triloba, Cornus spp., Crataegus spp., Lindera benzoin, Staphylea trifolia and Viburnum prunifolium .
Ostrya virginiana, Tilia americana, Ulmus spp. and species in the "other" category
(mostly species that do not attain large size, e.g., Lindera benzoin, Asimina triloba
and Viburnum prunifolium) account for another 37% of the increase.
Several species {Acer saccharum, Carpinus caroliniana, Fraxinus americana, Ostyra
virginiana and Tilia americana) have contributed greatly to the increase in the density
of the ungrazed plot from 1931 to 1984. However, species in the "other" category,
particularly Lindera benzoin, have contributed the greatest to this value. Thirty-four
percent of the total increase is attributable to these species.
Density values for Carya spp. and Quercus spp. on both grazed and ungrazed
plots from 1931-1932 to 1984 have either decreased or remained fairly constant in most
cases. Again, selective logging removed a portion of these stems (many of which had
been previously injured) although most had died naturally. However, Quercus rubra
density increased from 24.7 to 93.9 stems ha-1 on the combined Romey plots.
Basal area for all species combined has increased from 1931-1932 to 1984 on
all plots sampled; however, as with density, relative increases were greatest on the
grazed plots. Basal area decreased at the intermediate inventory only in the Romey
woods.
Basal area values increased 32°7o (from 18.0 to 23.8 m2ha~') on the combined
Romey plots (Table 2). The largest increase in basal area was on the Deam plot; basal
Ecology 221
area increased from 19.0 to 35.4 m2~\ an increase of 86% (Table 3). Basal area on
the ungrazed Hoffman plot increased from 30.2 to 36.7 m2ha~'.
Carya glabra and C. ovata increased in basal area by 6.9 m2ha'M' on the Romey
plots. This increase exceeds the total increase for these plots, because some species
exhibited a net decrease in the basal area (e.g., Quercus spp. which decreased in basal
area by 2.9 m2ha~ ').
In contrast to the Romey plots, Carya spp. do not account for the greatest in-
crease in basal area on the Deam plot. Only 1.8 m2ha_l of the basal area increase of
16.8 m2ha_l is attributable to Carya spp. Quercus spp. added 10.40 m2ha~' of basal
area; Q. rubra was responsible for over one-half of this increase by 0.9 and 2.7
m2ha_1, respectively, whereas Ulmus spp. basal area totals remained nearly constant.
Quercus rubra increased in basal area by 12.6 m2ha" ' on the ungrazed plot, which
is nearly double the increase for the entire plot. Concurrently, Tilia americiana, Ulmus
rubra, and Carya glabra decreased in basal area by 5.1, 2.7 and 2.2 m2ha_l,
respectively.
In 1932 on the combined Romey plots, Carya ovata, C. glabra, Acer saccharum
and Quercus rubra had Importance Values (IV's) of 34.0, 13.6, 12.2 and 10.6%, respec-
tively (Table 2). The IV of Carya ovata dropped to 24.9% in 1984; however, C. ovata
still has the highest IV. Carya glabra and Quercus rubra IV's also decreased in 1984,
while the IV of Acer saccharum increased to 22.3% (mainly due to an increase in
density). Ostrya virginiana IV increased the most from 1932 to 1984 of any species
present, with an increase from 2.6 to 21.3% (mainly due to an increase in density).
Carya ovata and C. cordiformis had the highest IVs of all species in 1931 on
the Deam plot (combined IVs 39.1%; Table 3). Only two other species (Quercus rubra
and Ulmus americana) had IVs of 10.0% or greater in 1931. In 1984, the IV for Carya
spp. was 12.0%; less than one-third of what it was in 1931. Quercus rubra showed
a slight increase in IV over this 53 year period. Ulmus spp. IV has remained the same;
however, the density for this genus has increased from 24.8 to 528.9 stems ha~ '. Ulmus
americana and U. rubra seedlings were combined in the U. rubra category due to the
difficulty in distinguishing between the two species at this early age. Acer saccharum
had the greatest IV increase from 1931 to 1984 (25.3% increase).
Only two species (Quercus rubra and Tilia americana) found on the ungrazed
Hoffman plot in 1931, had Importance Values greater than 10.0% (18.8 and 16.2%,
respectively; Table 4). Both of these species exhibited larger changes in IVs from 1931
to 1984; however, Quercus rubra IV increased to 31.0%, while Tilia americana IV
decreased to 9.0%. Fraxinus nigra also decreased in IV from 7.0 to 0.6%.
Size-class distributions for all species combined on grazed and ungrazed plots
appear similar in 1931 and 1984 (Figure 2). Stem numbers in most size-classes on the
ungrazed plots were greater than those on the grazed plots in 1931. In 1931, both
the grazed and ungrazed plots have a notable depletion of stems in the smallest size-
class and again in the 15.0-19.9 cm size-class. By 1984 this latter underrepresentation
of stems is prominent in the 25.0-29.9 cm size-class.
The size-class distribution for Acer saccharum and Ostrya virginiana combined
on the grazed plot for 1984 (Figure 3) is quite distinct from that of 1931, primarily
due to tremendous ingrowth of these species. Differences between these two yeas is
less striking on the ungrazed plot, although stem numbers are currently greater in the
smallest size-class. These two species made up nearly all of the stems in the lower
four classes, all species combined (Figure 2) in 1984.
Quercus spp. size-class distribution is similar for the grazed and ungrazed plots
in 1931, in that both distributions roughly resemble a bell-shaped curve (Figure 4).
In both cases no stems were found in the 1.0-4.9 cm size-class in 1931; however, there
were more stems in the classes from 5.0 to 49.9 cm on the ungrazed plot than on
222
Indiana Academy of Science
Vol. 94 (1985)
H V-
20 0 300 40 0 500 2600
size-class lower limit icm.dbhi
10 0 20 0 300 40 0
SIZE-CLASS LOWER LIMIT CM.D8H
500 >600
Figure 2. Size-class distribution (stems ha ')of all species combined in 1931 (and 1932)
and 1984 for grazed and ungrazed plots.
the grazed plot. There were approximately 70 stems ha~ ' of Quercus spp. in the 1 .0-4.9
and 5.0-9.9 cm size-classes combined on the grazed plots in 1984. This increase is due
to Quercus rubra ingrowth on Romey plot no. 66; Quercus spp. regeneration was absent
on the other plots.
Q D 1931
• • 1984
h — i — i — y
20 0 30 0 40 0
SIZE-CLASS LOWER LIMIT <CM OBH)
50 0 2:60 0
20 0 30 0 40 0 50 0
SIZE-CLASS LOWER LIMIT (CM. 08HI
Figure 3. Size-class distribution (stems ha+l) of Acer saccharum and Ostrya virgi-
niana in 1931 (and 1932) and 1984 for grazed and ungrazed plots.
Ecology
223
10 0 20 0 300 400 500
SIZE-CLASS. L0WEH LIMIT ICM.DBM)
Figure 4. Size-class distribution (stems ha ') of Quercus spp. in 1931 (and 1932) and
1984 for grazed and ungrazed plots.
Discussion
Lowered forest productivity is a known consequence of grazing (5). The effect
of grazing on various soil properties has been documented in many publications. Soil
physical properties (e.g., porosity and permeability) are especially affected (2 and 17).
Destruction of the forest floor, a vital part of the forest mineral and hydrologic cycles,
is also a consequence (3).
The current basal area of Deam woods (grazed) of 35.4 m2ha" ' is relatively high
compared to that for mature forests in the Central Hardwoods Region (1, 10 and 14),
although this woods had been grazed somewhat heavily in the early 1900s. But, such
comparisons are not entirely valid because of differences in the lower stem diameter
limit among studies (i.e., all stems) J> 1 cm in present study versus 10 cm in others).
This disparity in lower diameter limit leads to even more dubious comparisons of stem
densities. For example, the density in Deam woods of 3099 stems ha" ' is substantial-
ly higher than the average density of 284 stems ha- ' for old-growth forests in Indiana
(14). However, if only those stems J> 10.0 cm dbh are counted at Deam woods the
resulting density is 143 stems ha" '. Another reason for such discrepancies in basal area
and density between the present study and others cited is differences in stand age.
Natural thinning has probably occurred to a greater degree in the old-growth forests
previously mentioned compared to the younger Deam woods.
Basal area and density values are much lower at Romey woods partly because
site quality may be lowest of the three woods studied. Therefore, it appears that grazing
may have a longer and more detrimental effect on forest processes (reduced tree species
colonization, less growth, greater mortality, etc.) at Romey woods, although the graz-
ing intensity and/or high-grading may have also been severe enough to cause such
differences. Economically, less desirable timber species apparently have been favored
by the combination of poor site quality and past grazing. However, more research
is needed to evaluate site quality on all stands in this study.
Comparisons between plots at Deam and Romey woods are likewise difficult
because of initial differences in location, forest type, silvical condition, soils, etc.
224 Indiana Academy of Science Vol. 94 (1985)
Although we initially believed that the ungrazed plot (Hoffman woods) would serve
as a reasonable control, the 1931-32 compositional and structural data suggest that
this woods was also disturbed prior to plot establishment. Written records indicate
that this woods has been protected from grazing since the 1870s, but other disturbance
factors could have affected this woods.
Some authors (11, 14) have claimed that periods of stand disturbance could be
determined based on the size-class distribution of stems as depicted in Figures 2, 3,
4 (i.e., the size-class is plotted on the abscissa, the log of the number of stems in
that class is plotted on the ordinate; a plot of these values constitutes the negative
exponential distribution). Large deviations from the constantly decreasing straight line
supposedly indicate periods of disturbance; however, others (13 and 20) have objected
to such an affirmation. If this assumption is allowed, at least two major disturbances
are apparent in all plots prior to 1930, as shown by the substantial underrepresenta-
tion of stems in the 1.0-4.9 and 15.0-19.9 cm classes for the species combinations shown.
Although the lack of stems in the smallest size-class in 1931-1932 could be attributed
to grazing effects, this same phenomenon exists for the ungrazed plot. Possible reasons
for this parity include: (1) the ungrazed plot had actually been grazed in the late 1800s
to early 1900s; and/or (2) severe disturbance affected all stands similarly during the
same years. Photographs of the ungrazed plot in 1931 reveal that few small stems
existed in the understory, as in the grazed plots. Selective logging in all plots during
this period could have contributed to depletion of larger stems, but a more likely factor
is drought since the smallest trees seem to have been particularly susceptible. Climatic
records at the nearest weather station (Fort Wayne, Indiana) indicate that severe droughts
occurred periodically in the late 1800s to early 1900s (19). Such a climatic aberration
could explain some of the similarities in the size-class distributions between grazed
and ungrazed plots. There is also evidence that small fires had occurred in some of
these stands, according to Den Uyl's unpublished data.
The size-class distribution of Quercus spp. in both grazed and ungrazed plots
suggests that at least this component of these stands established within a relatively
narrow time period, i.e. the Quercus spp. are even-aged. This belief is based on a
comparison of the present results with Quercus size-class distributions of Schnur (15)
which represent even-aged stands at various ages over a range of site quality.
Initially, the increase in Ulmus spp. IV in all woods may seem surprising, con-
sidering the effect that Dutch elm disease and elm yellows has had on this genus in
Indiana (16). This increase results despite mortality of larger individuals because of
substantial ingrowth of U. americana and U. rubra. Similar density increases in these
species have been noted elsewhere (12).
Results from the 1981 survey reinforce some of Den Uyl's (8) findings. For in-
stance, he stated that Acer saccharum, Ulmus spp., Ostrya virginiana and Prunus serotina
colonization frequently occurred following grazing. These species are primarily light-
seeded or bird-dispersed and produce some seed each year to provide a constant supply
of propagules (8). These species are also fairly shade tolerant; therefore they can establish
under a dense canopy and persist, at least while in the seedling stage (9). Day and
Den Uyl (3) also state that unpalatable, 'weed' species (such as Ostrya virginiana) re-
main on the site and regenerate prolifically following grazing which excludes establish-
ment of more desirable timber species.
Openings of 0.10 to 1 acre (0.04-0.4 ha) are required for the reproduction of
most desirable tree species. Canopy densities of 70, 80 and 90% were present on Deam,
Romey and Hoffman plots, respectively at establishment (Table 1) which did not allow
sufficient sunlight to reach the forest floor for these shade intolerant species to establish
and persist. Quercus spp. probably did not reproduce on most of the plots for this
reason; furthermore, the Quercus rubra seedlings present on plot no. 66 were presumably
Ecology 225
the same individuals that Den Uyl noted in 1957 within openings created by the removal
of six large overstory trees in 1942 (8). Also, Den Uyl (8) showed photographs of
abundant Fraxinus americana reproduction on plot no. 67 of Romey woods. In 1984
only two individuals of this species were tallied on the entire 0.5 acre (0.2 ha) quadrat.
This decline in Fraxinus americana density is probably attributable to a decline in shade
tolerance as this species ages.
Due to site and historical differences among plots it is difficult to make specific
conclusions about long-term species response to release from grazing. Site quality is
an important factor to consider, since it undoubtedly affects the recovery of different
species to release from grazing. In order to better ascertain the effects of grazing on
forest composition and structure, it would be necessary to reestablish many more plots
with similar characteristics. Ideally, woodlots which contain contiguous grazed and
ungrazed plots should be utilized; however, rarely is such a condition available. Den
Uyl did establish some plots in woodlots that were divided by fence into both grazed
and protected sections; but, unfortunately these plots are no longer intact or they have
been severely perturbed since their establishment.
Literature Cited
1. Auten, J.T. 1941. Notes on old-growth forests in Ohio, Indiana and Illinois. U.S.
For. Serv. Exp. Stn. Tech. Note. No. 49, 8 p.
2. Chandler, R.F. 1940. The influence of grazing upon certain soil and climatic con-
ditions in farm woodlots. Jour. Amer. Soc. Agron. 32:216-230.
3. Day, R.K. and D. Den Uyl 1932. The natural regeneration of farm woods following
the exclusion of livestock. Purdue Univ. Ag. Exp. Sta. Bull. No. 368, Lafayette,
In. 47 p.
4. Den Uyl, D., O.D. Diller and R.K. Day. 1938. The development of natural
regeneration in previously grazed farmwoods. Purdue Univ. Ag. Exp. Sta. Bull.
No. 431. Lafayette, In. 28 p.
5. Den Uyl, D. 1944. The growth of timber in Indiana farmwoods, J. For. 42:169-174.
6. Den Uyl, D. 1947. Forest grazing in the Central States Region. Proc. Soc. Am.
For., pp. 255-261.
7. Den Uyl, D. 1958. A twenty year record of the growth and development of Indiana
woodlands. Purdue Univ. Ag. Exp. Sta. Res. Bull. No. 661, Lafayette, In. 51. p.
8. Den Uyl, D. 1961. Natural tree reproduction in mixed hardwood stands. Purdue
Univ. Ag. Exp. Sta. Res. Bull. No. 728, Lafayette, In. 19 p.
9. Fowells, H.A. (ed.) 1965. Silvics of forest trees of the United States. Ag. Hdbk.
No. 271, USDA, Washington, D.C., 762 p.
10. Held, M.E. and J.E. Winstead. 1975. Basal areas and climax status in mesic forest
systems. Ann. Bot. 39:1147-1148.
11. Johnson, F.L. and D.T. Bell. 1975. Size-class structure of three streamside forests.
Amer. J. Bot. 62:81-85.
12. Parker, G.R. and D.J. Leopold. 1983. Replacement of Ulmus americana L. in
a mature east-central Indiana woods. Bull. Torrey Bot. Club 110:482-488.
13. Robertson, P. A., G.T. Weaver and J. A. Cavanaugh. 1978. Vegetation and tree
species patterns near the northern terminus of the southern flood plain forest
Ecol. Monogr. 48:249-267.
14. Schmelz, D.V. and A. A. Lindsey. 1965. Size-class structure of old-growth forests
in Indiana. For. Sci. 11:258-264.
15. Schnur, G.L. 1937. Yield, stand, and volume tables for even aged upland oak
forests. USDA Tech Bull. No. 560, Wash. D.C., 87 p.
226 Indiana Academy of Science Vol. 94 (1985)
16. Schuder, D.J. 1955. Distribution of three important insect transmitted tree diseases.
Indiana Acad. Sci. Proc. 64:116-120.
17. Steinbrenner, E.C. 1951. Effect of grazing on floristic composition and soil pro-
perties of farm woodlands in southern Wisconsin. J. For. 49:906-910.
18. U.S. Dept. of Commerce, Bureau of Census. 1984. 1982 Census of agriculture.
Vol. 1 Geo. Area Pt. 14 Indiana, 443 p.
19. Visher, S.S. 1944. Climate of Indiana. Indiana Univ., Bloomington, In. 511 p.
20. West, D.C., H.H. Shugart, Jr., and J.W. Ranney. 1981. Population structure
of forests over a large area. For. Sci. 27:701-710.
Characteristics of Drumming Habitat of Ruffed Grouse in Indiana
Steven E. Backs,
Department of Natural Resources, Mitchell, Indiana 47446
Sean T. Kelly
U.S. Fish and Wildlife Service, Manchester, New Hampshire 03100
P. Decker Major
Department of Natural Resources, Mitchell, Indiana 47446
Brian K. Miller
Department of Environmental Protection, North Franklin, Connecticut 06254
Introduction
The drumming of the male ruffed grouse (Bonasa umbellus) is part of its ter-
ritorial and reproductive behavior. Drumming sites are the focal point of spring court-
ship activity and relatively easy to identify. Several studies have concentrated on describ-
ing habitat characteristics around drumming sites (Palmer 1963, Boag and Sumanik
1969, Stoll et al. 1975, Sousa 1978). Drumming sites are generally associated with dense
understories of young trees or shrubs (Boag 1976, Hale et al. 1982). Gullion (1977)
described optimum drumming habitat to be composed of 14,000-20,000 woody stems
per ha. The objective of this study was to describe the vegetative characteristics of
drumming sites used by ruffed grouse in Indiana.
Study Area
Drumming sites were studied on 3 areas. Thirty-two drumming sites were examined
on 517 ha of the Jasper-Pulaski State Fish and Wildlife Area in northwest Indiana.
Field work was conducted as part of an evaluation of ruffed grouse releases made
in 1970 and 1971 (Kelly 1971, Kelly and Kirkpatrick 1979). The area is composed of
two vegetative communities. The upland hardwood-brush community occurs on dry,
sandy ridges and is dominated by black, white, and red oaks (Quercus velutina, Q.
alba and Q. rubra). Brush consists of scrub oak, principally stunted black oak, sprouted
from burned hardwood areas. The lowland- woody association consists of moist areas
dominated by river brich (Betula nigra), quaking aspen (Populus tremuloides), and
pin oak (Q. paulstris).
Sixteen drumming sites were also examined on each of two study areas in
unglaciated, southcentral Indiana on Hoosier National Forest. One area was Happy
Hollow, 320 ha, located in Perry County; T4S, R1W, section 3. The other area, referred
to as the Maumee Grouse Study Area, located on the Jackson-Brown county line con-
sists of 335 ha; T7N, R2E, sections 11, 12, 13, and 14. Common upland species in-
clude red, black, white, and chestnut oaks (Q. prinus), American beech (Fagus
gradifolia), and hickories (Carya spp.). Common lowland species include ash (Frax-
inus spp.), yellow-poplar (Liriodendron tulipifera), elm (Ulmus spp.), sycamore (Plan-
tinus occidentalis), and river birch. Several pines (Pinus strobus, rubra, virginiana,
and echinata) are found in small plantations introduced by various public agencies.
Common understory species include blackberry (Rubus spp.), cherry (Prunus spp.),
sassafras (Sassafras albidum), ironwood {Qstrya virginiana), greenbriar (Smilax spp.),
flowering dogwood (Cornus florida), maples (Acer saccharum and rubrum), and sumac
(Rhus spp.).
Methods
Male grouse were located by listening for drumming and searching for the drum-
227
228 Indiana Academy of Science Vol. 94 (1985)
ming stage (the spot where a grouse habitually stands while drumming) from late-
March through mid-April. Good indications of an actively used drumming state are
an accumulation of fecal droppings on the stage, molted feathers, and a bare spot
of ground at the base of the log where leaves have been blown away by the drumming
performance. The physical characteristics of the drumming stage are generally not con-
sidered important (Gullion 1967, Boag and Sumanik 1969). Ruffed grouse are known
to use rock ledges, boulders, rock walls, moss mounds, upturned roots, and stumps
(Frank 1947, Bump et al. 1947). All stages used in this study were downed logs.
Characteristics of the vegetation directly surrounding used drumming logs were
compared to similar but unused logs located within 50 m of the used log and to the
surrounding vegetation sampled at 4 points 20 m from each used log in 4 cardinal
directions. The mean value of each vegetational parameter measured at the 4 20-m
points was used to represent the surrounding vegetation. Trees (woody species > 13
cm diameter at breast height, DBH) were tallied by point sampling at the drumming
stage with a 10-factor, basal area prism (Beers and Miller 1964). A similar point was
sampled at each unused log and at the 4 20-m points. A 0.002 ha circular plot (radius
= 2.5 m) centered at the drumming stage and the other sample points was used to
sample shrubs (wood species < 13 cm DBH). The herb layer vegetation was not sampled
since it is absent during the early spring and thus would not influence the selection
of drumming sites (Boag and Sumanik 1969, Palmer 1963). Tree and shrub frequen-
cies were reported as their proportional occurrence in the plots sampled. An analysis
of variance and Duncan's new multiple-range test (Steel and Torrie 1960) were used
to determine differences in the tree and shrub densities surrounding used logs, unused
logs, and 20-m plots.
Results and Discussion
Tree densities were similar (P >0.05; F = 1.21) at the three sample plots (Table
1). However, shrub densities were greatest at used drumming logs (P<0.01; F = 16.05)
and were the most important variable in determining drumming log use. High shrub
Table 1. Tree and shrub densities (stems per ha) surrounding used drumming logs,
unused logs, and at sample points 20 m from drumming logs used by ruffed grouse in
Indiana.
Variable Used Logs Unused Logs 20-m Plots
TREES
x 258 222 216 '
SE
Range
SHRUBS
x
SE
Range
OVERALL
x
SE
n
Range
'Any two means not underscored by the same line are significantly different; those underscored are not (P<0.05);
Duncan's New Multiple Range Test.
26.5
19.7
13.0
0-1,389
6-582
6-591
34.914
20,789
21,350
2,425.4
1,869.5
1,769.3
4,500-87,500
1,500-63,000
1,750-49,995
35,172
21,011
21,566
2,366.0
1,830.0
1,555.0
64
64
64
4,500-88,889
1,506-63,582
1,756-50,586
Ecology 229
densities were responsible for the overall woody stem densities being greatest at used
drumming logs (P<0.01; F = 16.96). The importance of a dense shrub layer in the
selection of drumming sites by ruffed grouse has been determined previously (Boag
and Sumanik 1969, Rusch and Keith 1971, Boag 1976, Stoll et al. 1979) and is further
supported by results of this study. The overall mean stem density for drumming logs
used in Indiana falls within the range of values reported elsewhere (Palmer 1963, Boag
and Sumanik 1969, Rusch and Keith 1971, Gullion 1977, Sousa 1978, Stoll et al. 1979,
Hale et al. 1982).
Species composition of the vegetation sampled generally reflected the overall com-
position of the study areas (Table 2). Oaks were the most common trees occurring
Table 2. Vegetative composition surrounding used drumming logs, unused logs, and
at sample points 20 m from logs used by ruffed grouse in Indiana.1
Used Logs Unused Logs 20-m Plots
Variable °/o % %
TREES
Quercus velutina 39 45 67
Quercus alba 47 67 77
Quercus palustris 23 17 45
Carya glabra 9 13 23
Quercus rubra 11 25 33
Populus spp. 12 8 24
SHRUBS
Prunus virginiana 36 25 41
Rubus spp. 34 25 53
Cornus florida 41 41 50
Sassafras albidum 45 48 72
Prunus serrotina 34 25 53
Smilax sqp. 23 30 33
Quercus alba 44 50 77
Ostrya virginiana 30 34 41
'Tree and shrub frequencies reported as their proportional occurrence in the plots sampled.
around used drumming logs in Indiana. Shrub species occurring at high frequencies
around used drumming logs were cherry, blackberry, flowering dogwood, sassafras,
and greenbriar. Although aspen is considered an important component of ruffed grouse
habitat in the Lake States (Gullion 1977) it occurred at relatively low frequency in
our sample plots. Overall, the vegetation surrounding used drumming logs reflected
the species composition of early serai or understory types indicative of the central hard-
wood forests of Indiana. Species composition is generally considered less important
in determining drumming site use than the physical structure of the vegetation (Stoll
et al. 1979, Hale et al. 1982).
Conclusions
Habitat around drumming logs used by ruffed grouse in Indiana is generally
characterized by high woody stem densities. The mean number of stems around 64
used drumming logs was 35,172 stems/ha, ranging from 4,500 to 88,889 stems/ha.
Stem densities around unused logs averaged 21,011 stems/ha ranging from 1,506-63,582
stems/ha. Stem densities at sample points 20 m from drumming logs averaged 21,566
stems/ha, ranging from 1,756-50,586 stems/ha. Differences in shrub densities separated
230 Indiana Academy of Science Vol. 94 (1985)
used from unused logs. Results from this study agreed with similar studies elsewhere,
indicating that the physical structure of the habitat, primarily the shrub layer, governs
drumming log selection.
Much of the work reported herein was supported by funds under Federal Aid
in Fish and Wildlife Restoration Act; Wildlife Research Project W-26-R, Indiana.
Jennifer Eckensberger is acknowledged for typing this manuscript.
Literature Cited
1. Beers, T.W. and C.I. Miller. 1964. Point sampling; research results, theory and
applications. Purdue Univ. Res. Bull. 786, 56 pp. West Lafayette, IN.
2. Boag, D.A. and K.M. Sumanik. 1969. Characteristics of drumming sites selected
by ruffed grouse in Alberta. J. Wildl. Manage. 33(3):621-628.
3. , 1976. The effect of shrub removal on occupancy of ruffed grouse drumm-
ing sites. J. Wildl. Manage. 40:105-110.
4. Bump, G., R.W. Darrow, F.C. Edminster, and W.F. Crissey. 1947. The ruffed
grouse: life history, propagation, management. New York State Cons. Dept. 915 pp.
5. Frank, W.J. 1947. Ruffed grouse drumming site counts. J. Wildl. Manage.
11(4):307-316.
6. Gullion, G.W. 1967. Selection and use of drumming sites by male ruffed grouse.
Auk 84:87-112.
7. , 1977. Forest manipulation for ruffed grouse. Trans. No. Amer. Wildl. and
Nat. Resour. Conf. 42:449-458.
8. Hale, P.E., A.S. Johnson, and J.L. Landers. 1982. Characteristics of ruffed grouse
drumming sites in Georgia. J. Wildl. Manage. 46(1): 1 15-123.
9. Kelly, S.T. 1977. Evaluation of a ruffed grouse reintroduction in northern Indiana.
M.S. Thesis, Purdue Univ., West Lafayette, IN.
10. and CM. Kirkpatrick. 1979. Evaluation of a ruffed grouse reintroduction
in northern Indiana. Wildl. Soc. Bull. 7(4):288-291.
11. Palmer, W.L. 1963. Ruffed grouse drumming sites in northern Michigan. J. Wildl.
Manage. 27(4):656-663.
12. Rusch, D.H. and L.B. Keith. 1971. Seasonal and annual trends in numbers of
Alberta ruffed grouse. J. Wildl. Manage. 35(4):803-822.
13. Sousa, P.J. 1978. Characteristics of drumming habitat of ruffed grouse (Bonasa
umbellus) in Grafton, Vermont. M.S. Thesis, Univ. of Vermont, Burlington, 134
pp.
14. Steel, R.G.D. and J.H. Terrie. 1960. Principles and procedures of statistics.
McGraw-Hill Book Co., New York, N.Y. 481 pp.
15. Stoll, R.J., M.W. McClain, R.L. Boston, and G.P. Honchol. 1979. Ruffed grouse
drumming sites characteristics in Ohio J. Wildl. Manage. 43(2):324-333.
The Foraging Ecology of Some Bats in Indiana
Virgil Brack, Jr.
Department of Forestry and Natural Resources
Purdue University, West Lafayette, Indiana 47907
Introduction
Twelve species of Chiroptera have been reported from Indiana (23), but Myotis
austroriparius (southeastern myotis) and Plecotus rafinesquii (Rafinesque's big-eared
bat) are extremely rare. Only 1 colony of Myotis grisescens (gray bat) is known from
Indiana (7). Pipistrellus subflavus (eastern pipstrelle) is relegated to southern Indiana
which was not glaciated by the most recent (Wisconsinan) glaciation. Nycticeius humeralis
(evening bat) is uncommon in Indiana, with only a few nursery colonies containing
adult females and young of the year having been located in Indiana (23). Lasiurus
cinereus (hoary bat) is widely distributed but rarely common at any locale. Males are
rare in the state (23). Lasionycteris noctivagans (silver-haired bat) is found in Indiana
only in spring and autumn as a migrant (23). During the summer, the sexes of Myotis
sodalis (Indiana bat) are allopatric within the state. Myotis keenii (Keen's bat), Myotis
lucifugus (little brown myotis), Lasiurus borealis (red bat), and Eptesicus fuscus (big
brown bat), occur throughout Indiana.
Several species of bats can frequently be found within the same area or same
habitat. This study was undertaken to determine the foods eaten, habitats or parts
of habitats used, and times of activity, of each of the 10 species. Results of 2 of these,
M. sodalis and M. keenii, will be reported upon elsewhere.
Materials and Methods
Bat Capture
Bats were captured during the season of reproduction (15 April to 15 August)
in wooded upland (14 sites; 89 net nights) and riparian areas (21 sites; 61 net nights)
throughout Indiana. Mist nets were "stacked" and run on a rope pulley system to
close off all flight space from the forest floor or stream surface up to the canopy.
Capture time and height, and the sex, age, and reproductive condition were noted
for each bat. Chi-square tests were used to determine randomness of activity during
the night (divided into the periods: dusk to 22:00 h/22:00 to 24:00 h/24:00 to 02:00
h/02:00 h to dawn), height of catch, and habitat (riparian/nonriparian) of catch.
Heights of capture correspond to the 3 foliage layers (22): shrub (< 0.6 m), canopy
(usually > 7.6 m, depending of the vegetation), and the understory or subcanopy.
Catch per habitat was tested by both catch per net night and by catch per net site.
Feces were sometimes collected from bats captured at caves.
Fecal Analysis
The analysis method used was that of Brack and LaVal (5). Briefly, insect parts
were identified from the feces, and quantified by an estimate of percent volume. When
the diets of 2 or more bats was combined each bat contributed equally to the combined
diet. An analysis of variance was conducted on an arcsine-transformation of the date
to compare diets among dates of sampling or sample groups. Statistical analyses
were completed on Digital Equipment Corporation PDP-1 1/70 computer systems using
a version of SPSS (24) from Northwestern University.
A diet diversity index (DDI) was calculated for each species, and for some species
by date, sex, and age of sample. The diversity index used was that of MacArthur
(21): DDI = l/]£ Pj2, where P,, P2 ... were the proportions of each insect order in the
diet.
231
232
Indiana Academy of Science
Vol. 94 (1985)
Results
Myotis lucifugus
Adult males were captured at caves during the summer but few individuals roosted
there. No females or juveniles were caught at caves until late in the season. Only 4
adult males, but 34 adult females and 19 juveniles, were caught outside the cave region.
County records were established for Porter, Jasper, Starke, and LaPorte counties.
The catch of M. lucifugus was similar in riparian and nonriparian habitat when
considering catch per site, but more bats were caught in riparian habitat when con-
sidering catch per net night (Table 1). In riparian habitat, catch was concentrated in
the under story; in nonriparian habitat, catch was too small to test (Table 2). Catch
was distributed evenly throughout the night (Table 3).
Myotis grisescens
Only 7 lactating females and 4 males were netted, all in riparian habitat. Two
were caught in the subcanopy layer and 9 in the shrub layer. The catch appeared bimodal
with bats captured early and late in the night, but the sample was too small to test.
A total of 84 fecal pellets, 48 from females and 36 from males, were analyzed. Males'
and females' diets were similar. Trichopterans formed 56.0% of the diet, coleopterans
23.3%, lepidopterans 11.3%, dipterans 5.8%, hymenopterans 1.2%, plecopterans 0.5%,
and homopterans 0.7%. Fewer homopterans were eaten by females (P = 0.040). The
diet diversity index (DDI) was 5.79 for both sexes and 5.18 when combined.
Lasiurus borealis
A total of 85 individuals were caught; 6 unsexed, unaged bats escaped from nets
before they could be removed. Four bats were caught at caves; 2 adult males and
2 juveniles. The adult male (N = 22) and female (N = 21) catch was nearly equal. Lasiurus
borealis was caught at more sites than any other species (Table 1). Catch was equal
in riparian and nonriparian habitat when considered by net site, but greater in riparian
habitat when considered by net night (Table 1). In riparian habitats the catch was
greatest in the subcanopy layer but equal in the subcanopy and canopy layers in
nonriparian habitat (Table 2). On 2 occasions, pastures dotted with small trees con-
tained large numbers of L. borealis foraging several times the height of existing vegeta-
tion. This bas was most frequently caught during the dusk and dawn periods, representing
a bimodal activity period (Table 3).
Table 1 . Bat catch by net night and by catch site in riparian (R) and nonriparian (NR)
habitats. Statistics are based on 150 net nights (61 riparian, 89 nonriparian) at 35 catch
sites (21 riparian, 14 nonriparian).
Species
Total Catch
Ni
amber of Bats Caught
N
umber of Sites Where
Caught
Proportion
Bats/Net
of
Night
Sites
R
NR
X2
P
R
NR
X2
P
M. lucifugus
0.3867
.4000
50
8
49.843
0.000
10
4
0.762
0.383
M. grisescens
0.0733
.1143
11
0
16.047
0.000
4
0
L. borealis
0.6133
.8000
56
36
15.557
0.000
18
10
0.214
0.643
L. cinereus
0.1200
.2857
4
14
2.539
0.111
4
6
1.667
0.197
E. fuscus
1.7133
.7714
110
147
0.484
0.487
14
13
0.747
0.388
P. subflavus
0.0733
.1429
11
0
16.047
0.000
5
0
N. humeralis
0.0333
.0286
0
5
0
1
L. noctivagans
0.0133
.0286
0
2
0
1
Ecology
233
Table 2. Bat catch at shrub (1), subcanopy (2), and canopy (3) levels in riparian,
nonriparian, and both habitats combined.
Species
Riparian
Nonriparian
Combined
1
2
3
X2
P
1
2
3
X2
P
X2
P
M. lucifugus
5
40
3
54.125
0.000
0
5
3
55.750
0.000
M. grisescens
9
2
0
0
0
0
L. boreal is
4
31
11
25.609
0.000
0
16
16
16.000
0.000
35.615
0.000
L. cinereus
0
3
2
0
6
7
6.615
0.037
9.000
0.011
E. fuscus
7
55
13
54.720
0.000
1
95
34
104.969
0.000
157.532
0.000
P. subflavus
0
9
2
0
0
0
N. humeralis
0
0
0
0
0
5
L. noctivagans
0
1
0
0
1
2
Feces, totaling 318 pellets, from 59 bats were analyzed. Coleoptera (42.5%) and
Lepidoptera (37.5%) were the major prey. Insects of the orders Diptera and Homoptera
were each 4.3% of the diet, Plecoptera 2.1%, Neuroptera 1.8%, Hymenoptera 0.9%,
and Trichoptera 0.5% of the diet. The following families of Coleoptera were iden-
tified in the feces: Scarabaeidae 10 times, Elateridae 8 times, Silphidae 3 times, and
Carabidae once. Curculionidae remains were identified 3 times; 2 of these were the
Asiatic oak weevil, Cyrtepistomus castaneus. The diets of males, females, and juveniles
were similar. There was no difference between the diets from bats captured in dif-
ferent years. Dietary variation of bats captured at widely separated localities was also
low, although consumption of Neuroptera varied (P = 0.011). DDI's varied between
2.00 and 6.13. The overall DDI was 5.07.
Lasiurus cinereus
Five adults (1 male), 12 juveniles, and 1 unsexed unaged bat were caught. County
records were established for Porter, Steuben, and Noble counties. The adult male is
only the second known from the state. There was no difference between the numbers
of bats caught in riparian and nonriparian habitats, either by site or by net night (Table
1). In nonriparian habitat catch was divided between the canopy and subcanopy; riparian
catch was too small to test (Table 2). Bats were caught throughout the night (Table 3).
Twelve feces were collected from an adult female who had eaten only hymenopteran
insects. Diets of 8 juvenile bats, determined from 37 fecal pellets, varied widely. Six
had eaten diets containing more than 90% coleopterans. The remainder of their diets
Table 3. Bat catch per species during four periods between dusk and dawn.
Sunset
22:00 h
24:00 h
02:00 h
to
to
to
to
Species
22:00 h
24:00 h
02:00 h
Sunrise
X2
P
M. lucifugus
18
19
16
9
3.935
0.269
M. grisescens
3
2
0
6
L. boreal is
34
18
10
20
14.585
0.002
L. cinereus
2
8
3
5
4.667
0.198
E. fuscus
112
69
22
42
74.233
0.000
P. subflavus
4
2
4
1
N. humeralis
0
3
1
1
L. noctivagans
1
1
0
2
234 Indiana Academy of Science Vol. 94 (1985)
were lepidopterans. Two bats ate predominantly lepidopterans (83.6 and 96.3%) but
both also consumed some coleopterans (3.8 and 15.0%). Carabidae (Order: Coleoptera)
were identified 6 times. Individual bats also ate insects belonging to the orders: Diptera
(5.0%), Homoptera (1.3%), and Orthoptera (1.0%). The DDI was ,2.42.
Eptesicus fuscus
This bat is common statewide and was most frequently caught (Table 1). Four
nursery colonies were located, 1 each in Shelby, Hamilton, St. Joseph, and Miami
counties. At the caves, a few males could be caught as they came to night roost.
Sometimes 1 or more bats would use the same roost spot night after night, beneath
which was a notable feces accumulation. The catch of E. fuscus was similar in riparian
and nonriparian habitats (Table 1), and in both habitats the catch was largest in the
subcanopy layer (Table 2). Most bats were caught in the 2 periods from dusk to 24:00
h, with the smallest catch from 24:00 to 02:00 h (Table 3).
Pipistrellus subflavus
Males were captured at caves during summer sampling; females and juveniles
were not. Only 11 individuals (5 females) were netted away from caves, all in riparian
habitat. Two males caught over the Salamonie River, Wabash County represent both
the northern most Indiana record and a county record. The sample was too small
to test, but most captures were in the understory (Table 2). The catch appeared
distributed throughout the night.
Feces were analyzed from 23 bats. The diet contained 33.0% dipterans (both
Chironomidae and Muscidae were each identified once), 19.7% trichopterans, 14.1%
coleopterans (Elateridae was identified 9 times; Curculionidae, 2 of which were Asiatic
oak weevils, 8 times; Scarabaeidae 6 times; and Silphidae 3 times), 13.6% lepidopterans,
12.0% homopterans, 3.0% hymenopterans, 2.6% neuropterans, and 0.1% plecopterans.
The DDI of males and females were similar; the combined DDI was 6.68.
Nycticeius humeralis
Two females and 3 juveniles, were caught in 1980 in a Montgomery County upland
woodlot; all were caught in the canopy layer after 22:00 h. The females ate 69.6%
coleopterans, 29.1% lepidopterans, and 1.2% homopterans, while the juveniles at 68.9%
coleopterans, 9.2% dipterans, 14.9% homopterans, 5.3% trichopterans, 1.5%
hymenopterans, and 0.2% hemipterans. The combined DDI was 5.26.
Lasionyceteris notivagans
Two adult males were caught in Miami County on 3 June 1981 from the canopy
layer of an upland woodlot. A third male was caught in Tippecanoe County on 18
June 1983 from the subcanopy of riparian habitat. Thus all 3 represent later springs
records than previously recorded in Indiana, i.e., 28 May (23). A juvenile was caught
on 8 September 1981 from the subcanopy of an upland woodlot. These four captures
were scattered throughout the night (Table 3).
Feces were collected only from adult males. All ate dipterans (55.2%), neuropterans
(22.1%), and lepidopterans (9.3%); one individual had also eaten insects belonging
to the Coleoptera, Trichoptera, and Hymenoptera. The DDI was 4.80.
Discussion
Myotis lucifugus has frequently been found foraging low over pond and stream
surfaces (13, 9, 2, 1, 10), and food habits studies have further substantiated this behavior
(9, 2, 1). In the present study, M. lucifugus frequented subcanopy riparian habitat,
and was active throughout the night. In Iowa (16) the species was active early but
almost totally inactive the latter half of the night.
Ecology 235
Chemiluminescently tagged M. grisescens in Missouri foraged largely in riparious
areas, just over the water surface (18). The habitat and height of captures in the pre-
sent study concur with those findings, as does the diet with that in Missouri (20),
emphasizing aquatic based prey.
Foraging by L. borealis has been reported mainly from high over trees and pastures
(19, 18). Prey reported previously (26, 27, 28, 8) and herein have been largely ter-
restrial. Inconsistent with this, more bats per net night were caught in riparian habitats.
One logical explanation for this discrepancy is that riparian captures, mostly in the
subcanopy, represent use of this space as a travel lane. As in Iowa (16), activity was
greatest during early evening.
Although homopterans were frequently a small part of the L. borealis diet,
Whitaker (28) and Brack et al. (8) found they sometimes constitute major parts of
the diet. Whitaker (28) also found larger percentages of Orthoptera in the diet. However,
similarities to past studies (26, 27, 28, 8), and comparisons among sex, age, and temporal
subgroups of this study indicate a relatively stable diet composed largely of terrestrial prey.
It is probable that L. cinereus, like L. borealis, frequents waterways primarily
as travel lanes. This is supported by present and past food habit studies (3, 4, 27,
28, 30, 8), and past foraging observations (18, 11, 30, 23). L. cinereus has been referred
to as a moth specialist (3, 4), although a variety of other prey has been reported (27,
28, 30, 8). The species has a robust jaw and a skull morphology suitable for eating
hard-bodied insects (12). In this study, most prey were hard-bodied; most individuals'
diets contained small percentages of soft-bodied (Lepidoptera) prey. Two bats ate
predominantly lepidopteran prey. In British Columbia (11) and Iowa (16) L. cinereus
was active late at night, temporally separating the foraging of the 2 Lasierus species.
Typically, the diet of E. fuscus contains large proportions of hard-bodied insects,
especially coleopterans (14, 25, 3, 4, 28). Since aquatic insect species are predominantly
soft-bodied, it appears that E. fuscus uses open understory waterways for travel and
feeds predominantly in uplands. Although catch was greatest in the understory, E.
fuscus also uses the canopy and higher air spaces while foraging (25, 11). In Iowa
(16) and British Columbia (11), as in Indiana, E. fuscus foraged predominantly early
in the evening.
Whitaker (28) reported a diet for P. subflavus similar to that reported here, with
a wide diversity of prey items, including terrestrial and aquatic species. In Missouri,
trichopterans predominated in the diet (20), and luminescently tagged bats foraged
over or near streams (18). Data from the present study complement those findings;
all captures were in the subcanopy and canopy of riparian habitat.
Limited observations (18, 23) indicate that N. humeralis frequents tree crowns
of open and early successional wooded pastures and floodplains. This bat has a cranial
and jaw morphology of intermediate robustness, appropriate for some types of hard-
bodied prey (12), and has been reported to eat largely Coleoptera, Homoptera,
Hymenoptera, and Hemiptera, as well as Lepidoptera and Diptera (27, 28). Though
again limited, the data collected on this species encourages a similar interpretation.
In general, L. noctivagans forages in or near woodlands adjacent to streams or bodies
of water (17), and has post dusk and predawn feeding periods (16, 15). Past dietary samples
are small but include representatives of the Lepidoptera, Hemiptera, Coleoptera, Diptera,
Trichoptera, and Isoptera (28, 29, 15). Similarly, small dietary samples in this study
contained neuropterans, and lepidopterans as major components.
In summary, 3 of the species of bats studies rely heavily upon a riparian environ-
ment. M. grisescens foraged low over water, M. lucifugus was caught in the understory,
and P. subflavus foraged around the riparian canopy and understory. The 2 Myotis
species eat aquatic prey. Lasiurus borealis, L. cinereus, and E. fuscus frequent the
236 Indiana Academy of Science Vol. 94 (1985)
riparian understory but do not forage there. They likely used it as a travel lane. E.
fuscus feeds on coleopterans and is frequently caught in the upland understory, while
both lasurines feed around and above woodland canopy. Because of a lack of data
in this and other studies, the foraging ecology of TV. humeralis and L. noctivagans
cannot be accurately characterized.
Acknowledgments
The majority of financial support was provided by the U.S. Forest Service, North
Central Forest Experiment Station. Many individuals provided field support, in par-
ticular Virgil R. Holmes spent many long hours in service. Bobby Witcher was a cons-
tant companion. George P. McCabe and his students provided statistical help. Russell
E. Mumford and Harmon P. Weeks provided equipment, encouragement, advice, and
constructive criticism, and read various parts of the manuscript. Research was con-
ducted under federal endangered permits PRT 2-4988 and PRT 2-9170 and appropriate
Indiana state permits.
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1. Anthony, E.L.P., and T.H. Kunz. 1977. Feeding strategies of the little brown
bat, Myotis lucifugus, in southern New Hampshire. Ecology, 58:775-786.
2. Bel wood, J.J., and M.B. Fenton. 1976. Variation in the diet of Myotis lucifugus
(Chiroptera: Vespertilionidae). Canadian J. Zool., 54:1674-1678.
3. Black, H.L. 1972. Differential exploitation of moths by the bats Eptesicus fuscus
and Lasiurus cinereus. J. Mamm., 53:598-601.
4. Black, H.L. 1974. A north temperate bat community. Structure and prey popula-
tion. J. Mamm., 55:138-157.
5. Brack, V., Jr., and R.K. LaVal. 1985. Food habits of the Indiana bat in Missouri.
J. Mamm., 66:308-315.
6. Brack, V., Jr., and R.E. Mumford. 1984. The distribution of Pipistrellus subflavus
and the limit of the Wisconsinan glaciation: an interface. Amer. Midland Nat.,
112:397-401.
7. Brack, V., Jr., R.E. Mumford, and V.R. Holmes. 1984. The gray bat {Myotis
grisescens) in Indiana. Amer. Midland Nat., 111:205.
8. Brack, V., Jr., S. Taylor, and V.R. Holmes. 1984. Bat captures and niche par-
ticipating along portions of three rivers in southern Michigan. Mich. Acad.,
16:391-399.
9. Buchler, E.R. 1976. Prey selection by Myotis lucifugus (Chiroptera: Vesper-
tilionidae). Amer. Nat., 110:619-628.
10. Fenton, M.B., and G.P. Bell. 1979. Echolocation and feeding behaviour in four
species of Myotis (Chiroptera). Canadian J. Zool., 57:1271-1277.
11. Fenton, M.B., C.G. Van Zyll DeJong, G.P. Bell, D.B. Campbell, and M. Laplante.
1980. Distribution, parturition dates, and feeding of bats in south-central British
Columbia. Can. Field-Nat., 94:416-420.
12. Freeman, P.W. 1981. Correspondence of food habits and morphology in insec-
tivorous bats. J. Mamm., 62:166-173.
13. Griffin, D.R. 1958. Listening in the dark: The acoustic orientation of bats and
men. Yale Univ. Press, New Haven, 413 pp.
14. Hamilton, W.J., Jr. 1933. The insect foot of the big brown bat. J. Mamm.,
14:155-156.
15. Jones, J.K., Jr., R.P. Lampe, C.A. Spenrath, and T.H. Kunz. 1973. Notes on
the distribution and natural history of bats in southeastern Montana. Occas. Papers
Mus. Texas Tech. Univ., 15:1-12.
Ecology 237
16. Kunz, T.H. 1973. Resource utilization: Temporal and spatial components of bat
activity in central Iowa. J. Mamm., 54:14-32.
17. Kunz, T.H. 1982. Lasionycteris notivagans. Mammal. Species, 172:1-5.
18. LaVal, R.K., R.L. Clawson, M.L. LaVal, and W. Caire. 1977. Foraging behavior
and nocturnal activity patterns of Missouri bats, with emphasis on the endangered
species Myotis grisescens and Myotis sodalis. J. Mamm., 58:592-599.
19. LaVal, R.K., and M.L. LaVal. 1979. Notes on reproduction, behavior, and abun-
dance of the red bat, Lasiurus borealis. J. Mamm., 60:209-212.
20. LaVal, R.K., and M.L. LaVal. 1980. Ecological studies and management of
Missouri bats, with emphasis on cave-dwelling species. Missouri Dept. Conserv.
Terrest. Series, 8:1-53.
21. MacArthur, R.H. 1972. Geographical ecology. Harper and Row, New York, 269
PP.
22. MacArthur, R.H., and J.W. MacArthur. 1961. On bird species diversity. Ecology,
42:594-598.
23. Mumford, R.E., and J.O. Whitaker, Jr., 1982. Mammals of Indiana. Bloom-
ington Univ. Press, Indiana, 537 pp.
24. Nie, N.H. et al. 1975. Statistical package for the social sciences. Second ed.
McGraw-Hill, St. Louis, Missouri, 675 pp.
25. Phillips, G.L. 1966. Ecology of the big brown bat (Chiroptera: Vespertilionidae)
in northwestern Kansas. Amer. Midland Nat., 75:168-198).
26. Ross, A. 1961. Notes of food habits of bats. J. Mamm., 42:66-71.
27. Ross, A., 1967. Ecological aspects of the food habits of insectivorous bats. Proc.
West. Found. Vertebr. Zool., 1:205-263.
28. Whitaker, J.O., Jr. 1972. Food habits of bats from Indiana. Canadian J. Zool.,
50:877-883.
29. Whitaker, J.O., Jr., C. Maser, and L.E. Keller. 1977. Food habits of bats of
western Oregon. Northwest Sci., 51:46-55.
30. Zinn, T.L., and W.W. Baker. 1979. Seasonal migration of the hoary bat, Lasierus
cinereus, through Florida. J. Mamm., 60:634-635.
Legal Game Harvest by Indiana Landowners Hunting without a License
John S. Castrale
Indiana Division of Fish and Wildlife
Mitchell, Indiana 47446
and
Robert E. Rolley and William J. Pfingsten
Indiana Division of Fish and Wildlife
Bloomington, Indiana 47401
Monitoring the harvest of game animals is important to wildlife management
agencies. Annual harvest figures provide information on the population status and
distribution of game species and the popularity of various game species to sportsmen.
This information can then be used by natural resources agencies to guide management
efforts.
Indiana state law requires all hunters to purchase a license to take game within
the framework of established seasons and regulations. Landowners and tenants who
hunt solely on their own land are exempt from purchasing a license but are still bound
by hunting regulations. Surveying licensed hunters to determine game harvest is relatively
straightforward because names and addresses can be obtained from receipt books main-
tained by vendors selling hunting licenses. The Indiana Division of Fish and Wildlife
samples licensed hunters annually in this manner to determine game harvest. Deter-
mining harvest by unlicensed hunters is more difficult but is necessary to obtain more
accurate estimates of the total harvest.
This paper reports on the relative contribution of unlicensed landowners to the
game harvest during the 1981 hunting season in Indiana. Unlike the survey of licensed
hunters which was initiated in the 1940s and has been conducted annually in its pre-
sent form since 1976 (Eisenhauer 1977a), the landowner survey is conducted at 5-year
intervals with the first beginning in 1976 (Eisenhauer 1977b). Besides providing harvest
estimates, the present survey gives information about the willingness of landowners
to allow hunting on their land.
Methods
Landowners surveyed were selected from a list of 73,000 farm operators main-
tained by the Agricultural Experiment Station at Purdue University in cooperation
with the Statistical Reporting Service of the United States Department of Agriculture.
This list contains farms with minimum annual sales of $1,000 in agricultural products.
There were approximately 89,000 farms in Indiana in 1981 (United States Department
of Agriculture 1982). A survey form was mailed to 12,196 Indiana landowners and
tenants. The 1976 landowner survey utilized a single mailing, so a similar procedure
was used in the present survey to maintain conformity. The postcard survey form con-
tained 8 questions dealing with land ownership, hunting by family members, and whether
hunting for small game and white-tailed deer (Odocoileus virginianus) was allowed
by the landowner (Castrale and Pfingsten 1982). A table was provided to record animal
harvest by species for each family member who hunted without a license on their
property.
Because sportsmen typically overestimate their harvests (Atwood 1956, Eisenhauer
1977c, Filion 1980), bias correction factors derived from a previous study (Pfingsten
1980) were used to arrive at final harvest estimates. Correction factors used were as
follows: squirrels (Sciurus niger, S. carolinensis), 0.588; eastern cottontail (Sylvilagus
239
240 Indiana Academy of Science Vol. 94 (1985)
floridanus), 0.543; northern bobwhite (Colinus virginianus), 0.469; ring-necked phea-
sant (Phasianus colchicus), 0.738; American woodcock (Scolopax minor), 0.471; and
ruffed grouse (Bonasa umbellus), 0.812. Correction factors for white-tailed deer are
calculated annually and for 1981 this figure was 0.832 (John C. Olson, personal
communication).
Statistical treatments included chi-square goodness-of-fit tests, chi-square tests
for independence and Spearman rank correlation coefficients.
Results and Discussion
Of 12,196 questionnaires mailed to farm operators, 3,095 (25.4%) were returned
and usable and form the basis of this report. Participation in hunting by farm households
showed a slight (37% vs. 34%) but significant (X2 = 8.49, df = 1, P <0.05) increase
over the previous survey (Eisenhauer 1977b). Hunting households averaged 1.69 hunters
in 1 98 1 . The number of households reporting hunting exclusively on their own land without
purchasing a license also showed a significant increase (21% vs 15%, X2 = 34.61,
df = 1, P<0.001) in 1981 over 1976. Households averaged 1.44 unlicensed hunters.
Projecting these figures, hunting landowners numbered 55,865 or 20% of Indiana sport-
smen who pursued deer and small game and 10% (27,562) of the state's estimated
283,682 deer and small game hunters were landowners hunting without a license.
Squirrels and eastern cottontails were the most commonly hunted game species
(Table 1). These mammals also experienced the highest seasonal harvests per landowner
which resulted in the greatest total harvests of all species. Squirrels and rabbits were
also popular with licensed hunters (Rolley 1984), however, so the contribution of lan-
downers to the total harvest of these species was actually the least of all species (Table
1). White-tailed deer were also frequently hunted by landowners, but the total kill
was much lower than most other species due to more restrictive bag limits. American
woodcock and ruffed grouse were of least interest to landowners, although the pro-
portional harvest of woodcock was high.
Landowners not purchasing a hunting license appeared to be less avid, or less
experienced hunters than licensed sportsmen. With the exception of deer, seasonal
harvests by landowners for each species averaged 65% (range = 58-82%) below licensed
hunters, assuming each household represented 1.44 hunters. The mean number of white-
tailed deer taken by landowners was higher than that of licensed hunters (Olson 1981),
which indicates more interest in deer, or possibly a more utilitarian attitude toward
sport hunting. Landowners more interested in hunting may buy a license in order to
increase their hunting opportunities or to contribute to the management of wildlife
Table 1 . Harvest statistics by landowners and tenants hunting only on their own property
during the 1981-1982 hunting season in Indiana.
Harvest by
State harvest
Seasonal
Seasonal
landowners
accounted for by
Landowners
harvest
harvest (no./
Species hunted
(no. ± SE)
landowners (%)
hunting (%)
(no. /landowner)
licensed hunter)
Squirrels
45,515 ± 4,049
4.5
15.5
3.30
5.59
Eastern cottontail
33,384 ± 2,699
4.9
19.5
1.92
3.39
Northern bobwhite
6,452 ± 620
6.6
7.2
1.01
1.66
American woodcock
2,357 ± 521
11.8
4.2
0.62
1.35
White-tailed deer
2,250 ± 264
12.4
11.4
0.27
0.14
Ring-necked pheasant
1,610 ± 429
7.3
5.4
0.34
0.66
Ruffed grouse
1,197 ± 407
5.0
3.7
0.37
1.42
Trom Rolley (1984) and Olson (1981).
Ecology
241
resources. Therefore, unlicensed hunters may hunt less often. Persons who purchase
a license may be more motivated to hunt even if they restrict their efforts to their
own property.
Changes in the harvests of deer and some small game species have been dramatic
between survey periods (Eisenhauer 1977b). The harvest of ruffed grouse has shown
the greatest increase (greater than an order of magnitude) since 1976. This species has
undergone natural range expansion supplemented by the establishment of new popula-
tions by transplanting (Backs 1984). Hunting opportunities for ruffed grouse have also
increased with 13 counties open to hunting in 1981 compared with 9 in 1976. Grouse
hunting in Indiana is a relatively young sport, and its popularity is rapidly increasing.
The estimated harvest of American woodcock by unlicensed hunters has almost tripl-
ed, although harvests by licensed hunters have declined about 50% between comparable
years. Deer populations have increased in recent years (John C. Olson, personal com-
munication), and the estimated number of white-tailed deer killed by landowners doubled
over 1976. Declines in the kill of northern bobwhite and ring-necked pheasants are
evident, and have been blamed on reduced populations due to unfavorable winters
in the late 1970s as well as to loss of habitat. The harvest of squirrels have shown
a substantial increase, while fewer rabbits were taken by landowners hunting without
a license.
Harvest figures show regional differences (Table 2) which are primarily due to
the relative abundance and distribution of each game species in the state. Tree squir-
Table 2. Regional distribution of game harvest by landowners hunting on their own
property. Values given are percentages of total harvest for each species.
Region
Northwest
Northeast
Central
Southwest
South-central
Southeast
(425 )a
(721)
(609)
(351)
(445)
(528)
Landowners
responding
(3,094)a
13.8
23.4
19.9
11.4
14.4
17.1
Non-licensed
hunters (897)
10.6
25.3
13.5
12.2
17.7
20.7
Species
harvested
Squirrels
(2,519)
6.4
15.6
6.0
10.1
39.2
22.6
Eastern
cottontail
(2,003)
11.0
19.7
8.6
19.4
25.6
15.8
Northern
bobwhite
(448)
14.5
2.5
3.1
47.8
20.8
11.4
White-tailed
deer (88)
12.5
29.5
4.5
5.7
23.9
23.9
American
woodcock
(163)
8.0
54.6
14.1
6.7
4.9
11.7
Ring-necked
pheasant (71)
43.7
0.0
28.2
1.4
8.5
18.3
Ruffed grouse
(48)
0.0
0.0
25.0
0.0
52.1
22.9
Sample size.
242
Indiana Academy of Science
Vol. 94 (1985)
rels are more common in the forested areas of southern Indiana. Changes in the regional
harvests of northern bobwhite reflect more severe declines suffered by this species in
northern Indiana (Castrale 1985). White-tailed deer populations are traditionally greatest
in south-central Indiana, but northeastern and southeastern populations appeared to
have increased. Complaints of deer damage to agricultural crops received by the Divi-
sion of Fish and Wildlife have become common from these areas. Ring-necked pheasants
are still principally harvested in the Northwest, but declines in this region as well as
the Northeast are apparent. Releases of pheasants by conservation clubs and individuals
throughout the state may help explain patterns of harvest in other areas. Ruffed grouse
are no longer being harvested strictly from south-central Indiana due to the opening
of other areas to grouse hunting.
Hunting of small game was allowed by 76% of the landowners responding but
only 51% permitted deer hunting. This category of response was the major reason
for the highly significant difference (X2 = 455, df = 3, P<0.001) in the patterns
of the permission categories for allowing hunting of deer and small game. Whether
or not hunting is allowed on a landowner's property is related to the amount of pro-
perty owned (Figures 1, 2). Large farm operators are more likely to allow hunting
of both deer (rho = 0.88, df = 6, P< 0.001) and small game (rho = 0.94, df =
6, P<0.001). With increasing farm size, landowners were more likely to allow small
game hunting by others (rho = 0.88, df = 6, P<0.001) and were more likely to hunt
deer themselves or allow friends of the family to hunt deer (rho = 0.88, df = 6,
P<0.001).
50
45
40
o
en
LU 7r
Q_ 35
Family & Friends
.->o.
30
,-o-
__._-.-cr"
/
p <r"
Others
'---</
/
0-
25-
50-
75-
100-
150-
200-
25
50
75
100
150
200
300
>300
FARM SIZE (ACRES)
Figure 1. Relationship of farm size with percentage of sampled landowners allowing
small game hunting by family and friends and other hunters.
Ecology
243
35
30
25
c_>
20
Fami ly & Friends
0-
25-
50-
75-
100-
150-
200
25
50
75
100
150
200
300
>300
FARM SIZE (ACRES)
Figure 2. Relationship of farm size with percentage of sampled landowners allowing
deer hunting by family and friends and other hunters.
Regional differences existed in a landowner's willingness to allow hunting of deer
(Table 3; X2 = 213, df = 10, P< 0.001) and small game (Table 4; X2 = 65, df =
10, P<0.001) on their property. Differences for deer hunting were primarily due to
a liberal hunting attitude by northeastern Indiana landowners and a restrictive attitude
by farm operators in central Indiana. A similar pattern is shown for small game hunting
with northwestern and central regions showing the fewest relative opportunities for
hunting. It is unclear why these regional differences exist.
Acknowledgments
We appreciate the computer assistance provided by personnel at LARS in West
Lafayette. John Olson calculated unpublished deer harvest figures. John Olson and
Table 3 . Landowner's willingness to allow white-tailed deer hunting on their property.
Values given are percentages of total responses by region.
Hunting
Region
permission
Northwest
Northeast
Central
Southwest
South-central
Southeast
response
(421)a
(715)
(599
(350)
(441)
(524)
No hunting
58.7
32.4
69.3
43.4
45.1
50.2
Family and
friends only
25.2
34.3
17.0
29.4
24.3
30.7
Permission
needed
13.8
27.1
11.4
20.3
22.4
15.6
Unrestricted
2.4
6.2
2.3
6.9
8.2
3.4
No. landowners responding.
244 Indiana Academy of Science Vol. 94 (1985)
Table 4. Landowner's willingness to allow small game hunting on their property. Values
given are percentages of total responses by region.
Hunting
Region
permission
Northwest
Northeast
Central
Southwest
South-central
Southeast
response
(425)a
(721)
(609)
(351)
(445)
(528)
No hunting
30.1
19.8
31.4
19.7
19.1
21.4
Family and
friends only
44.9
40.5
40.1
44.4
42.0
46.4
Permission
needed
22.1
33.6
25.8
30.8
31.9
28.8
Unrestricted
2.8
6.1
2.8
5.1
7.0
3.4
No. landowners responding.
Robert Feldt reviewed the manuscript and Jennifer Eckensberger typed the drafts. This
study was funded by Federal Aid to Wildlife Restoration in Indiana, Project W-26-R,
and the Indiana Division of Fish and Wildlife.
Literature Cited
1. Atwood, E.L. 1956. Validity of mail survey data on bagged waterfowl. J. Wildl.
Manage. 1:1-16.
2. Backs, S.E. 1984. The historic and present distribution of ruffed grouse in Indiana.
Proc. Indiana Acad. Sci. 93:161-166.
3. Castrale, J.S. 1985. Bob white quail spring population levels. Indiana Div. of Fish
and Wildl. Fed. Aid Prog. Rep. W-26-R-16. Job XX-F-3.
4. Castrale, J.S. and W.J. Pfingsten. 1982. Landowner-tenant small game harvest.
Indiana Div. Fish and Wildl. Fed. Aid Prog. Rep. W-26-R-13. Job XXIII-M-4.
5. Eisenhauer, D.I. 1977a. Small game harvest survey. Indiana Div. Fish and Wildl.
Fed. Aid Prog. Rep. W-26-R-8. Job XXIII-M-5.
6. Eisenhauer, D.I. 1977b. Landowner-tenant small game harvest. Indiana Div. Fish
and Wildl. Fed. Aid Prog. Rep. W-26-R-8. Job XXIII-M-4.
7. Eisenhauer, D.I. 1977c. Determine magnitude of bias. Indiana Div. Fish and Wildl.
Fed. Aid Prog. Rep. W-26-R-8. Job XXIII-M-2.
8. Filion, F.L. 1980. Human surveys in wildlife management. Pages 441-453 in S.D.
Schemintz (ed.). Wildlife management techniques, 4th ed. The Wildlife Society,
Washington, D.C. 686 p.
9. Olson, J.C. 1981. Deer harvest report card survey. Indiana Div. Fish and Wildl.
Fed. Aid Prog. Reg. W-26-R-12. Job XIII-B-1.
10. Pfingsten, W.J. 1980. Determine magnitude of response error. Indiana Div. Fish
and Wildl. Fed. Aid Prog. Rep. W-26-R-11. Job XXIII-M-2.
1 1. Rolley, R.E. 1984. Small game harvest survey. Indiana Div. Fish and Wildl. Fed.
Aid Prog. Rep. W-26-R-15. Job XXIII-M-5.
12. United States Department of Agriculture. 1982. Farm numbers hold steady. Indiana
Agric. Rep. 2(16):3.
The Natural Regions of Indiana
Michael A. Homoya, D. Brian Abrell, James R. Aldrich
and Thomas W. Post
Indiana Natural Heritage Program
Indiana Department of Natural Resources
Indianapolis, Indiana 46204
Introduction
Schemes recognizing regions of natural features have a long history in Indiana.
The surveyors and workers for the General Land Office (29) were possibly the first
to describe and map the major natural community types, such as prairie and forest,
found in presettlement Indiana. Starting in the early 1800s geologists were mapping
and describing geologic features, with occasional references to vegetation, while
naturalists were cataloging the flora and fauna. One of the first regionalizations of
Indiana biota was by W.S. Blatchley, who in 1909 defined three life zones of the state
using insect distribution as the criterion (11). Thirteen years later, the classic work
on the physiographic regions of Indiana was published by C.A. Malott (45). Since
then, a number of works have been published depicting regionalizations of various
natural features, including maps on the Forestal Areas of Indiana (20) and Floral Areas
of Indiana (21) by Deam, vegetation maps by Gordon (31), Potzger et al. (64), and
Lindsey et al. (41), and maps of faunal areas by Barnes (9) and Chandler (16). The
Natural Divisions of Indiana map by Lindsey et al. (43) was the first in Indiana to
delineate natural landscape units based on a combination of natural features (with
an emphasis on presettlement vegetation). Illinois (72) and Missouri (75) are two near-
by states that have used this concept to develop natural region classifications. The
present work is also a development of this concept.
A natural region is a major, generalized unit of the landscape where a distinctive
assemblage of natural features is present. It is part of a classification system that in-
tegrates several natural features, including climate, soils, glacial history, topography,
exposed bedrock, presettlement vegetation, species composition, physiography, and flora
and fauna distribution to identify a natural region. A section is a subunit of a natural
region where sufficient differences are evident such that recognition is warranted. The
text and map presented here describe and illustrate the twelve natural regions and twenty-
five sections determined by the authors.
In a practical sense, knowledge of the features of a natural region should help
one visualize the landscape and permit expectations about what can and cannot be
found in a region. For example, only in the Knobstone Escarpment Section of the
Highland Rim Natural Region can one expect to see a natural community with chestnut
oak and Virginia pine growing on a steep hillside composed of Mississippian shale
and siltstone. Conversely, one would not expect to see a calcareous fen natural com-
munity in the section.
Editor's Note: The Editor wishes to acknowledge not only the encouragement, but the patience, expertise, and
and critical reviews of the above manuscript by the following persons — Henry Gray, Marion Jackson, Ben Moulton,
John Patton and Damian Schmelz. Their combined efforts greatly assisted the authors in getting this manuscript
into its final form for publication in this centennial volume of the Proceedings.
The enclosed map, Plate 1, which accompanies this manuscript was made possible with the assistance of Henry
Gray, John Hollingsworth and William Moran. The original map by J.E. Switzer, upon which the map is based, was
printed previously in the following publications:
Kingsbury, R.C. 1970. An Atlas of Indiana. Dept. of Geography, Indiana Univ., Bloomington, IN. 94 p.
Switzer, J.E. 1937. The Geography of Indiana. Ginn and Co., Boston. 52 p.
245
246 Indiana Academy of Science Vol. 94 (1985)
Methods
No single criterion was used in determining natural regions although some single
feature may have been emphasized for mapping purposes. For instance, the boun-
daries of some natural regions may have been determined by the extent of the major
natural community present, e.g., Grand Prairie Natural Region, or by the area of
a dominant topographic feature, e.g., Shawnee Hills Natural Region. Although a single
feature is used to delimit some boundaries, it is the combination of natural features
that distinguish a natural region.
Species composition was an important criterion, especially when considering the
occurrence of rare and/or disjunct species, or species at the periphery of their range.
These species reveal much about the landscape, not only about the area where they
occur, but also about the area where they do not. For example, swamp chestnut oak
(Quercus michauxii), a southern species on the periphery of its range in the Bluegrass
Natural Region, does not occur in the adjacent Central Till Plain Natural Region..
Some significant difference in soil, glacial history, or other natural feature between
the two natural regions is implied by the absence of this species in the Central Till
Plain. Therefore, the distribution of this species was one criterion used to support
separation of the Bluegrass Natural Region from the Central Till Plain Natural Region.
A natural community is a group of organisms that are interrelated with each
other and their environment (80). They are identified by such natural features as soil
moisture and reaction, substrate, species composition, vegetation structure and
topographic position. An excellent discussion of natural communities and their classifica-
tion can be found in White and Madany (80). Although the present work is not in-
tended to be a treatise on natural communities, those occurring most frequently in
each natural region are discussed as are those restricted to or best developed in a region.
Most of the communities found in Indiana are discussed somewhere in the text, although
not in every region where they occur. For example, the fen natural community type
occurs throughout northern Indiana, but is described in some detail only in the North-
western Morainal region. Since fens are rather uniform compositionally, it would be
redundant to describe them in every natural region where they occur. If a natural com-
munity type is significantly different from one region to another, a description of the
community is given in the discussion of each region.
In describing features of a natural region, certain terms are used that need clarifica-
tion. Characteristic refers to an association of one or more natural features with another.
It may refer to a species commonly associated with a community (but not necessarily
restricted to it), or to a species that occurs uncommonly in a community type, especially
if it is restricted to it. For example, both the cliff clubmoss (Lycopodium porophilum)
and Bradley's spleen wort (Asplenium bradleyi) are indigenous species of sandstone
cliff natural communities in the Shawnee Hills. The former is regularly seen, but the
latter has been found only once. Both characterize the community. Since it is usually
difficult to identify a natural community by a single species, an assemblage of species
is listed to distinguish one community from the next.
The state is roughly divided into quadrants with northern and southern divisions
separated by U.S Highway 40 east of Indianapolis and U.S. Highway 36 west of
Indianapolis, and eastern and western divisions separated by U.S. Highway 31 north
of Indianapolis and State Route 135 south of Indianapolis. Species that are geographically
restricted are those found in only one section or region of a particular quadrant, yet
also occurring in at least one other quadrant of the state. For example, the blunt-lobed
grape fern {Botrychium oneidense) is a geographically restricted species of the
Muscatatuck Flats and Canyons Section, for it occurs in no other section of the southeast
quadrant. The species occurs elsewhere in the state, however, namely the northeast
Ecology 247
and northwest quadrants. State restricted species are indigenous to only one section
or region in the entire state, e.g. French's shooting star (Dodecatheon frenchii) is a
state restricted species known in Indiana only from the Crawford Upland Section of
the Shawnee Hills Natural Region. These distinctions are intended only to illustrate
disjunct or restricted occurrences of organisms and to help distinguish further one
natural unit from the next.
Except in a few instances, it is not stated whether natural communities listed
for a region are extant. The reader can assume that communities listed have current
examples, albeit in many cases small and/or highly degraded ones.
Because of their strong community association and relative lack of mobility, reptiles
and amphibians are some of the best community indicators of the fauna and are used
for that purpose here when appropriate. Most birds and mammals are normally highly
mobile and ubiquitous, and thus are used less frequently here as indicators, although
some good community indicators are known. All organisms (plant and animal) listed
in the text reflect documented occurrences of native populations but may or may not
be extant.
In the assignment of names for regions and sections, the traditional name identifying
a particular region was used when appropriate, e.g. Scottsburg Lowland physiographic
region became the Scottsburg Lowland Section of the Bluegrass Natural Region. In
some cases, a traditional name was altered to emphasize major characteristic natural
features, e.g. Mitchell Plain physiographic region became Mitchell Karst Plain Section
of the Highland Rim Natural Region. Names of topographic features were incorporated
into most names to help distinguish the area, e.g. Central Till Plain Natural Region.
Where possible, names were given to maintain continuity with similar classifications
in surrounding states, e.g. the Grand Prairie Natural Region adjoins the Grand Prairie
Natural Division in Illinois.
Boundary lines on the map (Plate 1 in envelope in back cover) do not necessarily
indicate an abrupt change in all natural features, i.e. all the distinctive features listed
for a region do not terminate at the line indicated, to be replaced by an entirely new
set of features. As there is a continuum from one natural community to the next,
so it is with natural regions.
A variety of sources was consulted for information detailing natural features of
the state. Physiographic works by Fenneman (25), Malott (45), Schneider (71), and
Quarterman and Powell (65) proved most useful. County soil surveys and the Map
of the Soil Associations of Indiana (44) were consulted for soils information. Regional
geologic maps published by the Indiana Geological Survey were invaluable for illustrating
bedrock and unconsolidated deposits. Wayne (77), and Wayne and Zumberge (78) were
major sources of information on glacial geology. Information on the flora, including
nomenclature, came from Deam (21) and the Indiana Natural Heritage Program (36).
The latter, along with Lindsey et al. (43), were good references for vegetation infor-
mation on specific sites, and survey notes of the General Land Office provided pre-
settlement information (29). Separate works on the state's fish, birds, mammals, and
herpetofauna by Gerking (30), Mumford and Keller (52), Mumford and Whitaker (53),
and Minton (50), respectively, proved invaluable. Several additional papers consulted
are cited in the text.
Description of Natural Regions
Region One — Lake Michigan Natural Region
This natural region is an entirely aquatic one that includes Indiana's portion of
a tremendous body of water, Lake Michigan. Formed from meltwater of the Wiscon-
sinan ice sheet, this large lake is so different from the rest of Indiana's natural features
248 Indiana Academy of Science Vol. 94 (1985)
that it deserves recognition as a separate natural region. It harbors (or formerly harbored)
a number of fish species found nowhere else in the state, including lake whitefish (Cor-
geonus clupeaformis), brook trout (Salvelinus fontinalis), lake trout (Salvelinus
namaycush), longnose sucker (Catastomus catastomus), slimy sculpin (Cottus cognatus),
four horn sculpin (Myoxocephalus quadricornis), and ninespine stickleback (Pungitius
pungitius). Unfortunately, many of these fishes have been replaced largely by exotics
either by accidental or by intentional introduction.
Region Two — Northwestern Morainal Natural Region
The glaciated area formed in part by the latest advances of the Lake Michigan
Lobe of the Wisconsinan ice sheet identifies this natural region. It is divided into three
sections: the Valparaiso Moraine Section, the Chicago Lake Plain Section, and the
Lake Michigan Border Section.
A tremendous diversity of natural communities is present for such a small region,
and floristically, no other natural region can compare in species diversity, at least on
an acre for acre basis. This is due in part to the merging of several major vegetation
types, these being the eastern deciduous forest, the tall grass prairie, and the northern
forest and wetlands. In addition, an interesting assemblage of Atlantic Coastal Plain
species, along with Lake Michigan shoreline endemics contribute to the diversity.
The region is heavily populated and industrialized, but because much of it is poor
agricultural land, and thus was never cultivated, high quality natural areas can be found
interspersed among factories, homes, landfills, and city streets. The region and its sec-
tions correlate with Illinois natural regions of similar names. Physiographic regions
identified by Malott (45) include the Valparaiso Moraine Section and the Calumet
Lacustrine Section of the Northern Moraine and Lake Region. Ecological studies of
the region include Cowles (18), Olson (54), Rohr and Potzger (69), Bacone and Campbell
(5), and Wilhelm (81).
Section 2A — Valparaiso Moraine Section
This section is identified by the presence of the Valparaiso Moraine, a moraine
characterized by a mostly knob-and-kettle topography in the east that grades into a
gently rolling till plain in the west. The soils generally are well drained, mostly calcareous
silty clay loams of the Markham, Elliott, Morley, Blount, and Pewamo series. The
eastern portion formerly was predominantly forested, while much of the western area
was prairie. Other natural community types include fen, bog, lake, marsh, savanna,
seep spring, and swamp.
The forest community on mesic sites is of special interest, for it marks the western
limit of the beech-maple community in the lower Lake Michigan region. Oak-hickory
forest characterize drier sites, and include white oak (Quercus alba), red oak (Q. rubra),
black oak (Q. velutina), shagbark hickory {Carya ovata), pignut hickory (C. glabra),
and black cherry (Prunus serotina). Bur oak (Quercus macrocarpa) and black oak
savannas occurred formerly but now are gone completely. The areas of prairie also
are gone, except for a few small remnants in pioneer cemeteries and railroad rights-of-
way. Species composition of these prairies is similar to those of the Grand Prairie
Region. One notable exception is the former presence of Mead's milkweed (Asclepius
meadii), as extirpated, state restricted species of this section.
Excellent examples of the fen natural community type occur on the moraine. These
normally unforested areas of mineral-rich seepage through muck commonly have a
high diversity of species that include Kalm's lobelia (Lobelia kalmii), shrubby cin-
quefoil (Potentilla fruticosa), Indian plantain (Cacalia tuberosa), tofieldia (Tofieldia
glutinosa), small white ladyslipper (Cypripedium candidum), parnassia (Parnassia glauca),
prairie dock (Silphium terebinthinaceum), fringed gentian (Gentiana crinita), marsh
Ecology 249
muhly (Muhlenbergia racemosa), and several Carex species, notably Carex leptalea
and C. sterilis. Bog communities are similar in composition to those of the Northern
Lakes Natural Region. Deep River is characteristic of streams of this section.
Section 2B — Chicago Lake Plain Section
This section is identified by the ridge-and-swale and lacustrine plain topography
that occurs between the Valparaiso Moraine and the Border Section along Lake Michigan.
It is located on the former site of Lake Chicago, and the ridge-and-swale topography
is a remnant of water-level fluctuations of that glacial lake. Almost all of the natural
communities are on sand substrates. Most of the sand is acid in reaction. Characteristic
soil associations include the Whitaker-Milford-Del Rey and Oakville Maumee-Brems.
Muck soils are scattered throughout.
Major natural communities of this section include marsh, lake, sand savanna,
sand prairie, and swamp, along with minor areas of various forest types. The sand
savana is primarily comprised of two types: the black oak (Quercus velutina) and the
black oak-pine (Pinus strobus, P. Banksiana) savanna. Almost pure stands of black
oak characterize the savannas throughout most of this section, whereas the black oak-
pine savannas are associated with the dune complex in the north part of the section.
Typical species of the savannas include little bluestem (Andropogon scoparius), Junegrass
(Koeleria cristata), goat's-rue (Tephrosia virginiana), lupine (Lupinus perennis), and
sedges (Carex muhlenbergii and C. pensylvanica). Sand prairie intergrades with the
savanna. Extensive areas of marsh once occurred throughout the section, especially
along the Little and Grand Calumet Rivers.
Many of the same animals found in the Kankakee Sand Section occur here also,
apparently owing to the similarities of natural communities. The Chicago garter snake
(Thamnophis sirtalis semifasciata) may be more common here than elsewhere in the state.
Section 2C — Lake Michigan Border Section
The three major natural features distinguishing this section are the beach com-
munity, the high dunes (especially the foredune community), and the pannes. All oc-
cur in the immediate vicinity and influence of Lake Michigan, and all are represen-
tative of natural communities bordering the Lake throughout much of its shoreline.
Sand is the major substrate, and the Oakville fine sand is the major soil series of
the high dunes. Various mucks occur in the interdunal depressions. Calcareous sand
occurs locally in the pannes.
The beach community occupies a narrow strip of sand between the edge of Lake
Michigan and the first line of dunes. It is an area of shifting sands where characteristic
pioneer species include sea rocket (Cakile edentula var. lacustris), beachgrass (Am-
mophila breviligulata), bug-seed (Corispermum hyssopifolium), spurge (Euphorbia
polygonifolia), and silverweed (Potent ilia anserina). The beach community grades into
the foredune of the high dunes complex. The foredune, like the beach, is on the wind-
ward side of the high dunes, but it is somewhat more stable than the beach because
of the presence of stabilizing plants, e.g. little bluestem (Andropogon scoparius), longleaf
reedgrass (Calamovilfa longifolia), red-osier dogwood (Cornus stolonifera), beach pea
(Lathyrus japonicus), aromatic sumac (Rhus aromatica), Pitcher's thistle (Circium pit-
cheri), bearberry (Arctostaphylos uva-ursi), prostrate juniper (Juniperus communis),
jack pine (Pinus banksiana), and gland leaf willow (Salix syrticola).
Forests of the lee side high dunes are characterized by a mixture of mesophytic
forest and savanna. White pine (Pinus strobus), red oak (Quercus rubra), white oak
(Q. alba), black oak (Q. velutina), basswood (Tilia americana), red maple (Acer rubrum),
white ash (Fraxinus americana), dogwoods (Cornus florida and C. rugosa), witchhazel
(Hamamelis virginiana), and wafer ash (Ptelea trifoliata) are characteristic species of
250 Indiana Academy of Science Vol. 94 (1985)
this area. The savanna component is similar to that of the Chicago Lake Plain Sec-
tion, except that conifers are more important.
Pannes are interdunal depressions composed of wet, calcareous sand typically
on the lee side of the first or second line of dunes from the lakeshore. They are
characterized by an unique floristic composition suggestive of a fen. Typical panne
species include Kalm's lobelia (Lobelia kalmii), fringed gentian (Gentiana crinita),
bladderwort (Utricularia cornuta), white upland aster (Aster ptarmicoides), rose gen-
tian (Sabatia angularis), loesel twayblade (Liparis loeselii), rush (Juncus balticus), cladium
(Cladium mariscoides), and sedges (Carex aurea, Rhynchospora capillacea, and Scleria
verticillata).
State restricted species from this section include beach grass, sea rocket, Pitcher's
thistle, gland leaf willow, white upland aster, spurge, russet buffaloberry (Shepherdia
canadensis), fringed polygala (Polygala paucifolia), Hooker's orchid (Platanthera
hookeri), and sedge (Carex richardsonii). The piping plover (Charadrius melodus) was
known in Indiana only from this section but is now extirpated.
Region Three — Grand Prairie Natural Region
This region is identified by the predominance of the tall grass prairie community
type. The name "Grand Prairie" is applied in reference to the large expanse of prairie
that occurred here and over much of northern Illinois. This area in Indiana is the
major eastern lobe of the Prairie Peninsula as illustrated by Transeau (76). The region
occupies a glaciated plain where a variety of unconsolidated deposits of Wisconsinan
age are present, including dune sand, lacustrine sediments, outwash plain sediments
(mostly sand and gravel), and till (end and ground moraines). The southern and eastern
borders of the region are defined by the Wabash River Valley and the Maxinkuckee
Moraine, and the Valparaiso Moraine marks the northern boundary.
This region is identified not only by what is present, but by what is not. Many
species characteristic of the eastern deciduous forest are noticeably absent here. Beech
(Fagus grandifolia) and sugar maple (Acer saccharum), the major components of the
beech-maple forest, are exceptionally rare species in this region. On a percentage basis,
this region is the most altered of all natural regions in the state. Only remnants of
the Grand Prairie are known to exist. Three sections are recognized: the Grand Prairie
Section, the Kankakee Sand Section, and the Kankakee Marsh Section. They occupy
parts of the Tipton Till Plain physiographic region and the Northern Moraine and
Lake physiographic region of Malott (45). Ecological studies in this region include
Finley and Potzger (26), Welch (79), Meyer (48), and Betz (10).
Section 3A — Grand Prairie Section
This section is distinguished by the predominance of loamy soil as opposed to
the sandy and highly organic soils of the other sections of Region Three. The swell
and swale topography in the northern part of the section is best ch tracterized by the
silty clay loam soils of the Brookston-Odell-Corwin Association. Some areas of muck,
particularly Carlisle muck, are present. The better drained soils in the south of the
section are characterized by Parr silt loam and the Elston-Shipshe-Warsaw Associa-
tion of well drained neutral to acid loam. Outwash and lacustrine deposits are characteriz-
ed by the Rennselaer-Darroch-Whitaker Association. This area was the epitomy of
the vast tall grass prairie of presettlement times. A great variety of prairie natural
community types must have existed, but little is known about the species composition
except what can be determined from small remnants in railroad rights-of-way and aban-
doned pioneer cemeteries. Characteristic species of prairies on well drained sites in-
clude little bluestem (Andropogon scoparius), big bluestem (A. gerardi), Indian grass
(Sorghastrum nutans), switchgrass (Panicum virgatum), side-oats grama (Bouteloua
Ecology 25 1
curtipendula), compass plant (Silphium laciniatum), prairie dock (Silphium terebin-
thinaceum), blazing star (Liatris pycnostachya), hairy sunflower (Helianthus mollis),
feverfew (Parthenium integrifolium), pale purple coneflower {Echinacea pallida), yellow
coneflower (Ratibida pinnata), leadplant (Amorpha canescens), rattlesnake master
(Erynigium yuccifolium), prairie clovers (Petalostemum candidum and P. purpureum),
prairie goldenrod (Solidago rigida), and prairie violet (Viola pedat if ida). The wet prairies
are characterized by cordgrass (Spartina pectinata), big bluestem, Culver's-physic
( Veronicastrum virginicum), water parsnip (Sium suave), golden alexander (Zizia aurea),
cowbane (Oxypolis rigidior), Carex spp., and bluejoint grass (Calamagrostis canaden-
sis). Other community types present include savanna, marsh, pond, bog (rare), and
forest, the latter mostly along stream courses and in oak groves. Animals characteristic
of this section include fox snake (Elaphe vulpina), prairie king snake (Lampropeltis
calligaster), smooth green snake (Opheodrys vernalis), plains garter snake (Thamnophis
radix), Franklin's ground squirrel (Spermophilus franklinii), western meadowlark
(Sturnella neglecta), upland sandpiper (Bartramia longicauda), and the extirpated prairie
chicken (Tympanuchus cupido). Typical streams of this section are low-gradient and
silty, e.g. Sugar Creek (Benton County) and Iroquois River.
Section 3B — Kankakee Sand Section
This area is characterized by the presence of predominantly prairie and savanna
natural community types associated with sandy soils. It consists mostly of dune sand
and outwash plain sediments. The dune areas are typically the Plainfield-Maumee-
Oshtemo Association of acidic to neutral sand and sandy loams. The outwash plains
consist of poorly drained sandy loams of the Maumee-Gilford-Sebewa Association and
well drained sandy loams of the Tracy-Door-Lydick Association. The sand prairie and
savanna communities are similar in species composition to the prairie of the Grand
Prairie Section except that, in addition, a number of sand-dwelling species are present.
These include porcupine grass (Stipa spartea), dropseed (Sporobolus clandestinus),
longleaf reedgrass (Calamovilfa longifolia), Junegrass (Koeleria cristata), prairie talinum
(Talinum rugospermum), puccoon (Lithospermum croceum), primrose violet (Viola
primulifolia), sedges (Carex gravida and C. cumulata), and dwarf-dandelion (Krigia
virginica). Savannas dominated by black oak (Quercus velutina) and prairie species
occur on the dunal areas. Typical associates of the savannas include sand prairie species
along with goat's-rue (Tephrosia virginiana), bracken fern (Pteridium aquilinum), lupine
(Lupinus perennis), sedge (Carex pensylvanica), bird's-foot violet (Viola pedata), black
huckleberry (Gaylussacia baccata), dryland blueberry (Vaccinium vacillans), and lowbush
blueberry (V. anqustifolium). Swales between the dunes might have any of several
possible natural community types, including wet prairie, marsh, swamp, wet sand flat,
and wet muck flat. A remarkable assemblage of plants with coastal plain affinities
is known from the wet sand/muck flat community, including bladderwort (Utricularia
radiata), panic grass (Panicum verrucosum), nutrush (Scleria reticularis), beak rush
(Psilocarya scirpoides), sedge (Fimbristylis caroliniana), yellow-eyed grass (Xyris caroli-
niana), bugleweed (Lycopus amplectens), and flax (Linum intercursum). Forest natural
communities occur primarily in the eastern part of the section, where white oak (Quer-
cus alba) and black oak (Q. velutina) are dominants. Pin oak flatwoods characterize
some of the swales in dunal areas. Fauna of the Grand Prairie Section are found in
this section also, along with species that thrive in sandy habitat, e.g. ornate box turtle
(Terrapene ornata), bull snake (Pituophis melanoleucus), glass lizard (Ophisaurus at-
tenuates), plains pocket gopher (Geomys bursarius), and lark sparrow (Chondestes
grammacus). A geographically restricted population of eastern mud turtles (Kinoster-
non subrubrum) occurs here. State restricted species of the section include bladder-
252 Indiana Academy of Science Vol. 94 (1985)
wort (Utricularia radiata), flax (Linum intercursum), St. John's-wort {Hypericum ad-
pressum), and sedge (Carex cumulata). Stream communities have all been altered greatly
by channelization.
Section 3C — Kankakee Marsh Section
This section is identified by the predominance of marsh, lake, and wet prairie
communities that existed along the Kankakee River in presettlement times. The marsh
was several miles wide on both sides of the river for almost its entire run in Indiana.
Extensive ditching beginning in the late 1800s has all but eliminated the natural wetlands.
The section is part of a large Wisconsinan glacial outwash plain, with a substrate of
acidic silt and sand. Characteristic soil series include Suman, Gilford, Maumee, and
Bourbon. Good examples of prairie and marsh are absent from the area today. Rem-
nants indicate that the wetlands were characterized by spatterdock {Nuphar advena),
watershield (Brasenia schreberi), swamp loosestrife (Decodon verticillatus), bluejoint
grass (Calamagrostis canadensis), reed canary grass (Phalaris arundinacea), common
reed (Phragmites communis), giant bur-reed (Sparganium eurycarpum), knotweeds
{Polygonum spp.), Spanish needles (Bidens spp.), arrowheads (Sagittaria spp.), and
sedges (Scirpus spp. and Carex spp.). A narrow border of forest along the river con-
tains characteristic floodplain species, e.g. silver maple (Acer saccharinum), red maple
(A. rubrum), black willow (Salix nigra), green ash (Fraxinus pennsylvanica), cotton-
wood (Populus deltoides), sycamore (Platanus occidentalis), river birch (Betula nigra),
and indigo bush (Amorpha fruticosa), the last geographically restricted here. Two plants
occurring as remarkable disjuncts include American snowbell (Styrax americana) and
climbing hempweed (Mikania scandens) . The northern weed shiner (Notropis texanus)
is state restricted here. The area was formerly a significant breeding habitat for waterfowl.
Region Four — Northern Lakes Natural Region
This natural region is identified by the presence of numerous fresh water lakes
of glacial origin. Approximate borders of the area are the southern edge of the Packerton
Moraine, the eastern edge of the Mississinewa and Salamonie Moraines north of the
Eel River, and the western edge of the Maxinkuckee Moraine.
This area was invaded from the northwest by the Lake Michigan Lobe of the
late Wisconsinan ice sheet, from the northeast by the Saginaw Lobe, and from the
east by the Erie Lobe. Consequently, the area is covered now with a thick and com-
plex deposit of glacial material which, in places, is over 450 feet thick. Glacial topography
also is complex and is characterized by knobs, kettles, kames, valley trains, and out-
wash plains. The diversity of soils include: loamy soils in the morainal areas and till
plains, typically the Miami-Crosier-Brookston-Riddles Association; neutral, clayey soils
in morainal areas of the southeastern portion of the section, typically the Morley-
Blount-Pewamo Association; and sandy loam soils on the outwash deposits, typically
by the Oshtemo-Fox Association and the Plainfield-Maumee-Oshtemo Association. Muck
soils, which are important components of wetland natural communities, include
Houghton, Edwards and Adrian series.
Natural community types are numerous, including bog, fen, marsh, prairie, sedge
meadow, swamp, seep spring, lake, and various deciduous forest types. Oak and hickory
species, especially red oak (Quercus rubra), white oak (Q. alba), black oak (Q. velutina),
shagbark hickory (Carya ovata), and pignut hickory (C. glabra) dominate the dry and
dry-mesic upland forests which once covered approximately one half of the region. Mesic sites
characteristically have beech (Fagus grandifolia), sugar maple (Acer saccharum), black
maple (A. nigrum), and tulip tree (Liriodendron tulipifera) as dominants. Floodplain
forests typically include sycamore (Platanus occidentalis), American elm (Ulmus
americana), red elm (U. rubra), green ash (Fraxinus pennsylvanica), silver maple (Acer
Ecology 253
saccharinum), red maple (A. rubrum), cottonwood (Populus deltoides), hackberry (Celt is
occidentalis), and honey locust (Gleditsia thacanthos).
Swamp communities commonly border lake and bog sites where red maple, silver
maple, green ash, American elm, black ash (Fraxinus nigra), and locally, yellow birch
(Betula luted), are typical associates. Swamps dominated by black ash typically are
associated with seep springs.
Bogs are more numerous here than in any other natural region. These communities
commonly consist of a floating mat of Sphagnum moss occupying a glacial depres-
sion. Distinctive bog plants include leatherleaf (Chamaedaphne calyculata), cranberry
(Vaccinium macrocarpori), bog rosemary (Andromeda glaucophylla), pitcher plant (Sar-
racenia purpurea), sundews (Drosera rotundifolia and D. intermedia), mountain holly
(Nemopanthus mucronata), tamarack (Larix laricina), Virginia chain fern (Woodwar-
dia virginica), grass-pink orchid (Calopogon pulchellus), rose pogonia orchid (Pogonia
ophioglossoides), sedges (Carex oligosperma and Rhynchospora alba), poison sumac
(Rhus vernix), and Sphagnum spp.
Areas of marsh commonly are associated with the lake community. Typical marsh
species include swamp loosetrife (Decodon verticillatus), cattails (Typha augustifolia and
T. latifolia), bulrush (Scirpus validus), marsh fern (Thelypteris palustris), marsh
cinquefoil (Potentilla palustris), and sedges, notably Carex stricta and C. lasiocarpa.
In deeper water bordering the marsh, the lake community begins, where such distinc-
tive species as spatterdock (Nuphar advena), water shield (Brasenia schreberi), fragrant
water-lily (Nymphaea tuberosa), pickerelweed (Pontederia cordata), hornwort
(Ceratophyllum demersum), wild celery (Vallisneria americana), pondweeds (Pot-
amogeton spp.), Virginia arrow-arum (Peltandra virginica), and sedge (Scirpis subter-
minalis) occur.
Wet sand flats and muck flats border some of the lakes and shallow basins. In
some places an unique flora of Atlantic Coastal Plain disjuncts is associated with these
communities, including sedges such as Psilocarya scirpoides, Fuirena pumila, Rhyn-
chospora macrostachya, and Eleocharis olivacea.
State restricted plants of this region include ginger-leaved pyrola (Pyrola asarifolia),
needle-and-thread grass (Stipa comata), knotted spikerush (Eleocharis equisetoides) ,
autumn willow (Salix serissima), and Deam's rockcress (Arabis missouriensis var. deamii).
Distinctive fauna of the region include spotted turtle (Clemmys guttata), massasauga
rattlesnake (Sistrurus catenatus), Blanding's turtle (Emydoidea blandingi), star-nosed
mole (Condylura cristata), cisco (Coregonus artedii), marsh wren (Cistothorus palustris),
swamp sparrow (Melospiza georgiana) and sandhill crane (Grus canadensis).
Typical streams are clear, medium to low-gradient, and have sandy gravel beds.
Good examples are Pigeon River, Elkhart River, upper Tippecanoe River and Fawn
River. Exemplary lakes include Olin Lake, Crooked Lake, Marsh Lake and Lake
Manitou. Ecological studies of the region include Scott (73), Potzger and Friesner (62),
Mills (49), Everman and Clark (24) and Aldrich (2).
Region Five — Central Till Plain Natural Region
This, the largest natural region in Indiana, is a formerly forested plain of Wiscon-
sinan till in the central area of the state. Aside from the Entrenched Valley Section,
it is topographically homogeneous, although several glacial features, especially moraines,
are common. The region is a major divide between the communities with strong northern
affinities and those with strong southern affinities, and the Entrenched Valley Section
is a concentrated melting pot of species with northern, southern, eastern, and western
affinities.
The three sections of the region are: the Entrenched Valley Section, characterized
by moderately thick loess over Wisconsinan till; the Tipton Till Plain Section,
254 Indiana Academy of Science Vol. 94 (1985)
characterized by loamy Wisconsinan till; and the Bluffton Till Plain Section, characterized
by clayey Wisconsinan till. Besides the predominant forest community types, areas
of prairie, marsh, fen, seep spring, bog, swamp, and lake are known.
This region occupies most of Malott's (45) Tipton Till Plain physiographic region
and portions of the Northern Moraine and Lake physiographic region. Ecological studies
of the region include Cain (13), Friesner and Potzger (28), Potzger (59), Ebinger and
Bacone (23), Petty and Harwood (55), Hollet and Jackson (33), and Post et al. (57).
Section 5A — Entrenched Valley Section
This section is quite unlike the other sections of the region. It is identified by
the deeply entrenched valleys along major drainages, particularly the Wabash, Sugar,
and Big Pine riverine systems. Bedrock is exposed in many places, and massive cliffs
are common. Pennsylvanian, Mississippian, Devonian, and Silurian sandstone, siltstone,
shale, and limestone are the predominant rock types. A variety of soils is present,
including poorly drained to well drained silt loams that are acid to neutral in reaction
and commonly covered with a moderately thick layer of loess. Representative soil series
include Fincastle, Russell, Miami, and Brookston. Upland forests, bottomland forests,
and flatwoods are the major natural community types present.
Except in the specialized cliff and ravine communities, the forest associations
are essentially the same as those of the Tipton Till Plain Section. Other natural com-
munity types present in the section include prairie, gravel-hill prairie, fen, marsh,
savanna, cliff, seep spring, and pond. The circumneutral seep spring is well represented
and possibly is more common here than elsewhere in the state. This relatively open
community typically is situated on the lower slopes of hills, particularly those border-
ing larger drainages, such as the Wabash River. Water oozes through a muck soil
in a diffuse manner, creating an environment where such plants as skunk cabbage
(Symplocarpus foetidus), marsh marigold (Caltha palustris), Pennsylvania saxifrage
(Saxifraga pennsylvanica), swamp woodbetony (Pedicularis lanceolata), jewelweed (Im-
portiens biflora), queen-of-the-prairie (Filipendula rubra), nannyberry (Viburnum lentago),
black ash (Fraxinus nigra), sedges (Carex bromoides, C. trichocarpa, and C. sterilis),
white turtlehead (Chelone glabra), roughleaf goldenrod (Solidago patula), and purple-
stem aster (Aster puniceus) are characteristic.
The cliff and ravine communities provide an environment for an interesting
assemblage of species, many of which occur as disjuncts that have northern affinities.
Two of these, white pine (Pinus strobus) and hemlock (Tsuga canadensis), give a boreal
appearance to the landscape. Other northern disjuncts include Canada yew (Taxus
canadensis), Canada blueberry (Vaccinium canadense), shinleaf (Pyrola elliptica), wild
sarsaparilla (Aralia nudicaulis), northern enchanter's nightshade (Ciracea alpina),
roundleaf dogwood (Cornus rugosa), false melic grass (Schizachne purpurascens), and
two-leaf Solomon's seal (Maianthemum canadense). Gravel hill prairies are state restricted
here. Along with typical prairie species, they also have geographical and state restricted
species, including many that have southern and western affinities. These include plains
muhly (Muhlenbergia cuspidata), western wallflower (Erysimum arkansana), narrowleaf
houstonia (Houstonia nigricans), gromwell (Lithospermum incisum), androsace
(Androsace occidentalis) , and post oak (Quercus stellata). This section marks the nor-
thern limit of several herpetofaunal species, including the cave salamander (Eurycea
lucifuga), zigzag salamander (Plethodon dorsalis), long-tailed salamander (Eurycea
longicauda), earth snake (Carphophis amoenus), and copperhead (Agkistrodon con-
tortrix). State restricted species of this section include pitcher sandwort (Arenaria patula),
forked aster (Aster furcatus), Forbe's saxifrage (Saxifraga forbesii), Canada yew, plains
muhly, and Canada blueberry. Streams of this section are typically medium-gradient,
relatively clear, and rocky, e.g. Sugar Creek, Big Walnut Creek, and Raccoon Creek.
Ecology 255
Section 5B — Tipton Till Plain Section
This section is a mostly undissected plain formerly covered by an extensive beech-
maple-oak forest. The soils are predominantly neutral silt and silty clay loams of the
Crosby-Brookston Association. The northern flatwoods community associated with
these poorly drained soils was ubiquitous but now is confined to scattered woodlots.
Species common within the community include red maple (Acer rubrum), pin oak (Quer-
cus palustris), bur oak (Q. macrocarpa), swamp white oak (Q. bicolor), Shumard's
oak (Q. shumardii), American elm (Ulmus americana), and green ash (Fraxinus penn-
sylvanica). In slightly better drained sites beech (Fagus grandifolia), sugar maple (Acer
saccharum), black maple (Acer nigrum), white oak (Quercus alba), red oak (Q. rubra),
shagbark hickory (Carya ovata), tulip poplar (Liriodendron tulipifera), red elm (Ulmus
rubra), bass wood (Tilia americana), and white ash (Fraxinus americana) are characteristic.
Other community types of this section include bog, prairie, marsh, seep spring, and
pond. A few fens are known, including the well studied Cabin Creek Bog (28). They
are similar in composition to fens elsewhere in the state. Because of the section's loca-
tion and the scarcity of specialized natural communities, there are no restricted species.
Section 5C— Bluffton Till Plain Section
This section is characterized by the predominance of clay-rich soils on a relatively
level till plain. This area, along with the Black Swamp, Northern Lakes and Northwestern
Morainal Natural Regions, was one of the last areas of Indiana to be occupied by
glacial ice, in this case, by the Ontario-Erie Lobe of the Wisconsinan ice sheet. A
distinct series of moraines is evident in this section, and the Union City Moraine marks
its southern border. As a consequence of the widespread presence of clayey till, much
of the area is poorly drained. The acid to neutral silty clay loams of the Blount-Pewano-
Morley Association characterize the region. Most of the natural communities are forested,
along with minor areas of bog, prairie, fen, marsh and lake communities. Composi-
tion of forest species is similar to the Tipton Till Plain Section, although swamp cotton-
wood (Populus heterophylla) which formerly occurred regularly in swamps here, was
and is rare on the Tipton Till Plain. A greater number of northern wetland species
occur in this section than in the others of the region, e.g. cottongrass (Eriophorum
gracile), northern St. John's-wort (Hypericum boreale), pitcher plant (Sarracenia pur-
purea), and sedges (Carex alopecoidea, C. laricina, and C. limosa). Interestingly, two
southern swamp species are known here as geographic restrictions, namely, swamp
St. John's-wort (Triadenum tubulosum) and log sedge (Carex decomposita).
Region Six — Black Swamp Natural Region
This region is the western lobe of a large lacustrine plain occupying the area once
covered by the ancient Lake Maumee. Lake Maumee, a predecessor to modern Lake
Erie, was created when the meltwater of the Ontario-Erie Lobe of the Wisconsinan
ice sheet was dammed by the Fort Wayne Moraine (45). The Lake, long since
abandoned, is now an almost featureless, naturally poorly drained plain. Soils are typical-
ly deep, acidic to neutral clay and silt loams of the Hoytville-Nappanee Association.
This area is the same as Malott's (45) Maumee Lacustrine Section of the Northern
Moraine and Lake Region.
Named the Black Swamp by early settlers, the predominant natural community
in the region consisted of swamp forest dominated by American elm (Ulmus americana),
black ash (Fraxinus nigra), and maples (Acer rubrum and A. saccharinum). This and
other natural community types are virtually non-existant in this region of Indiana to-
day, for extensive drainage has permitted an almost complete conversion of the land-
scape to agricultural uses. Other species known from the swamp forest and environs
256 Indiana Academy of Science Vol. 94 (1985)
included bur oak (Quercus macrocarpa), swamp white oak (Q. bicolor), white ash
(Fraxinus americana), shellbark hickory (Carya laciniosa), pawpaw (Asimina triloba),
and spicebush (Lindera benzoin). No flora and fauna are known to be restricted to
the region. Typical streams are low-gradient, silty and shallowly entrenched, e.g. Maumee
River.
Region Seven — Southwestern Lowlands Natural Region
This region, which is characterized by low relief and extensive aggraded valleys,
includes the area bounded in Indiana by the Shawnee Hills Natural Region to the east,
the Wisconsinan glacial border to the north, the Southern Bottomlands Natural Region
(along the Ohio River) to the south, and the Wabash River (north of Vincennes) to
the west. Similar terrain occurs across the Wabash and Ohio Rivers in Illinois and
Kentucky. Much of the region is nearly level, undissected, and poorly drained, although
in several areas the topography is hilly and well drained. This region, except for the
southern portion, was glaciated by the Illinoian ice sheet. The region is divided into
three sections: the Plainville Sand Section, the Glaciated Section, and the Driftless
Section. The extant natural communities are mostly forest types, although barrens were
formerly dominant in the Plainville Sand Section, and large areas of prairie occurred
in the Glaciated Section. All of this region occurs in the Wabash Lowland physiographic
region of Malott (45).
Ecological studies in the region include Lawlis (39), Lindsey (40), Ridgway (68),
McCoy (46), Schneck (70), Homoya (34), Aldrich and Homoya (4), and Green (32).
Section 7A — Plainville Sand Section
The Plainville Sand Section is a small but unique area of eolian sand dunes east
of the Wabash River and the White River. The sandy, acid soils are mostly in the
Princeton, Bloomfield, and Ayrshire series. The barrens natural community type, now
virtually gone from the landscape, was predominant on the ridges and well drained
sites, and swamp, marsh, and wet prairie occupied the swales (29). The barrens vegeta-
tion consisted mostly of prairie species, along with a collection of sand dwelling species
of western and southern affinities, including beard grass (Gymnopogon ambiguus),
Carolina anemone (Anemone carol in iana), tube penstemon (Penstemon tubaeflorus),
clustered poppy-mallow (Callirhoe triangulata), hairy golden-aster (Chrysopsis villosa),
narrowleaf dayflower (Commelina angustifolia), black hickory {Carya texana), sand
hickory (C. pallida), androsace (Androsace occidentalis), rose gentian (Sabatia cam-
panulata), sedge (Carex gravida), and fleabane (Erigeron pusillus). In a few degraded
remnants, one can still observe barrens vegetation, including little bluestem (Andropogon
scoparius), big bluestem (A. gerardi), Indian grass (Sorghastrum nutans), side-oats grama
(Bouteloua curtipendula), New Jersey tea (Ceanothus americanus), and blackjack oak
(Quercus marilandica). These areas also were inhabited by a prairie fauna. Species
geographically restricted here include bull snake (Pituophis melanoleucus), ornate box
turtle (Terrapene ornata), and six-lined racerunner (Cnemidophorus sexlineatus). The
biota of this section are similar to those of the Kankakee Sand Section of the Grand
Prairie Natural Region.
Section 7B — Glaciated Section
This section coincides with the Illinoian till plain of southwestern Indiana. The
soils are predominantly acid to neutral silt loams with a thick layer of loess, typically
the Iva, Cinncinati, Avon, Vigo, and Alford series. Natural communities are mostly
forest types, but several types of former prairie are known. The flatwoods community
type is common, but it is of different composition than the flatwoods in the Driftless
Ecology 257
Section, i.e. several species of southern affinity are uncommon or absent. Common
flatwoods species include shagbark hickory {Carya ovata), shellbark hickory (C
laciniosa), pin oak (Quercus palustris), shingle oak {Q. imbricaria), hackberry (Celtis
occidentalis), green ash {Fraxinus pennsylvanica), red maple {Acer rubrum), and silver
maple {A. saccharinum). Black ash {Fraxinus nigra) swamps are near their southern
limit here. This section appears to have had the largest amount of prairie south of
the Wisconsinan glacial border in Indiana. Little is known about the composition of
the prairie, but it probably was very similar to the prairies of the Grand Prairie Region.
Additional community types include swamp, marsh, pond, and low-gradient stream.
Typical examples of the latter are Eel River and Busseron Creek. The prairie kingsnake
{Lampropeltis calligaster) and the crawfish frog (Rana areolata) are characteristic species
of this region. Smallmouth bass {Micropteris dolomieu) and northern rock bass
(Ambloplites rupestris), common game fishes, are uncommon or absent in this section
and in the natural region.
Section 7C — Driftless Section
This section is south of the Illinoian glacial border, and is therefore placed in
the Interior Low Plateaus Physiographic Province. It is characterized by a topography
of low hills and broad valleys, in an area that has the longest growing season and
highest average summer temperature in the state. Most of the natural communities
are upland forest types, occupying well drained slopes underlain by soils of the Zanesville,
Wellston, and Tilsit series, which were formed in loess and weathered bedrock. Southern
flatwoods occupy the lacustrine plains and river terraces, which are characterized by
the McGary, Weinbach, Elkinsville, and Ginat series. Soils are predominantly acid
in reaction. Characteristic species of the flatwoods include cherry bark oak {Quercus
falcata var. pagodaefolia), sweetgum {Liquidambar styraciflua), shellbark hickory {Carya
laciniosa), pin oak {Quercus palustris), swamp white oak {Q. bicolor), Shumard's oak
{Q. shumardii), green ash {Fraxinus pennsylvanica), black gum {Nyssa sylvatica), and
locally, post oak {Quercus stellata). State restricted species of the flatwoods are Indian
pink {Spigelia marilandica), black quillwort {Isoetes melanopoda), and lesquerella {Les-
querella globosa). The barrens associated with the post oak flatwoods do not have
a typical prairie flora as do most other barrens communities. Instead, these xeric,
ephemerally wet sites characteristically are dominated by lichens, mosses, poverty grass
{Danthonia spicata), three-awn grass {Aristida ramosissima), spike-rush {Eleocharis ver-
rucosa), and rushfoil {Crotonopsis elliptica), the latter state restricted here. The upland
sites of this section are relatively dry oak-hickory dominated natural communities. The
occurrence and abundance of southern red oak {Quercus falcata), post oak {Q. stellata),
blackjack oak {Q. marilandica), and locally, chestnut oak {Q. prinus) help distinguish
the upland forests of this section from those of the Glaciated Section. At least one
acid seep spring community is known from this section. Other natural community types
include marsh, swamp, sandstone cliff, and low to medium-gradient stream.
Region Eight — Southern Bottomlands Natural Region
This natural region includes the alluvial bottomlands along the rivers and larger
streams in southwestern Indiana. It is distinguished from other bottomland regions
in the state by the presence of several species with affinities to the lower Mississippi
Valley and Gulf Coastal Plain. The Illinoian glacial border (see enclosed map) bisects
the region, thus placing the northern portion in the Central Lowlands Physiographic
Province and the southern portion in the Interior Low Plateaus Physiographic Pro-
vince. The glacial border has had little effect on the bottomland biotic communities;
therefore, the region is presented as one natural unit.
258 Indiana Academy of Science Vol. 94 (1985)
The soils are mostly neutral to acid silt loams, and include series such as Nolin,
Newark, Huntington, Linside, Stendal, and Bonnie. Much of the area is subject* to
frequent flooding (or did flood prior to the construction of control structures).
The natural communities of the region include bottomland forest, swamp, pond,
slough, and formerly marsh and prairie. The bottomland forest, the major community
of this region, is characterized by pecan {Carya illinoensis), sugarberry (Celt is laevigata),
swamp chestnut oak (Quercus michauxii), pin oak (Q. palustris), swamp white oak
(Q. bicolor), red maple {Acer rubrum), silver maple (Acer saccharinum), honey locust
(Gleditsia triacanthos), catalpa (Catalpa speciosa), shellbark hickory (Carya laciniosa),
sycamore (Platanus occidentalis), and green ash (Fraxinus pennsylvanica). The strongest
southern influence is reflected in the swamps and sloughs, where bald cypress (Tax-
odium distichum), swamp cottonwod (Populus heterophylla), water locust (Gleditsia
aquatica), pumpkin ash (Fraxinus tomentosa), and overcup oak (Quercus lyratd) occur.
Other distinctive southern species (many of which are restricted to this region)
include American featherfoil (Hottonia inflata), bloodleaf (Iresine rhizomatosa), acanthus
(Dicliptera brachiata), climbing dogbane (Trachelospermum difforme), milkweed
(Asclepias perennis), catbird grape (Vitis palmata), woolly pipe-vine (Aristolochia tomen-
tosa), sedge (Carex socialis), swamp privet (Forestiera acuminata), American snowbell
(Styrax americana), climbing hempweed (Mikania scandens), spiderlily (Hymenocallis
occidentalis), mistletoe (Phoradendron flavescens), and giant cane (Arundinaria
gigantea).
Distinctive southern animals include cottonmouth (Agkistrodon piscivorus),
hieroglyphic turtle (Pseudemys concinna hieroglyphica xfloridana hoyi), diamondbacked
watersnake (Nerodia rhombifera), eastern mud turtle (Kinosternon subrubrum), northern
copperbelly (Nerodia erythrogaster), swamp rabbit (Sylvilagus aquaticus), mosquitofish
(Gambusia affinis), harlequin darter (Etheostoma histrio — only one occurrence in In-
diana), and yellow-crowned night heron (Nyctanassa violacea).
The Patoka River is exemplary of a silt-bottomed, low-gradient stream characteristic
of this region. Other typical aquatic features include large bottomland ponds, especially
along the Wabash River, e.g. Foote Pond, Half Moon Pond, and Wabash Pond. The
Wabash, Ohio, and White Rivers themselves are considered a separate natural region.
Ecological studies in this region include: Cain (15), DenUyl (22), Lindsey (40),
Schneck (70), and Ridgway (68).
Region Nine — Shawnee Hills Natural Region
"Shawnee Hills" is a name given by Flint (27) to a physiographic region of the
Interior Low Plateaus in southwestern Indiana, southern Illinois, and western Ken-
tucky. Only the contiguous belt of rugged hills on the outer (southern and eastern)
periphery of the physiographic region denotes the Shawnee Hills Natural Region as
identified here. The region is divided into the Crawford Upland Section and the Escarp-
ment Section. Pennsylvanian and Mississippian bedrock, mostly sandstone, crops out
in many places to form distinctive cliffs and rockhouses. Except for small areas of
till in the northern portion, the region is driftless.
This region appears to represent general presettlement conditions better than any
other terrestrial region in the state. It is a rugged and generally sparsely populated
area. The majority of natural communities are upland forest types, although a few
sandstone and limestone glades, gravel washes, and barrens are known.
Ecological studies in this region include Potzger et al. (63), Petty and Lindsey
(56), Abrell and Jackson (1), Bacone et al. (6), and Badger and Jackson (8).
Ecology 259
Section 9A — Crawford Upland Section
The most distinctive features of this section are the rugged hills with sandstone
cliffs and rockhouses. Mississippian sandstone composes most of the cliffs in the eastern
portion of the section, as well as lower elevation outcrops to the west, whereas
Pennsylvanian sandstone (especially the Mansfield Formation) dominates the western
portion and higher hills. The well drained acid silt loams of the Wellston-Zanesville-
Berks Association are characteristic. The forest vegetation consists of an oak-hickory
assortment of the upper slopes, while the coves have a mesic component. Characteristic
upper slope species include black oak (Quercus velutina), white oak (Q. alba), chestnut
oak (Q. prinus), scarlet oak (Q. coccinea), post oak (Q. stellata), pignut hickory {Carya
glabra), small-fruited hickory (C. ovalis), shagbark hickory (C. ovata), and rarely,
sourwood (Oxydendrum arborewn). The cove forests, especially those associated with
rockhouses, most resemble the mixed mesophytic forest communities of the Mixed
Mesophytic Region of the Cumberland Plateau as defined by Braun (12). Characteristic
species include beech (Fagus grandifolia), tulip tree (Liriodendron tulipifera), red oak
(Quercus rubra), sugar maple (Acer saccharum), black walnut (Juglans nigra), white
ash (Fraxinus americana), and locally, yellow buckeye (Aesculus octandra), white
basswood (Tilia heterophylla), umbrella magnolia {Magnolia tripetala), hemlock (Tsuga
canadensis), and yellow birch (Betula luted). The sandstone cliff and rockhouse com-
munities provide an environment for several species with Appalachian affinities, e.g.
mountain laurel {Kalmia latifolia), mountain spleenwort (Asplenium montanum), sour-
wood, and umbrella magnolia. Distinctive species of the rockhouses include filmy fern
(Trichomanes boschianum), alumroot (Heuchera parviflora), Bradley's spleenwort
(Asplenium bradleyi), French's shooting star (Dodecatheon frenchii), and the
Appalachian gametophyte ( Vittaria sp.). A few examples of the acid seep spring community,
a type extremely rare in Indiana, occur in this section. The characteristic flora of these
bog-like environments includes cinnamon fern (Osmunda cinnamomea), royal fern (O.
regalis), sedges (Carex bromoides, C. lurida), small clubspur orchid (Platanthera clavellata),
black chokeberry (Aronia melanocarpa), winterberry (Ilex verticillata), tearthumb
(Polygonum arifolium), jewelweed (Impatiens biflora), crested wood fern (Dryopteris
cristata), and Sphagnum spp. The barrens community is (and probably was) a minor
component of this section. Only a few high quality remnants remain. Floristically,
they are similar to the glades and barrens of the Highland Rim Natural Region, although
missing many of the distinctive glade species. Sandstone glades are almost non-existent
in Indiana, but at least two small ones are known from this section. Characteristic
species of sandstone glades include little bluestem (Andropogon scoparius), slender
knotweed (Polygonum tenue), poverty grass (Danthonia spicata), farkleberry (Vaccinium
arboreum), goat's rue (Tephrosia virginiana), pineweed (Hypericum gentianoides),
pinweed (Lechea tenuifolia), and panic grass (Panicum depauperatum). Most of Indiana's
timber rattlesnake (Crotalis horridus) collections have come from this region and the
Brown County Hills Section of the Highland Rim Natural Region (51). Two interesting
mammals characteristic of this section are the smoky shrew (Sorex fumeus) and the
pygmy shrew (Sorex hoyi), which are restricted in Indiana to this region and the Highland
Rim (19).
Section 9B — Escarpment Section
This section includes the rugged hills situated along the eastern border of the
region. It is a blend of the Crawford Upland Section and the Mitchell Karst Plain
Section of the Highland Rim. Sandstone and sandstone derived soils (Wellston-Zanesville)
cap most of the hills, and the lower elevations present limestone and limestone-derived
soils (Crider, Hagerstown, Bedford, and Corydon). Sandstone cliffs and rockhouses
260 Indiana Academy of Science Vol. 94 (1985)
are virtually unknown, but, limestone crops out to form large cliffs, especially along
the Ohio River, and smaller stream courses. Karst features are not uncommon, especially
in the lower and middle elevations. The natural communities consist of various upland
forest types, especially dry-mesic and mesic. The species composition is similar to that
of the Crawford Upland Section, except that certain species, e.g. post oak (Quercus
stellata) and black oak (Q. velutina) commonly replace chestnut oak (Q. prinus) in
the dry sites, and some of the mesic cove species, especially those with Appalachian
affinities, are absent. Limestone glades and barrens occur in this section, but are not
nearly as common as in the Highland Rim region. Limestone cliff communities occur
mostly at the southern end of the section. Here, rare calciphiles such as alumroot
(Heuchera villosa), wall-rue spleenwort (Asplenium ruta-muraria), cleft phlox (Phlox
bifida var. stellaria), wild liveforever (Sedum telephioides), and black-seeded sedge (Carex
eburnea) occur. Eastern woodrats (Neotoma floridana) inhabit the crevices of cliffs
along the Ohio River, which is also a favorite roosting and nesting site for black vultures
(Coragyps atratus). Cave communities are common in this section, where some of the
largest caves in Indiana occur. They support an unique fauna, including a troglobitic
crayfish (Orconectes inermis) and the northern cavefish (Amblyopsis spelaea). Some
caves support large populations of hibernating bats, especially the endangered Indiana
bat (Myotis sodalis). Limestone gravel wash communities are well represented here,
and are similar to the same community type in the Highland Rim and Bluegrass Natural
Regions. The wild blue indigo (Baptisia australis) is apparently confined in Indiana
to this community type in this section. The typical aquatic features include normally
clear, medium and high-gradient streams, springs, and sinkhole ponds. The lower Blue
River is an exceptionally fine example of a larger stream in this section.
Region Ten — Highland Rim Natural Region
This natural region occupies in part the Highland Rim physiographic region of
the Interior Low Plateaus that occurs in a discontinuous belt from northern Alabama
through Tennessee, Kentucky, and into Indiana (65). The underlying strata are
predominantly of Mississippian age, although some Pennsylvanian aged strata crop
out in places. The region is unglaciated, except for relatively unmodified glaciated
areas at the northern and eastern boundary. A distinctive feature of this region is the
large expanse of karst topography, although several other major topographic features
are known including cliffs and rugged hills. Much of the area was forested in presettle-
ment times, but large areas of barrens occurred along with smaller areas of glade
(limestone and siltstone) and gravel wash communities.
This natural region is divided into three sections: the Mitchell Karst Plain Sec-
tion, the Brown County Hills Section and the Knobstone Escarpment Section, They
essentially occupy three of Malott's (45) physiographic regions: the Mitchell Plain,
the Norman Upland, and the Scottsburg Lowland.
Ecological studies in this region include Cain (14), Lindsey and Schmelz (42),
Potzger (58), McQueeny (47), Keith (38), Bacone et al. (7), Aldrich et al. (3), and
Homoya and Hedge (35).
Section 10A — Mitchell Karst Plain Section
The major feature of this section is the karst (sinkhole) plain. Several natural
community types are associated with this plain, including cave, sinkhole pond and
swamp, flatwoods, barrens, limestone glade and several upland forest types. The plain
is relatively level, although in some areas, especially near the section's periphery,
limestone cliffs and rugged hills are present. Caves are common. The soils are generally
well drained silty loams derived from loess and weathered limestone. Acid cherty Bax-
Ecology 261
ter silt loam is present mostly in the south (correlating somewhat with the barrens
community type), as is the netural to basic Corydon stony silt loam (correlating with
the limestone glade and cliff community type). Crider silt loam is a major soil throughout
most of the region. Possibly the largest area of barrens in Indiana was located in this
section. Species commonly found in remnants of this prairie-like community include
Indian grass (Sorghastrum nutans), big bluestem (Andropogon gerardi), little bluestem
(Andropogon scoparius), rattlesnake master (Eryngium yuccifolium), prairie dock
(Silphium terebinthinaceum), hairy sunflower (Helianthus mollis), prairie willow (Salix
humilis), clasping milkweed (Asclepias amplexicaulis) and Carex meadii. Most of
Indiana's limestone glades occur in this region, particularly in Harrison and Washington
Counties. This bedrock community, like the barrens, has a prairie flora with addi-
tional distinctive glade species including downy milk pea (Galactia volubilis var. mississip-
piensis), angle-pod (Gonolobus obliquus), axe-shaped St. John's-wort (Hypericum
dolabriforme), adder's tongue fern (Ophioglossum engelmannii), crested coral root
orchid (Hexalectris spicata), and heartleaf alexander (Zizia aptera). Gravel wash com-
munities composed of limestone and chert gravel border most streams. Characteristic
species include big bluestem, Indian grass, Carolina willow (Salix caroliniana), water
willow (Justicia americana), ninebark (Physocarpus opulifolius), pale dogwood (Cornus
obliqua), and bulrush (Scirpus americanus). Karst wetland communities are the major
aquatic features of the section. Southern swamp species are known from some of the sinkhole
swamps, including beakrush (Rhynchospora corniculata), log sedge (Carex decomposita),
giant sedge (C. gigantea), Virginia willow (Itea virginicia), and small buttercup (Ranun-
culus pusillus), and netted chain fern (Woodwardia areolata). Usual dominants of these
swamps are swamp cottonwood (Populus heterophylla), pin oak (Quercus palustris),
swamp white oak (Q. bicolor), red maple (Acer rubrum), and sweet gum (Liquidam-
bar styraciflua). Sinkhole pond communities normally have open water and marshy
borders with cattails (Typha latifolia), bulrush (Scirpus validus), bur-reed (Sparganium
androcladum), spatterdock (Nuphar advena), buttonbush (Cephalanthus occidentalis),
swamp loosestrife (Decodon verticillatus), bladderwort (Utricularia gibba) and Carex
comosa. Several forest communities are present in the section, but the western mesophytic
forest type predominates (41), in which white oak (Quercus alba), sugar maple (Acer
saccharum), shagbark hickory (Carya ovata), pignut hickory (C. glabra), and white
ash (Fraxinus americana) are typical. Near the glade communities, some xeric forest
occurs in which post oak (Quercus stellata), chinquapin oak (Q. muhlenbergii) and
blue ash (Fraxinus quadrangulata) are characteristic. Chestnut oak (Quercus prinus),
a very common component of the Brown County Hills Section and the Knobstone
Escarpment Section, is uncommon in this section. State restricted species include quillwort
(Isoetes engelmannii), netted chain fern, monkshood (Aconitum uncinatum), mannagrass
(Glyceria acutiflora), blackstem spleenwort (Asplenium resiliens), glade violet (Viola
egglestonii), and southern cavefish (Typhlichthys subterraneus). In karst areas, surface
streams are few. Typical examples include medium and high-gradient streams with rocky
bottoms, e.g. Indian Creek, Clear Creek, Buck Creek, and upper stretches of the Blue
River.
Section 10B — Brown County Hills Section
This section is characterized by deeply dissected uplands underlain by siltstone,
shale, and sandstone. The soils are well drained acid silt loams with minor amounts
of loess, specifically the Berks-Gilpin-Weikert Association. Bedrock is near the sur-
face, but rarely crops out. The natural communities are rather uniform in composi-
tion, with uplands dominated by oak-hickory, especially chestnut oak (Quercus prinus),
and ravines with mesic species, e.g. beech (Fagus grandifolia), red oak (Q. rubra),
262 Indiana Academy of Science Vol. 94 (1985)
sugar maple (Acer saccharum), and white ash (Fraxinus americana). Typically, upper
slopes have an almost pure stand of chestnut oak, a thick growth of greenbriar (Smilax
spp.), low growing shrubs (Gaylussacia baccata and Vaccinium vacillans), and a carpet
of sedges, notably Carex picta. The latter is essentially restricted in Indiana to this
section, and yet is ubiquitous here. Yellowwood {Cladrastis kentuckea) is known in Indiana
only from a small area of this section. The green adder's mouth orchid (Malaxis unifolia),
trailing arbutus (Epigaea repens), and large whorled pogonia orchid (Isotria verticillata)
are geographically restricted here except for single collections of the latter two in the
Knobstone Escarpment Section. One occurrence of an acid seep spring community is
known (58). Small, high-gradient ephemeral streams are common. Most larger streams
are predominantly medium to low-gradient streams, e.g. Guthrie Creek, and all forks
of Salt Creek.
Section IOC — Knobstone Escarpment Section
This section is similar to the Brown County Hills Section in terms of substrate
and topography, but is distinguished by floristic, faunistic, and compositional differences
of the forest communities. The major compositional difference is the presence of Virginia
pine (Pinus virginiana) in the upland forest communities. The pine is commonly a
co-dominant with chestnut oak (Quercus prinus) on many of the ridge crests and south-
facing slopes. American chestnut (Castanea dentata) was a dominant historically, given
the frequency that it was mentioned in the survey records of the General Land Office
and its continued presence today as stump sprouts. Its place has been taken by chestnut
oak. Carex picta, a species common in the Brown County Hills Section, is rare here.
Rock outcrops are few and are restricted to ridge tops. Glades with a shaly substrate
(fragments of siltstone, shale, and sandstone) are present but rare and normally occur
on south-facing slopes. They are typically rather sterile environments primarily because
of the unstable substrate and harsh climatic conditions. Typical associates include scat-
tered clumps of little bluestem (Andropogon scoparius), goat's rue (Tephrosia virgi-
niana), bird-foot violet (Viola pedata), and St. Andrew's cross (Ascyrum hypericoides).
Xeric forests of blackjack oak (Quercus marilandica), chestnut oak, and scarlet oak
(Q. coccinea) commonly border these glades. Species restricted in Indiana to this sec-
tion include stout goldenrod (Solidago squarrosa), rattlesnake-weed (Hieracium
venosum), bluegrass (Poa cuspidata), Virginia pine, red salamander (Pseudotriton ruber),
scarlet snake (Cemophora coccinea), and crowned snake (Tantilla coronata). Small,
and ephermeral high-gradient streams are the major aquatic features of this section.
Typical larger streams include Muddy Fork of Silver Creek, Buffalo Creek, Twin Creek
and Rush Creek.
Region Eleven — Bluegrass Natural Region
This natural region is identified and named not for a predominance of bluegrass
(Poa spp.), but for similarities of the physiography and natural communities to the
Bluegrass region of Kentucky. Traditionally, this portion of Indiana has not been con-
sidered a part of the Interior Low Plateaus Bluegrass Region as outlined by Fenneman
(25). However, several geologists have pointed out similarities in the Kentucky Bluegrass
Region and the Indiana area, including Malott (45) and Ray (66), the latter placing
them together in the Bluegrass part of the Interior Low Plateaus. Major portions of
three of Malott's (45) physiographic regions are included in the Bluegrass Natural Region:
the Dearborn Upland, the Muscatatuck Regional Slope, and the Scottsburg Lowland.
The three sections of this natural region, the Switzerland Hills Section, the Muscatatuck
Flats and Canyons Section, and the Scottsburg Lowland Section, approximate the area
of these physiographic units.
Ecology 263
Although the entire natural region has been covered by one or more of the pre-
Wisconsin ice sheets, today much of it is mantled by only a relatively thin veneer
of till. The northern boundary of the region approximates the southern terminus of
Wisconsinan glaciation. This boundary marks the northern limit in this region for several
southern plant species, as well as many herpetofaunal species (74).
Most of the natural region was originally forested, although a few glade, cliff,
and barrens remnants are known, as well as non-forested aquatic communities. Ecological
studies in the region include those of McCoy (46), Chapman (17), Potzger and Chandler
(60, 61), Reidhead (67), and Jackson and Allen (37).
Section 11A — Scottsburg Lowland Section
The main features of this section are the wide alluvial and lacustrine plains that
border the major streams, particularly the Muscatatuck River, the East Fork of White
River, Silver Creek, and their tributaries. Major soils are acid to neutral silt loams,
particularly of the Stendal, Atkins, Haymond, and Wilbur series. A sizable area of
eolian sand occurs just east of the East Fork of the White River, but no unique com-
munities or species are known to have been associated with it. Bedrock rarely crops
out, the major exception being the Falls of the Ohio near Clarksville. Predominant
natural communities are floodplain forest and swamp, although areas of upland forest
are included that grade into the Muscatatuck Flats and Canyons Section. The swamp
community is characterized by the occurrence of swamp cottonwood (Populus
heterophylla), red maple (Acer rubrum), pin oak (Quercus palustris), river birch (Betula
nigra), green ash {Fraxinus pennsylvanica), stiff dogwood (Cornus foemina), and button-
bush (Cephalanthus occidentalis). The slightly better drained floodplain forest adds
sweetgum (Liquidambar styraciflua), swamp chestnut oak (Quercus michauxii), swamp
white oak (Q. bicolor), American elm (Ulmus americana), black gum (Nyssa sylvatica),
beech (Fagus grandifolia), shellbark hickory (Carya laciniosd), and rarely, pecan (Carya
illinoensis) . Characteristic herbs include Carex muskingumensis, C. louisianica, Virginia
day flower (Commelina virgin ica), lizard's tail (Saururus cernuus), and woodreed (Cinna
arundinacea). The very rare southern pale green orchid (Platanthera flava var. flavd)
is geographically restricted here, as are the northern copperbelly (Nerodia erythrogaster
neglecta), and the eastern ribbon snake (Thamnophis sauritus sauritis). The northern
studfish (Fundulus catenatus) is known in Indiana only from streams in the far northern
portion of this section. State restricted plants include the extinct stipuled scurf-pea
(Psoralea stipulatd), and the extirpated Short's goldenrod (Solidago shortii). Wetland
features in this section include swamps, acid seep springs, low-gradient, silty-bottomed
streams and rivers and ponds. Were it not for the location of this section, it con-
ceivably could fit into the Southern Bottomlands Natural Region.
Section 1 1 B — Muscatatuck Flats and Canyons Section
This section consists primarily of a broad, relatively flat west sloping plain with
steep walled canyons entrenched by major streams. The plain is characterized best
by the presence of poorly drained, acidic Cobbsfork and Avonburg silt loam soils and
the occurrence of a southern flatwoods natural community type. These flatwoods
typically have beech (Fagus grandifolia), red maple (Acer rubrum), sweetgum (Liquidam-
bar styraciflua), pin oak (Quercus palustris), swamp chestnut oak (Q. michauxii), and
tulip tree (Liriodendron tulipifera). A few species are restricted geographically here,
including fox grape (Vitis labrusca), blunt-lobed grape fern (Botrychium oneidense),
swamp dewberry (Rubus hispidus), dwarf ginseng (Panax trifolium) and false lily-of-
the-valley (Maianthemum canadense). In canyons, cliffs and slopes of Silurian and
Devonian limestone provide an environment quite unlike the flats. These sites are com-
264 Indiana Academy of Science Vol. 94 (1985)
paratively rich floristically, and have a predominantly mixed mesophytic forest com-
postiion. Canada violet (Viola canadensis), longspur violet (V. rostrata), and crinkleroot
(Dentaria diphylla) are more common here than elsewhere in southern Indiana. American
pennywort (Hydrocotyle americana), wideleaf ladies' tresses (Spiranthes lucida), and
Carex pedunculata are restricted geographically here. Sullivantia (Sullivantia sullivan-
tii) and golden St. John's-wort (Hypericum frondosum) are known in Indiana only
from canyons in this section. The dusky salamander (Desmognathus fuscus) is a distinc-
tive species of this section and the Bluegrass Natural Region. Non-forested community
types include small areas of limestone gravel wash and limestone glade, the latter har-
boring the only Indiana occurrence of Michaux leavenworthia (Leavenworthia uniflora).
Minor areas of karst topography occur along valley borders. The major aquatic features
include medium-gradient streams with beds of pavement-like limestone, such as Graham
Creek, Big Creek, and the upper stretches of the Vernon Fork of the Muscatatuck River.
Section 11C — Switzerland Hills Section
This section is characterized by deeply dissected uplands composed of calcareous
shale and limestone of Ordovician age. Bedrock is near the surface, but cliffs are rare.
The area was glaciated, yet unconsolidated deposits are thin or absent. The Eden,
Switzerland, and Pate neutral silty clay loams are the dominant soils series. Most of
the natural communities are forested, although a few barrens remnants are known.
A mixed mesophytic forest type is well represented, especially in the ravines. These
forests should not be confused with the mixed mesophytic forests of the Cumberland
Mountains as described by Braun (12), for there is little similarity in terms of floral
composition, bedrock, soils, etc. Characteristic tree species include beech (Fagus gran-
difolia), white ash (Fraxinus americana), sugar maple (Acer saccharum), white oak
(Quercus alba), chinquapin oak (Q. muhlenbergii), red oak (Q. rubra), shagbark hickory
(Carya ovata), blue ash (Fraxinus quadrangulata), tulip tree (Liriodendron tulipifera),
Ohio buckeye (Aesculus glabra), and black walnut (Juglans nigra), with occasional
occurrences of yellow buckeye (Aesculus octandra), and white bass wood (Tilia
heterophylla). Historical evidence indicates that this area, especially along the Ohio
River, possibly may be the only location where black locust (Robinia pseudoacacia)
is native in the state (20, 29). Although no indigenous plant species is unique to this
section, two species are more common here than elsewhere in the state, namely a fox-
glove (Penstemon canescens), and Kentucky viburnum (Viburnum molle). The ravine
salamander (Plethodon richmondi) is essentially restricted in Indiana to this section.
Rocky, gravel-bottomed, medium-gradient streams such as Laughery Creek and
Whitewater River, typify the major aquatic features of the region.
Region Twelve — Big Rivers Natural Region
This aquatic natural region includes those rivers (or portions of rivers) where
the average flow is 7000 cubic feet per second or greater. This includes all of the Ohio
River bordering Indiana, the White River up to the confluence of its two forks, and
the Wabash River from its mouth to near Attica in Fountain County. These rivers
provide an environment for several species not found in smaller riverine systems, e.g.
the lake sturgeon (Acipenser fulvescens), shovelnose sturgeon (Scaphirhynchus platoryn-
chus), alligator gar (Lepisosteus spatula), shortnose gar (Lepisosteus platostomus), ship-
jack herring (Alosa chrysochloris), smallmouth buffalofish (Ictiobus bublaus), goldeye
(Hiodon alosoides), mooneye (Hiodon tergisus), and the blue sucker (Cycleptus
elongatus).
Mussel species distinctive of the Big Rivers Region include the fat pocketbook
pearly mussel (Potamilus capax), white cat's paw pearly mussel (Epioblasma sulcata
Ecology 265
delicata), tubercled-blossom pearly mussel (E. torulosa torulosa), pink mucket pearly
mussel (Lampsilis orbiculata), and Sampson's pearly mussel (Epioblasma sampsoni —
extinct). The alligator snapping turtle (Macroclemys temmincki), and the hellbender
(Cryptobranchus alleganiensis) are characteristic species of this region, but currently
are very rare if not absent. At least one vascular plant is state restricted to this region,
that being riverweed (Podostemum ceratophyllum).
Acknowledgments
Special appreciation is directed to the many individuals who contributed to the
creation of this paper. Those individuals who provided helpful suggestions and criticisms
include John Bacone, Lee Casebere, James Gammon, Henry Gray, Cloyce Hedge,
Max Hutchinson, James Keith, Alton Lindsey, Sherman Minton, Larry Morse, Robert
Petty, Richard Powell, John Schwegman, John Whitaker and John White. Technical
assistance was provided by Tammy Carrigg, Terri Engle, Marilyn Glander, Hank Huff-
man, Nancy Lax Kozar, Tim Renner, Bonnie Thomas, Jerrie Worthy, and especially
by the principal author's wife, Barbara Homoya. A special thanks to the Department
of Natural Resources, especially the Division of Nature Preserves, for the interest and
support of this project.
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57. Post, T.W., J. A. Bacone, and J.R. Aldrich. 1985. Gravel hill prairies of Indiana.
Proc. Indiana Acad. Sci. 94: (in press).
58. Potzger, J.E. 1934. A notable case of bog formation. Amer. Midland Nat.
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59. . 1935. Topography and forest types in a central Indiana region. Amer.
Midland Nat. 16:212-229.
60. . and L. Chandler. 1950. Beech in the forest about Laughery Creek Valley.
Proc. Indiana Acad. Sci. 59:82-94.
61. and L. Chandler. 1952. Oak forests in the Laughery Creek Valley, Indiana.
Proc. Indiana Acad. Sci. 62:129-135.
62. and R.C. Friesner. 1943. An ecological survey of Berkey Woods: A rem-
268 Indiana Academy of Science Vol. 94 (1985)
nant of forest primeval in Kosciusko County, Indiana. Butler Univ. Bot. Stud.
6:10-14.
63. , R.C. Friesner and CO. Keller. 1942. Phytosociology of the Cox Woods:
A remnant of forest primeval in Orange County, Indiana. Butler Univ. Bot. Stud.
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64. , M.E. Potzger and J. McCormick. 1956. The Forest primeval of Indiana
as recorded in the original U.S. land surveys and an evaluation of previous inter-
pretations of Indiana vegetation. Butler Univ. Bot. Stud. 13:95-111.
65. Quarterman, E. and R.L. Powell. 1978. Potential ecological/geological natural
landmarks of the Interior Low Plateaus. U.S. Department of the Interior,
Washington, D.C. 738 p.
66. Ray, L.L. 1974. Geomorphology and Quaternary geology of the glaciated Ohio
River Valley — a reconnaissance study. U.S.G.S., Prof. Paper 826, 77 p.
67. Reidhead, V.A. 1984. A reconstruction of the presettlement vegetation of the
middle Ohio Valley region, pp. 386-426, in Patrick J. Munson, Ed. Experiments
and Observation on Aboriginal Wild Plan Food Utilization in Eastern North
America. Prehistory Research Series. Indiana Historical Society 6.
68. Ridgway, Robert. 1872. Notes on the vegetation of the Lower Wabash Valley.
Amer. Naturalist 6:658-665.
69. Rohr, F.W. and J.E. Potzger. 1951. Forest and prairie in three Northwestern
Indiana counties. Butler Univ. Bot. Stud. 10:61-70.
70. Schneck, J. 1876. Catalogue of the flora of the Wabash Valley below the mouth
of White River, and observations thereon. Ann. Rept. Indiana Geol. Survey
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Features of Indiana. Indiana Academy of Science, Indianapolis, Indiana.
72. Schwegman, J.E., principal author. 1973. Comprehensive plan for the Illinois
Natural Reserves System. Part 2. The natural divisions of Illinois. Illinois Nature
Preserves Commission, Rockford, 111. 32 p.
73. Scott, Will. 1905. The Leesburg Swamp. Proc. Indiana Acad. Sci. 15:209-226.
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fauna of Illinois and Indiana. Amer. Midland Nat. 58:341-351.
75. Thorn, R.H. and J.H. Wilson. 1980. The natural divisions of Missouri. Transac-
tions of the Missouri Acad. Sci. 14:9-23.
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Indiana Academy of Science, Indianapolis, Indiana.
78. and J.H. Zumberge. 1965. Pleistocene geology of Indiana and Michigan,
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Lakeshore. Indiana Dunes National Lakeshore Research Program. Report 80-01.
262 p.
Successional Relationships of Pine Stands at Indiana Dunes
Eric S. Menges and Thomas V. Armentano
Holcomb Research Institute
Butler University
Indianapolis, Indiana 46208
Originally presented as
Demographic and Community Aspects of White Pine and Jack Pine
in Lake Michigan Dune Ecosystems
Introduction
Controversy about plant succession, its pathways, characteristics, rates, and
mechanisms, continues to be a significant part of plant ecology today (e.g., 5, 10,
22, 25, 26). The first major formulation of succession as a theory (7) was based largely
on research at Indiana Dunes. Cowles (6) recognized that the parallel sand ridges,
each marking the southern border of the Lake Michigan shoreline for a definable period
following regional deglaciation, provided a unique opportunity to interpret the tem-
poral dynamics of vegetation development. Later, Olson (23, 24) more quantitatively
analyzed vegetation and soil patterns at the dunes and advanced successional theory.
Cowles hypothesized that long-term changes in the vegetation composition of a
site could be related to amelioration of harsh physical conditions, and that improve-
ment in soil properties mediated changes by successive plant communities. Olson (23,
24) emphasized that at Indiana Dunes, several plant communities thought by Cowles
to be serai might be more or less permanent, largely because of nutrient limitations.
Recently, the role of periodic fire in maintaining the integrity of a variety of dune
communities has been emphasized (3, 13, 32).
Within the Indiana Dunes area, individual species that are rare or at range limits
have been considerable interest (32). These include two pine species, white pine (Pinus
strobus) and jack pine {Pinus banksiana). Both species, particularly jack pine, are found
at Indiana Dunes near their range limits (1 1) and are considered to be boreal relicts (32).
Both Cowles and Olson considered pine stands to be successional at the dunes,
in keeping with their apparent role at other sites. Cowles concluded that upland pine
communities were usually replaced by oak-dominated communities. He did not com-
ment specifically on the fate of lowland pine stands. While Olson (24) convincingly
demonstrated that conversions of oak to sugar maple as hypothesized by Cowles were
unlikely, he also concluded that oak would usually "quickly replace" pine in upland
sites. The present paper summarizes community and population characteristics of pine
stands at Indiana Dunes, and interprets the data in relation to the successional status
of the two species.
Our study was conducted in 1984, about 35 and 85 years after observations by
Olson and Cowles, respectively. Although successional theory has evolved since these
studies, interpretation of successional dynamics in forests still depends largely on in-
ferences from stand structure and compositions at a single point in time. Long-term,
replicable data sets are rare. Although we were unable to relocate plots established
by Olson, our sampling stations were established at nearby locations closely similar
in vegetation and site properties. In several cases we analyzed the same stands sampled
by Olson (pers. comm.).
Methods
To evaluate white and jack pine populations, the present distribution of both
species at Indiana Dunes National Lakeshore (IDNL) and Indiana Dunes State Park
269
270 Indiana Academy of Science Vol. 94 (1985)
(IDSP) was determined. Outside these properties, few undisturbed populations of either
species exist on dunes habitat within Indiana. Low altitude aerial photographs (1:400)
and previous reports by Wilhelm (32) and Krekeler (18) were used to locate stands.
All sites likely to contain populations of either pine species were visited by the authors.
For purposes of this investigation, a population was defined as a grouping of >15
individuals of a species within 0.4 ha.
Field work was done during the 1984 growing season. Pine stands were sampled
using 100 m2 (5.64 m) circular quadrats. Quadrats were stratified randomly along
transects to efficiently cover intrastand variability. For stands limited in extent, con-
tiguous square quadrats or complete sampling were used. All quadrat centers were
marked with metal pins.
All trees (woody stems 2.5 cm dbh or larger) in the quadrat were measured for
diameter at breast height (dbh). We also recorded the presence of herbaceous species
in each quadrat. Community attributes were sampled in June and July, and follow-up
surveys were conducted in late August and September. Vouchers were collected.
Nomenclature follows Wilhelm (32).
We sampled pine trees more intensively, adding trees outside quadrats to increase
sample size to 80 or more when possible. Heights of pine seedlings and samplings
(< 2.5 cm dbh) in quadrats were measured. Increment cores were obtained from IDNL
pines in late summer 1984. However, complete age-structures were taken for only two
of the smaller populations of each species. Additional populations were partially cored
nonrandomly to assure coverage of a range of sizes.
Community analysis considered species presence in quadrats, and weighted all
species (trees, shrubs, herbs) equally. We included species with two or more occur-
rences in our samples. A polythetic divisine clustering technique called TWINSPAN
(two-way indicator species analysis) was used to group floristically similar quadrats
and co-occurring species. This technique, described elsewhere (12, 16), is considered
to give particularly lucid placement of samples within a dendogram, and also forms
divisions that may reflect secondary gradients (12). Community relationships also were
interpreted, using detrended correspondence analysis (DC A). This iterative procedure
ordinates species and samples simultaneously, and is effective in removing the arch
distortion characteristic of many other multivariate techniques (12).
Results
Distribution of Pine Stands
The largest populations of white pine remaining in the lakeshore area are located
within state park boundaries. A total of seven white pine populations are located in
the dunes area (Table 1). Population SP-7, located at the eastern end of the state
park near the Keiser Survey Unit (KE-1), is the largest, consisting of 84 individuals
(Figure 1, Table 1). This population is among the most diverse in terms of tree sizes.
The second largest population is located in the Keiser Survey unit of the IDNL, less
than 0.5 km east of the SP-7 and consists of 81 individuals located in a mesic pocket
behind primary dunes. The remaining populations studied are far smaller in size (Table
1). These sites currently support mixed hardwood pine forests in mesic pockets or on
dune slopes.
Nine populations of jack pine were selected for sampling (Figure 1, Table 1);
these ranged in number of trees from less than 50 to over 300. Several populations
were in interdunal depressions (pannes) that hold temporary standing water in the spring,
or were located adjacent to permanent ponds. Others were found on open slopes, in
woodlands on dune-complexes (sensu 32) or in mixed-hardwood stands on slopes or
DA-1
5 WP, >300 JP (most sapling
and tree-sized)
DA-2
22 WP (Nearly all old growth)
DA-3
52 JP (all old growth)
Miller Dunes —
east of steel mills, near slag ponds
MD-1
47 JP
MD-2
-300 JP
Keiser — east of
state park road parking lot
KE-1
81 WP
State Park
SP-1
17 WP, >300 JP
SP-7
84 WP
Ecology 271
Table 1 . Location and Status of Studied Jack Pine (JP) and White Pine (WP) Populations
in Indiana Dunes
Site Population Size Comments
Ogden Dunes — West Beach Unit, east of town of Odgen Dunes
OD-5 34 WP, many JP Old-growth woods on east facing dune slope, most
JP on north and east edges; WP scattered in interior
OD-1 >300 JP Open pine stand, edge of panne just behind lakefront
dune
West Beach — West Beach Unit, west of Odgen Dunes, east of parking lot, just behind primary dunes
WB-1 >300 JP (most tree-sized) Open pine stand on well-drained dunes
WB-2 >300 JP (most seedlings) Open pine stand adjacent to panne and ponds
Dune Acres — west of town of Dune acres, along shore (DA-1); in mesic pocket, south of DA-1 (DA-2); and east
of Mineral Springs Road, ca 1 km north of Cowles Bog (DA-3)
Open pine stand on lakefront dune
Old-growth forest in mesic pocket between dunes
Closed swamp forest adjacent to marsh
Dunes, upslope from pond
Dune flat adjacent to panne and pond
Mesic pocket and adjacent dune ridge
Open lakefront dunes and mixed woods
Mixed secondary woods, west of State Park road,
ca 1 km south of lake front.
in dune flats. Only one of the sampled populations was located in IDSP. Several other
populations in IDSP were not sampled.
Our inventory of IDNL populations reveals that outside of Pinhook Bog (over
15 km to the southwest), no natural stands of jack or white pine are located east of
the white pine population KE-1 or south of U.S. Route 12. Several additional stations
of planted white pine were not considered in this study. These additional sites contain
too few individuals of either species to meet our definition of a population.
All pine populations occurred on areas mapped in soil surveys (30, 31) either
as dune land or Oakville fine sand. Soils are composed of fine sand with some medium
sand and fine gravel, and have little or no horizon development. They are extremely
low in moisture-holding capacity, with neutral to acid pH.
Community Analysis of Pine Stands
Classification of Pine Stands. Pine stands at Indiana Dunes were divided by
TWINSPAN into four groups (Figure 2). In labeling such stands, we relied both on
field observations, known autecologies of major species, and previous work at Indiana
Dunes (6, 18, 24, 32).
(1) Jack-pine-dominated, open panne communities with an incomplete canopy,
located near temporary or permanent bodies of water (WB-2, MD-2, OD-1).
These stands had moderately high similarity to each other.
(2) Jack-pine-dominated woodlands and dunes in upland areas (WB-1, MD-1,
SP-1, DA-1). These dune-complex areas were generally extensive, structural-
ly heterogeneous areas. Canopies were mostly open. The four stands were
very similar to each other compositionally.
272
Indiana Academy of Science
Vol. 94 (1985)
«
« o
c e
z
O <t
Z l
<t *
z "
-I
1
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Figure 1. Location of white and jack pine populations analyzed in this study.
Ecology 273
% Similarity (2W/A+B) Within Groups
en
en
A
■P>.
co
CO
Ol
i
O
i
Ol
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1
Ol
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0D5 KE1 .
WB1 MD1 SP1 DA1 .
WB2 MD2 OD1
Figure 2. Cluster diagram of Indiana Dunes sites, based on TWINSPAN classification
analysis, with similarities within and between groups calculated by city-block distance
(2W/A + B where A are species in one quadrat, B those in the second, and W those in
common; distance was averaged over all quadrats).
(3) Mixed hardwood-pine forests, less protected or drier than the following group.
The two sites included were OD-5, a slope forest with both pine species,
and KE-1, a pocket behind large curving dunes with white pine and hard-
wood trees. We followed previous convention in describing these areas as
"mesic pockets."
(4) Mesic or wet-mesic forests on dune flats. Two are mixed hardwood-pine
stands: DA-3, a swamp forest with jack pine, and SP-7, a mixed hardwood
forest with white pine. They contain species interpreted as indicating
mesophytism or association with wet soils including Fraxinus americana, Acer
rubrum and Nyssa sylvatica.
A TWINSPAN classification at the quadrat level provided additional detail on what
species are significant indicators of various groups of floristically similar quadrats.
Within pine stands at Indiana Dunes, the major division was between closed
forest/woodland areas and more open woodlands and dune formations (Figure 3).
Many other species, particularly shade-intolerant trees (e.g., Populus deltoides), wetland
plants (e.g., Hypericum kalmianum), and dunes forbs (e.g., Artemisia caudata), were
found only rarely in closed forests. Many other species, however, are characteristically
restricted to forests, including Acer rubrum, Prunus serotina, and Sassafras albidum.
The major TWINSPAN division within closed forests distinguishes wet-mesic forests
on dune flats (DA-3, SP-7) from upland mesic forests and woodlands (Figure 3). In
open areas, the major division also results from apparent moisture (Figure 3). Areas
adjacent to standing water, often pannes (OD-1, WB-1), support shade-intolerant
moisture-loving species such as Hypericum kalmianum and Sabatia angu/aris. The op-
posing species are characteristic of dry, open dunes and woodlands.
Further divisions in the cluster analysis often can be attributed to more local
factors. For example, the driest open areas are subdivided (at level 3) into open forma-
tions dominated by dune grasses and annuals versus more shrubby thicket areas. A
division of quadrats within the mesic pocket DA-2 reflects canopy gaps that favor
274
Indiana Academy of Science
Vol. 94 (1985)
LEVEL OF DIVISION
*
Sites
Quads
(# Q)
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Figure 3. Cluster diagram of IDNL quadrats, based on TWINSPAN classification
analysis. Indicated within dendrogram are environmental differences between branches.
Along the left are shown the sites to which the quadrats belong and the number of
quadrats found in each cluster.
gap-phase trees over shade-tolerant shrubs and herbs. Although some subsequent divi-
sions reflect finer distinctions in moisture, cover, and species diversity, other clusters
are not associated with recognizable environmental factors.
Ecology
275
Ordination of Pine Stands. The major axis of variation in the DCA ordination
reflects crown cover: it extends from open sites (dry and wet) to closed forests. Quadrats
from OD-1 to SP-7 define the endpoints of this axis, which passes respectively through
wet open areas, dry open areas, drier forests and woodlands, and mesic forests (Figure
4). The second major axis separates forests stands only, from the swamp forests at
DA-3 to the mesic pocket at DA-2. SP-7 and KE-1 occupy intermediate positions.
This ordination emphasizes the distinctness of several sites: the mesic pocket DA-2,
the swamp forest DA-3, and the panne site MD-2. Other sites feature a compositional
range. For example, OD-1 contains quadrats in temporary pannes with unique floristic
elements, and also quadrats on sandy ridges similar to the upland dunes sites (Figure
4). A compositional continuum is also evident in the upland dune areas, from quite
open areas all the way to fairly mesic forests. Some sites (KE-1, MD-1) have rather
narrow ranges within the continuum, but WB-1, SP-1, DA-1, and OD-5 all span a
broad compositional range. SP-1 is the most heterogeneous, with some forest area
similar to the mesic pocket KE-1, and other open areas similar to OD-1.
DCA uses both sample (quadrat) and species distributions in its iterative analysis;
thus, a map of species centroids is directly comparable to the sample map. In Figure
5, some species centroids are indicated. Their distribution is generally similar to sam-
ple distribution, with first-axis variation showing a gradient from open (Typha latifolia,
Opuntia humifusa) to closed (Osmunda claytoniana), and second-axis reflecting mesic
(Viburnum acerifolium) to wet-mesic {Quercus palustris) species. The species distribu-
0D1
Wet & Open
Figure 4. Location of sample quadrats on first two axes of DCA ordination. Lines
encompass all quadrats belonging to sites indicated in bold letters; major environmental
factors are also indicated.
276
Indiana Academy of Science
Vol. 94 (1985)
tion shows more intermediate centroid placement for species occupying a range of
habitat conditions. For example, Rhus radicans and Pinus banksiana, both located
near the center of the ordination, range from wet-mesic to dry forests and woodlands
to more open dune areas, although Pinus banksiana is notably absent from most closed
forests (Figure 5).
CO
X
<
AXIS 1
Figure 5. Location of major species centroids on first two axes of DCA ordination.
Lines encompass all species locations.
The preceding discussion was based on sample and species locations or graphs
of the first two axes of DCA, i.e., those explaining the greatest amount of variation.
The major effect of the third axis is to better separate some quadrants in WB-1 from
the majority of those in SP-1 and DA-1. Species with strong correlations in the direc-
tion of this separation include open-sand specialists such as Opuntia humifusa and
Populus deltoides, as well as some weedy elements (e.g., Saponaria officinalis). We
believe this compositional gradient reflects erosional damage from heavy recreational
use of the West Beach Unit.
Structure of Pine Stands
Five jack pine populations occurred in areas with little other tree cover. WB-2
has no other tree-size vegetation, while at MD-1, MD-2, and OD-2, only a few in-
dividuals of other tree species occurred. These four sites all have less than 650 dm2
basal area/ha. Reproduction of jack pine is especially dense in the panne areas, but
few seedlings of other species were sampled.
At WB-1, jack pine accounted for 81% of sampled stems greater than 2.5 cm
in diameter. Nearly all other species were understory trees or shrubs (Prunus virgi-
niana, Amelanchier sp., Ptelea trifoliata) but seedlings were relatively sparse. At DA-1,
Ecology
277
▲ ▲ Pinus banksiana
• • Quercus velutina
O O Tilia amencana
■ ■ Amelanchier sp.
D---Q Others
Figure 6. Forest structure at DA-1.
44% of the tree-sized stems present were jack pine, as were many smaller trees and
seedlings (Figure 6). Reproduction of black oak, witch hazel {Hamamelis virginiana),
and basswood is restricted to the highest parts of the dune slope.
Mixed jack and white pine populations are present in the pine woodlands and
mixed forests at SP-1 and OD-5. At both sites, white pine, along with black oak at
SP-1, comprise the largest trees. However, white pine seedlings and midsized and smaller
trees are scarce compared to jack pine and other species (Figure 7). Jack pine reproduc-
tion is found largely in more open areas with cottonwood, red cedar {Juniperus virgi-
niana), and sand cherry (Prunus pumila).
In forests at the closed end of the first DCA axis, the forest structure is different.
White pine dominates the mesic pockets KE-1 and DA-2, as well as the mixed secon-
dary woods SP-7. In KE-1, it is the most common tree in every tree size class except
the two smallest. The smallest tree-size classes are dominated by understory tree species
(witch hazel, sassafras, and Amelanchier sp.), with few canopy species represented.
Seedlings are quite dense at KE-1, consisting mainly of hardwoods. In the mesic pocket
DA-2, white pine are mainly large, with no seedlings and only one sapling. In con-
trast, the next largest tree species, red oak, has been reproducing well (Figure 8). This
stand has a high basal area (4,260 dmVha), half again as great as KE-1. Small stems
of white ash (Fraxinus americana), red maple, and basswood are present, although
seedling density is low for all species.
SP-7 has a substantial number of smaller white pine, although seedlings are sparse.
278
Indiana Academy of Science
Vol. 94 (1985)
A —
SP1
A — -A Pinus banksiana
/V-A Pinu3 strobus
9 « Quercus velutina
V V Hamamelis virginiana
■ ■ Amelanchier sp.
Dp- -O Others
20 25
DBH (cm)
Figure 7. Forest structure at SP-1
35
25
20
O 15
Z
10
DA2
— A Pinus strobus
• Quercus rubra
— O Quercus alba
A Prunus serotina
X K Hamamelis virginiana
x- X Sassafras albidum
D -Q Others
5 10 15 20 25 30 35 40 45 50 55 60 65 70 75
DBH (cm)
Figure 8. Forest structure at DA-2.
Ecology 279
All sizes of white oak are widely distributed, whereas many young black gum and
red maple are found mainly in the wetter areas of this stand. Shrub density at SP-7
is the highest of any of the sites sampled, with six species occurring in 20% or more
of sampled segments.
At DA-3, a swamp forest, neither small jack pine nor reproduction were noted.
The larger adult jack pine have very small crowns high on the bole and some appeared
to be senescent. In contrast, small trees and seedlings of several hardwood species,
especially red maple (Acer rubrum), are common. Basal area is quite high (3400 dmVha)
in this stand.
Size Distributions of Pines
Among the natural populations of white pine in the IDNL region, diameter distribu-
tions ranged from 2.5 cm (the minimum by definition), to a 75.1 cm tree found at
DA-2. Sapling and seedlings were less common, with only 50 individuals under 2.5
cm dbh encountered in the lakefront — less than a quarter of trees tallied.
Size distribution of white pine within individual populations varied (Figure 9).
The population in one mesic pocket (DA-2) consisted mainly of trees ranging from
15 to 75 cm, and completely lacked small trees, saplings, or seedlings. In contrast,
the third mesic pocket at Keiser (KE-1) lacked the largest size classes (> 65 cm) but
contained some seedlings. Many of these, however, were diseased. We found few recruits
into small tree classes at Keiser. Overall, recruitment of white pine in mesic pockets
is poor.
The other three white pine populations sampled consisted primarily of medium-
sized or small trees (Figure 9). All contained small numbers of saplings and/or seedl-
ings, probably insufficient at present to maintain population levels in the future. Perhaps
the most unusual stand was found at SP-7, where small white pine form a scattered
but consistent understory beneath part of a mixed hardwood forest.
Jack pine exists as a small tree in the lakeshore areas, with increasing numbers
from larger to smaller size classes. Over 40% of tree-sized individuals were 7.5 cm
dbh or less. Reproduction, as interpreted by seedling and sapling occurrence, was com-
mon, and these size classes contained twice the number of trees. At all locations, small
trees and juveniles were uncommon under closed canopies.
The distribution of jack pine sizes was much less variable than that of white pine
(Figure 10). Seven of the nine sampled populations were dominated numerically by
seedlings or samplings, with small trees (2.5-7.5 cm dbh) making up the majority of
> 2.5 cm dbh individuals. Populations MD-2, WB-2, and OD-1, all located near pannes
or ponds, lacked larger trees and were numerically dominated by seedling size classes.
Drier sites with abundant reproduction and medium-sized dominants were found at
WB-1 and MD-1. The two populations with somewhat lower levels of reproduction
and relatively large trees were at OD-5 and DA-1. Both sites contained trees in fairly
open areas, dominated entirely by jack pine, but grading into nearly closed woodland
with a mixture of hardwoods, some overtopping the pines.
An eighth population, SP-1, was similar to OD-1 and DA-1 in physiognomy and
site, but reproduction was poor. The jack pine population, located in a swamp forest
at DA-3, consists of medium-sized and larger trees, nearly all with meager crowns
located far from the ground.
Age Structures and Long-term Growth of Pine Populations
Jack pine populations at IDNL differ in age. Although the small population at
Miller Dunes (MD-1) consists of trees ranging up to 73 years of age (Figure 11), most
trees originated between 15 to 30 years ago. A conspicuous gap in the age distribution
indicates that no trees presently found were recruited between 1928 and 1949. This
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Indiana Academy of Science
Vol. 94 (1985)
w
No. Trees
O r-
cn co *$ in to r^
1 l 1 1 1 1
<
/
No. Trees
o
Q
O
No. Trees
cn n rj- id co r^
No. Trees
o o o
Figure 9. Size distribution of white pine populations sampled in the Indiana Dunes region.
age gap, however, is not reflected in a similar gap in size structure.
Jack pine populations in the wetter areas are both smaller and younger than at
MD-1. The age of the 20 trees cored at MD-2 ranged only up to 24 years. The largest
tree at OD-1 was 64 years old, but most were probably much younger. At MD-2,
trees have been continuously recruited since the recent origin of the population.
Ecology
281
E
0-50
51-100
101+
2.5-7.5
7.6-12.5
12.6-17.5
17.6-22.5
22.6-27.5
27.6-32.5
32.6+
No. Trees
o o o
No. Trees
o o o
No. Trees
X CO
V
Figure 10. Size distribution of jack pine populations sampled in the Indiana Dunes region.
The other two jack pine populations that were partially sampled for age, WB-1
and DA-1, consist of trees with age distributions similar to MD-1. Maximum ages
are 65 years in WB-1 and 58 years in DA-1. Both have a preponderance of individuals
between 15 and 20 years of age. Another parallel is apparent in these three popula-
282
Indiana Academy of Science
Vol. 94 (1985)
CO
LU
LU
CC
h-
o
CC
LU
OQ
D
10r-
10
20
30
40
50
60
70
80
90 100
TREE AGE
Figure 11. Age distribution of jack pine at MD-1.
tions: a striking gap in the number of individuals recruited in the 1930s and 1940s
(Table 2).
Table 2. Apparent Absence of Jack Pine Recruitment over Two Decades as Determined
by Increment Coring, Indiana Dunes National Lakeshore
Year of First Seedling
Entire
Youngest
Oldest
No. Trees
Population
Oldest
Pre- 1935
Post-1935
Population
Cored
Sampled
Tree
Recruit
Recruit
MD-1
19
Yes
1911
1928
1949
MD-2
20
No
1960
—
1964
WB-1
27
No
1919
1930
1949
DA-1
25
No
1926
1932
1949
OD-5
2
No
1914*
—
—
OD-1
2
No
1920*
—
—
♦Largest trees found at these sites were cored and aged.
Overall, jack pine size and age distributions are highly correlated, suggesting a
lack of suppression. Jack pine size is a good predictor of age with < orrelation coeffi-
cients about 0.7 in 3 of 4 populations. Regression slopes indicate that diameter in-
creases by 0.69 to 1.27 cm in an average year. This high growth rate, if maintained
for 60 years, places lakeshore jack pines into a "good" site index (11).
White Pine. A complete age distribution for white pine is available from DA-2,
a mesic pocket site on the lakefront (Figure 12). This population includes the largest
(75.1 cm dbh) and oldest (162 years) white pines sampled. The latter individual has
a fire scar dating from about 1879, the approximate date that the five next largest
pines were established. All other white pines in DA-2, with one exception, are 68-111
years old; the number present slowly declines in younger age classes. Only one living
tree originated within the last 64 years — a 19-year-old tree established on an encroaching
dune.
Ecology 283
10T
0)
LU
UJ
oc
h-
u_
O 5
QC
LU
m
z
. n
m m
, probable scar date
□ ,
10 20 30 40 50 60 70 80 90 100 110 120 130 140 150 160 170 180 190
TREE AGE
Figure 12. Age structure of white pine population at DA-2.
The history of population KE-1, the only other white pine population available
for aging, was similar in history to DA-2. The oldest tree was 132 years old; peak
recruitment occurred between 80 and 110 years ago; and only two trees sampled were
younger than 60 years. While seedlings exist in this population, poor recruitment of
canopy trees since 1900 suggests that high seedling mortality has prevailed.
The size of white pine at DA-2 is fairly closely related to age (r = 0.69, p <
0.001), and the average year resulted in an increment of 0.47 cm dbh. This growth
is near the bottom of a range cited for dominant white pine (11). Growth at Keiser
has been slower and more variable, and the age size correlation (r = 0.30) is not
significant.
Discussion
Significant differences in the structure of the pine stands and the makeup of the
pine populations reflect both the highly diverse dune environments and the distinct
roles of the two species. Most evident is that white pine is less widely distributed and
less vigorous than jack pine at Indiana Dunes. This distinction is curious, given that
jack pine is at its southern range limit and is uncommon in most other southern Lake
Michigan dunes (6; person, observ. of authors), whereas white pine is or was found
widely in the region. However, industrial and urban development have destroyed areas
that supported stands of both species (29, 32). White pine was highly valued as lumber,
and merchantable trees may have been largely removed from the dunes area by the
early twentieth century.
Based on site physiognomy, and confirmed by quantitative vegetation analyses,
jack and white pine are components of many community types in the dunes. Based
on ordinations, cluster analyses, and analysis of forest structure, five types of pine
communities can be distinguished: lowland pannes, pine woodlands, dry-mesic mixed
slope forests, mesic pockets, and swamp forests. Jack pine is lacking only in mesic
pockets, while white pine is absent in lowland pannes, dune-complex woodlands, and
some swamp forests. The pine woodlands [pine dune of Krekeler (18)] are most
widespread. Such woodlands are one of several intergrading community types in the
dune-complex (32). Lowland pannes [pine bottoms, (6)] and mesic pockets are the
most floristically distinct [see (6) for a detailed discussion]. The swamp forests are
a heterogeneous group; our stands resemble the hydromesophytic forest/conifer swamp
and pin oak flat classifications of Wilhelm (6). Pines were heavily logged in these com-
munities (6) and current stands may be recovering from that disturbance.
The ordering of community types from open to closed formations also suggests
the successional sequences proposed by Cowles (6, 7) and Olson (24). Pines are generally
284 Indiana Academy of Science Vol. 94 (1985)
assumed to play a serai role at the dunes. What evidence does our data provide to
suggest that those community types represent a successional series? We examined the
evidence separately for jack pine and white pine communities.
Jack Pine. Size and age distribution and seedling recruitment of jack pine suggest
that at several sites the species is a long-term feature of the dunes landscape. Jack
pine should survive in the near future at all but one (DA-3) of the studied sites, although
it may share dominance with basswood and black oak in some local areas. Reproduc-
tion and dominance are particularly impressive in the wet open areas lacking signifi-
cant numbers of other tree species. Jack pine establishment is commonly highest in
moist, open situations (4, 11, 27).
The role of fire in jack pine persistance at the dunes is unclear. Although over
most of its range the species carries serotinous cones, this is not generally true of dunes
populations. In most areas, however, jack pine success is related to periodic burning
(3, 8, 28), although Cowles (6) thought that fire would promote a shift from pines
to oaks in dune systems. The availability of unforested areas with a mineral soil seed-
bed at the dunes appears to allow continued recruitment and survival of jack pine
without fire. Fire currently occurs at fairly high frequencies in oak stands characteristical-
ly lacking either pine species at Indiana Dunes (13), but little information has been
summarized for other communities.
Among the several tree survivorship models suggested to have general interpretative
value, the negative exponential or "reverse J" size distribution is thought to represent
a population in which mortality is constant over a range of sizes, and reproduction
is sufficient to maintain population levels (20, 21). White pine recruitment is quite
low, causing important deviations from a negative exponential curve. In contrast, jack
pine distribution at Indiana Dunes is very closely modeled by the negative exponential
(r2 = .922, P < 0.001), strong evidence that jack pine self-replacement is likely. Rapid
and consistent jack pine growth at four dunes sites suggests its lack of shade tolerance
(11) and the lack of success of competing species. We suspect that stands dominated
by jack pine, those with abundant reproduction, have been characterized generally
by open canopies.
Jack pine populations also contain all ages, a typical condition for self-reproducing
stands (1, 19). Although jack pine has established periodically during this century,
no survivors were found that established between 1928 and 1949. The recruitment gap
may be related to the effect of drought on germination and subsequent establishment,
but pollution stress also may have been involved.
The forest structure of jack pine stands, its continued recruitment in most stands,
the present density of juvenile individuals, and a healthy growth rate all suggest that
jack pine is self-replacing in many Indiana Dunes stands. Conditions appear too harsh
for more shade-tolerant species to be important invaders.
Although Cowles (6, 7) emphasized the great resistance of the dune systems to
change, he stated that "probably the oaks follow the pines, but the evidence on which
this is based is not voluminous" (7, p. 174). He excluded exposed dune crests from
this shift; there, pines might persist (6). Olson (24) suggested that first-generation pine
stands would be "rapidly replaced" by black oak. He compared "invading" stands
of young pine with decadent, adjacent stands at OD-5. However, 34 years later, hard-
woods are still absent from jack-pine-dominated portions of these stands; the pine
species being replaced are largely white pine. Thus, jack pine replacement by oaks,
if it occurs, takes place only quite gradually. In contrast to assertions of both Cowles
and Olson, jack pine upland communities are largely stable, compositionally.
Cowles suggested that oaks should be excluded by the hydric conditions of pine
bottoms, but did not speculate on succession there. Downing (9) described pannes as
temporary phases in dune formation and Olson (24) implied as much in his model
Ecology 285
of community change. However, we agree with Wilhelm (32) that jack-pine-dominated
pannes have a high degree of stability.
White Pine. Unlike jack pine forests, white pine forests appear to be a temporary
stage of vegetation succession throughout Indiana dunes. White pine appears to be
in demographic decline at all sites, because more shade-tolerant species dominate all
but the upper vegetation stratum. Only at SP-7 is there evidence that white pine is
competing successfully. Depending on site, replacement by oaks (black, red, white),
basswood, or red maple is likely, although dense populations of understory trees (especial-
ly witch hazel) and shrubs in mesic and wet-mesic forests may slow the transition.
The growth rates of adult white pine at two sites have been slow and variable, sug-
gesting that competition may be inhibiting vigor.
The low levels of current white pine recruitment are long-standing, judging by
age distribution in residual stands. At one mesic pocket, fire may have eliminated most
white pine trees established before 1879, while providing suitable conditions for recruit-
ment. This evidence is consistent with previous work showing that white pine establish-
ment often responds well to fire (14, 17), although other disturbances such as clearing
can initiate regeneration. Further recruitment is sometimes possible in the absence of
major disturbance (15). Data on fire frequency at the Indiana Dunes area, with the
exception of oak stands (13), is not available.
The poor recruitment of white pine in the last half-century, the low numbers
of seedlings currently established, and the slow growth rate of adults suggest that white
pine forests may indeed be shifting toward hardwood domination. These results agree
with Cowles's (6) and Olson's (24) assessment of white pine stands at IDSP.
Acknowledgments
We wish to thank Jane Molofsky and Jeanette Daniel for their assistance in the
field, and Jane Molofsky for her help in data analysis. Jerry Olson contributed useful
ideas during several field trips. This research was supported by the Denver Office of
the Air and Water Quality Division of the National Park Service, and by Holcomb
Research Institute.
Literature Cited
1. Abrams, M.D. 1984. Uneven-aged jack pine in Michigan. Journal of Forestry
82(5):306-307.
2. Bacone, J.A., R.K. Campbell, and G.S. Wilhelm. 1980. Presettlement vegetation
of the Indiana Dunes National Lakeshore. Proc. Second Conference on Scien-
tific Research in the National Parks, San Francisco. Vol. 4. Resource Analysis
and Mapping. National Park Service, Washington, D.C. 364 pp.
3. Carlton, T.J. 1982. The composition, diversity, and heterogeneity of some jack
pine (Pinus banksiana) stands in northeastern Ontario. Canadian Journal of Botany
60(12):2629-2636.
4. Cayford, J.H., Z. Chrosciewicz, and H.P. Sims. 1967. A Review of Silvicultural
Research in Jack Pine. Canada Department of Forestry and Rural Development,
Forestry Branch, Departmental Publication No. 1173.
5. Connell, J.H. and R.O. Slatyer. 1977. Mechanisms of succession in natural com-
munities and their role in community stability and organization. Am. Nat.
111:1119-1144.
6. Cowles, H.C. 1899. The ecological relations of the vegetation on the sand dunes
of Lake Michigan. Botanical Gazette 27:95-117, 167-202, 281-308, 361-391.
7. Cowles, H.C. 1901. The physiographic ecology of Chicago and vicinity; a study
286 Indiana Academy of Science Vol. 94 (1985)
of the origin, development, and classification of plant societies. Botanical Gazette
31:73-108, 145-182.
8. Cwynar, L.C. 1977. The recent fire history of Barron Township, Algonquin Park,
Canadian Journal of Botany 55(11): 1524-1538.
9. Downing, E.R. 1922. A naturalist in the great lakes region. Univ. of Chicago
Press, Chicago, IL (cited in Wilhelm 1980).
10. Drury, W.T. and I.C.T. Nisbet. 1973. Succession. J. Arnold Arbor. (Harvard
Univ.) 54:331-368.
11. Fowells, H.A. 1965. Silvics of Forest Trees of the United States. U.S. Depart-
ment of Agriculture, Forest Service. Agriculture Handbook 271.
12. Gauch, H.G., Jr. 1982. Multivariate Analysis in Community Ecology. Cambridge
University Press, Cambridge.
13. Henderson, N.R. and J.N. Long. 1984. A comparison of stand structure and
fire history in two black oak woodlands in northwestern Indiana. Bot. Gaz.
145:222-228.
14. Henry, J.D. and J.M.A. Swan. 1974. Reconstructing forest history from live and
dead plant material — An approach to the study of forest succession in southwest
New Hampshire. Ecology 55(4):772-783.
15. Hibbs, D.E. 1982. White pine in the transition hardwood forest. Canadian Jour-
nal of Botany 60(10): 2046-205 3.
16. Hill, M.O. 1979. TWINSPAN— A FORTRAN program for arranging multivariate
data in an ordered two-way table by classification of the individuals and attributes.
Ithaca, NY. Cornell University (Cited in Guach 1982).
17. Horton, K.W. and G.H.D. Bedell. 1960. White and Red Pine: Ecology, Silviculture,
and Management. Canada Department of Northern Affairs and National Resources,
Forestry Branch. Bulletin 124. Ottawa, Ontario, Canada.
18. Krekeler, C.H. 1981. The Biota of the Indiana Dunes National Lakeshore. Chapter
3. In M. Reshkin, W.E. Keifer, C.H. Krekeler, N.V. Weber, and L. Brunansky
(eds.), Ecosystem Study of the Indiana Dunes National Lakeshore, Volume Two.
Indiana Dunes National Lakeshore Research Program, Report 81-01. U.S. Depart-
ment of the Interior, National Park Service, Midwest Region, pp. 3-1 to 3-346.
19. Larsen, W.C. 1982. Structure, Biomass, and Net Primary Productivity for an
Age-Sequence of Jack Pine Ecosystems. Ph.D. Dissertation, Michigan State Univer-
sity. (Cited in Abrams 1984.)
20. Lorimer, C.G. and L.E. Frelich. 1984. A simulation of equilibrium diameter
distributions of sugar maple {Acer saccharum). Bulletin of the Torrey Botanical
Club 11 1(2): 193-199.
21. Meyer, H.A. and D.D. Stevenson. 1943. The structure and growth of virgin beech-
birch-maple-hemlock forests in northern Pennsylvania. Journal of Agricultural
Research 67:465-484. (Cited in Lorimer and Frelich 1984.)
22. Odum, E.P. 1969. The strategy of ecosystem development. Science 164:262-270.
23. Olson, J.S. 1951. Vegetation-substrate relations in Lake Michigan sand dune
development. Ph.D. Dissertation, University of Chicago, Department of Botany,
Chicago, IL.
24. Olson, J.S. 1958. Rates of succession and soil changes on southern Lake Michigan
and sand dunes. Botanical Gazette 119(3): 125-170.
25. Peet, R.K. and N.L. Christensen. 1980. Succession: a population process. Vegetatio
43:131-140.
26. Pickett, S.T.A. 1982. Population patterns through twenty years of old field suc-
cession. Vegetatio 49:45-59.
27. Shirley, H.L. 1945. Reproduction of upland conifers in the Lake States as af-
fected by root competition and light. American Midland Naturalist 33(3):537-612.
Ecology 287
28. Swain, A.M. 1973. A history of fire and vegetation in northeastern Minnesota
as recorded in lake sediments. Quaternary Research 3(3):383-396.
29. Swink, F.A. and G. Wilhelm. 1979. Plants of the Chicago Region. 3rd edition.
Morton Arborteum. Lisle, IL.
30. U.S. Department of Agriculture. 1981. Soil Survey of Porter County, Indiana.
31. U.S. Department of Agriculture. 1972. Soil Survey of Lake County, Indiana.
32. Wilhelm, G.S. 1980. Report on the Special Vegetation of the Indiana Dunes Na-
tional Lakeshore. Indiana Dunes National Lakeshore Research Program, Report
80-01. U.S. Department of the Interior, National Park Service.
The Roots of Ecology in Indiana
Edwin R. Squiers
Department of Biology and Environmental Science
Taylor University, Upland, Indiana 46989
If history is the interaction between places and people, then there can be little
doubt that Indiana has an honored "place" in the history of ecology in North America.
In fact, Indiana is the place where modern ecological science finds its roots. The Lake
Michigan dunes of northwest Indiana served as the site for the pioneering research
of Henry Chandler Cowles. In his 1899 doctoral dissertation entitled "The Ecological
Relations of the Vegetation on the Sand Dunes of Lake Michigan," Cowles (1) defin-
ed the science for succeeding generations of ecologists:
"The province of ecology is to consider the mutual relationships between plants
and their environment."
Influenced by the ideas of European biogeographers such as E. Warming (2) and
A.F.W. Schimper (3), and geologists, especially T.C. Chamberlin, Cowles filled the
first several pages of his dissertation with his vision of the new science.
"The ecologist employs the methods of physiography, regarding the flora of a
pond or swamp or hillside not as a changeless landscape feature, but rather as
a panorama, never twice alike."
"Any plant society is the joint product of present and past environmental condi-
tions, and perhaps the latter are much more important than most ecologists have
thought."
"The ecologist, then, must study the order of succession of the plant societies
in the development of a region, and he must endeavor to discover the laws which
govern the panoramic changes. Ecology, therefore, is a study in dynamics."
Cowles recognized the division between community ecology (synecology) and popula-
tion ecology (autoecology). Of the former, he writes:
"The species characteristic of each formation must be discovered, together with
the facts and laws of their distribution. The progressive changes that take place
and the factors in the environment which cause these changes must be discussed."
and of the latter:
". . . it is the author's purpose to discuss the adaptations of the plants to their
dune environment, paying especial attention to those species which show a large
degree of plasticity, and which are found growing under widely divergent
conditions."
It is notable that Cowles chose the dunes of northwest Indiana as the site for his
study because he felt "that nowhere else could many of the living problems of ecology
be solved more clearly; that nowhere else could ecological principles be subjected to
a more rigid test." Thus, it is the rare ecology text that does not identify Indiana
as the "place" where ecology finds its roots in North America.
With ecology firmly rooted in Indiana as "place," let me address the question
of "person." It would be especially convenient at this point, to be able to say that
Cowles was a "Hoosier" (He was not.) or that he attended Indiana University (He
did not.) or that he taught at Butler or Purdue or Taylor or Hanover or any one
of a number of Indiana's fine old colleges and universities (No luck there either.).
289
290 Indiana Academy of Science Vol. 94 (1985)
Indiana's connection with the roots of ecology as "person" is more subtle, though
no less real. Again, we must look carefully at Cowles' dissertation. There, toward
the end of the introduction, Cowles gratefully acknowledges the "kindly interest and
cooperation shown by his associates . . . especially Head Professor John M. Coulter,
through whose influence the author was directed along lines of ecological research."
It seems, that Cowles' ideas were shaped by both a "place," the Indiana dunes, and
a "person," John M. Coulter.
John M. Coulter was a "Hoosier." Before accepting the position at the Univer-
sity of Chicago, Coulter had been associated with Hanover College (graduating in the
class of 1870), Wabash College and Indiana University. John was one of the founding
fathers of the Indiana Academy of Science, elected President in 1886-7 and made a
Fellow in 1893. Throughout his life, Dr. Coulter remained interested in the Academy
and its affairs, returning on several occasions to address the membership. One of John
Coulter's books, "Plant Relations" (4), firsst published in 1899, can be considered the
first North American ecology textbook. This volume offers modern plant ecologists
a fascinating look at the beginnings of their discipline. In the Preface to the 1901
edition, Coulter cites the "recent rapid development of the subject" and adds addi-
tional material, including several photographs, from Cowles research. It is notable
that John was not the only Coulter to make an impact on the Indiana Science, his
brother Stanley was also elected a Fellow of the Academy in 1893 and served as Presi-
dent in 1895-96. Stanley Coulter would later become the Dean of the Purdue Univer-
sity School of Forest Science.
The interaction between John Coulter and Henry Cowles reminds us again of
the importance of the relationship between professor and student in shaping the history
of science. Cowles had begun his graduate work at the University of Chicago in
geography, when Coulter, recognizing his potential, encouraged him first, to join the
fledging Department of Botany and finally to study the ecology of the Indiana dunes.
After completing his doctorate, Cowles remained at the University of Chicago as a
master teacher. One of his students would later write of him:
"No teacher brought his students more directly to nature. He was a master in
the field. ... He was at his genial best around campfires in the evening. It is
given to few men to found a new science and to live to see it well established." (5)
The "pedagogical genealogy" of American plant ecologists, as outlined by Sprugel
(6) in 1980, confirms Henry Chandler Cowles' extraordinary role in the development
of ecology in North America. Figure 1, though far from a complete listing, illustrates
the magnitude of Cowles' influence as a teacher. If Cowles is the "father of modern
ecology" then surely John M. Coulter, Indiana Academy of Science President and
Fellow, must be considered the "grandfather" of the science.
Although Henry Cowles never published in the Proceedings of the Indiana Academy
of Science, his ideas about ecology and succession influenced the research of Indiana
scientists. As early as 1905, Will Scott wrote the following concerning his research
on the Leesburg Swamp:
"One of the main purposes has been to test the theories and factors proposed
by Warming and Cowles. His (Cowles) most important conclusion is that plant
societies are intimately associated with the physiography of a region and as the
topographic forms change from one form to another the plant societies are also
modified." (7)
The Indiana Academy of Science, through John Coulter, left its mark on Henry
Cowles and Cowles would return the favor many times. For example, the 1917 edition
of the Proceedings contains a paper by M.S. Markle entitled "A Comparison of the
Ecology
COULTER Chamber 1 in
291
Trans eau
Braun
Shreve
Coital
Olmsted /\ Egler Bug 1 1 V\BauheMiire
Lindeman vj -w\
HcCormick food Wistendahl
Skiers
E. Qdum
Whit taker
Mi 1 lam
BornianK L Cooper
Woo dwell Bliss
Figure 1. Some of the "pedagogical descendents" of John M. Coulter and Henry
Chandler Cowles (after Sprugel (6)).
Plant Succession on Hudson River Limestone with that on Niagra Limestone, Near
Richmond, Indiana" (8). Millard Markle was a graduate student of Cowles at the
University of Chicago from 1910 to 1915. Markle would spend 58 years in active ser-
vice to the Academy, serving as its President in 1945 and authoring "The History
of Plant Taxonomy and Ecology in Indiana" in 1966 for Indiana's Sesquicentennial
celebration.
On 8 October 1965, the Ecology Section of the Indiana Academy of Science was
formally approved and at the 1966 annual meeting the first papers, a total of four,
were read. Today, the Section is alive and well with a membership of more than 330
and with participation at annual meetings averaging more than 20 presentations per
year. We've come along way John, I think you'd be proud.
I stand before you today as a plant ecologist, Chairman on the Ecology Section
of the Indiana Academy of Science at this centennial meeting, because of the influence
of a "place," the Indiana dunes, and a "person" John M. Coulter, through his stu-
dent Henry Chandler Cowles, through his student William S. Cooper, through his
student Murray F. Buell, and through his students Jack McCormick, Ralph E. Good,
and Warren A. Wistendahl. If John M. Coulter is the "grandfather" of ecology, then
I am his "great, great, great grandson." Thus, I find myself connected to the roots
of ecology in Indiana.
Literature Cited
1. Cowles, H.C. 1899. The ecological relations of the vegetation on the sand dunes
of Lake Michigan. Doctoral Dissertation. University of Chicago.
2. Warming, E. 1895. Plantsamfund. Copenhagen.
3. Schimper, A.F.W. 1898. Pflanzengeographie auf physiologischer. Grundlage, Jena.
4. Coulter, J.M. 1898. Plant relations. D. Appleton and Company, New York.
292 Indiana Academy of Science Vol. 94 (1985)
5. Markle, M.S. 1966. The history of plant taxonomy and ecology in Indiana. Proc.
Ind. Acad. Sci. 76:142-150.
6. Sprugel, D.G. 1980. A "pedagogical geneaology" of American plant ecologists.
Bull. Ecol. Soc. Amer. 61:197-200.
7. Scott, W. 1905. The Leesburg Swamp. Proc. Ind. Acad. Sci. 14:209-226.
8. Markle, M.S. 1917. A comparison of the plant succession on Hudson River
Limestone with that on Niagra Limestone near Richmond, Indiana. Proc. Ind.
Acad. Sci. 28:109-113.
ENGINEERING
Chairperson: David D. Chesak
Box 883
St. Joseph's College
Rensselaer, Indiana 47978
(219)866-7111
Chairperson-Elect: William Stanchina
Department of Electrical Engineering
Notre Dame University
Notre Dame, Indiana 46556
(219)239-5693
ABSTRACTS
The IAS Engineering Section: A Brief History. David D. Chesak, St. Joseph's College,
Rensselaer, Indiana 47978. A survey of Engineering Section activity and some
of the people involved will be made.
Stress Corrosion Cracking of Sensitized Austenitic Stainless Steels in Boric Acid Solu-
tion Containing Sulfur Oxyanions. S. Dhawale, Department of Chemistry, Indiana
University East, Richmond, Indiana 47374 and G. Crangnolino and D.D. Macdonald,
Ohio State University, Columbus Ohio. The stress corrosion cracking of Type
304 stainless steel in boric acid solution containing thiosulfate or tetrathionate at room
temperature was studied using the slow strain technique. The minimum concentration
of each species required for stress corrosion cracking was determined in experiments
at open circuit potentials. Studies on the potential dependence of stress corrosion cracking
showed that severe stress corrosion cracking occurs over a narrow range of potential near
the corrosion potential. Scanning electron microscopy was used to determine the resulting
corrosion morphology. No stress corrosion cracking was observed for 304 L and 316
L stainless steels under applied potential conditions.
An electrochemical method was used to study the degree of sensitization of 304
stainless steel and the effect of heat treatments.
Engineering and Science Education's Dilemma: Inadequate Science Programs in the
Public School System. Andrew Hollerman, Department of Physics, Purdue Univer-
sity, West Lafayette, Indiana 47907. The role of science education in today's society
has been changing in the last several years. The rapid increase of technology has caused
many educators to begin to doubt the quality of science training in our public educa-
tional system. The shortcomings of present science programs will be discussed. Per-
sonal experiences will be cited.
Prediction of the Variation of Azeotropic Compostion Using the Gibbs-Konovalov
Theorem. Scott Oblander and W.W. Bowden, Department of Chemical Engineer-
ing, Rose-Hulman Institute of Technology, Terre Haute, Indiana 47803. It has
long been known that the assumption of the simple Margules equation
ln7, = AX.2
7. = activity coefficient of component i
293
294
Indiana Academy of Science
Vol. 94 (1985)
X. = mole fraction of component j in liquid
A = empirically-determined constant
leads to the following simple equation
X,(T2)-'/2
X,(T,)-'/2
_L_xlnPXIi)
A2 P°2(T2)
1 lnP°(T.)
A, P°2(T.)
(1)
where
P°,P° = vapor pressures of 1 and 2 at T,, T2
A,,A2 = Margules constant at T,,T2
This paper investigates how well the Gibbs-Konovalov theorem predicts the variation
of azeotropic composition with pressure and temperature. The Gibbs-Konovalov theorem:
If a two-phase boundary curve passes through an extreme value the composition
of the two phases must be identical at that point.
The equation used to determine the extreme-point is as follows:
P = XJ.P1 + X2T2P°2 (2)
where
P = total pressure
X,X2 = mole fractions
7,, 72 = activity coefficients
P°,P°2 = vapor pressures
The activity coefficients were assumed to be related to composition and temperature
by the NRTL equation:
ln7,
X2
r2lG2
r,2G,
l(X, + X2G2,)2
(X2 + X,G,2);
(3)
ln72 = X,
T|2G,;
T2,G;
l(X2 + X,G12)2 (X, + X2G2,)2
(4)
Gji = expC-o-Tjj)
r:: = Sij-gjj
RT
(a)
(b)
(5)
Tij = gJi-gii
RT
(c)
The vapor pressures P° were related to temperature through the Antoine equation
Bi
logP°= A.
C +t
(6)
At a given temperature the conditions for an extremum in pressure in Equation (2)
Engineering 295
were investigated using ISML routines available on the Rose-Hulman VAX 780. At
a given pressure the conditions for an extremum in temperature in Equation (2) were
investigated. Calculated and experimental results are compared.
The PVT Behavior of Compressed Liquids. Dennis West and W.W. Bowden, Depart-
ment of Chemical Engineering, Rose-Hulman Institute of Technology, Terre Haute,
Indiana 47803. Since about 1895 the 'Tait' equation
V = Vs [1 - C(t)ln <5^±1>]
s B(t) + Ps
V = unit volume of liquid at Pressure P
Vs = unit volume of saturated liquid at Ps
C(t), B(t) = empirically-determined functions of temperature
has been used to correlate the PVT properties of compressed liquids. In this paper
it is shown that the following simpler, more physically-meaningful equation with more
easily-determined constants correlates the PVT data on 3 common liquids at least as
well as the Tait equation:
P-P
V = Vs (1 - J)
PCK
P-Ps
K = _^c_ = a(t) + b(t)Pr + c(t)Pr2
V-V
V
s
p = p/p
r c
Pc = critical pressure
a,b,c = empirically-determined functions of temperature.
Evaluation of Landsat Thematic Mapper Data for Classifying Forest Lands
Paul W. Mueller, Roger M. Hoffer, and John E. Jacobson
Department of Forestry and Natural Resources
and Laboratory for Applications of Remote Sensing (LARS)
Purdue University
West Lafayette, Indiana 47907
With the launch of Landsat- 1 in July of 1972, man entered a new era for obtain-
ing information about earth resources. Landsat- 1 was the first unmanned satellite design-
ed specifically for collecting data about earth resources on a global, repetitive, multispec-
tral basis.
Technology developed rapidly during the seventies for processing and analysis
of the digital multispectral scanner data that was collected by Landsat. There was a
great deal of interest in the multispectral data — commonly referred to as MSS data —
and many applications were developed. Two more Landsat satellites with MSS sensors
were launched before the end of the decade.
Another milestone in Earth resource observations occurred in July 1982 when
the fourth satellite in the Landsat series was launched. In addition to a MSS sensor,
a new improved sensor called the Thematic Mapper (TM) was carried aboard Landsat-4.
The TM sensor has improved spatial resolution and spectral dimensionality as com-
pared to the MSS sensor (see Table 1). The MSS sensor collects data in only four
Table 1. Comparison of Landsat Scanning Sensors. Adapted from (2).
Thematic Mapper
Multispectral Scanner
(TM)
(MSS)
Spectral Band
Wavelength
Spectral
Ground
Wavelength
Spectral
Ground
Designation
Range
Region
IFOV
Range
Region
IFOV
1
0.45-0.52 fim
Visible Blue
30 m
0.5-0.6 nm
Visible Green
80 m
2
0.52-0.60 Mm
Visible Green
30 m
0.6-0.7 ^m
Visible Red
80 m
3
0.63-0.69 nm
Visible Red
30 m
0.7-0.8 nm
Near Infrared
80 m
4
0.76-0.90 nm
Near Infrared
30 m
0.8-1.1 tim
Near Infrared
80 m
5
1.55-1.75 ^m
Middle Infrared
30 m
6
2.08-2.35 /xm
Middle Infrared
30 m
7
10.40-12.50 ftm
Thermal Infrared
120 m
spectral bands — two in the visible and two in the near infrared region of the elec-
tromagnetic spectrum-whereas the TM sensor collects data in seven spectral band —
three in the visible, one in the near infrared, two in the middle infrared, and one
in the thermal infrared region. Because of the relatively low level of energy emitted
in the thermal infrared region, the spatial resolution of this band is 120 meters — much
larger than the other TM or MSS bands. The resolution, expressed as instantaneous
field of view (IFOV), for the remaining six bands of the TM sensor is 30 meters as
opposed to approximately 80 meters for the MSS sensor.
Many studies, including those by Hoffer et al. (4), Kalensky and Scherk (5),
and Strahler et al. (7), have shown that MSS data is useful for classifying geographic
areas into broad cover types. Given the improvements of TM data, the purpose of
this study was to determine the utility of TM data for classifying a predominantly
forested area into broad cover types. The objectives were twofold:
297
298 Indiana Academy of Science Vol. 94 (1985)
1) Evaluate the utility of wintertime Thematic Mapper data for classifying forest
and other broad cover types using supervised training statistics and a minimum
distance classifier.
2) Determine the value of different wavelength bands and combinations of bands
for classifying the various cover types.
Procedures
The TM data were obtained by Landsat 4 on December 18, 1982. The study area
was composed of St. Regis Corporation land in Baker County, Florida. Reference
data used to interpret the TM data included 1:58,000 color infrared aerial photographs
obtained on January 24, 1983, and a forest stand map that included stand boundaries,
species, and ages. This map and the associated information was provided by the St.
Regis Forest Resource Information System (FRIS) Center. Field visits (August and
October 1984) to the study area by the authors provided a better understanding of
the characteristics of the forest and other cover types present. Comparisons of the
reference data and the spectral cluster maps proved to be very beneficial when analyz-
ing and interpreting the TM data.
The study area was predominantly forested. Major forest types in the area were
slash and longleaf pine (the former often in plantations), and also pondcypress and
mixed hardwoods principally occurring in shallow ponds or bays (1). A small number
of agricultural areas were located within the study area and a small amount of exposed
water was present.
After viewing the aerial photographs, the St. Regis forest stand map, and a gray-
scale printout of the TM data, it was determined that all the land cover types of the
study area could be divided into six broad cover type classes, called information classes.
The informational classes of interest included: three classes of pine forest — Young (0
to 5 years), Medium-Aged (6 to 10 years), and Older (11 or more years); Deciduous
Forest; Agricultural Areas; and Water. Because of spectral variability within some of
these informational classes, it was determined that nine spectral classes were needed
in order to adequately represent the informational classes defined.
As indicated previously, at any one instant of time, the Thematic Mapper scan-
ner on the Landsat satellite measures the reflectance and thermal emission in each
of seven wavelength bands over a resolution element (or pixel) that represents an area
on the ground of 30 meters by 30 meters (120 meters by 120 meters for band 7). These
measurements provide the sets of data values that define the spectral patterns of the
various cover types on the ground. In order to use a computer to classify satellite
spectral data, the analyst must "train" the computer to recognize specific spectral
patterns and then classify the data having these defined spectral patterns into the in-
formational classes of interest. Such computer classification is based upon statistical
pattern recognition theory — a well-documented body of knowledge used in many
disciplines (8).
The first step in computer classification involves the definition of a set of train-
ing data that statistically represents the informational classes of interest. This step is
one of the most critical parts of the entire classification procedure (3).
In our analysis, we started by studying the St. Regis forest stand map and color
infrared photographs and selecting potential training areas. Each training area involved
a single cover type. Several training areas were defined for each cover type, so every
spectral class in the study area would be represented in the training data set. The digital
format TM data were then displayed on a Comtal Vision One/20 digital display unit
as a color infrared composite (the digital equivalent of a color infrared photograph).
Engineering 299
The pixel coordinates of potential training areas were then designated. Each training
area consisted of several contiguous pixels, and at least three such training areas were
defined for each spectral class. Additional training areas were defined, if necessary,
so that a minimum of 70 pixels (10 times the maximum number of wavelength bands
used) would be included in the training statistics for each class, in-so-far as possible.
The statistical characteristics of the training areas were then defined using the
LARSYS software system. These statistics included the mean and covariance matrix
of the seven bands for each spectral class (6), and provided the information necessary
for computer classification of the various informational classes.
The next step involved the actual classification of the TM data. The classification
process involves the use of an algorithm to compare the training data statistics to the
reflectance and emission values measured by the TM scanner for each pixel in the
entire data set. Several classification algorithms are available within the LARSYS soft-
ware. For this study, we used the relatively simple and fast minimum Euclidean distance
classification algorithm of the CLASSIFYPOINTS processor. A detailed description
of this processor and the entire LARSYS software system is documented by Phillips (6).
To test quantitatively the accuracy of the classifications, a set of "test areas"
were defined. Each test area consisted of a block of pixels thought to be representative
of the six informational classes present. (Thus, a test area is very similar to a training
area, but is used for an entirely different purpose.) A systematic statistical sampling
procedure was used to define the test data locations so that the training areas and
test areas were obtained from mutually exclusive locations in the data set. Fifty-three
test fields totalling 2372 pixels were thus defined for this study.
In order to evaluate the utility of the various TM wavelength bands for purposes
of computer classification, a method to assess the information content of each wavelength
band and band combination was required. Part of the LARSYS software (i.e.
SEPARABILITY) involves a "feature selection" technique which allows the analyst
to determine the optimum combination of bands to use, given any set of one through
"n" wavelength bands. Transformed divergence (TD), a statistical distance measure,
is calculated between all possible pairs of spectral classes for the specific combination
of wavelength bands being considered. When the TD is large (e.g. values above 1900;
maximum is 2000), there is a high probability that the two spectral classes can be
discriminated and a correct classification will result (8).
For this study, the "best" combinations of wavelength bands for each set of
the one through seven bands of TM data were defined using the average and minimum
TD values. Large average and minimum TD values were desirable as this indicated
that the classes were spectrally separable. Generally, only the minimum TD values
defined for each pair of spectral classes representing different informational classes
(rather than spectral classes within the same informational class) were utilized.
Based upon the Transformed Divergence results for determining the optimum
one through seven wavelength band combinations, seven separate classifications of
the data were then obtained, and the results were quantitatively summarized using
the test fields that had been previously defined. The key point here is that the same
training and test data were used for each of the seven classifications — the only variables
were the number and combinations of wavelength bands utilized.
Results and Discussion
The "best" channel combinations and their average transformed divergences are
summarized in Table 2. The performances for each of the seven classifications were
assessed using the classification results for the test fields. Table 3 is the LARSYS-
generated classification performance matrix for the "best" combination of four bands,
300
Indiana Academy of Science
Vol. 94 (1985)
Table 2. "Best" wavelength band combinations selected and their associated average
and minimum transformed divergences.
Number
CHANNELS
Bands
TRANSFORMED DIVERGENCL
Minimum
Average
5
4,5
4,5,7
3,4,5,7
3,4,5,6,7
2,3,4,5,6,7
1,2,3,4,5,6,7
5321
1781
1837'
1938'
1950'
1952'
1955'
1761
1980
1987
1991
1992
1993
1993
'Lower transformed divergence did occur between two spectral classes within the same information class.
namely, bands 3, 4, 5, and 7, showing how the test pixels were classified. Such a matrix
was generated for each of the seven classifications. The classification results for all
classifications are summarized in Table 4.
Table 3. Classification Performance Matrix for the "Best" Combination of Four Bands.
(Fl = Young Pine Forest, F+ = Medium- Aged Pine Forest, F* = Older Pine Forest,
FD = Deciduous Forest, AA = Agricultural Areas, WW = Water)
FOREST
LABORATORY FOR APPLICATIONS OF REMOTE SENSING OCT. 31, 1984
RESULTS 4
PURDUE UNIVERSITY
08 48 01 AM
LARSYS VERSION 3
CLASSIFICATION STUDY
430540387 CLASSIFIED OCT. 31, 1984
CLASSIFICATION WRITTEN ON DISK
CHANNELS USED
Channel 3
Spectral Band
0.63 TO 0.69 Micrometers
Calibration Code =1 CO = .0
Channel 4
Spectral Band
0.76 TO 0.90 Micrometers
Calibration Code =1 CO = .0
Channel 5
Spectral Band
1.55 TO 1.75 Micrometers
Calibration Code =1 CO = .0
Channel 7
Spectral Band
10.40 TO 12.50 Micrometers
Calibration Code =1 CO = .0
SPECTRAL
INFORMATION
CLASSES
SPECTRAL INFORMATION
CLASS
CLASS
CLASS CLASS
1 Fl
Fl
6
Al AA
2 F +
F +
7
A2 AA
3 F*
F*
8
A3 AA
4 FD
FD
9
w WW
5 FDC
FD
TEST CLASS PERFORMANCE
NUMBER OF SAMPLES CLASSIFIED INTO
INFORMATION
NO OF
PCT.
CLASS
SAMPS
CORCT
1 Fl
351
71.8
2 F +
235
98.3
3 F*
1141
99.1
4 FD
432
95.4
5 AA
207
97.1
6 WW
6
83.3
TOTAL
2372
Fl
F +
F*
FD
AA
WW
252
0
0
0
99
0
3
231
0
0
0
0
0
5
1131
5
0
0
0
5
15
412
0
0
6
0
0
0
201
0
0
0
0
1
0
5
261
241
1147
418
300
Overall performance ( 2232/ 2372) = 94.1
Average Performance By Class ( 545.0/ 6) = 90.8
10103 CPU TIME USED WAS 3.813 SECONDS
(LARSMN)
Engineering
301
Table 4. Summary of Classification Results.
Classification Performance (%)
Number
of
Test
Pixels
Number of TM Wavebands
(Specific TM Wavebands)
1
(5)
2
(4,5)
3 4
(4,5,7) (3,4,5,7)
5
(3-7)
6
(2-7)
7
(1-7)
Young Pine
Forest
351
68.9
72.4
72.4
71.8
66.4
66.4
64.7
Medium-Aged
Pine Forest
235
70.6
97.4
97.4
98.3
98.3
98.3
99.1
Older Pine
Forest
1141
81.3
98.9
98.9
99.1
99.6
99.6
99.5
Deciduous
Forest
432
93.8
95.4
95.6
95.4
95.1
95.1
94.9
Agriculture
Areas
207
81.2
97.1
97.1
97.1
96.1
96.1
- 96.6
Water
6
83.3
83.3
83.3
83.3
83.3
83.3
83.3
Overall
Performance
2372
80.7
94.0
94.1
94.1
93.4
93.4
93.2
Average
By Class
79.9
90.8
90.8
90.8
89.8
89.8
89.7
The overall performance for the classifications was high in all cases, except when
only one TM wavelength band was used. Disregarding the one-band classification, the
classification performance values for the individual informational classes were also
very high except for the Young Pine Forest and Water classes. The low performance
for the Young Pine Forest class is due to the fact that a significant number of pixels
were being misclassified into the Agricultural Areas class. This is not surprising since
the class Young Pine Forest includes recently harvested areas which consist of residual
understory vegetation mixed with bare soil. This is spectrally similar to the situation
often found in Agricultural Areas where agricultural crops and bare soil are mixed.
This confusion is illustrated in Table 3 where Fl is the Young Pine Forest information
class and AA is the Agricultural Areas information class. The relatively low classifica-
tion performance for water stems from the fact that there was very little exposed water
in the study area. With LARSYS, the test field must be rectangular. In the process
of selecting a rectangular test field for a small, non-rectangular water body, one pixel
(of six) was apparently an edge pixel — a mixture of two spectral classes — and was
therefore misclassified. The small number of Water test pixels is directly related to
the small amount of exposed water in the study area.
Conclusions
The results of this study show that:
Forest and other broad cover type groups can be classified with a high degree
of accuracy using wintertime Landsat Thematic Mapper data.
Even the relatively simple minimum distance classification algorithm achieved highly
accurate classification results for the six informational classes defined.
The 1.55-1.75 /mi middle infrared wavelength band was found to be the single
most useful band for discrimination between the spectral classes defined.
302 Indiana Academy of Science Vol. 94 (1985)
The "best" combination of two wavelength bands included a band in the near
infrared (0.76-0.90 /mi) and a band in the middle infrared (1.55-1.75 /mi) portion
of the electromagentic spectrum.
The 10.4-12.5 /im thermal infrared wavelength band appears to provide signifi-
cant additional information for the classification process.
The "best" combination of four wavelength bands included one band from each
of the four major portions of the spectrum— visible, near infrared, middle in-
frared, and the thermal infrared.
Acknowledgments
The authors would like to express their appreciation to St. Regis Corporation
for their cooperation and help. This project was supported in part by NASA Contract
NAS-26859.
Literature Cited
1. Avers, P.E., and K.C. Bracy. Soils and physiography of the Osceola National
Forest. U.S. Department of Agriculture Forrest Service, Southern Region. 94.
2. Freden, S.C., and F. Gordon, Jr. 1983. Landsat Satellites. Chapter 12 in: Col-
well, R.N. (ed.), Manual of Remote Sensing. American Society of Photogrammetry,
Falls Church, Virginia, pp. 517-570.
3. Hoffer, R.M. 1981. Computer-aided analysis of remote sensor data: magic, mystery,
or myth? Proceedings of Remote Sensing for Natural Resources: An Interna-
tional View of Problems, Promises, and Accomplishments, University of Idaho,
Moscow, Idaho, pp. 156-179.
4. Hoffer, R.M., and LARS Staff. 1973. Techniques for computer-aided analysis
of ERTS-1 data, useful in geologic, forest and water resource surveys. Proceedings
of the Third Earth Resources Technology Satellite- 1 Symposium, NASA Goddard
Space Flight Center, Washington, D.C., Volume 1, Section A. pp. 1687-1708.
5. Kalensky, Z., and L.R. Scherk. 1975. Accuracy of forest mapping from Landsat
computer compatible tapes. Proceedings of the 10th International Symposium
on Remote Sensing of Environment, Ann Arbor, Michigan, pp. 1159-1167.
6. Phillips, T.L. (ed.). 1973. LARSYS users manual. Laboratory for Applications
of Remote Sensing, Purdue University.
7. Strahler, A.H., T.L. Logan, and N.A. Bryant, 1978. Improving forest cover
classification accuracy from Landsat by incorporating topographic information.
Proceedings of the 12th International Symposium on Remote Sensing of Environ-
ment, Ann Arbor, Michigan, pp. 927-942.
8. Swain, P.H. 1978. Fundamentals of pattern recognition in remote sensing. Chapter
three in: P.H. Swain and S.M. Davis (eds.), Remote Sensing: The Quantitative
Approach. McGraw-Hill, Inc. pp. 136-187.
ENTOMOLOGY
Chairperson: Paul Robert Grimstad
Department of Biology
University of Notre Dame
Notre Dame, Indiana 46556
(219)239-5493
Chairperson-Elect: James Haddock
Department of Biological Sciences
Indiana University-Purdue University
at Fort Wayne
2101 Coliseum Boulevard East
Fort Wayne, Indiana 46805
(219)482-5254
ABSTRACTS
Effect of Barley Yellow-dwarf Virus Infection of Wheat and Oats on the Life Cycle
of Rhopalosiphum padi (L.). Jaime E. Araya and John E. Foster, Department of
Entomology and the U.S. Department of Agriculture, Purdue University, West Lafayette,
Indiana 47907. The life cycle of the bird cherry oat aphid, Rhopalosiphum padi
(L.), was studied in the laboratory comparing specimens carrying Barley Yellow-dwarf
virus (BYDV, PAV isolate) and virus-free aphids. Sections of leaves of wheat cultivars
'Abe' and 'Caldwell,' and oats 'Clintland 64' and 'Porter,' infected with BYDV and
virus-free, were used to rear the aphids in Petri dishes at 18 ± 1°C. Daily observations
were recorded for pre-and reproductive periods, life duration, adult life, total number
of progeny produced, mean progeny produced, and mean number of nymphs per day
during the reproductive period of all treatments. The data were analyzed separately
for each crop by ANOVAs and the Student-Neuman Keuls' test was used to separate
means (P = 0.05).
The data showed the aphids had a shorter life period and adult life in virus-
infected wheat plant material. There were also differences for life duration when analyz-
ing wheat cultivars x BYDV-infection. Virus infection in wheat increased the reproductive
capacity of the aphid. No significant differences were detected when using oats. Fur-
ther studies are needed to clarify the epidemiological relationships of all strains of
BYDV, their vectors, and plant cultivars.
Efficiency of Pollen Traps with Various Sized Trap Screens. William E. Chaney,
R.P.E., Extension Apiculturist, Purdue University, West Lafayette, Indiana
47907. In recent years beekeepers have become interested in trapping the pollen
pellets from incoming foraging bees for a variety of reasons. These reasons include:
1) Trapping pollen for sale 2) Trapping pollen to feed to different hives or the same
hive at a later date 3) Prevent the hive from becoming pollen-bound 4) Preventing
pesticide contaminated pollen from being stored in the hive.
Five sizes of wire mesh were tested in identical traps randomly assigned to a dif-
ferent hive of approximately equal strength. One of these meshes is the commonly
recommended size. The study was replicated in three locations. The trapped pollen
was collected regularly and weighed. Halfway through the experiment the traps were
randomly reassigned within the five hives in each location.
None of the four meshes tested was found to be better than the currently recom-
303
304 Indiana Academy of Science Vol. 94 (1985)
mended size. The size of the pollen pellet influenced the effectiveness of the various
sized meshes. The size of the pollen pellet was determined mostly by the plant foraged
and by the habits of individual foraging bees.
Effect of Viruliferous and Non-viruliferous Rhopalosiphum padi (L). Aphids on Winter
Wheat. B.H. Chen, J.E. Foster, and H.W. Ohm. Departments of Entomology, U.S.
Department of Agriculture and Department of Agronomy, Purdue University, West
Lafayette, Indiana 47907. The bird-cherry oat aphid, Rhopalosiphum padi (L.)
is capable of damaging cereal crops by direct feeding and by transmitting the barley
yellow dwarf virus (BYDV). Experiments were conducted to determine the effect of
viruliferous and non-viruliferous R. padi on two wheat cultivars, Caldwell and Abe,
and one wheat germplasm line, Elmo, in the greenhouse. R. padi without carrying
any isolate of BYDV and those with PAV isolate were used for non-viruliferous and
viruliferous infestation respectively. Results indicated that both viruliferous and non-
viruliferous R. padi significantly affected tiller number, kernel number, and kernel
weight per plant of Abe and Elmo. The non-viruliferous and viruliferous aphids reduced
the weight of kernels per plant of Abe 37% and 48%, respectively. No significant
reductions in these yield components were found on Caldwell plants infested with non-
viruliferous aphids. Caldwell was shown to have a measure of tolerance to R. padi
and/or BYDV while Abe was shown to be susceptible.
Mass Rearing the Bird Cherry Oat Aphid, Rhopalosiphum padi (L.). C. Kudagamage
and J.E. Foster, Department of Entomology and the U.S. Department of Agriculture,
Purdue University, West Lafayette, Indiana 47907. Breeding cereal crops for
resistance to Rhopalosiphum padi (L.) and or barley yellow dwarf virus (BYDV) disease
could provide a cheap means of control of the aphid and BYDV without adversely
affecting the environment. In a resistance breeding program methods should be available
for conveniently rearing the aphids.
Most studies on laboratory rearing have been directed towards finding the effect
of host plant and temperature on the reproduction and survival of R. padi. However,
in the literature, studies on the temperature effects on the biology of the aphid shows
considerable variation of results by different workers. Therefore we decided to in-
vestigate the effect of temperature and light on mass rearing of bird cherry oat aphid.
We investigated the effect of five temperature regimes 13, 18, 20, 28°C and two
photophase and scotophase periods 12:12, 14:10 h on prereproductive period (time
taken for the aphids to reach reproductive stage) and fecundity. The optimum
temperature and photophase: scotophase for rearing R. padi was determined to be
20°C and 14:10 h respectively. At this temperature and photophase the mean progeny
production was high and pre-reproductive period was short.
Assessment of Numbers of Striped Cucumber Beetle Adults and Frequency of Feeding
Injury on Muskmelon Cultivars. G.L. Reed and D.K. Reed, Fruit and Vegetable Insects
Research Laboratory, Agriculture Research Service, USDA, Vincennes, Indiana
47591. Field plantings of 74 muskmelon cultivars were evaluated to compare relative
differences in attraction and feeding injury by adult striped cucumber beetles. Seedling
and early vining stage of muskmelon plants were observed for numbers of beetles and
evidence of feeding injury to leaves and stems. Considerably more beetles were observed
on the cultivars Cobmelon, Tamdew and White-rinded honey dew, Charentais Improved
and Ogen. Lower frequencies of feeding injury were observed on the stems of cultivars
Milwaukee Market, Seneca Delicious, Campo, Early May, and Early Delicious and
on the leaves of the cultivar Seneca Delicious.
Entomology 305
Relationship of Probing Behavior of Sitobion avenae (Fabricius) to Transmission of
Luteoviruses Causing Cereal Yellow-dwarf Diseases. H.V. Scheller, R.H. Shukle,
E.S. Furgason and J.E. Foster, NATO scholar, Departments of Entomology, Elec-
trical Engineering and U.S. Dept. of Agriculture, Purdue University, West Lafayette,
Indiana 47907. The probing behavior of the English grain aphid, Sitobion avenae
(Fabricious), on oats (Avena sativa, var. Clintland 64) has been studied by means of
an electronic impedance monitoring system. This system records characteristic waveforms
due to changes in the impedance of the aphid/plant connection associated with behavioral
elements such as salivation, phloem contact, non-phloem ingestion, and phloem inges-
tion. Interpretation of recorded waveforms has been confirmed by determining the
position of aphid stylets through histological sectioning when characteristic waveforms
are produced.
Aphids carrying the PAV strain of barley yellow dwarf virus were given access
to noninfected oat plants for limited periods of time. Some aphids were hindered in
making phloem contact, other were manipulated to produce multiple probes. Plants
were subsequently tested for the presence of virus by means of enzyme-linked immunosor-
bent assay (ELISA). The association between elements of feeding behavior of 5. avenae
and the transmission of cereal yellow-dwarf virus will be discussed.
Identification of a Pectinase in Larvae of the Hessian Fly, Mayetiola destructor (Say).
R.H. Shukle, H.V. Scheller and J.E. Foster, Department of Entomology; NATO
scholar, and U.S. Dept. of Agriculture, Purdue University, West Lafayette, Indiana
47907. The Hessian fly, Mayetiola destructor (Say), is a major pest of wheat in
the United States, Europe and other parts of the world. We have shown that larvae
of this insect possess a pectinase (a polygalacturonase) enzyme that is presumedly in-
volved in the breakdown of cell wall and intercellular matrix material in the wheat
plant. Polygalacturonase activity can be demonstrated in extracts of the salivary glands
and midgut of larvae by an electrophoretic method using pectin-acrylamide gels. The
presence of this enzyme has been further confirmed by reducing sugar assays using
polygalacturonic acid (PGA) as the substrate. Optimum pH for hydrolysis of PGA
by this enzyme appears to be 7.5. Larvae of five biotypes of M. destructor have been
examined to date, and all appear to possess polygalacturonase activity.
The possible association of a pectinase enzyme with resistance in wheat to Hes-
sian fly infestation either through a hypersensitive response by the wheat plant, through
changes in the chemical composition of cell wall and intercellular carbohydrates, or
through the presence of enzyme inhibitors in the plant's tissues will be discussed.
Preference of the Bird Cherry Oat Aphid, Rhopalosiphum padi (L.) on Hessian Fly-
infested Wheat and Effects on its Biology. V. thirakhupt and J.E. Foster, Depart-
ment of Entomology and the U.S. Department of Agriculture, Purdue University, West
Lafayette, Indiana 47907. It has been observed frequently in the greenhouse that
the Hessian fly-infested wheat plants are also infested with the bird cherry oat aphids
much more often than the healthy or resistant plants. The studies were prompted and
experiments were planned for confirmation of this observation. The Hessian fly biotype
D and biotype D-susceptible wheat varieties — Blueboy, Knox 62, Monon and Seneca —
were used as hosts in comparisons with the non-infested plants of the same varieties.
Under controlled environmental chamber (20 ± 1°C and 14:10 hours photoperiod),
R. padi showed significant preferences, providing both free-choice and no-choice tests,
on the Hessian fly-infested plants of the three varieties to the non-infested ones, but
not on Knox 62. When the aphids were confined on both plants, there were indica-
tions that the infested plants provided better conditions to favor their performances.
306 Indiana Academy of Science Vol. 94 (1985)
The most striking effects were on the reproduction and longevity with the least on
time to maturity. However, the varietal differences existed and it should be noted that
R. padi nymphs died before reaching maturity and, thus, failed to establish on the
Hessian fly-infested Blueboy.
Anecdotal History of Entomology in Indiana
John J. Favinger
Indiana Department of Natural Resources
Indianapolis, Indiana 46204
The history of entomology has been covered in various degrees from time to time
in Proceedings of the Indiana Academy of Science (Everman, 1917; Davis, 1932, Mont-
gomery, 1955; Deay, Luginbill, Ulman, Wilson, Young, 1955). I also had available
an unpublished manuscript prepared by Dr. B. Elwood Montgomery in 1966.
The science of entomology has been an integral part of the Indiana Academy
of Science since its founding. At least six of the charter members can be considered
to have entomology as a primary or secondary discipline. Six presidents have had en-
tomology as their principal scientific interest but numerous other entomologists have
held responsible offices or committee assignments in the first one hundred years of
the Academy's existence. There was a paper on "Indiana Entomology," by P.S. Baker,
at the first meeting and the Academy has continued to be an important influence and
forum for Indiana Entomology since that time. Only four of the 100 meetings of the
Academy have been without entomological papers and a high percentage of these have
been published in full in the Proceedings. An informal session of entomologists began
in the middle 1930s and this was organized as an official division in 1946.
Early travelers commented on the abundance of bedbugs, fleas, mosquitoes and
gnats in various parts of Indiana as early as its beginnings as a state in 1816.
Thomas Say who was part of the "Boatload of Knowledge" that came down
the Ohio and up the Wabash to New Harmony where Robert Owen and William Maclure
intended to establish their Utopian experiment in communal living. Although the ex-
periment failed the group's influence made New Harmony a center for culture and
science for many years to come.
Thomas Say was expert in many facets of natural history. He studied all groups
of animals, described thousands of insects in many orders, and was the leading con-
chologist of this time.
Say died in 1834 and is buried at New Harmony. The student entomological society
at Purdue is named for Say and the Thomas Say Foundation administered by the En-
tomological Society of America continues to publish important monographs and treatises.
After Say's death there was little organized entomological work in Indiana for
50 years. Although there were occasional articles dealing with insect pests published
in farm papers and, beginning in 1851, the Reports of the Indiana State Board of
Agriculture. The first State Chemist, Harvey W. Wiley, who later became the father
of the Food and Drug laws, suggested in the 1879 Report that entomologists should
study the habits and methods of reproduction of many agricultural pests and to pro-
vide some way to arrest their almost marvelous fertility. Beginning in the 1855 report,
after F.M. Webster came to Indiana there were more frequent and well illustrated
articles on insect problems confronting the farmer.
The year 1884 was a banner year for Indiana entomology. Francis Marion Webster
was appointed a special agent of the Bureau of Entomology of the United States Depart-
ment of Agriculture and stationed as a consultant in entomology at its Purdue
Agricultural Experiment Station. With little formal education Webster became one
of the outstanding economic entomologists of the late 19th and early 20th centuries.
He wrote Bulletin -1 of the Purdue Agricultural Experiment Station on the hessian
fly and was an early advocate of cultural controls. Webster was a charter member
of both the Indiana Academy of Science and the old American Association of Economic
Entomologists serving as president of the latter organization in 1897. In 1906 Webster
307
308 Indiana Academy of Science Vol. 94 (1985)
became head of the Cereal and Forage Crop Insect section of the U.S.D.A. and for
seven years was headquartered at Purdue. He died in 1916 a few days after being
elected president of the Entomological Society of America.
Also in 1884, James Troop came to Purdue as Professor of Horticulture and
Entomology. A native of New York he earned a BS and MS at Michigan State College.
He taught at Michigan State before coming to Purdue where he taught the first formal
courses in entomology.
When the General Assembly created an Office of State Entomologist in 1899
primarily because of the rapid spread of San Jose scale by means of infested nursery
stock, Troop was given this additional responsibility by Governor James Mount and
served two terms of four years each.
When Horticulture and Entomology at Purdue became separate departments, Pro-
fessor Troop became head of Entomology, but remained very active in the Indiana
Horticultural Society, where he was affectionately known as the "Grand Old Man
of Hoosier Horticulture." He was designated Professor Emeritus in 1920 when J.J.
Davis became head of the department, but he continued to teach until 1929. Professor
Davis kept him on the staff until his death in 1941. Daddy Troop, as he was called
in my undergraduate days, came to the office occasionally driving an old tan Buick
in his own fashion.
David Starr Jordan was not an entomologist but was well versed in many areas
of zoology especially ichthyology. Jordan, a founder and first president of the Indiana
Academy of Science, along with John Caspar Branner fostered Willis Stanley Blatchley's
interest in nearly all sciences. The first course in entomology at Indiana University
was taught by Branner in 1886 with three students, Blatchley, Charles Boleman and
Jerome McNeil, each of whom became a recognized authority in some phase of
entomology.
W.S. Blatchley can be considered the first Hoosier entomologist. He was born
in Connecticut but came with his family to Indiana at the age of one. He graduated
from Indiana University in 1887 and was awarded a M A in 1891, teaching science
at the Terre Haute High School in the meanwhile. Blatchley began publishing while
still an undergraduate. In all he published more than 200 titles, 80 on entomology
and the rest on a wide variety of subjects, including geology, birds, reptiles, batrachians
and plants. He described 14 new genera and subgenera and 470 new species and varieties
primarily in the Coleoptera, Othoptera and Hteroptera. The Department of Geology
and Natural Resources in Indiana was for many years headed by a State Geologist
which had become an elective office. Blatchley, with considerable backing from Academy
members, was nominated by the Republicans in 1894, was elected that fall and reelected
three times.
After being defeated in the 1910 election, Blatchley spent the rest of his life writing,
collecting, etc. He distributed his books through his own publishing company, the
Nature Publishing Co., Indianapolis. Many of his nature books were autobiographical.
Probably his best known work was the "Coleoptera of Indiana" published as Bulletin
No. 1 of the Indiana Department of Geology and Natural Resources during his last
few months as State Geologist. The "Coleoptera" was distributed free to libraries
and many schools throughout the state. When copies became scarce and in great de-
mand 15-20 years later, Blatchley wrote letters to recipients of the volume offering
to buy copies in good condition for $5.00 each. I purchased a copy in almost mint
condition for $75.00 75 years after publication and considered it a bargain.
Blatchley was thrifty, even miserly, in many respects but from his own funds
he established a pension for his life-long secretary to be administered by J.J. Davis
but to be kept secret from his two sons because they would raise hell.
E.B. Williamson graduated from Ohio State in 1898 and served as assistant curator
Entomology 309
of insects at the Carnegie Museum in Pittsburg. He returned to Bluffton, Indiana and
eventually succeeded his father as president of the Wells County Bank. Meanwhile
on a part-time basis became a note authority on the Odonata. He also became interested
in hybridizing iris and gained world-wide recognition in that field. Longfield Iris Gardens
had Indiana Nursery Certificate #\ for many years.
Williamson published his first paper in 1898 and in 35 years published 123 scien-
tific papers mainly on dragonflies but also on birds, fishes, and other groups. He was
also an associate curator of Odonata at the University of Michigan Museum of Zoology
1916-1928 and research associate from 1928 until his death in 1933.
He made many expeditions to Central and South American and is credited by
C.C. Deam as getting him seriously interested in botany.
Benjamin Wallace Douglass was appointed State Entomologist by Governor J.
Frank Hanley when regulatory work in entomology was moved to Indianapolis in 1907.
Douglass was an expert photographer and skillful writer but must have gained his
entomological expertise by osmosis. He had attended medical school in Indianapolis
and had worked for C.C. Deam at the State Board of Forestry. The four annual reports
written during his tenure are filled with accounts of insect pests, plant diseases and
horticultural advice illustrated with excellent photographs.
The State Entomologist was appointed for a four year term and Douglass' extended
2 years into Governor Thomas R. Marshall's term. Douglass was not appointed and
went into the tree surgery business with one of his assistants, Frank N. Wallace. He
later operated an orchard in Brown County and continued to write for farm magazines
like, The Country Gentleman.
Douglass was succeeded by C.H. Baldwin who continued to publish excellent and
informative annual reports. Baldwin assembled an especially competent staff. Harry
Dietz and Harold Morrison wrote the "Coccidae or Scale Insects of Indiana" which
was illustrated by R.E. Snodgrass, also a staff member.
Dietz later worked for the Federal Horticultural Board but returned to Indiana
in 1920 to be Frank Wallace's chief assistant for 10 years. After attending graduate
school at Ohio State, he later became chief of pesticide research for Grasselli Chemicals
(DuPont).
Morrison, like Dietz, a native Hoosier, also later worked for the Federal
Horticultural Board and was insect curator at the U.S. Museum and a world expert
on scale insects.
Snodgrass was a meticulous illustrator as well as an accomplished caricaturist
and cartoonist. He became world famous for "Anatomy and Philosophy of the
Honeybee" and texts on arthropod morphology and physiology. He was associated
with the Bureau of Entomology and Plant Quarantine and the University of Maryland.
He maintained an association with the U.S. Museum after retirement in 1945 and was
mentally alert and physically fit for the next 20 years.
Frank N. Wallace was first hired for his accounting skills by Ben Douglass although
he had no more formal training in this field than he did in entomology. He was a
fast learner and very adaptable and had a unique way with people. Shortly after Ben
Douglass was replaced by C.H. Baldwin, Douglass and Wallace formed the State Forestry
Company in Indianapolis, a tree surgery and maintenance service. It was in this capacity
that Dean Stanley Coulter of Purdue, long time member of the State Forestry Com-
mission and later of the Conservation Commission recommended Wallace to Gene
Stratton Porter. Mrs. Porter, the then famous novelist and naturalist, needed someone
to supervise the rehabilitation of the trees at her new estate on Sylvan Lake. Wallace
married Lorene Miller, who was Mrs. Porter's secretary. When it came time for Governor
Samuel M. Ralston to appoint a successor to C.H. Baldwin, Mrs. Porter recommended
Wallace.
310 Indiana Academy of Science Vol. 94 (1985)
Wallace served as State Entomologist for 43 years (1915-1958), probably the longest
tenure for a chief plant regulatory official anywhere in the United States. The four
year term was omitted in new legislation creating the Indiana Department of Conser-
vation in 1919. Wallace is the only person without a college degree ever to have served
as president of the Indiana Academy of Science (1940). For many years he was the
principal lobbyist to obtain the legislative appropriation to publish the Proceedings
of the Academy.
Wallace was in great demand as a slide lecturer on the many aspects of nature
study and the Indiana State Parks. He served as one of the Central Plant Board's
representatives to the first meeting of the National Plant Board in 1925.
John June Davis probably had more influence on the history of entomology in
Indiana than any other person. He graduated from the University of Illinois in 1907
and first worked for S.A. Forbes, State Entomologist both in extension and research.
He was an authority on the taxonomy of aphids and later on the biology and control
of white grubs after becoming head of the Cereal and Forage Crops Insect Laboratory
at Lafayette. When Japanese beetle was discovered on the East Coast he was appointed
head of the laboratory at Riverton, New Jersey.
In 1920 Davis was appointed head of the Entomology Department at Purdue
and held this position until his retirement in 1956. At Purdue he taught, conducted
research and extension work, traveling widely in state and out. He was an inspira-
tional teacher and taught his favorite course, introductory entomology, for his entire
tenure.
He planned and began the development of a comprehensive insect collection and
personally was responsible for the acquisition of many fine collections, like that of
Blatchley and many others. Davis was an innovator in many other ways. He was respon-
sible for many new programs and special courses in entomology adapted to the needs
of forestry and pharmacy students. He arranged for the first meeting of North Central
Entomologists in 1921 which was attended by 13 entomologists from 4 states. When
this group again met at Purdue in 1930 there were nearly 100 in attendance.
Davis also initiated the first conference of Indiana entomologists which was held
at Purdue in the fall of 1923. He started the 4-H insect collection competition which,
from a meager beginning in 1925, has expanded into a major project over most of
the state. Glen Lehker, as Indiana's first full-time extension entomologist, came into
the program some 10 years later and further developed the program which has given
many individuals a start in the profession of entomology.
Perhaps the most lasting contribution of J.J. Davis was in making the field of
structural pest control a respectable profession. The famous Purdue Pest Control Con-
ference was initiated with 68 attendees in 1937 and now has some 600 participants
each year.
Nearly all colleges and universities in Indiana have had an entomologist on staff
either in a biology or zoology department even if no formal entomology courses were
taught.
Indiana University has had a number of distinguished teachers as well as students
in entomology. Frank Young discussed a number of these in his History of Biology
at Indiana University two years ago at Notre Dame.
Notre Dame has gained prominence in entomology in the last 27 years since George
Craig joined the faculty in 1957. The Vector Biology Laboratory and the Laboratory
for Arborvirus Research and Surveillance are world famous and have trained students
for responsible positions in this highly specialized field.
Ball State University, noted for mosquito and tick research, has had a number
of entomologists both in the Department of Physiology and Health Science and Depart-
ment of Biology. The late Russell E. Siverly was the author of "Mosquitoes of In-
Entomology 3 1 1
diana" which was published by the Indiana State Board of Health which now has
its own staff of entomologists dealing with public health aspects of the discipline.
There have been many changes in the field of entomology in the last one hundred
fifty years since the death of Thomas Say. Say was a taxonomist and interested primarily
in classification. Later the economic aspects became increasingly important and life
histories and control measures were studied. All of these are still important but there
are also highly specialized areas like molecular biology and DNA research that were
unknown just a few years ago.
Literature Cited
1. Daily, W.A. and F.K. Daily. 1984. History of the Indiana Academy of Science
1885-1984, Ind. Acad. Sci. Indianapolis, IN. 249 p.
2. David, J.J. 1932. Entomologists and Entomology in Indiana. Proc. Ind. Acad.
Sci. 41:43-70.
3. Deay, H.O. 1955. Entomology at Purdue, Proc. Ind. Acad. Sci. 64:152-157.
4. Everman, Barton Warren. 1917. A Century of Zoology in Indiana. Proc. Ind.
Acad. Sci. 26:189-224.
5. Luginbill, Phillip. 1955. Federal Entomology in Indiana. Proc. Ind. Acad. Sci.
64-161-164.
6. Mallis, Arnold. 1971. American Entomologists. Rutgers U. Press, New Brunswick,
N.J. 549 p.
7. Montgomery, B. Elwood. 1955. Entomology Before 1854, Proc. Ind. Acad. Sci.
64:142-147.
8. 1966. One Hundred Fifty Years of Entomology in Indiana, Unpublished
manuscript, 27 p.
9. Ulman, Paul T. 1955. Regulatory Entomology in Indiana, Proc. Ind. Acad. Sci.
64:158-160.
10. Wilson, M. Curtis. 1955. Entomological Pioneers in Indiana, Proc. Ind. Acad.
Sci. 64:148-151.
11. Young, F.N. 1955. Work at Other Institutions and by Private Individuals Since
1854. Proc. Ind. Acad. Sci. 64:165-172.
Indiana Gypsy Moth Survey — A History
Philip T. Marshall
Indiana Department of Natural Resources
Vallonia, Indiana 47281
and
James A. Clark
Indiana Department of Natural Resources
Indianapolis, Indiana 46204
Introduction
Since gypsy moth's, Lymantria dispar L., (Lepidoptera, Lymantriidae), escape
from a botanist in Medford Massachusetts in 1869, this forest defolitor has gradually
spread west. Currently, the defoliation to timberlands of the United States occurs north-
ward from western Pennsylvania, northeastern West Virginia, northern Virginia,
Maryland, and Delaware through the New England states to Canada (6). While the
gypsy moth caterpillars were eating their way through 14 states and over 52 million
acres of forest land, man has unknowingly aided gypsy moth in their spread to other
states (2). Currently, man and his vehicles have introduced gypsy moth to all states
east of the Mississippi River and to several states west of the Mississippi including
all west coast states. Realizing that gypsy moth would be introduced to noninfested
states as man moved and travelled, the United States Department of Agriculture, Animal
and Plant Health Inspection Service (APHIS) began cooperative surveys to detect gypsy
moth in these states. The cooperative gypsy moth survey in Indiana began in 1972,
and this paper presents a history of the survey from 1972 through 1984.
Methods and Materials
The gypsy moth survey uses the gypsy moth pheromone trap. This trap is a delta
trap (tent-like) approximately 9.4" x 4.0" x 4.0". The trap is made of plastic coated
paperboard with two internal sides covered with Tack-Trap and the third side used
to attach the pheromone bait. The traps are orange, tan or green in color. The trap
will hold 15-20 male moths (3).
The gypsy moth pheromone is called disparlure. Between 1972 and 1980, the racemic
form of the pheromone was used. Starting in 1981, the improved 'plus' form of the
pheromone has been used. The racemic form came in a green plastic dispenser approx-
imately 1 " x 1 ". The 'plus' form is dispensed from a tan plastic dispenser 1 " x 1/8".
The pheromone is released at a constant rate over the trapping period (3).
The gypsy moth survey begins with a detection survey, and then, if a male moth
is trapped, a delimitation survey is conducted the following year. If more than one
moth is caught in one trap or when several traps in one localized area have one or
more moths, an egg mass survey is conducted in the fall of the survey year. One addi-
tional part of the gypsy moth survey is mass trapping. Mass trapping is used to follow-
up aerial spray programs and in areas where patterns of male moth catches indicate
an infestation has started but no other life stage has been found.
In the detection survey, traps are placed according to two grid systems — one trap
per three square miles (one trap every 1.7 mile) and one trap per 25 square mile (one
trap every 5 mile, 5 mile grid). The USDA, APHIS uses the one trap per three square
mile grid, and the Division of Entomology uses the one trap per 25 square mile grid.
The grid system is rotated in each county each year to prevent surveying the same
area each year and to achieve a complete survey of all land area after three years.
313
314 Indiana Academy of Science Vol. 94 (1985)
In some years, APHIS has intensified the detection survey to one trap per one square
mile in some areas of the counties that they survey.
In addition to the grid system of the detection survey, traps are placed in special
sites such as campgrounds, interstate rest areas, motels, truck stops, national campers
association meetings, nature preserves, classified forests, federal installations, univer-
sities, and homes of people newly moved into Indiana from the northeast.
The delimitation survey is conducted at a greater density of traps per square mile.
Generally, 25 traps per square mile is used; however 32 or 81 traps per square mile
may be used. The nine square mile area around the trap that caught a gypsy moth
is trapped at the above density. An additional 16 square mile area surrounding the
nine square mile area may be trapped at nine traps per square mile (5). When several
gypsy moths are detected in close proximity to each other, the delimitation grid pat-
terns will be modified and combined to efficiently delimit all catches of the gypsy
moth. When only one moth is caught in a county, the nine square mile area may
be reduced to a four square mile area at 25 traps per square mile with the detected
moth at the center of the four square mile area.
The egg mass survey is a general survey of all the area around the point where
a gypsy moth was trapped. Personnel of the USDA, APHIS and the Division of
Entomology search the environment for egg masses. They also contact people in the
areas trying to locate anyone who may have moved there from the generally infested
area of the northeastern United States. If egg masses are found, they are destroyed,
and the area is defined as an infestation and will be placed in a control program the
following year.
Mass trapping is conducted on a grid system of three traps per acre (1920 traps
per square mile) or one trap every 120 feet. It is confined to small areas because of
the quantity of traps needed.
In the cooperative survey, the USDA, APHIS selected certain counties each year
to survey, and the Division of Entomology surveyed all remaining counties. From 1972
through 1980, APHIS would survey 1/3 of the counties in the state. The selection
of the counties was rotated each year so that after three years, APHIS had surveyed
all counties of the state once, and the Division of Entomology has surveyed all counties
once. Since 1981, APHIS has surveyed the counties where gypsy moth has been trap-
ped the previous year, and the Division of Entomology has surveyed all remaining coun-
ties. In 1984, the survey changed to target the placement of the traps in areas where
gypsy moth had a high probability of being introduced. In 1984, APHIS surveyed
all counties where gypsy moth was detected in 1983 and the counties with major
metropolitan areas. The Division of Entomology surveyed all remaining counties targeting
the traps into the cities.
Personnel involved in the gypsy moth survey are given maps showing the grid
system indicating where traps are to be placed. The traps are placed as close as possi-
ble to the grid point on the map. The traps are placed on the siies of trees, posts,
or poles. The location of each trap is recorded on a trap record form by trap number,
county, township, range, section number, city or other name for the trap location
such as the name of the campground. Directions to the trap are recorded, and a sketch
map is drawn on the trap record form to help locate the trap.
The traps are placed across the state during June with all traps to be in place
by the first of July. Traps in the detection survey, generally, are not checked during
the survey. However, traps in delimitation surveys are periodically checked, and traps
in a mass trapping survey are checked regularly. All traps are removed during August,
and the number of gypsy moths and their location are reported to the USDA, APHIS
and the Division of Entomology. All moths found for the first time in a county are
Entomology 3 1 5
submitted to the USDA to be confirmed for official record of first find. The locations
of moths are plotted on maps to observe the distribution of the gypsy moth. These
maps aid in identifying the start of infestations and in the planning for the following
year's survey.
Results of the survey are summarized annually and reported to the USDA, APHIS
and Forest Service, the Indiana Department of Natural Resources, Divisions of
Entomology and Forestry, and the National Gypsy Moth Management Board. The
report is also published in the Indiana Pest Informer, a newsletter on forest insect and
diseases.
Results
Since 1972, the gypsy moth survey has placed 72,168 traps in the state (Table
1). Personnel of the USDA, APHIS have placed 52,211 traps, and personnel of the
Division of Entomology have placed 19,166 traps. An additional 791 traps have been
placed by members of the National Campers and Hikers Association.
Table 1. The number of gypsy moth traps set in Indiana by year and cooperators.
Cooperators
Year
Federal
State
Other'
Total
1972
883
1640
2523
1973
1622
2
51
1673
1974
2031
849
94
2974
1975
1602
1193
2795
1976
1413
1919
36
3368
1977
3991
1355
5346
1978
3465
1233
4698
1979
4902
1257
6159
1980
5371
1227
200
6798
1981
4678
1819
300
6797
1982
3827
2313
6140
1983
9063
1209
10272
1984
9363
3152
110
12625
Total
52211
19166
791
72168
'Primarily set by the National Campers and Hikers Association.
'Records unavailable.
The number of traps placed in the state started to increase in the late 70s. This
increase was in response to the increased introduction of gypsy moth in Indiana from
the increasing population of gypsy moth in the northeastern United States during this
time (Table 1) (6). During 1979-1982, the number of traps placed in the state was
at a constant level over 6,000. Then in 1983 and 1984, the number of traps placed
almost doubled. This increase in traps placed was due to the use of mass trapping
in areas where a gypsy moth infestation had been found, to increased intensity of
the grid system in some counties of the state, and to the increased use of delimitation
trapping around the increased number of gypsy moth catches of 1982 and 1983 (Table 2).
During the thirteen years of the survey, traps have been placed in every county
of the state, except for five years. In 1972, 1973, 1978, 1981, and 1984, traps were
not placed in 1, 3, 4, 2, and 1 counties, respectively.
1973 was the first year gypsy moth was found in Indiana. One male moth was
found in Lake County (Table 2). Surveys in 1974 and 1975 did not catch gypsy moths;
thus, the first find of gypsy moth was a 'hitchhiker.'
316 Indiana Academy of Science Vol. 94 (1985)
Table 2. The number of gypsy moth males trapped in Indiana by year and county.
Year County Number of moths
1973 Lake 1
1977 Whitley 1
1980 1. Allen 1
2
1
1
4
1
10
iv;;i i Allrn 2
1
1
20
1
1
2
32
4
1.
Allen
2.
Elkhart
3.
Franklin
4.
Hendricks
5.
Vigo
6.
Wayne
1.
Allen
2.
Bartholomew*
3.
Boone*
4.
Elkhart
5.
Lake
6.
LaPorte*
7.
Tippecanoe*
8.
Vigo
9.
Wayne
!.
Allen
2.
Bartholomew
3.
Blackford*
4.
Brown*
5.
Elkhart
6.
Fulton*
7.
Hancock*
8.
Hendricks
9.
Jefferson*
10.
Johnson*
11.
Kosciusko*
12.
LaPorte
13.
Marion*
14.
Monroe*
15.
Montgomery*
16.
Morgan*
17.
Noble*
18.
Putnam*
19.
St. Joseph*
20.
Tippecanoe
21.
Wayne
2.
Bartholomew
3.
Elkhart
4.
Greene*
5.
Hamilton*
6.
Hendricks
7.
LaGrange*
8.
Lake
9.
LaPorte
10.
Marion
11.
Monroe
12.
St. Joseph
64
!'.-•.; Mlrn 5
14
1
1
372
1
2
2
1
20
1
4
11
1
1
2
1
1
21
8
3
473
1983 1. Allen 1
7
29
1
2
1
1
1
1
35
1
n
91
Entomology
317
Table 2. — Continued
Year
County
Number of moths
1984
1 . Allen
2. Decatur*
3. DeKalb*
4. Elkhart
5. Fulton
6. Hamilton
7. Jackson*
8. Johnson
9. Kosciusko
10. LaGrange
11. Lake
12. Marion
13. Marshall*
14. Monroe
15. Orange*
16. St. Joseph
17. Wabash*
18. Wayne
19. Whitley
11
1
I
13
1
1
1
4
4
3
8
14
2
1
1
23
1
1
1
92
*New county record for that year.
The second gypsy moth was caught in 1977 in Whitley county. Again, this moth
was a 'hitchhiker' and no infestation developed.
1980 was the first year when more than one moth was found and more than
one county had gypsy moth (Table 2). Since 1980 when 10 moths from 6 counties
were found, gypsy moth has been found in Indiana every year. In 1981, 64 moths
were found in 9 counties with 4 of the counties being new county records. In 1982,
473 moths were found in 21 counties with 14 of the counties being new county records.
In 1983, 91 moths were trapped in 12 counties with 3 counties being new county records.
And in 1984, 92 moths were found in 19 counties with 7 new county records (Table
2, Figure 1).
The survey has trapped 732 male moths from 35 different counties since 1972
(Table 3, Figure 2). Most of these gypsy moth catches have been one moth in one
trap in one location. These single catches are 'hitchhikers' that did not develop into
infestations. And, yearly survey records indicate 13 counties are more likely to have
gypsy moth introduced and trapped from them. These counties are Allen, Elkhart,
Hamilton, Hendricks, LaGrange, Lake, LaPorte, Marion, Monroe, St. Joseph, Tip-
pecanoe, Vigo, and Wayne (Table 2 & 3). The major metropolitan areas of the state
occur in or next to these counties, and this is one reason why these counties are prone
to gypsy moth introduction.
The survey has detected and located six infestations. This number may increase
after the 1984 multiple-catches of gypsy moth are delimited to determine if an infesta-
tion has started (Table 3).
The first infestation was found in Vigo county in 1981. The 1980 survey found
4 moths in a subdivision called Krislynn Woods near Tecumseh. In 1981, the survey
trapped 31 moths in this areas. Egg mass surveys in 1981 found 63 egg masses around
one home in the subdivision. Residents of this home had moved there from an infested
area of New Jersey.
Also in 1981 a second infestation was found in Elkhart county in the city of
318
Indiana Academy of Science
Vol. 94 (1985)
PULASKI
PULTON
ELKHART
KOSCIUSKO
LAG*A**Gt« snu*i*
Moeu
OCKALB
WW(TL£V J
CASS
CA«90u.
;P£C*NOt
WABASH
TON Lw
'IPTQN
WEi-i-S
aDamS
a
DELAWARE
RasCXX-P^
Figure 1. Locations where gypsy moth was trapped in 1984.
Goshen. Surveys in 1980 found 2 moths in Elkhart county. In 1981, 20 moths were
found with 18 moths being found in Goshen. Surveys in 1982 found 372 moths and
80 egg masses.
Entomology
319
Table 3. List of counties where gypsy moth males have been trapped including total
moths trapped, year first trapped, number of consecutive years trapped and number of
infestations.
County
Total moths
First year
Consecutive
Years
Number
Infestations
1.
Allen
20
1980
2.
Bartholomew
19
1981
3.
Blackford
1
1982
4.
Boone
1
1981
5.
Brown
1
1982
6.
Decatur
1
1984
7.
DeKalb
1
1984
8.
Elkhart
436
1980
9.
Franklin
1
1980
10.
Fulton
2
1982
11.
Greene
1
1983
12.
Hamilton
3
1983
13.
Hancock
2
1982
14.
Hendricks
4
1980
15.
Jackson
1
1984
16.
Jefferson
1
1982
17.
Johnson
22
1982
18.
Kosciusko
5
1982
19.
LaGrange
4
1983
20.
Lake
11
1973
21.
LaPorte
6
1981
22.
Marion
60
1982
23.
Marshall
2
1984
24.
Monroe
3
1982
25.
Montgomery
1
1982
26.
Morgan
2
1982
27.
Noble
1
1982
28.
Orange
1
1984
29.
Putnam
1
1982
30.
St. Joseph
55
1982
31.
Tippecanoe
10
1981
32.
Vigo
36
1980
33.
Wabash
1
1984
34.
Wayne
9
1980
35.
Whitley
2
1977
732
Table 4: Explanation of symbols for figure 1
One male moth in one trap in one location.
More than one trap containing one male moth in one location.
Multiple male moths in one trap in one location.
One or more of the following in one location — one male moth per trap and multiple male moths
per trap.
320
Indiana Academy of Science
Vol. 94 (1985)
Figure 2. Counties where gypsy moth has been trapped since the survey began in 1972.
The infestation in Bartholomew county was in the city of Columbus. One moth
was caught in 1981. In 1982, 14 moths were caught in the same area. Egg mass surveys
in 1982 found 5 old egg masses on a boat trailer belonging to a family who had recently
Entomology 321
moved to Columbus from Connecticut. No viable egg masses were found, and with
mass trapping this infestation has died-out (Table 2).
The infestation in Johnson county (Table 3) was found in Camp Atterbury at
a national meeting of the Campers and Hikers Association. This infestation was mostly
'hitchhiking' moths, and the infestation died-out from mass trapping and ground sprays.
The two infestations in Marion county were classified infestations based on the
pattern of trapped moths from one year to the next. In 1982, 11 moths were found
in the two areas. This increased to 35 in 1983. In both areas egg mass surveys were
negative. Mass trapping has been used in each area, and only one of the two areas
had gypsy moth trapped from it in 1984.
Gypsy moth has been trapped from four state parks — Brown County, Chain-O-
Lake, Clifty Falls, and Shakamak, one state recreation area — Paynetown (Monroe Reser-
voir), several private and county campgrounds, on or near the campuses of Notre Dame,
Purdue, and Indiana Universities, rest areas on interstates, and classified forests. All
locations where multiple catches have been made can be linked to someone moving
and carrying gypsy moth on their cars or RVs and their personal property into Indiana.
Nurserystock from an infested northeastern nursery has also carried gypsy moth into
Indiana.
Discussion
The gypsy moth survey has found that Indiana can easily have this defoliator
introduced into the state and its forests. The survey has also found that gypsy moth
is more likely to be introduced in cities and large metropolitan areas where movement
of man is more likely to occur. Thus, the recent change in the survey to target traps
into these areas. The survey has also found that man's vehicles and other property
are the primary means of carrying gypsy moth into Indiana.
Although cities and metropolitan areas may have a greater chance of introducing
gypsy moth into Indiana, the rural areas of the state must not be forgotten. This is
especially important for south central Indiana where the major forest areas of the
state occurs (4). In this area, gypsy moth has been found in Brown County State Park,
Paynetown Recreation Area, and the city of Bloomington. Should gypsy moth infesta-
tions start in this area, a major natural resource of Indiana is threatened.
The patterns of gypsy moth catches within a year and between years indicate
that the current survey has done a good job in detecting the introduction of gypsy
moth to Indiana. The survey has located many single catches of gypsy moth and subse-
quently shown that these single catches were not the start of an infestation. The detec-
tion survey and following delimitation survey have located six infestations with four
of the six infestations eradicated and two under a control program. This early and
efficient detection of gypsy moth will provide many years before a gypsy moth infesta-
tion becomes an established population that could spread from within the state.
As found in research on Dutch Elm Disease, an introduced pest to the United
States like gypsy moth, efficient and intensive surveys to detect Dutch Elm Disease
resulted in a greater length of service before elms were infected and killed and in an
overall reduction in the cost of controlling the disease (1). This same intensive survey
effort for gypsy moth in the noninfested states can provide similar benefits by lengthening
the time to establish populations, by reducing costs of control and by defining in-
troductions of gypsy moth in such a manner as to allow better match of control methods
to the particular situation. Therefore, to protect the valuable forest resource and the
wooded urban environments of Indiana, the gypsy moth survey should continue at
the same or greater intensive level.
322 Indiana Academy of Science Vol. 94 (1985)
Literature Cited
1. Cannon, W.N., Jr. and D.P. Worley. 1976. Dutch elm disease control: perfor-
mance and costs. USDA, For. Ser. Res. Pap. NE-345, 7 pp.
2. Personal communication. U.S. Forest Service, State and Private Forestry, Forest
Pest Management, Morgantown, W.V., Oct. 1984.
3. Schwable, C.P. 1979. Using pheromone traps to detect and evaluate populations
of the gypsy moth: gypsy moth handbook. USDA, Agric. Handbook No. 544,
11 pp.
4. Spencer, John S., Jr. 1969. Indiana's timber. USDA, For. Ser. Res. Bui. NC-7,
61 pp.
5. USDA. 1980. Gypsy moth and browntail moth program manual. Animal and
Plant Health Inspection Service, 34 pp.
6. USDA. 1984. Gypsy moth suppression and eradication projects: final environmental
impact statement. USDA: Forest Service and APHIS, Washington, D.C.
Insects and Other Arthropods of Economic Importance in Indiana in 1984
Robert W. Meyer
Department of Entomology
Purdue University, West Lafayette, Indiana 47907
Introduction
The winter of 1983-1984 was harsh. In addition to depressed temperatures — a
minus 29 degrees F. was recorded on 24 December in Hobart and the same temperature
was recorded in English on 21 January — there was often little snow cover. This com-
bination probably reduced alfalfa weevil and Mexican bean beetle populations, the
latter already drastically reduced by high temperatures in the summer of 1983.
The spring was cool and wet. Planting began in the northern third of the state
the first week in May but was delayed until the third week in the southern half. Early
in June drought conditions prevailed over much of the state, lasting for much of the
summer; such rains as occurred were usually light and localized. Fortunately the sum-
mer temperatures were moderate, preventing a recurrence of the damage the crops,
especially corn, suffered in 1983.
Other factors affected this year's crops. According to Indiana Weekly Weather
and Crops (which provided most of the weather information above) 41% of the corn
ground and 46% of the soybean ground was prepared by plowing, 45°7o and 46%
were conservation tillage, and 14°7o of the corn and 8% of the soybeans were planted
no-till. An estimated 6,000,000 acres of corn and 4,200,000 acres of soybeans were
planted.
Corn and Small Grains
The western corn rootworm (Diabrotica virgifera) is generally Indiana's most costly
agricultural pest; an estimated 2,400,000 acres were treated in 1984 at a cost of
$24,000,000. As usual, not all of the treating was necessary and some untreated areas
should have been. In 1983 the government, in an effort to reduce corn surplusses,
offered growers grain if they reduced their corn acreages, the so-called payment-in-
kind program. Forty percent of the acres normally planted to corn were taken out
of production, incidentally reducing the acres producing corn rootworms. The average
number of beetles/stalk, counted late July and early August in visits to 225 fields,
was 0.97 in 1983; in 1984 the figure was 0.64, with district averages ranging from
0.43 go 0.87. Silk clipping rarely reached economic levels, if ever.
The first first-instar in a Tippecanoe Co. field regularly surveyed for this insect
was collected on 8 June, not unusually late, and the first adult reported in the state
was collected on 3 July in Parke Co., the normal date for its appearance.
Counts of the northern corn rootworm (D. barberi) averaged 0.07/stalk over the
state, as determined by the survey described above.
The fall, 1983, corn survey put European corn borer {Ostrinia nubilalis) larvae
at 84/100 stalks, the state average. Adults this year observed as early as 30 May in
Knox Co., but were probably present earlier as second instar larvae were collected
by 13 June in Jackson Co. The peak flight of the first generation moths to blacklight
traps occurred before the middle of June, when corn averaged less than 15 inches.
The second flight peaked the first 2 weeks of August, by which time most of the corn
had silked. Flights were not large, at least by 1983 standards when daily catches in
some traps exceeded 500 whereas this year's catches generally did not reach 300/week.
The fall survey this year of 300 fields in 60 counties found the average number of
live larvae to be 99/100 stalks, unevenly distributed. Most of the larger populations
323
324 Indiana Academy of Science Vol. 94 (1985)
were in the northern districts, which is normal, with 6 counties averaging more than
2 larvae/stalk. Only 1 county elsewhere — Jackson — averaged more than 2/stalk.
The disease crazy top was more common this year than in other years.
Minor pests in corn in 1984 were the following.
Corn leaf aphids (Rhopalosiphum maidis) did not build up to the high numbers
expected with moisture stresses; they were present as usual but at non-economic numbers.
Billbugs (Undetermined) required treatment in a muck field of 90 acres in LaPorte
Co.
Japanese beetles (Popillia japonica) were more common this year than last, and
when that happens there are usually a few fields of corn that require treatment to
prevent silk clipping. Most of this type of damage occurred in the NW and NC districts.
Black cutwork (Agrotis ipsilori) was infrequently reported from corn.
Grasshoppers (Several species) and yellow woollybears (Diacrisia virginica) were
both more common than usual, and sometimes did conspicuous damage to corn at
field edges either alone or in combination. The latter was occasionally responsible for
serious silk clipping.
A survey of 385 certified seed fields in 62 counties in the spring of 1984 (con-
ducted cooperatively by the Indiana Crop Improvement Association, the Agricultural
Research Service of the USDA, and Purdue's Entomology Department) yielded the
following data on the Hessian fly (Mayetiola destructor). The mean percent infested
of all wheats surveyed — including wheats with no resistance to the fly — was 1 .4; mean
puparia/100 stems for the same set: 2. Both of these figures were increases over last
year. The most commonly planted wheat with H6 resistance — Caldwell — was infested
at the rate of only 0.6%; all cultivars with H6 resistance together averaged only 0.4%
infested. Those with no sources of resistance averaged 7.2% infested.
The English grain aphid (Sitobion avenae) was common on small grains this year
in the southern half of the state, far outnumbering the bird-cherry oat aphid
{Rhopalosiphum padi). Neither was considered economic.
Cereal leaf beetle {Oulema melanopus) normally occurs in numbers only in Harrison
Co. This year adults were first swept from alfalfa on 25 April, an egg was seen on
barley on 1 May and early instars on 22 May, in trace numbers only, and only in
Harrison Co.
Forage Legumes and Soybeans
Aside from occasional, and usually field-edge, feeding by grasshoppers and/or
yellow woollybears, soybeans were relatively free from insect attack this year. Mexican
bean beetle {Epilachna varivestis) adults are usually swept early from alfalfa. This year
none was. Adults were rare, and immatures were seen only in a few fields in Jennings
Co. in soybeans. Green cloverworms (Plathypena scabra) were often present, but only
in trace numbers. Japanese beetles were sometimes numerous enough to do conspicuous
but non-economic feeding not confined to the northern districts as silk feeding is. A
soybean leafminer (Odontota horni) was present in trace numbers in soybeans in the
NW district. Bean leaf beetles (Cerotoma trifurcata) were swept from alfalfa about
mid-May at the rate of 40-60/100 sweeps in the WC district, and they were occasionally
numerous in soybeans later. They were seldom at economic numbers in soybeans.
The alfalfa weevil (Hypera postica) was rarely a problem in alfalfa even in the
southern third of the state. This was due both to good growth, enabling the plants
to tolerate more feeding, and low numbers of larvae. At early bud stage (about 15
May) when alfalfa averaged 70 or more centimeters, larvae averaged fewer than 1.5/stem
in the SW, 0.5/stem (as a result of disease) in the SC district. The cold, open winter
may have reduced adult numbers, and it certainly destroyed all the fall-laid eggs.
Entomology 325
Potato leafhopper (Empoasca fabae), usually the most serious pest of alfalfa in
Indiana, was rarely a problem during 1984. One estimate places treated acreage at
about 10% in the northern alfalfa-growing belt, much less in the southern. Only the
third cutting was affected.
Eggs of the variegated cutworm {Peridroma saucia) were first observed on white
plastic flags in alfalfa in Harrison Co. on 1 May. Since they hatched the next day
they must have been deposited several days before. They were not there 7 days earlier.
They have been collected as early as 7 April in the same field. The species occasionally
builds up in alfalfa and is often a garden pest.
Vegetable Insects
Garden insects were generally at lower-than-usual levels in 1984. Exceptions follow.
The European corn borer was a serious pest in sweet corn, in beans grown for process-
ing and in green peppers. There was an instance of this species also in onion tops,
which is rather unusual. The corn earworm (Heliothis zed) was a serious pest in late
sweet corn as well as in tomatoes. A pheromone trap in Tippecanoe Co. at its peak
caught 227 adults in 1 night.
Apparently it was a good year for the squash bug (Anasa tristis) and problems
with squash vine borer (Melittia satyriniformis) have remained fairly constant.
Fruit Insects
All of the data on fruit tree insects, unless otherwise noted, are based on catches
in 5 pheromone traps in Knox Co. operated by Thomas Mouzin of the USDA. The
year's total catch is used to compare 1984 with previous years — not the best system
but the best available.
Codling moth (Cydia pomonella) catches of 245 were half those of the 8-year
mean (1976-1983) of 513, with weak peaks at the end of May and the end of August.
The 1984 total of 971 male redbanded leafroller (Argyrotaenia velutinana) was
less than the 8-year average of 1429, with peaks in mid-April, mid- to end of June
and mid-August.
The 1984 catch of 307 obliquebanded leafroller {Choristoneura rosaceana) was
near the 7-year average of 316, with a peak at mid-June and a lesser at the end of
September.
The catch of 220 leafminers (Phyllonorycter sp.) is double the 4-year average
of 100. Knox Co. totals do not however reflect conditions occurring in the rest of
the state. Economic or near economic infestations were seen in the NE counties of
LaGrange and Adams, the EC counties of Wayne and Delaware, and the C district
county of Madison. Adults in those counties were in flight the latter half of July.
Oriental fruit moth (Grapholitha molesta) catches of 1372 were somewhat smaller
than the 8-year mean of 1748. Larger numbers flew at the end of May, most of July
and at the end of August and the beginning of September.
This is only the fourth year that pheromones have been used in Knox Co. (rather
than live females) to attract males of the lesser peachtree borers (Synanthedon pictipes).
The 3-year average of 1849 was exceeded by this year's 2045. Peaks occurred in mid-
June, mid-July and late August-early September.
The peachtree borer (Synanthedon exitiosa) catch of 289 exceeds the 5-year average
of 191; there was a mid-July peak.
Trapping of San Jose scale (Quadraspidiotus perniciosus) was begun in 1982 when
143 were collected. Half that many came in 1983. This year 1000-plus (too many to
be counted accurately) came from 9-15 July, and 1066 were collected from 8-14 Oct.
This year's total came to 3079.
326 Indiana Academy of Science Vol. 94 (1985)
Insects of Ornamental Trees and Shrubs
The ten insects most frequently seen by nursery inspectors during 1984 — using
data supplied by the office of the State Entomologist — are listed here. 1. Fall web-
worm, (Hyphantria cunea); 2. Japanese beetle; 3. Bronze birch borer, (Agrilus anx-
ius); 4. Honeysuckle aphid, (Hydaphis tartaricae); 5. Fletcher scale; (Lecanium flet-
cheri); 6. Yellownecked caterpillar, (Datana ministra); 7. Maple bladdergall mite, (Vasates
quadhpes)', 8. Oystershell scale, (Lepidosaphes ulmi); 9. Mimosa webworm, {Homadaula
anisocentra), and 10. Euonymus scale, (Unaspis euonymi).
In general, the State Entomologist noted that reports of both aphids and borers
were nearly double those of 1983. Apparently, the cool wet spring brought forth enough
new growth for the aphids to flourish. He attributed the increase of borers to the
stresses on trees caused by the drought of 1983 and the harsh winters that bracketed it.
Man and Animals
The following generalizations were provided by Medical Entomologist Michael
Sinsko, Indiana State Board of Health, and they reflect the situation as of 31 October.
Mosquito activity at best was spotty, with total activity down again due to a paucity
of breeding sites. There were no reported cases of St. Louis encephalitis, 1 1 cases of
LaCross encephalitis (about average) and no cases of eastern equine encephalitis. There
were 5 confirmed cases of Rocky Mountain spotted fever.
More difficult to categorize are several pest arthropods. Fleas, especially cat fleas
(Ctenocephalides felis) had a good year. Head lice (Pediculus humanus capitis) were
again common and scabies mites (Sarcoptes scabiei var. hominis) infestations seem
to have reached a plateau. House fly activity is particularly difficult to estimate. Local
conditions — the presence of a poultry operation for instance — may be a major nuisance
in a year otherwise not noted for flies. Best estimates classify 1984 as an average year
for house flies.
Judging from the number of complaints about them it must have been a good
year for yellow jackets.
Cheryl Towell provided the following. Over the fly season this year, face flies
(Musca autumnalis) averaged 7/ face, about half the average during the last 2 years.
Horn flies (Haematobia irritans) were about average at 28/side.
Beneficial Insects
Adult alfalfa weevils were difficult to obtain in large enough numbers to estimate
accurately the amount of parasitization by Microctonus aethiopoides, our most com-
mon adult parasite. More than 2300 larvae were reared, however, to estimate the activity
of Bathyplectes anurus and B. curculionis. On a district basis, the NC averaged 7%,
the NE, 14 and the WC 8%, almost entirely by B. curculionis. B. anurus is known
to be present all over the state; it is not known why it isn't more common at least
in the WC district, which is surveyed frequently enough. The SW average totalled
44%, 19% due to B. anurus, 25% to curculionis. The SC average was 61%, 56%
due to anurus and only 5% due to curculionis. The 28% due to anurus and 18%
due to curculionis adds up to 46% parasitized in the SE district. Sampling was done
on a weekly basis when larvae were present in sufficient numbers to be readily swept,
especially in the WC, the SW and the SC districts. B. anurus is the more common
under normal circumstances in the earliest samples, curculionis in the later. The SC
is exceptional, and is probably due to the presence of disease. The fungus Erynia sp.
especially in Harrison Co. during the last 4 or 5 years has decimated weevil larval
populations, especially the last to hatch. In summary, the state average including both
Entomology 327
species and all surveyed districts (the NW, C and EC districts were not surveyed) was
43.5%.
The red coccinellids collected on 10 sticky traps in a Tippecanoe Co. corn field
are counted each year as a population estimate. The most commonly collected is Col-
eomegi/la maculata; this year 538 were collected, the most ever collected, and that
does not cover the hibernation flight which sometimes occurs (the corn was harvested
too early to permit that count). The ratio of Coleomegilla maculata: Hippodamia con-
vergens.H. tredecimpunctata:Cycloneda sanguinea this year was 84:9:0:7. The same
ratio among the coccinellids observed during the fall corn insect survey was 96:4:0:0,
based on seeing only 156 C. maculata on the 7500 stalks surveyed.
Annual Changes in Flea Populations on Three Domestic Pets, 1978-1984
Jack R. Munsee
Department of Life Sciences
Indiana State University
Terre Haute, Indiana 47809
Introduction
There are about 1900 species and subspecies of fleas worldwide (4). In Indiana,
Whitaker (6) has listed 31 species from wild mammals; in addition, two other species
have been collected, one from barn swallows and another from man and domestic
animals. Few among the many species of fleas are of direct concern to humans. Those
that bother humans in modern societies are species that live on or in association with
domestic animals or pets. Of direct concern to owners of the latter are the fleas that
infest dogs and cats, especially if these pets share the living quarters of their owners.
Because fleas may not always have contact with their normal hosts, in heavy infesta-
tions humans may serve as temporary sources for blood upon which adult fleas feed
exclusively. Fleas on pets that live outdoors are not as likely to become a serious threat
to either pets or owners.
This report is an outgrowth of efforts to control fleas on two neutered pet dogs
and subsequently those infesting a neutered male cat. It was thought that if fleas were
removed from the pets regularly in order to prevent large population build-ups, insec-
ticides would not be needed. The purpose of control was to prevent development of
large populations of fleas, rather than attempt to eliminate all fleas.
Methods and Materials
Although the practice of regular weekly grooming and collecting fleas began earlier,
in 1978 recording of data started and continued through part of August 1984, covering
a period of six years and eight months. At the onset of flea infestations, particular
attention was given to combing as part of grooming. Also, since the pets not only
differed in size but in kind of pelage, different techniques were used in removing fleas.
The part retriever female dog weighs about thirty pounds (13.6 kg), has straight, black
hair which is dense over most of her body. The miniature male poodle, weighs about
eleven pounds (4.95 kg), has tightly curled, tan hair through which a comb cannot
be drawn. The grey tiger male cat has long, fine hair which forms a dense coat over
most of his body.
On the retriever and cat, collecting was begun with a regular comb (7 teeth/cm).
Some fleas were taken from hair mats removed with the comb. To remove fleas from
the mats, forceps were used to transfer them to 70% ethanol. (Experience proved that
it was important to keep the fleas in the mat and not allow them footing on any un-
broken surface. While in the mat they seldom jumped, but from the comb's surface
or finger they quickly sprung aloft). In addition, fine-toothed combs (12 teeth/cm)
were used and were the most efficient collecting tool. Besides becoming entangled in
hair mats, fleas were often wedged between the combs' teeth. They were then forced
out with a thin blade into the alcohol. With the retriever, an alcohol wash-bottle was
also used. In dense black hair, dark brown fleas can easily escape detection, but when
seen they were doused with the alcohol. Besides being entangled they were mildly
anesthetized making removal with blunt forceps easy. Forceps and alcohol wash-bottle
were used to remove fleas from the poodle. Apparently, flea behavior includes positive
thigmotropism. By pressing the hair upon the skin, a flea nearby would wedge itself
329
330 Indiana Academy of Science Vol. 94 (1985)
into the hair mesh so formed, and was easy to collect with the forceps. Fleas were
readily removed from the poodle with forceps, especially on ventral posterior areas
whf e the skin was mostly hairless.
Combing time varied on the pets, continuing on each until no fleas were seen.
Combing began anteriorly, proceeding posteriorly on the dorsal surfaces and the pro-
cess repeated on the ventral areas with the animals lying on their sides. By means
of a stereo microscope (30X), collected fleas were sexed and numbers of each sex recorded
by the week. During periods of population build-ups, collecting was done on a daily
basis. Specimens for each year were stored in vials with 70% alcohol. When several
successive checks for fleas revealed their absence, collecting stopped. Collecting resumed
when any of the pets were observed scratching themselves, or if during the weekly
grooming, blood clots appeared in the hair mats. The appearance of clots on their
sleeping pads also prompted the resumption of collecting. The presence of adult fleas
was a certain indication that collecting should resume.
As indicated in the Introduction, use of insecticides was not anticipated in con-
trolling fleas. However, it was deemed necessary to apply an insecticide twenty-one
different times during the six-year period, 1978 to 1984. Sevin™ as powder or spray
and Durakyl™ were applied to one or all three of the pets and/or to their sleeping quarters
on the following dates:
1978 August and September 1982 July, August (3)*, September and
October
1979 August and September 1983 June, July (2), August and
September
1980 None 1984 June, July (2), and August (2)
1981 December *Number of applications per month
Using methods described above, it was possible to maintain a flea-host relation-
ship among the pets that did not involve humans as temporary hosts. Additional steps
taken to maintain this relationship included regular shaking and sweeping of sleeping
pads and cleaning of sleeping quarters. Also, all debris combed from the pets was
caught on the grooming pad and removed from the house.
Results
Most of the fleas were collected during the second halves of the years (Figures
1 and 2). In the spring of 1983, however, more fleas were taken from the cat than
from the dogs (Figure 2). A total of 4,549 fleas were collected and sex determinations
made. Of the total, 2,937 were females and 1,612 were males resulting in a sex ratio
of about 1.8:1.0. The species of flea collected in this study was Ctenocephalides felis
felis (Bouche) as determined by Whitaker and Benton (personal communication). Among
specimens submitted for examination by these investigators, the genal spines were not
consistently subequal, nor was the distal end of the manubrium of the male clasper
typically that of C. f. felis. With some fleas, Whitaker noted that the manubrium
was expanded, somewhat similar to the condition found in C. can is /(Curtis), the dog
flea. Another characteristic that aided in the determination was head length. That of
C. f felis is relatively longer than C. canis. Geary (3) collected C. /. felis from three
times as many sources as C. canis which attests to its more widespread distribution
than the latter species.
In his study of ectoparasitic insects, Marshall (3) found that females usually
predominate in natural populations. Although he determined that the sex ratio of the
majority of ectoparasitic insects is parity at emergence, fleas represent an exception.
Entomology
331
500
400
300
200
100
500
400
300
200
100
500
400-
_
300 -
200 -
100 -
1984
■ MALE
□ FEMALE
t r
t 1 1 r
1983
1982
t 1 1 r
JAN FEB MAR APR MAY JUN JUL AUG SEP OCT NOV DEC
Figure 1. Number of fleas collected monthly from two domestic dogs, 1978-1984.
One factor offered to account for this is that male fleas, being more active and smaller
than females, tend to separate more readily from the host. Also, he believes that males
332
60 r
40-
20
Indiana Academy of Science
Vol. 94 (1985)
T-1984
■ MALE
□ FEMALE
T , 1 — f
_n jl|
t 1 1 r
60
40
20
T-1983
T r
60
40
20
T-1982
t r
p
I
liUi
60
40
20
T-1981
t 1 1 1 1 1 r
JAN FEB MAR APR MAY JUN JUL AUG SEP OCT NOV DEC
Figure 2. Number of fleas collected monthly from a domestic cat (T), 1981-1984.
are less able to withstand adverse conditions of nutrition and climate. He further noted
that fleas are seasonal breeders. Another factor is that a sampling bias may exist that
is caused by one sex spending more time on the host or in the home than the other.
Finally, he noted that the sex ratio may vary with the season.
In the present study, female fleas outnumbered males 1.8:1.0. Although collec-
tions for some years show one and sometimes two months where the number of males
Entomology 333
exceeds that of females, the preponderance of females over males is clear (Figures
1 and 2). This suggests that there is a certain amount of stability in the sex ratio.
Chumakova and Kozlov (1) tested the stability of sex ratios in three species of
fleas. They found the ratio stable in flea progeny as a function of the age of female
fleas, when the progeny fed on different host species, and when progeny fed on dif-
ferent generative states of the host.
An attempt was made to correlate area meteorological data with numbers of fleas
collected on a monthly basis for each of the six years and eight months. No significant
correlation was observed when numbers of fleas were compared with monthly means
of temperature, rainfall, and relative humidity. The highest correlation was found with
temperature (r = 0.3). It was thought that significant correlation with one or more
of these weather parameters would aid in explaining the changes in seasonal abun-
dance as well as annual fluctuations in numbers of fleas shown in Figures 1 and 2.
Cole (2) studied the effects of temperature on the sex ratio in Xenopsylla cheopis
(Rothschild), the rat flea. He found no significant correlation between percentage of
females, collected weekly for forty-five weeks, and rainfall, relative humidity, or satura-
tion deficiency. However, there was high correlation (r = 0.8) with the sex ratio and
weekly temperatures.
Throughout the flea control period represented in this report, a serious effort
was made to avoid using insecticides. When used most often, 1982-1984, population
increases were more pronounced than during previous years. In order to provide relief
to the pets and to keep the host-flea relationship at tolerable levels for them, insec-
ticides were applied as indicated. While numbers of fleas were reduced, distribution
patterns from year to year appear to be less affected by these applications (Figures
1 and 2). Insecticides were used when needed rather than according to schedule.
Conclusions
Fleas of dogs and cats that live in the home can become a serious nuisance to
human occupants if not actively controlled. In this study, from January 1978 through
August 1984 , the attempt was to prevent fleas from building up large, uncontrollable
populations on the pets without using insecticides. The intent was to maintain a
manageable population of fleas by removing them from the pets regularly, mostly on
a weekly basis. During the period indicated, on twenty-one occasions, however, it became
necessary to use insecticides in addition to mechanical removal and sanitary control
of fleas.
Results of tallies of fleas over the years suggest a seasonal basis for observed
changes in their populations. However, no significant correlation between numbers
of fleas and mean monthly temperatures, rainfall, or relative humidity was found.
With few exceptions, however, the sex ratio in which females outnumbered male fleas
was consistent from year to year. Overall, females outnumbered males about 1.8:1.0.
In this study during which fleas and their hosts were largely protected from the
influence of the changes in weather and in the seasons, it appears that annual changes
in the flea populations are intrinsically controlled on a cyclical basis which is indepen-
dent of climatological influences. This explanation could account for the presence of
fleas during the latter half of the year and their low numbers or absence in late winter
and spring.
I wish to thank Dr. John O. Whitaker, Jr., Indiana State University and Dr.
Allen Benton, State University College of New York at Fredonia for their collabora-
tion in determining the species of flea collected in this study. I am also indebted to
several members of the Indiana State University staff for assistance in preparing this
paper. They are: Mrs. Lucinda Roberts, graphic artist; Mr. Anthony Brentlinger,
334 Indiana Academy of Science Vol. 94 (1985)
photographic specialist; Mr. Milton Firestone, Computer Center; Mr. William Gustin,
I.S.U. Climatic Station; and Miss Kathi Paton, Life Science Department, for typing
the manuscript.
Literature Cited
1. Chumakova, I.V., and M.P. Kozlov. 1979. Stability of the sex ratio and its
significance in the reproduction of fleas (Aphaniptera). Entomol. Rev.
58(2):244-247.
2. Cole, L.C. 1945. The effect of temperature on the sex ratio of Xenopsylla cheopis
recovered from live rats. Public Health Reports 60(45 ):1337-1354.
3. Geary, J.M. 1959. The fleas of New York. Cornell University, Agric. Expt. Sta.
Memoir 355. Ithaca, N.Y.
4. James, M.T. and R.F. Harwood. 1969. Herm's Medical Entomology, 6th edition.
The Macmillan Co., N.Y., 484 p.
5. Marshall, A.G. 1981. The sex ratio in ectoparasitic insects. Ecol. Entomol.
6(2):155-174.
6. Whitaker, J.O., Jr. 1982. Ectoparasites of mammals of Indiana. Ind. Acad. Sci.
Monograph No. 4.
Control of Vegetable Insects with Neem Seed Extracts
David K. Reed1 and Gary L. Reed2
Agricultural Research Service, USDA
Vincennes University
Vincennes, Indiana 47591
Introduction
Plant products have a great potential for providing new and novel materials for
pest management. The Neem tree, Axadirachta indica, has provided researchers with
materials which appear promising against a variety of organisms (9). This remarkable
tree, which grows in hot and arid parts of the world, has been known for centuries
to possess unique properties (3-4-5), among them, the ability to ward off insects and
other pests. Neem seed is used for many practical purposes, and very little fractiona-
tion is necessary to provide materials with insecticidal, antifeedant, or growth modify-
ing activity. Many parts of the tree are currently, and have for centuries been, used
in medicine and cosmetics, an indication of the safety of these botanical materials.
This paper reports on results of experiments conducted to access the efficacy of
crude formulations of neem seed against economically important insect pests of
vegetables.
Materials and Methods
Experiments were conducted at Vincennes, IN during 1982-84. The neem for-
mulations used were either a liquid formulation made up of an ethyl alcohol extract
of neem seed flour as a 1:1 dilution, or a dust formulation made up of defatted ground
neem seed in kaolin. The liquid formulations of neem had previously been found to
be effective as antifeedants against striped cucumber beetle Acalymma vittatum (F,)
(STCB) (6), and 2 of its principal components, azadirachtin and salannin, were shown
to deter feeding of STCB and spotted cucumber beetle, Diabrotica undecimpunctata
howardi Barber, in greenhouse experiments (7). In these tests, Triton B-1956® 3 was
added at a 0.075% concentration to the liquid neem formulations.
Greenhouse tests — Muskmelon, var. Saticoy, seedlings were raised to a 2-leaf stage
in 64 cup trays and thinned to 8 rows of 4 plants each. After cotyledon leaves were
removed, treatments were applied to plants in the rows which had been randomly
assigned. The experiment was replicated 3 times by treating 3 trays, each randomized
differently, and placing them into separate 50x50x50-cm screen cages. The greenhouse
was maintained at 29.5 ± 5°C, 60 RH ± 10% and 15:9 LD photoperiod regime.
Dust was applied to individual plants with a puff duster whose nozzle was inserted
into a 100-ml plastic cup placed over each plant to prevent cross contamination. STCB
(50/cage) were immediately introduced into the cages. Plants were examined at 2-day
intervals and damage was rated from 0 (no damage) to 6 (complete destruction or
consumption of foliage).
Field tests — Sweet corn, 1982 and 1984. Sweet corn var. Silver Queen was planted
in 8 x 1.8-m plots replicated 3 times in 1982 and 4 times in 1984. Silks were treated
as they emerged by atomizing liquid formulations onto each ear to run-off using a
Forestry tree paint sprayer in 1982 and a Solo backpack sprayer in 1984. Ears were
treated 8 times in 1982 and 6 times in 1984 on an approximate 3-day schedule. All
marketable ears were harvested 1 day after the last application and examined for corn
earworm, Heliothis zea (CEW), and damage. In 1982, carbaryl and in 1984, Ammo®
(cypermethrin), a synthetic pyrethroid was used as a standard insecticide.
335
336 Indiana Academy of Science Vol. 94 (1985)
Eggplant — 1983. Eggplant var. Dusky was planted in 16 x 1.8-m plots replicated
3 times. Sprays were applied with a high clearance sprayer consisting of a 1-row boom
composed of 1 central nozzle over the plants with a 2 dropped nozzles. Weekly
applications were made (July 25-Sept 8) using 75 psi and 19 gpa. Dust treatments were
applied with a Hudson plunger type puff duster. Ammo was used as a standard. Damage
by flea beetle, Epitrix fuscula Crotch (FB) was rated on Aug 29 by applying a 2.5
cm2 template over 5 randomly selected leaves from each of 10 plants in a row and
counting the number of feeding holes. Colorado potato beetles Leptinotarsa decemlineata
Say (CPB) were counted at weekly intervals. Marketable fruits were harvested Aug
29 and Sep 9, 1983.
Potatoes — 1982. "Superior" potatoes were planted in 16 x 1.8 m plots replicated
4 times. Treatments were applied using a tractor mounted boom sprayer with 1 central
and 2 dropped nozzles at 65 psi and 21 gpa. Monitor® was applied as a standard.
Applications were made weekly from June 2 until July 8. Weekly counts were made
of CPB adults and larvae.
Cabbage — 1983. A fall cabbage crop, var. Danish Ballhead was planted on June
23 and transplanted into the field on Aug 8 in 16 x 1.8 m plots replicated 4 times.
Weekly treatments were applied with the same equipment as used on eggplant and
insect counts were made on weekly intervals from Sep 2 until Oct 14.
In all of the field experiments, a randomized complete block arrangement was
used. Data from all experiments were transformed (x + 1) and submitted to ANOV
and DNMR.
Results and Discussion
Greenhouse tests. In the experiment using neem seed formulations in kaolin (Table
10, the untreated plants were almost immediately consumed by STCB, but all dust
treatments afforded some protection. Even kaolin alone provided some deterrent activity
as long as other food was available. This avoidance by feeding beetles was probably
due to physical factors alone and was easily overcome by starvation. Loss of activity
by the higher dosages of neem after 3 days could be due to a lack of coverage after
leaf growth, and treatments on a 2-3 day interval would be needed for continued pro-
tection, particularly in the absence of alternate food. Pure neem seed flour (100%),
when applied to young seedlings, was very phytotoxic but no such phytotoxicity was
observed with the 20% dosage, which maintained some effect up to 6 days after
treatment.
Table 1 . Antifeedant activity of neem seed dust formulations against striped cucumber
beetle adults on muskmelon seedlings in the greenhouse.
Damage
rating1 at
indicated day after treatment
Material
Neem
Dosage
100%
1
oa
2
0a
3
1.00ab
5
2.67a
6
2.67ab
Neem
20%
0a
0.08a
0.33a
1.58a
1.58a
Neem
20% (Celite)
0a
0.33a
1.50ab
2.50a
3.00b
Neem
Neem
10%
5%
oa
oa
0.08a
0a
0.67ab
0.58ab
1.42a
1.92a
2.00ab
3.08ab
Neem
2%
0.67a
0.75a
1.67ab
3.17a
4.17ab
Kaolin
—
0.33a
1.25b
2.75b
4.61
4.75ab
Untreated
—
5.33b
6.00c
6.00c
6.00b
6.00b
'Rating = 0-no feeding and 6-complete consumption or destruction.
2Means followed by the same letter are not significantly different (P = 0.05) by Duncan's New Multiple Range Test.
%
Damaged Ears
Dosage
1982
1984
.8%
3.4a'
—
.2%
29.2b
8.75ab
.4%
26.2b
16.70bc
0.6 lb/ A
—
1.85a
—
69. 9C
23.60c
Entomology 337
Table 2. Efficacy of neem seed extract against corn earworm on sweet corn.
Materials
Carbaryl
Neem
Neem
Ammo
Untreated
'Means followed by the same letter are not significalty different (P = 0.05) by Duncan's New Multiple Range Test.
Field tests — Sweet corn, 1982 and 1984. Results of the trials on sweet corn are
presented in Table 2. During both years, the standard insecticides used provided excellent
control of CEW as expected. The neem formulations gave a significantly greater level
of control than the untreated but this level would not satisfy the requirements of a
commercial grower. In some instances, in the neem treatments, the observed damage
was very slight and the young larvae were either dead or not found. Such damage
would be tolerated in a home garden situation. There appeared to be little difference
between the 2 neem dosages so that increasing the dosage would not increase efficacy
to any extent.
Table 3. Efficacy of neem seed extract against flea-beetle (FB) and Colorado potato
beetle (CPB) on eggplant.
No. FB
No.
CPB'
Total wt
Total no.
Material
Dosage
holes/cm
Adult
Larvae
mkt. fruit (g)
mkt. fruit
Neem spray
Neem dust
.2%
20%
1.2a-'
8.3b
11.3
6.3
0.3a
8.3b
964 la
2723b
35. 7a
11.3b
Ammo
.06 lb
ai/A
0.4a
7.3
4.7a
117263
42. 3a
Untreated
—
10.4b
11.3ns
10.7b
2877b
9.3b
'Mean no./5 plants.
2Means followed by the same letter are not significantly different (P = 0.05) by Duncan's New Multiple Range Test.
Eggplant — 1983. As shown in Table 3, Ammo, the standard insecticide was
extremely effective against FB and CPB larvae. However, neem spray was just as ef-
fective against both of these insects, both being significantly better than the untreated.
Also, there was no difference between number and weight of marketable fruit between
the 2 treatments. None of the treatments appeared to control adult CPB, possibly
due to new infestations moving in from adjacent plots. Neem dust was not effective
against either FB or CPB and this was reflected in the number and weight of marketable
fruit.
Potatoes — 1982. CPB larvae were controlled by neem spray when applied to
potatoes (Table 4). As with eggplant, however, adults were not controlled by either
Table 4. Efficacy of neem seed extract against Colorado potato beetle (CPB) on potatoes.
1982.
Mean no. for 5 plants
Material Dosage Adults Larvae
6.0 1.0a'
4.3 1.3a
6.3ns 6.0b
'Means followed by the same letter are not significantly different (P = 0.05) by Duncan's New Multiple Range Test.
Neem
.2%
Monitor
.75 lb ai/A
Untreated
—
338 Indiana Academy of Science Vol. 94 (1985)
neem or the standard insecticide. Again, this may have been due to migration and
not to lack of toxicity of the insecticide.
Cabbage — 1983. During the fall crop, the major pest of cabbage is cabbage looper
Trichoplusia ni (Hubner) (CL). Although neem spray was not as effective as the syn-
thetic pyrethroid against CL on cabbage (Table 5), it did provide significantly greater
control than the untreated. Whether activity of neem is related to direct toxicity or
to a form of repellency is unknown at the present time.
Table 5. Efficacy of neem seed extracts against cabbage looper (CL) on cabbage. 1983.
Material Dosage Mean no. CL larvae/5 plants
Ammo .06 lb ai/A 1 .3 '
Neem .2% 13. 3b
Untreated — 53. 3C
'Means followed by the same letter are not significantly different (P = 0.05) by Duncan's New Multiple Range Test.
One of the major insect antifeedants isolated from neem kernels, azadirachtin,
has been shown to possess growth regulator activity against insects (1 and 8). The
reduction in larval development was not related to feeding inhibition. Azadirachtin
in both of these reported studies apparently interfered with the molting hormone pools
and affected normal ecdysis. Neem extracts were also shown to have a phagodeterrent
effect on a flea beetle, Phyllotreta striolata (F.) in the laboratory (2). Our research
substantiates this report. These are only a few of the many references to neem effec-
tiveness against insects, and indicate the great potential that this material may have
in pest management. Although it does not have the immediate, highly toxic activity
of many pesticides, its activity against a variety of insect ciders, its mammalian safety
and its environmentally non-disruptive nature should make it an ideal candidate for
use in vegetable insect control. Where efficacy is not great enough to produce a com-
mercial crop, home gardeners, because of their acceptance of greater injury levels,
may be able to utilize neem effectively. Although neem sprays appear to be more effective
as antifeedants, further work may be warranted with the dust formulations, particularly
against certain insects.
Acknowledgment
This research was conducted at Vincennes, Indiana, USDA Laboratory and
Southwest Purdue Agricultural Center, in cooperation with Indiana Agricultural
Experiment Station, Purdue University, where both authors hold adjunct appointments
in the Department of Entomology. Neem fractions were obtained from the Biologically
Active Natural Products Laboratory, ARS-USDA, Beltsville, Maryland.
Footnotes
1 . Present address: Asian Parasite Laboratory, c/o American Embassy, Seoul Korea,
APO San Francisco 9630.
2. Present address: Oregon State University, Columbia Basin Agricultural Research
Station, Box 105, Hermston, OR 97838.
3. This article reports the results of research only. Mention of a proprietary product
does not imply an endorsement or a recommendation for its use by USDA.
Entomology 339
Literature Cited
1. Kubo, I. and Kloche, J. A. 1982. Azadirachtin, insect ecdysis inhibitor. Agric.
Biol. Chem. 46:1951-1953.
2. Meisner, J. and Mitchell, B.K. 1982. Phagodeterrent effect of neem extracts and
azadirachtin on flea beetles, Phyllotreta striolata (F.). Z Pflkrankh. Pflschutz
89:463-467.
3. Radwanski, S. 1977. Neem tree 1. Commercial potential, characteristics and
distribution. World Crops and Livestock 29:62-65.
4. . 1977. Neem tree 2. Uses and potential uses. World Crops and Livestock
29:111-113.
5. . 1977. Neem tree 3. Further uses and potential uses. World Crops and
Livestock 29:167-168.
6. Reed, D.K., Jacobson, M., Warthen, J.D. Jr., Uebel, E.C., Tromley, N.J., Jurd,
L. and Freedman, B. 1981. Cucumber beetles antifeedants: Laboratory screening
of natural products. US Dep. Agri. SEA Tech. Bull. No. 1641. 13 pp.
7. Reed, D.K., Warthen, J.D. Jr., Uebel, E.C. and Reed, G.L. 1982. Effects of
two triterpenoids from neem on feeding by cucumber beetles (Col-
eoptera.Chrysomelidae). J. Econ. Entomol. 75:1109-1113.
8. Rembold, H., Sharma, G.K., Czoppelt, Ch., and Schmutterer, H. 1982.
Azadirachtin: A potent insect growth regulator of plant origin. Z. Ang. Ent.
93:12-17.
9. Warthen, J.D. Jr. 1979. Azadirachta indica: a source of insect feeding inhibitors
and growth regulators. US Dep. Agri. Rev. Manuals. ARM-NE-4, 21 pp.
Checklist of Adult Carabid Beetles Known from Indiana
John Richard Schrock
Association of Systematics Collections
University of Kansas
Lawrence, Kansas 66045
A followup of Munsee's 1964 study of insects on unreclaimed stripmines (8) placed
at least 32 species of adult ground beetles on Vermillion County spoilbanks. To ad-
dress the question "What percent of known Indiana carabids are represented in this
environment?" it was necessary to compile a checklist of ground beetles identified
from within the State.
Indiana is fortunate to have W.S. Blatchley's turn-of-the-century classic Coleoptera
of Indiana. Blatchley (1) listed 382 species of ground beetles, 351 of which are still
recognized under a valid species name today in the North America Beetle Fauna (NABF)
checklist (5).
In 1941, E.L. Montgomery and J.M. Amos surveyed the beetles of the Clark
County State Forest but published only the non-carabids (6). N.M. Downie added
new "Records of Indiana Coleoptera" in 1956 (2), again in 1958 (3) and in 1967 with
C.E. White (4). This increased the known Indiana carabid fauna by 21 species recognized
today.
The NABF Project checklist (5) clarified much of the synonymy and recorded
223 species from Indiana. However in many genera, species were recorded from most
of the surrounding states but not for Indiana and 181 of the valid species found in
Blatchley were not recorded in the NABF checklist for Indiana.
To develop a fuller checklist, the identified collections at Purdue University, Il-
linois Natural History Survey and the Snow Entomological Museum were examined
and a list of the holdings at Indiana University was incorporated. N.M. Downie pro-
vided a list of carabids (from his large personal collection) not found in the other
collections and clarified the status of "Bembidion intermedium Kirby," "Elaphrus
riparius Linneaus," and " Pentagonica flavipes Leconte." Most synonyms were traced
using the NABF list. Dr. George Ball placed six problematic names, updated the
systematics, and suggested the names Amara trivialis Roucu., PU, and Aniilinus falli
Bar, PU, are either unpublished or in error. Species from the Munsee (7) and Schrock
(8) surveys were added.
A total of 465 species is recorded for Indiana. In 1910, Blatchley listed 382 species
and suggested an additional 74 might be found within the State based on records from
neighboring states. Twenty-five of these are now in the present list. However, today
there are an additional 124 species of ground beetles found in neighboring states in
the NABF list that may have ranges extending across the Indiana border.
Since ground beetles are a large group of common beetles highly selective in habitat
and therefore important indicator species (9), this carabid checklist should provide
a useful inventory reference in future environmental studies. And cataloguing our cur-
rent holdings of carabids should make it easier to tally the remaining species yet to
be found within the State.
I would like to thank Dr. Jack R. Munsee for aid in repeating the stripmine
study. Dr. George Ball identified spoil bank specimens and both Dr. Ball and N.M.
Downie scanned this list for any major inaccuracies. Dr. Carl Krekeler of Valparaiso
University checked the Pseudanophthalmus listings. Any remaining errors however,
are mine. Examination of museum collections was made possible with the kind help
of: Dr. Wallace LaBerge and Steve Heydon, Illinois Natural History Survey; Dr. George
341
342 Indiana Academy of Science Vol. 94 (1985)
Byers, Snow Entomological Museum; and Dr. W. P. McCafferty and R.D. Waltz,
Purdue University. Dr. Frank Young tallied the holdings of the Indiana University
collection. Without the use of the computer word processing facilities at the Associa-
tion of Systematics Collections granted by Dr. Stephen Edwards, the management of
this list would have been extremely laborious.
CHECKLIST OF ADULT CARABID BEETLES KNOWN FROM INDIANA
DC = N. M. Downie Collection
IL = Illinois Natural History
Survey Collection
IU = Indiana University
KU = Snow Entomological Museum
University of Kansas
PU = Purdue University
B = Blatchley's Coleoptera of Indiana (1)
C = NABF Checklist (5)
Dl, D2, D3 = Records of Indiana Coleoptera
I, and II and III by Downie (2), Downie (3),
and Downie and White (4), respectively
M = Munsee's 1966 stripmine collection, reported in Schrock (8)
S = Schrock's 1981 stripmine survey (8)
1. A bacidus permundus (Say) IU,PU; C,D1
( = Pterostichus permundus Say) B
2. Acupalpus alternans LeConte C
3. Acupalpus cams (LeConte) PU;B
4. Acupalpus hydropicus (LeConte) PU;B
5. Acupalpus indistinctus (Dejean) DC
6. Acupalpus partiarius (Say)
( = Agonoderus partiarius Say) B
( = Tachistodes partiarus (Say)) PU
7. Acupalpus pauperculus (Dejean)
( = Agonoderus pauperculus Dejean) B
( = Tachistodes pauperculus (Dejean)) PU
8. Acupalpus rectangulus Chaudoir C
9. Acupalpus testaceus (Dejean)
( = Agonoderus testaceus Dejean) B
( = Tachistodes testaceus (Dejean)) PU
10. Agonum aeruginosum (Dejean)
( = Circinalia aeruginosus (Dejean)) KU
( = Platynus aeruginosus (Dejean)) PU;B
11. Agonum affine Kirby IL
( = Platynus affinis (Kirby)) PU;B
12. Agonum albicrus (Dejean)
( = Agonum albicrum Dejean) D3
( = Platynus albicrus (Dejean) PU;B
13. Agonum anchomenoides (Randall) IL
( = Platynus anchomenoides (Randall)) PU
14. Agonum basale LeConte C
( = Platynus basalis (LeConte)) PU;B
Entomology 343
15. Agonum collare Say C
( = Platynus collaris Say) B
16. Agonum corvus (LeConte)
( = Platynus corvus LeConte) PU;B
17. Agonum crenistriatum (LeConte)
( = Platynus crenistriatus LeConte) PU;B
18. Agonum cupripenne (Say)
( = Platynus cupripennis (Say)) PU;B
19. Agonum decorum (Say)
( = Platynus decorus (Say)) PU;B
( = Platynus obscurus LeConte) PU
20. Agonum dilutipenne Motschulsky PU
21. Agonum errans (Say)
( = Platynus errans (Say) PU;B
( = Platynus errans subcordatus LeConte) PU
( = Platynus subcordatus LeConte) B
22. Agonum excavatum (Dejean)
( = Platynus excavatus (Dejean)) PU;B
23. Agonum extensicolle (Say)
( = Platynus extensicollis (Say)) PU;B
( = Platynus extensicollis viridis (LeConte)) PU;B
24. Agonum ferreum Haldeman IL
( = Platynus ferreus (Haldeman)) PU;B
25. Agonum formosum Sturm C
26. Agonum gratiosum Mannerheim PU;C
( = Platynus ruficornis LeConte) PU;B
27. Agonum lutulentum (LeConte) IL;C
( = Platynus lutulentulus LeConte) PU;B
28. Agonum melanahum (Dejean) KU
( = Platynus melanarius (Dejean)) PU;B
29. Agonum moerens (Dejean) C
( = Platynus moerens Dejean) B
30. Agonum mutatum Gemminger & Harold IL,PU;D1
( = Platynus atratus LeConte) B
31. Agonum nutans (Say)
( = Platynus nutans (Say)) PU;B
32. Agonum octopunctatum (Fabricius)
( = Platynus octopunctatus (Fabricius)) PU;B
33. A go nu m pallipes Fabricius C
( = Platynus limbatus (Say)) PU;B
34. Agonum picticorne (Newman) D3
( = Platynus picticornis Newman) PU
35. Agonum placidum (Say)
( = Platynus placidus (Say)) PU;B
36. Agonum propinquum Gemminger & Harold
( = Platynus piceus (LeConte)) PU
37. Agonum punticeps Casey
( = Platynus pusillus LeConte) B
38. Agonum punctiforme (Say)
( = Platynus punctiformis (Say)) PU;B
39. Agonum quadhmaculatum (Horn) C
( = Platynus quadrimaculatus Horn) PU;B
344 Indiana Academy of Science Vol. 94 (1985)
40. Agonum rubripes LeConte C
( = Platynus rubripes (LeConte)) PU;B
41. Agonum rufipes (Dejean) DC
42. Agonum striatopunctatum (Dejean) C
( = Platynus nutans striatopunctatus (Dejean)) PU
( = Platynus striatopunctatus Dejean) B
43. Agonum tenue (LeConte)
( = Platynus tenuis LeConte) PU;B
44. Agonum thoreyi Dejean
( = Platynus gemellus LeConte) PU;B
( = Platynus picipennis (Kirby)) PU;B
45. Amara aenea DeGeer DC
46. Amara alpina Paykull
( = Curtonotus argutus Casey) PU
47. Amara angustata Say B
48. Amara apricaria (Paykull) DC
49. Amara avida (Say) PU;B
( = Leiocnemis avida Say) PU;D1
50. Amara basillaris (Say) PU;B
51. Amara calij ornica (Dejean)
( = Celia californica (Dejean)) PU
52. Amara chalcea Dejean B
( = Celia chalcea Dejean) PU
53. Amara confusa LeConte
( = Amara protensa Putzeys) PU;B
54. Amara convexa LeConte
( = Amara polita LeConte) PU;B
55. Amara crassispina LeConte DC
56. Amara cupreolata Putzeys PU;B
57. Amara erratica Duftschmidt B
( = Celia erratica (Sturm)) PU
58. Amara exarata Dejean PU;B,C
( = Bradytus exaratus Dejean) PU
59. Amara familiaris Duftschmid DC
60. Amara hyper bo rea Dejean
( = Curtonotus elongatus (LeConte)) PU
61. Amara interstitialis Dejean B
62. Amara impuncticollis Say PU;B
63. Amara latior Kirby B
( = Bradytus latior Kirby) PU
64. Amara littoralis Manner heim PU
( = Amara fallax LeConte) PU;B
65. Amara musculis (Say) B
( = Celia musculis (Say)) PU
66. Amara obesa (Say) PU;B
( = Percosia obesa Say) IU
67. Amara pallipes (Kirby)
( = Triaena pallipes Kirby) D2
68. Amara patruelis Dejean PU
69. Amara pennsylvanica (Hay ward) B
( = Curtonotus pennsylvanicus Hayward) PU
Entomology 345
70. Amara quenseli Schoenherr
( = Amara remotestriata Dejean) B
71. A mar a rubrica (Haldeman) B
( = Celia rubrica Haldeman) PU
72. Amara sinuosa Casey
( = Amara subaenea LeConte) B
73. Amara torrida Panzer
( = Curtonotus infaustus LeConte) PU
74. Amerinus linearis (LeConte) PU
( = Bradycellus linearis LeConte) B
75. Amphasia interstitialis (Say) PU;C
( = Anisodactylus interstitialis Say) B
76. Amphasia sericea (Harris)
( = Anisodactylus sericea Harris) B
( = Pseudamphasia sericea (Harris)) PU
77. Anatrichis minutus Dejean IU;B
78. Anillinus affabilis Brues PU
79. Anillinus fortis Horn B
80. Anillinus indianae Jeannel C
81. Anisodactylus agricola Say PU;B,C
82. Anisodactylus carbonarius (Say) PU;B
83. Anisodactylus caenus Say PU;B,D1
84. Anisodactylus discoideus Dejean B
( = Anadaptus discoideus (Dejean)) IU,PU
85. Anisodactylus dulcicollis LeFerte' PU
( = Triplectrus dulcicollis LaFerte') D2
86. Anisodactylus furvus LeConte PU;B,C
87. Anisodactylus harrisi LeConte PU;B,C
88. Anisodactylus kirbyi Lindroth DC
89. Anisodactylus melanopus Haldeman KU,PU;B,D1
90. Anisodactylus merula Germar C
91. Anisodactylus nigerrimus Dejean IU,PU;B
92. Anisodactylus nigrita Dejean B
( = Anisodactylus interpunctatus Kirby) PU;B
93. Anisodactylus nivalis Horn
( = Anadaptus parvulus Casey) PU
94. Anisodactylus ovularis Casey C
( = Triplectrus ovularis Casey) PU
95. Anisodactylus rusticus Say IL;B,C
( = Triplectrus rusticus (Say)) IU,PU
96. Anisodactylus sanctaecrucis Fabricius PU
( = Anisodactylus baltimorensis (Say)) PU;B
97. Anisodactylus similis LeConte
( = Anisodactylus semipunctatus LeConte) PU
98. Anisodactylus verticalis LeConte B
99. Apenes lucidula (Dejean) KU,IU; D1,S
100. Apenes sinuata Say B,C
101. Apristus subsulcatus Dejean B
( = Apristus cordicollis LeConte) B
102. Ardistomis puncticotlis (Dejean) IU,PU;B,C
103. Ardistomis viridis (Say) PU;B,C,D1
346 Indiana Academy of Science Vol. 94 (1985)
104. Aspidoglossa subangulata (Chaudoir) IU,PU;B,C
105. Atranus pubescens (Dejean) PU;B,C,D1
106. Axinopalpus biplagiatus Dejean B,C
107. Axinopalpus calif ornicus Motschulsky C
108. Badister flavipes LeConte C
109. Badister flavipes laticeps Blatchley C
( = Badister laticeps Blatchley) B
110. Badister maculatus LeConte PU;B,C
111. Badister neopulchellus Lindroth DC
112. Badister notatus Haldeman PU;B,C
113. Badister ocularis Casey C
( = Badister micans LeConte) B
114. Badister parviceps Ball C
115. Badister pulchellus LeConte B,C
116. Badister reflexus LeConte PU:B,C
117. Badister transversus Casey PU;C
118. Bembidion affine Say IU,PU;B
119. Bembidion americanum Dejean PU;B
120. Bembidion anguliferum (LeConte) PU;B
121. Bembidion cannula Chaudoir IL,PU;B
122. Bembidion chalceum Dejean PU;B
123. Bembidion concretum Casey DC
124. Bembidion confusum Hayward PU;B
125. Bembidion cordatum (LeConte) PU;B
126. Bembidion coxendix Say PU;B,C
127. Bembidion fortestriatum Motschulsky
( = Bembidion cautum (LeConte)) PU
128. Bembidion frontale (LeConte) PU
( = "Bembidion assimile Gyllenhal") B
129. Bembidion fugitans Casey C
130. Bembidion graciliforme Hayward PU;B
131. Bembidion grapei Gyllenhall C
( = Bembidion picipes (Kirby)) PU;B,D1
132. Bembidion honestum Say B
133. Bembidion inaequale Say PU;B,D1
134. Bembidion lacunarium Zimmermann C
135. Bembidion laevigatum Say PU;B,D1
136. Bembidion minax Casey C
137. Bembidion nigrum Say PU;B,D1
138. Bembidion nitidum Kirby B
139. Bembidion obscurellum Motschulsky D3
140. Bembidion patruele Dejean IL,PU;C
( = Bembidion fraternum LeConte) B
141. Bembidion pedicellatum LeConte IU,PU;B,D1
142. Bembidion planum (Haldeman) PU
( = Bembidion guexi Chaudoir) B
143. Bembidion postremum Say DC
144. Bembidion punctatostriatum Say PU;B
145. Bembidion quadrimaculatum oppositum Say DC
( = Bembidion quadrimaculatum (Linnaeus)) PU;B,D1
146. Bembidion rapidum LeConte DC
147. Bembidion semistriatum Haldeman B
Entomology 347
148. Bembidion tetracolum Say DC
149. Bembidion transparens Gebler C
150. Bembidion variegatum (Say) KU,PU;B
( = Bembidion postfasciatum Hamilton) B
151. Bembidion versicolor (LeConte) IL,IU,KU,PU;B
152. Blethisa quadricollis Haldeman PU;B,C
153. Brachinus adustipennis Erwin C
154. Brachinus alternans Dejean B,C
( = Brachinus deyrollei LaFerte') B
( = Brachinus tormentarius LeConte) B
155. Brachinus americanus LeConte B,C
156. Brachinus cordicollis Dejean C
157. Brachinus cyanipennis Say C
158. Brachinus cyanochroaticus Erwin C
159. Brachinus fulminatus Erwin C
160. Brachinus fumans Fabricius C
161. Brachinus janthinipennis Dejean C
162. Brachinus medius Harris C
163. Brachinus ovipennis LeConte C
164. Brachinus quadripennis Dejean C
165. Brachinus sublaevis Chaudoir C
166. Brachinus tenuicollis LeConte C
167. Bradycellus atrimedius (Say)
( = Tachycellus atrimedius Say) B
( = Triliarthrus atrimedius (Say)) PU
168. Bradycellus badipennis (Haldeman)
( = Tachycellus badiipennis Haldeman) B
( = Triliarthrus badiipennis (Haldeman)) PU
169. Bradycellus nigriceps LeConte C
170. Bradycellus nigrinus Dejean
( = Tachycellus nigrinus Dejean) B
171. Bradycellus rupestris Say IU;B,C
( = Stenocellus ruprestris (Say)) PU
172. Calathus gregarius Say IU,KU,PU;B,C,S
173. Calathus opaculus LeConte KU,PU;B,C,S
174. Callida punctata LeConte IU;B,C
175. Calosoma calidum Fabricius IL,PU;B,C
176. Calosoma externum Say IL,PU;B
177. Calosoma frigidum Kirby B,C
178. Calosoma sayi Dejean PU
( = Calosoms alternans sayi Dejean) D3
179. Calosoma scrutator Fabricius IU,PU;B,C
180. Calosoma wilcoxi LeConte PU;B,C
181. Carabus limbatus Say IL,KU,PU;B,M,S
182. Carabus maender Fischer IL;D3
183. Carabus nemoralis Muller IU,PU
184. Carabus serratus Say IL,KU,PU;B,C,D1
185. Carabus sylvosus Say PU;B
186. Carabus vinctus Weber IU,KU,PU;B,C
187. Chlaenius aestivus Say PU;B,C
188. Chlaenius brevilabris LeConte B,C
189. Chlaenius cordicollis Kirby C
348 Indiana Academy of Science Vol. 94 (1985)
190. Chlaenius emarginatus Say IU,KU;C,S
( = Anomoglossus emarginatus Say) B
191. Chlaenius erythropus Germar B,C
192. Chlaenius impunctifrons Say B,C
193. Chlaenius laticollis Say IU;B,C
194. Chlaenius leucoscelis Chevrolat PU;B,D1
195. Chlaenius lithophilus Say IL;C
( = Brachylobus lithophilus Say) B
196. Chlaenius niger Randall B,C
197. Chlaenius nemoralis Say IU,B,C
198. Chlaenius pensylvanicus Say B,C
199. Chlaenius platyderus Chaudoir C
( = Chlaenius diffinis Chaudoir) PU;B
200. Chlaenius prasinus Dejean B,C
201. Chlaenius purpuricollis Randall IL;B,C
202. Chlaenius pusillus (Say) C
( = Anomoglossus pusillus Say) B
203. Chlaenius tomentosus Say IL;B,C
204. Chlaenius tricolor Dejean C
205. Chlaenius sericeus Forster B,C
206. Chlaenius solitarius Say B,C
207. Clivina americana Dejean PU;B
208. Clivinia bipustulata (Fabricius) IU,KU,PU;B
209. Clivina dentipes Dejean KU,PU;B
210. Clivina impressifrons LeConte PU;B,C
211. Clivina postica LeConte D3
212. Clivinia puntigera LeConte PU;B,C,D1
213. Clivina rubicunda LeConte B,C
214. Clivina rufa LeConte PU;B,C,D1
215. Colliurus pensylvanica (Linnaeus)
( = Casnonia pensylvanica Linnaeus) IU;B
216. Coptodera aerata Dejean KU;B,C
217. Cratacanthus dubius (Beauvois) IL,PU;B,C
218. Cyclotrachelus convivus (LeConte)
( = Evarthrus convivus LeConte) KU,PU;C,S
219. Cyclotrachelus furtivus (LeConte)
( = Eumolops furtiva (LeConte)) PU
( = Evarthrus furtivus (LeConte)) B
220. Cyclotrachelus obsoletus (Say)
( = Evarthrus obsoletus Say) KU,PU;C,S
( = Pterostichus obsoletus Say) B
221. Cyclotrachelus seximpressus (LeConte)
( = Evarthrus seximpressus (LeConte)) KU,PU;B,C,S
222. Cyclotrachelus sigillatus (Say)
( = Evarthrus sigillatus (Say)) PU;B,D1
( = Evarthrus americanus Dejean) B
( = Evarthrus orbatus (Newman)) PU;B
222. Cyclotrachelus sodalis (LeConte)
( = Evarthrus sodalis LeConte) KU,PU;B,C,D1,M,S
223. Cymindis americana Dejean KU;B,S
224. Cymindis limbata (Dejean)
( = Pinacodera limbata (Dejean)) KU;B,C,D1,S
Entomology 349
225. Cymindis neglecta Haldeman DC
226. Cymindis pilosa Say B
227. Cymindis platicollis (Say)
( = Pinacodera platicollis (Say)) PU;B,C,D1
228. Dicaelus ambiguus LaFerte' IU,KU,PU;B,C,D1,S
229. Dicaelus dilatus sinuatus Ball
( = Dicaelus dilatus Say) PU ; B , C
230. Dicaelus elongatus Bonelli KU,PU;B,C,S
231. Dicaelus furvus Dejean KU;B,C,S
( = Dicaelus ovalis LeConte) B
232. Dicaelus furvus carinatus Dejean PU
233. Dicaelus politus Dejean IU,PU;B,C
234. Dicaelus purpuratus Bonelli IU,KU,PU;B,C,S
235. Dicaelus sculpt His intricatus LeConte
( = Dicaelus sculptilis Say) B,C
236. Dicaelus teter Bonelli IL;B
237. Diplocheila assimilis (LeConte)
( = Rembus assimilis LeConte) PU
238. Diplocheila impressicollis Dejean PU;B
( = Diplocheila impressicollis alternans) Casey B
( = Diplocheila laticollis LeConte) B
( = Rembus laticollis LeConte) PU
239. Diplocheila major LeConte PU;C
( = Diplocheila laticollis major LeConte) B
240. Diplocheila obtusa LeConte B,C
( = Rembus obtusa LeConte) PU;D1
241. Diplocheila striatopunctata LeConte IU;C
242. Discoderus parallelus (Haldeman) PU;B
243. Dromius piceus Dejean B,C
( = Dromius picipes [sic] Dejean) Dl
244. Dyschirius erythrocerus LeConte PU;B,C,D1
245. Dyschirius globulosus Say IU,PU;B
246. Dyschirius haemorrhoidalis Dejean PU;B,C
247. Dyschirius integer LeConte
( = Dyschirius nigripes LeConte) PU;B
248. Dyschirius longulus LeConte PU;B,C
249. Dyschirius pilosus LeConte
(- Dyschirius hispidus LeConte) PU;B,D1
250. Dyschirius sphaericollis Say PU;B,C,D1
251. Dyschirius terminatus LeConte PU;B,C,D2
252. Elaphropus dolosus (LeConte)
(=Tachys dolosa (LeConte)) PU
( = Tachys dolosus (LeConte)) B
253. Elaphropus ferrugineus (Dejean)
( = Tachys ferrugineus (Dejean)) PU;B
254. Elaphropus granarius (Dejean)
( = Tachys granaria (Dejean)) PU
( = Tachys granarius (Dejean)) B
255. Elaphropus incurvus Say C
( = Tachys incurva (Say)) PU
( = Tachys incurvus (Say)) B
350 Indiana Academy of Science Vol. 94 (1985)
256. Elaphropus parvicornis Notman C
257. Elaphropus tripunctatus Say C
( = Tachys tripunctatus Say) B
258. Elaphropus vernicatus Casey C
259. Elaphropus vivax (LeConte)
( = Tachys capax LeConte) B
( = Tachys vivax (LeConte)) PU;B
260. Elaphropus xanthopus (Dejean)
( = Tachys xanthopus (Dejean)) PU;B
261. Elaphrus calif ornicus Mannerheim DC
( = Elaphrus riparius Linnaeus) D2
262. Elaphrus cicatricosus LeConte PU;B,C
263. Elaphrus clairvillei Kirby B
264. Elaphrus fuliginosus Say PU;B,C
265. Elaphrus laevigatus LeConte B
266. Elaphrus lecontei Crotch IL
267. Elaphrus lindrothi Goulet
268. Elaphrus ruscarius Say IU,PU;B
269. Episcopellus autumnalis (Say) PU;C,D2
( = Harpalus autumnalis Say) B
270. Euphorticus pubescens (Dejean) D3
27 1 . Euryderus grossus (Say)
( = Nothopus grossus Say) B
( = Nothopus valens Casey) PU
( = Nothopus zabroides LeConte) PU;D1
272. Galerita bicolor Drury KU;B,S
273. Galerita janus Fabricius IU,KU;B,C,S
274. Geopinus incrassatus (Dejean) IU,PU;B,C
275. Gynandropus hylacis (Say) PU;B,C
276. Harpalus actiosus Casey PU;C
277. Harpalus af finis Schrank
( = Harpalus viridianeus Beauvois) PU
278. Harpalus bicolor (Fabricius) KU,PU;S
( = Harpalus compar LeConte) PU;B
279. Harpalus caliginosus (Fabricius) IL,IU,KU,PU;B,M
280. Harpalus erraticus Say PU;B
281. Harpalus erythropus Dejean KU,PU;B,C,M
282. Harpalus fallax LeConte DC
283. Harpalus faunus Say PU;B,C
( = Harpalus convivus LeConte) B
284. Harpalus fulgens Csiki C
( = Harpalus nitidulus Chaudoir) PU;B,D1
285. Harpalus fuliginosa Duftschmid DC
286. Harpalus herbivagus Say PU;B
287. Harpalus indianus (Csiki) C
( = Harpalus testaceus LeConte) B
( = Pharalus indianus Csiki) PU;D1
( = Pharalus testaceus (LeConte)) PU
288. Harpalus laticeps LeConte KU;B,C,D1
289. Harpalus lewisi LeConte DC
290. Harpalus longicollis LeConte KU;C,S
( = Harpalus vagans LeConte) B
Entomology 351
291. Harpalus mob His Casey PU
292. Harpalus par at us Casey DC
293. Harpalus pensylvanicus DeGeer PU;B,C
( = Harpalus longior Kirby) B
294. Harpalus pleuriticus Kirby DC
295. Harpalus protractus Casey PU;C
296. Harpalus viduus LeConte PU;B
297. Helluomorphoides ferrugineus (LeConte) D3
298. Helluomorphoides praeustus bicolor Harris PU;C
299. Helluomorphoides texanus LeConte B,C
300. Lebia abdominalis Chaudoir B,C
301. Lebia analis Dejean PU;B,C,D1
302. Lebia atriventris Say IL,IU;B,C
303. Lebia bivittata Fabricius B,C
304. Lebia collaris Dejean C
305 . Lebia divisa LeConte PU ; D 1
306. Lebia fuscata Dejean PU;B,C,D1
307. Lebia grandis Hentz IU;B,C
308. Lebia lobulata LeConte PU;B,C,D1
309. Lebia marginicollis Dejean KU;C,D3
310. Lebia ornata Say KU;B,C,S
311. Lebia pectita Horn C
312. Lebia pleuritica LeConte D3
313. Lebia pulchella Dejean IU;D3
314. LebiapumilaDejean B,C
( = Lebia rhodopus Schwarz) B,D2
315. Lebia solea Hentz C
( = Lebia scapularis Dejean) IU;B
316. Lebia tricolor Say C,D3
317. Lebia viridipennis Dejean B,C
318. Lebia viridis Say I U ; B ,C
319. Lebia vittata Fabricius B,C
( = Lebia furcata LeConte) B
320. Leptotrachelus dorsalis Fabricius B
321. Loxandrus agillis Dejean B
322. Loxandrus brevicollis LeConte B,C
323. Loxandrus cincinnatiensis Casey C
324. Loxandrus duryi Wright C
325. Loxandrus erraticus Dejean B
326. Loxandrus extendus Allen C
327. Loxandrus gibbus Allen C
328. Loxandrus minor Chaudoir B,C
329. Loxandrus nitidulus LeConte C
330. Loxandrus rectus Say IU;B
331. Loxandrus robustus Allen C
332. Loxandrus velocipes Casey C
333. Loxandrus velox Dejean B
324. Loxandrus vulneratus Casey C
335. Metabletus americanus Dejean B
336. Micratopus aenescens (LeConte) PU
337. Microlestes nigrinus Mannerheim PU;D1
( = Blechrus nigrinus (Mannerheim)) B
352 Indiana Academy of Science Vol. 94 (1985)
338. Microlestes pusio LeConte
( = Blechrus pusio LeConte) B
339. Mioptachys flavicauda (Say)
( = Tachymenis flavicauda (Say)) KU
( = Tachys flavicauda (Say)) IU,PU;B
340. Myas coracinus Say B,C
( = Trigonognatha coracinus (Say)) PU
341. Nebria lacustris Casey PU;C,D3
342. Nebria pallipes Say PU;B,C
343. Nomius pygmaeus (Dejean) PU
344. Notiobia nitidipennis (LeConte)
( = Anisotarsus nitidipennis LeConte) PU;B,D1
( = Anisodactylus nitidipennis LeConte) PU
345. Notiobia picea (LeConte) C
( = Anisodactylus sayi Blatchley) PU;B
( = Eurytrichus piceus LeConte) IU
346. Notiobia terminata Say C
( = Anisodactylus terminatus (Say)) PU;B
( = Anisodactylus agilis (Dejean)) PU
347. Notiophilus aeneus Herbst IU,KU,PU;B,C,S
348. Notiophilus novemstriatus LeConte PU;C
349. Notiophilus semistriatus Say IU,KU,PU;B,C,S
350. Olisthopus parmatus (Say) PU;C
351. Oodes amaroides Dejean B
352. Oodes americanus Dejean B
( = Oodes fluvialis LeConte) B
353. Oodes parallelus Say KU
( = Lachnocrepis parallelus Say) B
354. Omophron americanum Dejean IL,PU;B
355. Omophron nitidum LeConte PU;B,C
356. Omophron robustum Horn PU;B,C
357. Omophron tesselatum Say IL,KU,PU;C
358. Panagaeus crucigerus Say PU;B,C
359. Panagaeus fasciatus Say IU,PU;B,C
360. Paratachys oblitus Casey C
361. Paratachys proximus (Say)
( = Tachys proximus (Say)) IU,KU,PU;B
362. Paratachys pumilus Dejean
( = Tachys coruscus LeConte) PU;B
363. Paratachys scitulus (LeConte)
( = Tachys scitulus LeConte) PU;B
364. Pasimachus elongatus LeConte IL,PU;B
365. Pasimachus depressus Fabricius PU;B
366. Pasimachus punctulatus Haldeman PU;B,C
367. Pasimachus sublaevis Beauvois C
( = Pasimachus sublaevis Bonelli) B
368. Patrobus longicornis Say IL,IU,PU;B
369. Pentagonica picticornis Bates DC
( = Pentagonica flavipes (LeConte)) PU;B,D1
370. Pericompus ephippiatus (Say) PU;D1
( = Tachys ephippiatus (Say)) PU;B
Entomology 353
371. Perigona nigriceps (Dejean) PU;C,D2
( = Perigona pallipennis (LeConte)) PU;B
372. Platynus angustatus Dejean PU
373. Platynus brunneomarginatus Mannerheim PU
374. Platynus caudatus LeConte B,C
( = Rhadine caudata LeConte) PU
375. Platynus cincticollis Say B
376. Platynus decent is Say
( = Platynus sinuatus (Dejean)) IU,PU;B
( = "Platynus decens" Say) PU,B
377. Platynus hypolithos (Say) IL,IU,PU
( = Platynus hypolithus Say) B
378. Platynus opaculus LeConte B
379. Platynus parmarginatus Hamilton PU;B,C
380. Platynus tenuicollis LeConte
( = Platynus reflexus LeConte) PU;B
381. Plochionus timidus Haldeman B
382. Polyderis laevus (Say)
( = Tachys laevus (Say)) PU;B
383. Pseudapt inus pygmaeus (Dejean) D3
384. Pseudanophthalmus barri Krekeler C
385. Pseudanophthalmus blatchleyi Barr C
386. Pseudanophthalmus chthonius Krekeler C
387. Pseudanophthalmus emersoni Krekeler C
388. Pseudanophthalmus eremita (Horn) C
( = Anophthalmus eremita Horn) B
389. Pseudanophthalmus leonae Barr C
390. Pseudanophthalmus shilohensis Krekeler C
391. Pseudanophthalmus shilohensis boonensis Krekeler C
392. Pseudanophthalmus shilohensis mayfieldensis
Krekeler C
393. Pseudanophthalmus tenuis Horn IL,PU;C
( = Anophthalmus tenuis Horn) B
394. Pseudanophthalmus tenuis blatchleyi Barr C
395. Pseudanophthalmus tenuis jeanneli Krekeler C
396. Pseudanophthalmus tenuis morrisoni Jeannel C
397. Pseudanophthalmus tenuis strict icollis Jeannel C
398. Pseudanophthalmus youngi Krekeler C
399. Pseudanophthalmus youngi donaldsoni Krekeler C
400. Pterostichus adoxus (Say) PU;B
401. Pterostichus adst rictus Eschscholtz
( = Bothriopterus luczoti (Dejean)) PU
( = Pterostichus luczotii Dejean) B
402. Pterostichus bispiculatus Casey C
403. Pterostichus caudicalis Say B
( = Melanius caudicalis (Say)) PU
404. Pterostichus chalcites Say KU;C,S
( = Poecilus chalcites Say) PU
( = Pterostichus sayi Brulle) B
405. Pterostichus corvinus Dejean B
( = Melanius corvinus (Dejean)) PU
354 Indiana Academy of Science Vol. 94 (1985)
406. Pterostichus coracinus Newman B
( = Euferonia coracina (Newman)) PU
407. Pterostichus ebeninus Dejean B
( = Melanius ebeninus Dejean) PU
408. Pterostichus femoralis Kirby PU;B
409. Pterostichus haldemani (LeConte) IU;B,C
( = Lophoglossus haldemani (LeConte)) PU
410. Pterostichus honestus Say B
( = Gastrellarius honestus (Say)) PU
411. Pterostichus leconteianus Lutshnik IU,PU
( = Pterostichus erythropus Dejean) B
412. Pterostichus luctuosus Dejean B
( = Melanius luctuosus (Dejean)) PU
413. Pterostichus lucublandus Say B
( = Poecilus lucublandus Say) PU
( = Pterostichus convexicollis Say) KU
414. Pterostichus melanarius (Illiger) D3
415. Pterostichus moestus Say
( = Refonia moesta (Say)) PU
416. Pterostichus mutus Say B
( - Dysidius mutus (Say)) PU
417. Pterostichus obscurus Say B
( = Gasterosticta obscura (Say)) PU;D1
418. Pterostichus ohionis Csiki
( = Dysidius purpuratus LeConte) PU;D1
( = Pterostichus purpuratus LeConte) B
419. Pterostichus patruelis Dejean B
( = Micromaseus patruelis (Dejean)) PU
420. Pterostichus pensylvanicus LeConte B
421. Pterostichus relictus Newman B,C
( = Euferonia relicta (Newman)) PU
422. Pterostichus rostratus Newman C
423. Pterostichus scrutator (LeConte) B
( = Lophoglossus scrutator LeConte)
424. Pterostichus stygicus Say IU,KU,PU;B,C,S
( = Euferonia stygica (Say)) PU
425. Pterostichus tartaricus (Say) B,C
( = Lophoglossus tartaricus (Say)) PU
426. Pterostichus tristis Dejean PU
427. Scaphinotus andrewsi germari (Chaudoir)
( = Cychrus andrewsi germari Chaudoir) B
428. Scaphinotus elevatus flammeus Haldeman
( = Scaphinotus elevatus Fabricius) IL
( = Cychrus elevatus Fabricius) B
429. Scaphinotus tricarinatus Casey
( = Scaphinotus andrewsi tricarinatus Casey) PU
430. Scaphinotus unicolor (Fabricius) PU;C
43 1 . Scaphinotus unicolor hews Harris C
( = Cychrus unicolor Oliver) B
432. Scarites substriatus Haldeman KU,PU;B,M
433. Scarites subterraneus Fabricius IL,KU,PU;B,S
434. Schizogenius amphibius Haldeman B
Entomology 355
435. Schizogenius ferrugineus Putzeys PU;B,D1
436. Schizogenius lineolatus Say PU;B,D1
437. Selenophorus ellipticus Dejean PU;B
438. Selenophorus gagatinus Dejean KU,PU;B,C,D1,S
439. Selenophorus granarius Dejean DC
440. Selenophorus opalinus LeConte PU;B
44 1 . Selenophorus pedicularius Dej ean PU ; B , C , D 1
442. Sericoda bogemanni (Gyllenhal)
( = Platynus bogemanni (Gyllenhal)) PU;B
( = Agonum bogemanni Gyllenhal) KU
443. Sphaeroderus lecontei Dejean IL,IU,KU,PU;S
( = Cychrus lecontei Dejean) B
444. Sphaeroderus stenostomus Weber C
( = Sphaeroderus stenostomus indianae Blatchley) PU
( = Cychrus stenostomus indiane Leng) B
445. Stenocrepis cupreus (Chaudoir) C
( = Oodes cupreus Chaudoir) B
446. Stenocrepis quatuordecimstriata (Chaudoir) C
( = Oodes 14-striatus Chaudoir) B
447. Stenolophus carbonarius (Dejean) KU,PU;B,S
448. Stenolophus comma Fabricius IL,IU,PU
449. Stenolophus conjunctus (Say) PU;B
450. Stenolophus dissimilis Dejean B,C
451. Stenolophus fuliginosus Dejean PU;B
452. Stenolophus fuscatus Dejean PU
( = Stenolophus plebejus fuscatus Dejean) B
453. Stenolophus lecontei Chaudoir
( = Agonoderus lecontei Chaudoir) PU
( = Agonoderus pallipes "(Fabricius)") IU,PU,B
454. Stenolophus lineola (Fabricius)
( = Agonoderus lineola (Fabricius)) PU;B
455. Stenolophus ochropezus Dejean IU,PU;B
456. Stenolophus plebejus Dejean B
457. Stenolophus rotundicollis Haldeman
( = Stenolophus scitulus Casey) PU
458. Synuchus impunctatus (Say) IU
( = Calathus impunctatus Say) B
( = Pristodactyla impunctata (Say)) PU;D1
459. Tachyta nana inornata Say C
( = Tachyta nana (Gyllenhal)) PU
( = Tachys nanus (Gyllenhal)) B
460. Tetragonoderus fasciatus Haldeman IU;B,C
461. Trechus chalybeus Dejean B
462. Trichotichnus dichrous Dejean KU;C,S
( = Harpalus dichrous Dejean) PU;B
463. Trichotichnus vulpeculus (Say)
( = Harpalus vulpeculus Say) PU;B
464. Xestonotus lugubris (Dejean) PU
( = Anisodactylus lugubris Dejean) B
465. Zuphium americanum Dejean D3
The following Indiana "records" are highly doubtful (G. Ball, personal
cor-
356 Indiana Academy of Science Vol. 94 (1985)
respondence): Bembidion oblongulum (Mannerheim), PU, is probably B. wingatei Bland;
Pinacodera russata Newman, C, is now Cymindis complanata Dejean which does not
likely occur farther north than Alabama; and Scaphinotus elevatus tenebricosus Roeschke,
PU.
Literature Cited
1. Blatchley, W.S. 1910. An Illustrated Descriptive Catalogue of the Coleoptera or
Beetles Known to Occur in Indiana. The Nature Publishing Co., Indianapolis,
Indiana. 1385. p.
2. Downie, N. M. 1956. Records of Indiana Coleoptera, I. Proceedings of the Indiana
Academy of Science. 66:115-124.
3. Downie, N. M. 1958. Records of Indiana Coleoptera, II. Proceedings of the Indiana
Academy of Science. 66:115-124.
4. Downie, N. M. and C. E. White. 1967. Records of Indiana Coleoptera, III. Pro
ceedings of the Indiana Academy of Science. 76: 308-316.
5. Erwin, T., D. R. Whitehead and G. E. Ball. 1977. Family 4. Carabidae, The Ground
Beetles. In: Checklist of the Beetles of North and Central America and the West
Indies. Flora and Fauna Publications, Gainesville, Florida. 68 p.
6. Montgomery, B. E. and J. M. Amos. 1941. Contributions to a list of the Coleoptera
of the Clark County State Forest. Proceedings of the Indiana Academy of Science.
50:251-258.
7. Munsee, Jack R. 1966. The Ecology of Ants of Stripmine Spoil Banks. Ph.D. Disser-
tation. Purdue University, West Lafayette, Indiana 243 p.
8. Schrock, John R. 1983. The Succession of Insects on Unreclaimed Coal Strip Mine
Spoil Banks in Indiana. Ph.D. Dissertation, University of Kansas, Lawrence, Kansas.
207 p.
9. Thiele, H.-U. 1977. Carabid Beetles in Their Environment. Springer- Verlag, Berlin.
369 p.
A Checklist of the Aquatic Coleoptera of Indiana
Charles E. White, Frank N. Young, and N.M. Downie
Department of Biology
Indiana University, Bloomington, Indiana 47405
The only readily available listing of the aquatic Coleoptera of Indiana is that
of Blatchley in his Coleoptera of Indiana (1910). A number of species and genera
have been added and changes in nomenclature have occurred. The late Charles E.
White initiated the present list some years ago, but it has remained unpublished. The
families of Coleoptera considered to be truly aquatic are as follows: Gyrinidae,
Dytiscidae, Haliplidae, Noteridae, Helophoridae, Hydrochidae, Hydraenidae,
Hydrophilidae (except for subfamily Sphaeridinae), Psephenidae, Dryopidae, and
Elmidae. However, others such as the Heteroceridae, Limnichidae, and some groups
of Chrysomelidae and Curculionidae are associated with aquatic situations. The
Helodidae and Ptilodactylidae have aquatic larvae, but have not been included.
$4 % ♦ ♦
It is now going on 75 years since W.S. Blatchley's "An Illustrated Descriptive
Catalogue of the Coleoptera or Beetles (Exclusive of the Rhynchophora) Known to
Occur in Indiana; With Bibliography and Descriptions of New Species" appeared as
Bulletin No. 1 of the Indiana Department of Geology and Natural Resources in 1910.
This 1,386-page book with 590 figures and one map is still one of the two comprehen-
sive works on the Coleoptera, or beetles covering North America, the other being the
five volume series of M.H. Hatch, the Beetles of the Pacific Northwest.
In the aquatic families, Blatchley in 1910 listed the following: Haliplidae, 2 genera,
11 species; Dytiscidae (including Noteridae) 25 genera, 96 species; Gyrinidae, 3 genera,
14 species; Hydrophilidae (including Helophoridae, Hydrochidae, and Hydraenidae),
26 genera, 69 species; Parnidae (including Psephenidae, Dryopidae-Parnidae, and
Elmidae), 7 genera, 12 species. Some of the genera and species were presumptive and
not represented by actual Indiana specimens. The following list enlarges Blatchley's
list and attempts to bring the nomenclature up to date.
The voucher material for the following list is largely in the Purdue University
Laboratory of Insect Diversity collection or in the collections of the authors. The col-
lection of the late Charles E. White is in the Florida State Collection of Arthropods
in Gainesville, Florida.
HALIPLIDAE
Haliplus borealis LeC.
Haliplus cribrarius LeC.
Haliplus fasciatus Aube
Haliplus immaculicollis Harris
( = ruficollis/Blatchley)
Haliplus longulus LeC.
Haliplus ohioensis Wallis
(probably = lewisii/Blatchley)
Haliplus subguttatus LeC.
Haliplus pantherinus Aube
DYTISCIDAE
Laccophilus maculosus maculosus Say
357
358 Indiana Academy of Science Vol. 94 (1985)
Laccophilus proximum proximus Say
Laccophilus undatus Aube
Laccophilus fasciatus rufus (Melsh.)
Hydrovatus pustulatus pustulatus (Melsh.)
Hydrovatus indianensis Blatchley
Demopachria convexa (Aube)
Uvarus granarius (Aube)
Uvarus lacustris (Say)
Uvarus suburbanus (Fall)
Liodessus affinis affinis (Say)
Liodessus fuscatus (Crotch)
Liodessus flaviocollis (LeC.)
Bidessonotus inconspicuus (LeC.)
(= Bidessus pulicarius/Blatchley)
Celina bubbelli Young
Celina imitatrix Young
Peltodytes 12-punctatus (Say)
Peltodytes edentulus (LeC.)
Peltodytes lengi Roberts
Peltodytes muticus (LeC.)
Peltodytes sexmaculatus Roberts
Peltodytes litoralis Matheson
Peltodytes pedunculatus (Blatchley)
Peltodytes dunavani Young
Celina bubbelli Young
( = angustatus/Blatchley)
Celina imitatrix Young
Hygrotus sayi J. B-Browne
( = punctatus//Say)
Hygrotus turbidus (LeC.)
Hygrotus dispar (LeC.)
Hygrotus impressopunctatus Schall.
Hygrotus acaroides (LeC.)
Hygrotus laccophilinus (LeC.)
Hygrotus dissimilis (Harris)
Hygrotus nubilus (LeC.)
Deronectus rotandatus (LeC.)
( = depressus Fabr. of authors)
Deronectes griseostriatus (DeG.)
Falloporus triangularis (Fall) (Monroe Co.)
Hydroporus (Heterosternuta)
Hydroporus laetus Leech
Entomology 359
Hydroporus ohionis Fall
Hydroporus pulcher LeC.
Hydroporus wickhami Zaitzev
(= concinnus/Zaitzev Lee.)
Hydroporus (Neoporus)
Hydroporus spurius LeC.
Hydroporus venustus LeC.
Hydroporus undulatus Say
Hydroporus consimilis LeC.
Hydroporus mixtus LeC.
Hydroporus dimidiatus G. & H.
Hydroporus mellitus LeC.
Hydroporus vittatipennis G. & H.
Hydroporus striatopunctatus Melsh.
Hydroporus shermani Fall
Hydroporus sericeus LeC.
Hydroporus hybridus Aube
Hydroporus clypealis Sharp
Hydroporus semiflavus Fall (Monroe Co.)
Hydroporus solitarius Sharp
Hydroporus vitiosus LeC.
Hydroporus blanchardi Sherman
Hydroporus psammodytes Young
Hydroporus aequus Fall
Hydroporus filiolus Fall
Hydroporus pagus Fall
Hydroporus oblitus Aube (sensu H.C. Fall)
Hydroporus despectus Fall (Monroe Co.)
Hydroporus pseudovilis Young
(= Hydroporus vilis/Blatchley)
Hydroporus (Hydroporus s. str.)
Hydroporus dichrous Melsh.
Hydroporus brevicornis Fall
Hydroporus americanus Aube
Hydroporus melsheimeri Fall
Hydroporus dentellus Fall
Hydroporus notabilis LeC.
Hydroporus rufilabris Sharp
Hydroporus tenebrosus LeC.
Hydroporus tristis Payk.
Hydroporus signatus youngi Gordon
Hydroporus niger Say
Hydroporus despectus rusticus Sharp
Hydroporus striola Gyll.
Laccornis
(= Agaporus)
Laccornis conodeus LeC.
Laccornis difformis LeC.
Laccornis deltoids Fall
Agabus (Gaurodytes)
360 Indiana Academy of Science Vol. 94 (1985)
Agabus seriatus seriatus (Say)
Agabus semivittatus (LeC.)
Agabus aeruginosus Aube
Agabus falli Guignot
(= sharpi Fall)
Agabus disintegratus (Crotch)
( = taeniolatus//Blatchley)
Agabus scapularis Mann.
(= anthracinus Mann.)
Agabus gagetes Aube
Agabus stagninus (Say)
Agabus punctatus Melsh.
Agabus punctulatus Aube
(= aeneolus Crotch)
Agabus semipunctatus (Kirby)
Agabus congener Thunb.
Agabus ambiguus (Say)
(= reticulatus Aube)
Agabus phaeopterus (Kirby)
Agabus erichsoni G. & H.
Agabus obtusatus (Say)
Agabus confusus (Blatchley) (S. Indiana)
(= Rhantus confusus Blatchley)
Agabus leptapsis (LeC.)
Agabus erythropterus (Say)
Agabus (Eriglenus)
Agabus bifarius (Kirby)
Ilybius biguttulus (Germ.)
Ilybius ignarus LeC.
Ilybius fraterculus LeC.
Ilybius oblitus Sharp
Ilybius angustior (Gyll.)
Ilybius confusus Aube?
Agabetes acunductus (Harris)
Matus bicarinatus (Say)
Matus ovatus ovatus Leech
Copelatus glyphicus (Say)
Copelatus chevrolati renovatus Guignot
Coptotomus (Fabr.)
Coptotomus longulus LeC.
Coptotomus liticus Hilsenhoff
Coptotomus lenticus Hilsenhoff
Neoscutopterus angustus (LeC).
Rhantus sinuatus LeC.
Rhantus tostus LeC.
Entomology 361
Rhantus zimmermanni Wallis
( = bistriatus//Blatchley)
Colymbetes sculptilis (Harris)
Dytiscus fasciventris Say
Dytiscus verticalis Say
Dytiscus hyridus Aube
Dytiscus harrisii Kirby
Hydaticus modestus Sharp
( = stagnalis//Blatchley)
Hydaticus piceus LeC.
Acilius semisulcatus Aube
Acilius mediatus (Say)
Acilius fraternus (Harris)
Acilius sylvanus Hilsenhof
Thermonectus nigricollis ornaticollis (Aube)
Thermonectus basillaris basillaris (Harris)
Graphoderes liberus (Say)
Graphoderes fasciatocollis (Harris)
Graphoderes modestus Sharp
Cybister fimbriolatus fimbriolatus (Say)
NOTERIDAE
Suphisellus puncticollis Crotch
Suphisellus bicolor bicolor (Say)
Suphisellus bicolor punctipennis (Sharp)
Hydrocanthus atricolor (Say)
( = texanus Sharp)
GYRINIDAE
Dineutus ciliatus (Forsb.) (Owen Co.)
( = vittatus//Blatchley)
Dineutus nigrior Roberts
Dineutus discolor Aube
Dineutus emarginatus Say
Dineutus horni Roberts
Dineutus assimilis Kirby
( = americana Say)
Gyrinus minutus (Fabr.)
Gyrinus ventralis Kirby
Gyrinus aeneolus LeC.
Gyrinus affinis Aube
Gyrinus analis Say
Gyrinus lugens LeC.
362 Indiana Academy of Science Vol. 94 (1985)
Gyrinus plicifer LeC.
Gyrinus fraternus Coup.
Gyrinus aquiris Lee.
Gyrinus dichrous LeC.
Gyrinus piceolus Blatchley
Gyrinus borealis Aube
Gyrinus pectoralis LeC.
Gyretes compressus LeC. (Owen & Greene Co.)
HELOPHORIDAE
Helophorus oblongus LeC.
Helophorus nitidulus LeC.
Helophorus linearis LeC.
Helophorus lacustris LeC.
Helophorus lineatus Say
Helophorus tuberculatus Gyll.
Helophorus ventralis Mots.
( = obsoletesulcatus Mots.)
HYDROCHIDAE
Hydrochus scabratus (Mulsant)
Hydrochus inaequalis Lee.
Hydrochus subcupreus Randall
Hydrochus rufipes Melsh.
Hydrochus setosus Leech
Hydrochus ouelleti Leech
Hydrochus squamifer LeC.
Hydrochus excavatus LeC.
Hydrochus granulatus Blatchley
Hydrochus currani Brown
Hydrochus brevitarsis Knisch
Hydrochus undulatus Hellman
Hydrochus neosquamifer Hellman
LIMNEBIIDAE
(= HYDRAENIDAE)
Ochthebius foveicollis LeC.
Ochthebius putamensis Blatchley
Ochthebius cribricollis LeC.
Gymnocthebius nitidus (LeC.)
Hydraena pensylvanica Kies.
Hydraena punctata LeC.
Hydraena angulicollis Notm.
Hydraena quadricurvipes Perkins (Brown Co.)
Hydraena ancylis Perkins (Monroe Co.)
Limnebius discolor (Casey) (Monroe Co.)
Entomology 363
HYDROPHILIDAE
Hydrophilus triangularis Say
Dibolocelus ovatus (G. & H.)
Tropisternus lateralis nimbatus (Say)
Tropisternus glaber (Herbst)
Tropisternus blatchleyi modestus D'Orch.
Tropisternus collaris striolatus (LeC.)
Tropisternus mixtus (LeC.)
Tropisternus natator D'Orch.
Tropisternus sublaevis LeC?
Hydrochara obtusata (Say) (n. Indiana)
Hydrochara soror Smetana
Hydrochara spangleri Smetana (Monroe Co.)
Hydrochara leechi Smetana
Chaetarthria pallida (LeC.)
Chaetarthria atra (LeC.) (Monroe)
Berosus pugnax LeC.
Berosus pantherinus LeC.
Berosus aculeatus LeC.
Berosus infuscatus LeC. (Posey, Monroe, Crawford Cos.)
Berosus ordinatus LeC.
( = pennsylvanica Knisch)
Berosus exiguus Say (Monroe Co.)
Berosus peregrinus Herbst.
Berosus fraternus LeC. (Posey, Monroe, Crawford Cos.)
Berosus striatus (Say)
Derallus altus LeC. (Posey Co.)
Laccobius agilis Rand.
Laccobius punctatus Melsh.
Hydrobius fuscipes (Linn.)
Hydrobius globosus (Say)
Hydrobius tumidus LeC.
Hydrobius melaneum Ger.
Scherchopsis tesselatus (Ziegler) (Tippecanoe, Porter Cos.)
(= Hydrobius tesselatus/VBlatchley)
Paracymus (Creniphilus)
Paracymus despectus (LeC.)
Paracymus subcupreus (Say)
Paracymus confusus Wooldridge (Posey Co.)
Paracymus digestus (LeC.)
Crenitulus (Creniphilus)
364 Indiana Academy of Science Vol. 94 (1985)
Crenitulus suturalis (LeC.) (Monroe Co.)
(= Creniphilus suturalis//Blatchley)
Ancaena limbata (Fabr.)
( = infuscatus//Blatchley)
Crenitis longulus (Fall) (Monroe Co.)
Enochrus
( = Philhydrus/VBlatchley)
Enochrus pygmaeus nebulosus (Say)
Enochrus diffusus (LeC.) (Crawford Co.)
Enochrus ochraceus (Melsh.)
Enochrus consortus Green (Posey Co.)
Enochrus horni (Leech)
( = hamiltoni//Blatchley
Enochrus cinctus (Say)
Enochrus perplexus (LeC.)
Helochares maculicollis Mulsant
Helocombus bifidus (LeC.)
Cymbiodyta lasustris LeC.
Cymbiodyta vindicata Fall
Cymbiodyta blanchardi Horn
Cymbiodyta chamberlaini Smetana
Cymbiodyta semistriatus (Zimm.)
(= fimbriata Melsh.)
Sphaeridium scarabaoides (Linn.)
Sphaeridium bipustulatum (Fabr.)
Sphaeridium lunutatum (Fabr.)
Phaenonotus exstriatum (Say)
( = estriatum//Blatchley)
Genyon navicularis (Zimm.)
Cercyon analis (Payk.)
Cercyon pubescens LeC.
Cercyon pygmaeus (Iliger)
Cercyon nigriceps (Marsh.)
Cercyon quisquilius (Linn.)
Cercyon maculatus (Melsh.)
Cercyon convexiusculus Steph.
( = lugubris/VBlatchley)
Cercyon tristis (Illiger)
Cercyon haemorrhoidalis (Fabr.)
Cercyon terminatus (Marsh.)
( = melanocephala//Blatchley
Cercyon unipunctatus (Linn.)
Entomology 365
Cercyon praetextatus (Say)
Cercyon indistinctus Horm
Cercyon navicularis Zimm.
Cercyon atricapillus (Marsh.)
Cercyon minusculus Melsh.
Cercyon roseni Knish
Cercyon pygmaeus (111.)
Cercyon connivens Fall
Cercyon herceus Smetana
Cercyon erraticus Smetana
Cercyon mendax Smetana
Cercyon assecla Smetana
Cercyon occallatus (Say)
Cercyon lateralis (Marsh.)
Cryptopleurum minutum (Fabr.)
Cryptopleurum subtile Sharp
Cryptopleurum americanum Horn?
Pemelus costatus (LeC.)?
DRYOPIDAE
Helichus basalis LeC. (Parke County)
Helichus fastigiatus (Say)
Helichus lithophilus (Germ.)
Helichus striatus LeC. (Parke and Tippecanoe Co.)
ELMIDAE
Stenelmis crenata (Say)
Stenelmis decorata Sanderson
Stenelmis maerkelii Motsch.
(= sulcatus Blatchley)
Stenelmis musgravei Sanderson
Stenelmis quadrimaculatus Horn
Stenelmis sandersoni Musgrave
Stenelmis sexlineata Sanderson
Stenelmis vittipennis Zimmerman
Macronychus glabrata Say
HETEROCERIDAE
Centuriatus
( = Heterocerus//Blatchley)
Centuriatus auromicans (Kies.)
Lanternarius
(= Heterocerus Blatchley)
Lanternarius brunneus (Melsh.)
Lanternarius mollinus (Kies.)
Lanternarius parrotus Pacheo
366 Indiana Academy of Science Vol. 94 (1985)
Neoheterocerus
( = Heterocerus//Blatchley)
Neoheterocerus angustatus (Chev.)
Neoheterocerus pallidus (Say)
(= ventralis Melsh.)
Neoheterocerus sandersoni Pacheco
Dampfius
( = Heterocerus/VBlatchley)
Dampfius collaris (Kies).
Dampfius undatus (Melsh.)
Tropicus
( = Heterocerus//Blatchley)
Tropicus pusillus (Say)
LIMNICHIDAE ( = Byrrihidae//Blatchley, ex parte)
Limnichus obscurus (LeC.)
Limnichus ovatus (LeC.)
Limnichus nitidulus (LeC.)
Limnichus punctatus (LeC.)
Latrochus laticeps Csy.
CHRYSOMELIDAE
Macroplea nigricornis (Kby.)
Donacia aequalis Say
Donacia biimpressa Melsh
Donacia cincticornis Newm.
Donacia distincta LeC.
Donacia fulgens Lee.
Donacia hirticollis Kby.
Donacia hypoleuca Lac.
Donacia megacornis Blatch.
Donacia palmata Oliv.
Donacia parvidens Schffr.
Donacia piscatrix Lac.
Donacia porosicollis Lac.
Donacia proxima Kby.
Donacia pubescens LeC.
Donacia pubicollis Suffr.
Donacia quadricollis Say
(= curticollis Knab)
Donacia rufescens Lac.
Donacia rugosa LeC.
Donacia subtilis Kunze
Donacia tuberculifrons Schffr.
Plateumaris
( = Donacia//Blatchley)
Entomology 367
Plateumaris diversa (Schffr.)
Plateumaris emarginata (Kby.)
Plateumaris falvipes (Kby.)
Plateumaris metallica (Ahr.)
Plateumaris sulciocollis (Lac.)
Sominella
(= Donacia//Blatchley)
Sominella harrisi (LeC.)
CURCULIONIDAE (in part, aquatic weevils)
Tanysphyrus lemnae (Fab.)
Bagous americanus LeC.
Bagous atratus Blatch.
Bagous bituberosa LeC.
Bagous lengi Tanner
Bagous magister LeC.
Bagous nebulosus LeC.
Bagous obliquus LeC.
Bagous planatus LeC.
Bagous pusillus LeC.
Bagous restrictus LeC.
Bagous tanneri O'Brien
Bagous transversus LeC.
Lissorhoptrus oryzphilus Kusch.
Lissorhoptrus simplex (Say)
Brachybamus electus Germ.
Notiodes (Endalus/VBlatchley)
Notiodes limatulus (Gyll.)
Notiodes ovalis (LeC.)
Onychylis nigrirostris (Boh.)
Stenopelmus rufinasus Gyll.
Lixellus lutulentus (Boh.)
(= ASnchodemus angustus/VBlatchley)
Listronotus (In part = Hyperodes of Blatchley)
Listronotus appendiculatus (Boh.)
Listronotus callosus LeC.
Listronotus caudatus (Say)
Listronotus debilis (Blatch.)
Listronotus delumbis (Gyll.)
Listronotus dietzi O'Brien
Listronotus dorsalis (Dietz)
Listronotus echinatus (Dietz)
368 Indiana Academy of Science Vol. 94 (1985)
Listronotus frontalis LeC.
Listronotus grypidoides (Dietz)
Listronotus humilis (Gyll.)
Listronotus maculicollis (Kby.)
Listronotus montanus (Dietz)
Listronotus nebulosus LeC.
Listronotus porcellus (Say)
Listronotus poseyensis (Blatch.)
Listronotus sordidus (Gyll.)
Listronotus sparsus (Say)
Listronotus squamiger (Say)
Listronotus tuberosus LeC.
Catalogs, reviews, revisions, and other papers
since Blatchley (1910) useful in classification
Balfour-Browne, Jack. 1947. A revision of the genus Bidessonotus Regimbart
(Coleoptera: Dytiscidae). Trans. Royal Ent. Soc. London 98(9):425-448, 12 figs.
Blatchley, W.S. and C.W. Heng. 1916. Rhychrphora or Weevils of North Eastern
America. Indianapolis: Nortmas Publishing Co. L82 pp.
Brinck, Per. 1945. Nomenklatorische Studien uber Dytischiden. III. Die Klassifikation
der Cybisterinen. Lunds Universitets Arsskrift (N.F. Avd. 2) 41(4): 1-20, 1 fig.
(= Handlingar Kungl. Fysiografiska Sallskapets (N.F.) 56(4):l-20, 1 fig.).
Brown, Harley P. 19 A catalog of the Coleoptera of America north of Mexico
Family Elmidae. U.S. Dept. Agric, Agric. Handb. No. 529-550, 23 pp. Ibid.
Family: Dryopidae, 8 pp.
Darlington, Jr., P.J. 1936. A list of the West Indian Dryopidae (Coleoptera) with
a new genus and eight new species including one from Columbia. Psyche
43(2-3):65-83, 1 pi.
D'Orchymont, A. 1921. Le genre Tropisternus I (Col. Hydrophilidae). Ann. Soc. Ent.
Belgique 61:349-374.
. 1922. Le genre Tropisternus II (Col. Hydrophilidae). Ann. Soc. Ent. Belgique
62:11-48, 4 figs.
Fall, H.C. 1919. The North American species of Coelambus. John D. Sherman, Jr.,
Mt. Vernon, N.Y., 20 pp.
1922. A review of the North American species of Agabus together with a descrip-
tion of a new genus and species of the tribe Agabini. John D. Sherman, Jr.,
Mt. Vernon, N.Y., 36 pp.
1923. A revision of the North American species of Hydroporus and Agaporus.
John D. Sherman, Jr., Mt. Vernon, N.Y. 219 pp.
Leng, Charles W. 1920. Catalog of the coleoptera of America north of Mexico. John
D. Sherman, R., Mt. Vernon, N.Y., x/ + 470 pp. (with supplements).
Marx, Edward, J.K. 1957. Review of subgenus Donacia in Western Hemisphere. Bull.
Mus. Nat. Hist. Vol. 112, pp. 195-278.
Matheson, Robert. 1912. The Haliplidae of North America, north of Mexico. J. N.U.
Ent. Soc. 20:156-193, 6 pi., 2 figs.
Matta, James F., G. William Wolfe. 1981. A revision of the subgenus Heterosternuta
Strand of Hydroporus Clairville (Coleoptera: Dytiscidae). Pan-Pacific Ent.
57:176-218, 80 figs.
Musgrave, Paul N. 1935. A synopsis of the genus Helichus Erichson in the United
States and Canada with descriptions of new species (Coleoptera: Dryopidae). Proc.
Ent. Soc. Wash. 37:137-145, 1 pi.
Entomology 369
O'Brien, C.W. and G.J. Wibner. 1982. Annotated checklist of the weevils of North
America. Mem. No. 34 Amer. Entom. Inst. Ann Arbor, MI.
Pacheco, Francisco. 1978. A catalog of the Coleoptera of America north of Mexico.
Family Heteroceridae. Agriculture Handbook No. 529-47. Wash. D.C.: U.S. Dept.
Agriculture, X and 8 pp.
Roberts, Chris H. 1913. Critical notes on the species of Haliplidae of America north
of Mexico with descriptions of new species. J. N.Y. Ent. Soc. 21:91-123.
Sanderson, Milton W. 1938. A monographic revision of the North American species
of Stenelmis (Dryopidae: Coleoptera). Univ. Kansas Sci. Bull. 25:635-717, 2 pi.
Smetana, Ales. 1974. Revision of the genus Cymbiodyta Bed. (Coleoptera:
Hydrophilidae). Mem. Ent. Soc. Can. No. 93, iv + 112 pp., 147 figs.
. 1978. Revision of the subfamily Sphaeridiinae of America north of Mexico (Col-
eoptera: Hydrophilidae). Mem. Ent. Soc. Canada No. 105, 292 pp., 336 figs.
(colored frontispiece).
. 1980. Revision of the genus Hydrochara Berth. (Coleoptera: Hydrophilidae).
Mem. Ent. Soc. Canada No. Ill, iv+100 pp., 77 figs, (colored frontispiece)
Wallis, J.B. 1933. Revision of the North American species (north of Mexico) of the
genus Haliplus, Latreille. Trans. Royal Can. Inst. 19(1): 1-76, 38 figs.
1939. The genus Graphoderus Aube in North America (north of Mexico). Can.
Eng. 71:128-130.
. 1939. The genus Ilybius Er. in North America (Coleoptera: Dytiscidae). Can.
Ent. 71:192-199.
Winters, Fred C. 1926. Notes on the Hydrobiini (Coleoptera: Hydrophilidae) of Boreal
America. Pan-Pacific Ent. 3:49-58.
1927. Key to the subtribe Helocharae Orchym. (Coleoptera: Hydrophilidae) of
Boreal America. Pan Pacific Ent. 4:19-29.
Wolfe, G. William and James F. Matta. 1981. Notes on nomenclature and classifica-
tion of Hydroporus subgenera with the descriptions of a new genus of Hydroporini
(Coleoptera: Dytiscidae). Pan-Pacific Ent. 57:149-175, 36 figs.
Young, Frank N. 1953. The types of Hydradephaga in the W.S. Blatchley collection,
with generic reassignments and synonymies (Coleoptera: Noteridae, Dytiscidae,
Grinidae, Haliplidae). Can Ent. 85(3): 113-1 19.
. 1961. Pseudosibling species in the genus Peltodytes (Coleoptera: Haliplidae).
Ann. Ent. Soc. Amer. 54:214-222, 12 figs.
. 1963. The Nearctic species Copelatus Erichson (Coleoptera: Dytiscidae). Quart.
J. Florida Acad. Sci. 26:56-77, 11 figs.
. 1957. A key to the genera of American Bidessine water beetles with descriptions
of three new genera (Coleoptera: Dytiscidae, Hydroporinae). Coleopt. Bull.
21(3):75-84.
1978. A new predaceous water beetle from the eastern United States (Coleoptera:
Dytiscidae). Coleopt. Bull. 32:189-191, 5 figs. {Hydroporus psammodytes) .
1979. A key to the Nearctic species of Celina with descriptions of new species
(Coleoptera: Dytiscidae). J. Kansas Ent. Soc. 52:820-830, 9 figs. {Celina hubbelli
and imitratrix).
1979. Water beetles of the genus Suphisellus Crotch in the Americas north of
Colombia (Coleoptera: Noteridae). Southw. Nat. 24:409-429, 21 figs.
Zimmerman, James R. 1970. A taxonomic revision of the aquatic beetle genus Lac-
cophilus (Dytiscidae) of North America. Mem. Amer. Ent. Soc, No. 26:1-275,
332 figs.
Zimmerman, James R. and Robert L. Smith, 1975. The genus Rhantus (Coleoptera:
Dytiscidae) in North America. Part I. General account of the species. Trans. Amer.
Ent. Soc. 101:33-123, 102 figs.
ENVIRONMENTAL QUALITY
Chairperson: William Beranek
Indianapolis Center for Advanced Research
120 E. 38th Street
P.O. Box 647
Indianapolis, Indiana 46223
(317)264-2827
Chairperson-Elect Horst Siewert
Department of Natural Resources
Ball State University
Muncie, Indiana 47306
(317)285-5790
ABSTRACTS
The Determination of the Removal Rate of Specific Chemicals by the Indianapolis
Wastewater Treatment System. William Beranek, Jr. and Elizabeth DuSold,
Indianapolis Center for Advanced Research, Inc., 611 North Capitol Avenue, In-
dianapolis, Indiana 46206. The rate of removal of toxic chemicals from a municipal
wastewater treatment facility is a critical value for policy makers determining the in-
dustrial discharge concentrations into a sewer system.
Increased attention to the removal rate now is occurring because of its regulatory
use in adjusting the national categorical industrial discharge limits to the special condi-
tions present in specific municipal wastewater treatment facilities.
Due to the constantly changing heterogeneous chemical composition of the in-
fluent of the facility and to the changing retention times of the flow of material through
the facility, reliable measurement of the removal rates are very difficult.
This paper reviews the removal rate measurements at the Indianapolis Advanced
Wastewater Treatment facilities and discusses the significance of the measurements.
A Superfund Risk Assessment in Indiana: A Case Study of the Columbia City Site.
William Beranek, Jr., Elizabeth DuSold, John Merrill and Marten St. Clair,
Indianapolis Center for Advanced Research, Inc., Beranek Associates, Inc., and Califor-
nia Institute of Technology. The Wayne Waste Oil site in Columbia City, Indiana
is currently on the National Priority List of the U.S. Environmental Protection Agen-
cy of sites requiring a risk assessment under the Comprehensive Emergency Response,
Liability and Compensation Act. This is due to the presence of chemicals close to
an aquifer used as drinking water by a community of 5,000 people.
The methods and results of the risk assessment which were performed between
April 1983 and August 1984 are presented. The methods include chemical sampling
and measurement, groundwater flow measurement, geological strata evaluation and
draw-down pumping testing.
The Ratio of PM-10 to TSP in the Atmosphere. William Beranek, Jr. and David
Jordan, Indianapolis Center for Advanced Research, Inc., 611 North Capitol Avenue,
Indianapolis, Indiana 46206. The U.S. Environmental Protection Agency is pro-
posing to change the indicator pollutants of the National Ambient Air Quality stan-
dard from total suspended particulates (TSP) to the fraction of particulate matter smaller
371
372 Indiana Academy of Science Vol. 94 (1985)
than ten microns in aerodynamic diameter (PM-10). Marion County currently has a
non-attainment status for TSP, although 1983 readings showed no primary violations.
In order to estimate the ambient air quality levels in Marion County of this new
standard, the Indianapolis Air Pollution Control Division, with the support of
Indianapolis corporations through the Indianapolis PM-10 Task Force, since January
1983 has been monitoring PM-10 at four locations.
This paper reviews the results of the study and discusses the implication in the
context of the proposed changes in the federal regulations on particulates.
Evaporation Rates of Organic Liquids at Various Wind Speeds and Temperatures.
Howard E. Dunn, Benjamin P. Miller, Charles B. Macer and Michael E.
Klausmeier, Departments of Chemistry and Physics, Indiana State University Evansville,
Evansville, Indiana 47712. In previous papers the authors have investigated com-
puter model predictions of downwind concentrations of toxic gases from continuous
sources and from instantaneous releases. An additional case of importance is the calcula-
tion of a region to be evacuated resulting from the evaporation of a toxic liquid spill.
A review of the literature revealed minimal information pertaining to the calculation
of evaporation rates of liquids at various wind speeds and temperatures.
A wind tunnel was designed and constructed for the purpose of measuring the
desired evaporation rates. The weight loss from an evaporation dish can be measured
at intervals for wind speeds of three to twenty miles per hour at commonly encountered
atmospheric temperatures above freezing.
Results obtained have been compared with similar results reported by other in-
vestigators and with empirical correlations reported for evaporation rates. Completed
results from this evaporation rate study will be incorporated in a computer model to
predict an evacuation zone for protection of the population from a spill of any one
of several dangerous volatile chemicals.
Herbicide (Alachlor, Atrazine, Linuron and Paraquat) Residues in Deer
Mice Inhabiting Conventional and Minimum Tillage Row-crop Fields
Denise Benson, Claude D. Baker, and Bill J. Forsyth
Department of Biology
Indiana University Southeast
New Albany, Indiana 47150
and
John S. Castrale
Indiana Division of Fish and Wildlife
Mitchell, Indiana 47446
The acreage of cropland in the United States incorporating conservation tillage
methods has increased steadily from 14% in 1973 to over 24% in 1982 (12). During
1982, reduced tillage practices were utilized on 34% of Indiana's 13 million acres of
cropland (10). Based primarily on economic advantages and improved technology, it
is predicted that conservation tillage in some form will be used on 60% of the nation's
cropland by the year 2010 (12).
Although a variety of practices qualify as reduced or conservation tillage, all have
in common less disturbance to the soil with greater amounts of crop residues left on
the soil surface. In most situations, chemical control of weeds substitutes for mechanical
tillage. With minimum tillage (also referred to as no-till and zero tillage), weed control
is solely by herbicides, and chemical applications and planting can be combined in
the same operation.
The environmental consequences of this shift in agricultural practice are just begin-
ning to be explored. With minimum tillage, soils are less prone to compaction and
soil loss on sloped land can be reduced by as much as 90% (12). Besides maintaining
soil productivity, reduced soil erosion should result in decreased siltation of waterways
and decreased air-borne soil particles. Benefits to wildlife from conservation tillage
have also been envisioned and recent studies bear this out (11, 23, 25).
A potentially detrimental impact of conservation tillage practices is the greater
use of chemical pesticides. With reduced tillage, more vegetation residue remains on
the soil surface interfering with herbicide incorporation. Thus, chemical application
rates may need to be increased to maintain their effectiveness. Contact herbicides,
such as paraquat and glyphosate, are unique to minimum tillage operations and should
be the focus of research attention.
The purpose of this study was to determine if 4 commonly used herbicides or
their metabolites could be detected in deer mice (Peromyscus maniculatus), the most
common inhabitant of cultivated cropland in Indiana and much of the Midwest (17).
A secondary objective was to determine if these herbicides may be having detrimental
physiological effects on this rodent under natural field conditions. Alachlor, atrazine,
and linuron were chosen because they are used extensively in the production of corn
and soybeans, both in conventional and minimum tillage situations. Paraquat was chosen
because it is the major contact herbicide used in zero tillage practices. Although these
chemicals have been tested extensively on laboratory rodents and birds, few studies
have examined the impacts of agricultural chemicals on wildlife species under natural
field conditions (21).
Materials and Methods
Deer mice were taken from corn and soybean fields using snap traps baited with
peanut butter and oats. All fields were commercially farmed and information about
373
374
Indiana Academy of Science
Vol. 94 (1985)
Table 1 . Agricultural pesticide use and deer mice trapped from cultivated fields in Scott
County, Indiana, 1983. Capitalized chemicals are trade names.
Chemical application
rate (per acre)
Dates
Deer mice
Field
Spraying
Planting
Trapping
captured
Conventional corn
CRA
alachlor (3 qts.)
atrazine (2 lbs.)
Amaze (7 lbs.)
1 Jun
2 Jun
13-14 Jul
3
LSm
atrazine (1.5 lbs.)
butylate (4 qts.)
26 Apr
2 Jun
13-14 Jul
5
PFA
alachlor (2 qts.)
atrazine (2 lbs.)
carbofuran (15 lbs.)
9 Jun
9 Jun
19, 22 Jul
11
Conventional soybeans
LAG
alachlor (3 qts.)
linuron (2 qts.)
12 Jun
9 Jun
19, 22 Jul
14
H1G
alachlor (1 qt.)
linuron (1 qt.)
1 Jun
25 May
14-15 Jul
22
No-till corn
MON
alachlor (2 qts.)
atrazine (2 lbs.)
carbofuran (15 lbs.)
paraquat (1.5 pts.)
1 Jun
19 May
3, 5 Aug
15
BRO-C
Bicep (3 qts.)
carbofuran (9 lbs.)
paraquat (2 qts.)
24 May
23 May
3 Aug
13
No-till soybeans
BROS
Dual (1 qt.)
linuron (0.5 qt.)
paraquat (2 pts.)
3 Jul
3 Jul
3 Aug
5
KSm
alachlor (2 qts.)
linuron (2 qts.)
paraquat (1 pt.)
16 Jun
15 Jun
2 Aug
5
planting and spraying dates and chemicals used (Table 1) were obtained directly from
farmers. Fields were located in Scott County of southeastern Indiana, where soils are
primarily silt loams derived from glacial till. The topography is flat to moderately rolling.
Conventionally tilled corn and soybean fields had been plowed or disked in the
spring before planting. No-till cornfields were slot-planted directly into the previous
year's residues. A slot-planter uses a knife-like implement to make a narrow furrow
in which the seed is deposited. No-till soybeans had been planted to winter wheat the
previous fall, and were slot-planted with soybeans directly into residues following wheat
harvest in early summer. More detailed crop histories are given elsewhere (6).
Deer mice were trapped over a 4-night period in each field, and mammals cap-
tured were individually bagged, labeled, frozen, and transported to laboratories at Indiana
University Southeast. They were analyzed using thin-layer chromatographic (TLC) techni-
ques described below. It was necessary to pool 3-6 mice to obtain enough material
for each analysis. For histological analyses, mouse tissues were fixed in alcoholic for-
malin, and later dehydrated, cleared, and embedded in paraffin blocks. Sections ob-
tained from these blocks were attached to slides, stained with H & E, and mounted
with Permount. The sections were then visually scanned for evidence of histological
abnormalities.
Environmental Quality 375
Alachlor (= Lasso)
Alachlor (2-chloro-2,6 diethyl-N-(methoxymethyl)-acetanilide) is a preemergent her-
bicide manufactured by the Monsanto Corporation for the control of annual grasses
and certain broadleaf weeds in soybeans and corn. The concentrate most commonly
available at retail outlets contains 4 lbs. of alachlor per gallon. Following application,
the active ingredient persists in the soil for 6-10 weeks (5).
For alachlor, whole, skinned mice were homogenized in a blender and extrac-
tions obtained using Method 1A of the Pesticide Analytical Manual, Vol. II (19). The
2,6 diethylanilide residue obtained in this manner was concentrated "in vacuo" to
0.5 ml, subsequently dissolved into lOyul of chloroform, and spotted on fluorescent
silica gel TLC plates. Using a solvent system of 4:1 benzene-ethyl acetate, principal
yellow spots would appear at Rf 0.85 if alachlor was present in the tissue samples.
Atrazine (- AAtrex)
Atrazine (2-chloro-4-ethylamino-6-isopropylamino-s-triazine), a substitute s-triazine
available from CIBA-GEIGY Corporation in several trade name formulations such
as AAtrex 4L, is a selective herbicide utilized for season-long weed control in corn
and other crops. The retail product contains 4 lbs. of atrazine per gallon.
For atrazine analysis, homogenized mouse livers were extracted with anhydrous
methanol, and the resulting filtered solution was hydrolyzed with IN HC1 (23). Following
separation into phases with the addition of n-hexane, the aqueous bottom layer was
drawn off, neutralized with NFLOH, and then evaporated on a rotavapor to 0.5 ml.
The resulting residue was dissolved into 10/tl of chloroform, spotted on fluorescent
silica gel TLC plates, and developed in a solvent system consisting of 4:1 benzene-
ethyl acetate. Atrazine, if present, would be located at the Rf value of 0.27. The limit
of detection using this method was 0.03 ppm with approximately 70% recovery of
parent material.
Linuron (= Lorox)
Linuron (3-(3,4-dichlorophenyl)-l-methoxy-l-methylurea), distributed by DuPont
for selective weed control, is retailed as Lorox Weed Killer, a wettable powder con-
taining 50% linuron, and Lorox L Weed Killer, an aqueous suspension containing
41% linuron (13).
For these extractions, whole, skinned mice were homogenized in a blender, in-
itially extracted with anhydrous ether, and hydrolyzed with IN HC1 (4). The resulting
solution was then adjusted to a pH greater than 11 with NaOH, and from this solu-
tion, p-chloroaniline was re-extracted into a small volume of ether, dried with magnesium
sulfates, and then evaporated almost to dryness with a rotavapor. The resulting concen-
trate was dissolved in 9:1 petroleum ether-acetone, spotted on fluorescent silica gel
plates, and subsequently developed in the following solvent system: 35 ml methanol,
17.5 ml isoamylalcohol, 35 ml benzene, and 12.5 ml 2N HC1. If present, the p-
chloroaniline derivative of linuron would appear at the Rf value of 0.85. The reported
lower limit of detectability is 0.1 micrograms (4). Our recorded percent recovery of
standard reference materials was 91.7%.
Paraquat
Paraquat (l,l'-dimethyl-4,4 dipyridilium dichloride), a non-selective contact her-
bicide, is distributed by Chevron as Paraquat CL containing 2 lbs. of the paraquat
cation per gallon.
Whole, skinned, homogenized mice were extracted according to methods published
in the Pesticide Analytical Manual, Vol. II (19). The filtered solution was then concen-
376 Indiana Academy of Science Vol. 94 (1985)
trated "in vacuo" to 0.5 ml, spotted on fluorescent silica gel plates, and developed
in the following sequential solvent system which was designed specifically for the detection
of paraquat and its metabolites (1). The TLC plate remained in Solvent A which con-
sisted of 1:1:2:1 benzene-amyl alcohol-methanol-lN HCI for seven minutes and was
then immediately placed in Solvent B which consisted of 40:9:1 acetonitrile-H20-ammonia
until the solvent had traveled the entire length of the TLC plate. Paraquat, if present,
remains at Rf 0.19. Related compounds have been reported at the following locations
(1): QUINA-Rf 0.34, monopyridone-Rf 0.49, monoquat-Rf 0.54, and dipyridone-Rf 0.79.
Results and Discussion
A total of 76 deer mice was utilized in 17 assays for residues of the 4 herbicides
(Table 2). Five assays indicated the presence of herbicides or their metabolites. Nine
additional mice were histologically examined for lung and liver damage, and 2 individuals
showed evidence of liver abnormalities (Table 2).
Table 2. Summary of herbicide residue determinations and histological examinations
of deer mouse livers.
No.
. fields
No. mice
Frequency of samples
Frequency of
Herbicide
represented
sampled
with herbicide residues
liver damage
Alachlor
6
32
2/7
0/4
Atrazine
3
18
0/4
0/2
Linuron
2
11
0/3
3/3b
0/2
Paraquat
2
15
2/3
Pooled samples include 3-6 individual deer mice.
Metabolites uncertain.
Alachlor
Alachlor residues were detected in mice from two fields, a conventionally tilled
cornfield (CRA) and a no-till soybean field (KSm). UV spectrophotometric analyses
of the sample residues revealed principal wavelength peaks at 234 nm. Using p-
chloroaniline as a reference, the recorded residues were calculated to be 0.0003 ppm
(CRA) and 0.0001 ppm (KSm). These recorded levels are extremely low and far below
the reported sensitivity levels of 0.01 to 0.02 ppm for this compound (18).
Toxicology studies (5) of the effects of alachlor on rats indicated relatively high
acute oral LD50s ranging from 100 mg/kg to 5800 mg/kg for various formulations of the
retail product Lasso. Alachlor produced tumors in some laboratory mice when fed at levels
greater than 260 mg/kg/day over the entire lifetime of the experimental animals (5). All
4 deer mice histologically examined from fields in which alachlor was used showed
normal liver appearance. The presence of detectable alachlor in some individuals is
unlikely to create an environmental problem due to the large dosages required to in-
duce tumors. In conventional as well as reduced tillage fields, it is improbable that
these levels would ever be approached.
A trazine
Atrazine residues were not detected in any samples, nor did the livers of 2 mice
examined appear abnormal. The acute oral toxicity of atrazine (AAtrex 4L) in rats
was 1886 mg/kg in males and 1075 mg/kg in females (8). Besides this relatively high
level of toxicity, atrazine is rapidly excreted from the body (8) posing little threat to
nontarget rodents and their predators. Atrazine accounts for almost 25% of all her-
bicides applied to crops in the United States, and may be metabolically transformed
Environmental Quality 377
by both plants and animals into a mutagenic substance (12, 20). Atrazine has been
shown to affect the behavior of rats by altering their circadian rhythms (18).
Linuron
Linuron was not detected in any tissue samples. Dupont (13) reported an acute
oral LD50 for linuron of 1906 mg/kg for male mice and 2873 mg/kg for females.
Linuron fed to mice at dietary levels of 50 and 150 ppm for two years produced no
measurable chronic effects, but an extremely high dietary level of 1500 ppm produced
hepatocellular adenomas in female mice (13). In a reproduction study, linuron produced
a high incidence of deformed embryos at a feeding rate of 200 mg/kg (14). No general
toxic, reproductive, or teratogenic effects were noted in a 3-generation rat study at
a dietary level of 125 ppm (13).
Paraquat
In utilizing the sequential TLC procedure for the detection of paraquat and its
metabolites, no immediate evidence of paraquat was found at Rf 0.19-0.21. However,
identical streaks with an Rf range from 0.54-0.72 were noted on plates developed from
a no-till corn (BRO-C) and a no-till soybean field (BRO-S) (Figure 1). In addition,
the tissue extractions from the no-till cornfield produced plates with compact spots
located at Rf 0.89. The known degradation products of paraquat falling in this range
would be monoquat (Rf 0.54) and dipyridone (0.79). Considering that definite resolu-
tion was not obtained, it cannot be stated conclusively that these represent paraquat
metabolites. The livers of 2 of 3 additional mice examined from both fields showed
signs of damage, mainly changes in appearances of fatty cells which is a characteristic
response of the liver to a wide variety of toxic compounds, including paraquat (24).
In contrast to the relatively low toxicities of the other herbicides considered in
this study, paraquat is moderately toxic and known to damage epithelial tissues of
the skin, nails, cornea, liver, kidney, the gastrointestinal tract, and the respiratory
tract (24). Such injuries may be reversible in all but the lung where a severe pulmonary
reaction to paraquat in often fatal (24). Intraperitoneal injections were toxic at 17-21
mg/kg in rats. Acute oral mammalian toxicities ranged from an LD50 of 5 mg/kg
in hares (15) to 115 mg/kg in rats (24). This difference due to the mode of administra-
tion is attributed to poor absorption of paraquat through the gastro-intestinal tract
(9). Parquat is also a strong skin irritant with a reported acute dermal LD50 of about
85 mg/kg in rats (24). Deer mice we examined displayed no skin lesions or loss of
hair. Chronic administration of small doses of paraquat produced no clinical signs
for several weeks (24). Thereafter, signs of illness developed in the form of anorexia,
weight loss, and dyspnea. The animals usually died within 10 days of the onset of
the symptoms (24). Paraquat has been noted to have many mutagenic and embryo-
toxic properties (3, 24). Additional gross and microscopic morphological changes aris-
ing from paraquat ingestion in rats included: loss of body weight, teratogenic effects
in embryos, damage to the liver and kidney, lung weight increase and considerable
pulmonary fibrosis, smaller spleen and thymus, heavier adrenals with abnormal histology,
lowered white and red blood cell counts, degenerative changes in the testes, corneal
opacification, and other changes as well (2, 24).
Conclusions
The lack of detectable concentrations of atrazine and linuron or their metabolites,
the low frequency and extremely low concentrations of alachlor, and the absence of
apparent liver damage in deer mice taken from fields in which these herbicides were
used, indicate little cause for immediate environmental concern about their regulated
378
Indiana Academy of Science
Vol. 94 (1985)
Rf0.89 • • •
Rf 0.72
Rf 0.54
Initial Spots
BROS
BROC
Figure 1 . Tracings of TLC plates from tissue sample extractions of deer mice taken
from 2 paraquat-treated fields (BRO-S, BRO-C) indicating metabolites of uncertain
origin. Paraquat, if present, would have migrated directly above the initial spots.
use on agricultural lands. Unlike many insecticides, herbicides rarely persist i n the
environment for more than a few days or weeks (16). The results for paraquat, however,
suggest that metabolites may be present in deer mice 31-71 days after field application
and may be responsible for observed liver damage. This could result in elevated rates
of mortality in deer mouse populations in no-till fields, although population levels
and short-term mortality rates were found to be similar in conventional and minimum
tillage fields in southern Indiana (6, 7).
Further research is warranted to determine the exact origin of metabolites found
in this study, as well as to obtain better estimates of the incidence of liver damage
in deer mice inhabiting row-crop fields. It would seem prudent to encourage use of
alternative contact herbicides (e.g., glyphosate) for no-till farming that may pose less risk.
Acknowledgments
Dave Fellows, Ben Nassim, and Rick Speer assisted in various aspects of this
study. Special thanks go to Robert Feldt. Facilities were provided by the Departments
Environmental Quality 379
of Biology and Chemistry at Indiana University Southeast. The various chemical com-
panies (Chevron, CIBA-GEIGY, DuPont, and Monsanto) willingly supplied informa-
tion on the herbicides. Primary funding resulted from a grant to the senior author
by the Indiana Division of Fish and Wildlife. Additional funding came from Federal
Aid to Wildlife Restoration in Indiana, Project W-26-R.
Literature Cited
1. Abou-Donia, M.B. and A. A. Komell. 1978. Sequential thin-layer chromatography
of paraquat and related compounds. J. Chromatog. 152:585-588.
2. Bauer, C.A. 1983. The effects of paraquat on various reproductive and growth
parameters in first and second generation bobwhite quail. Ph.D. Diss., Indiana
State University, Terre Haute. 70 p.
3. Benigni, R., M. Bignami, A. Carere, G. Conti., L. Conti, R. Crebelli, E.
Doglotti, G. Gaulandi, A. Noveletto, and V. A. Ortali. 1979. Mutational
studies with diquat and paraquat in vitro. Mutat. Res. 68:183-193.
4. Bleidner, W.E. 1954. Application of chromatography in determination of micro-
quantities of 3-(p-chlorophenyl)-l,l-dimethyl-urea. J. Agric. Food Chem.
12:682-684.
5. Brandt, E.J. 1984. Personal communication from Environmental Affairs Depart-
ment, Monsanto Agricultural Products Co., St. Louis.
6. Castrale, J.S. 1984. Impacts of conservation tillage practices on farmland wildlife
in southeastern Indiana: population levels and habitat use. Indiana Div. Fish and
Wildl. Fed. Aid Prog. Rep. Proj. No. W-26-R-15. Job No. XXXV-M-2,3.
7. Castrale, J.S. 1984. Impacts of conservation tillage practices on farmland wildlife
in southeastern Indiana: pesticide levels. Indiana Div. Fish and Wildl. Fed. Aid
Prog. Rep. Proj. No. W-26-R-15. Job No. XXXV-M-4.
8. CIBA-GEIGY Corporation 1984. Toxicology Data. AAtrex Herbicides. Personal
communication from Agricultural Products Division. Greensboro, N.C.
9. Clark, D.G., T.F. McElligott, and E.W. Hurst. 1966. The toxicity of para-
quat. Brit. J. Ind. Med. 23:126-132.
10. Conservation Tillage Information Center. 1983. 1982 National survey of con-
servation tillage practices. Conservation Tillage Information Center, Fort Wayne,
Indiana 83 p.
11. Cowan, W.F. 1982. Waterfowl production on zero tillage farms. Wildl. Soc.
Bull. 10:305-308.
12. Crosson, P. 1982. Conservation tillage and conventional tillage: A comparative
assessment. United States Environmental Protection Agency. EPA Rep. No.
600/3-82-027. 72 p.
13. DuPont de Nemours & Co. (Inc.). 1984. Technical Data sheet for linuron. Per-
sonal communication from Biochemical Department. Wilmington, Delaware.
14. Khera, K.S., C. Whalen, and G. Trivett. 1978. Teratogenicity studies on linuron,
malathion, and methoxychlor in rats. Toxicol. Appl. Pharmacol. 45:435-444.
15. Milhaud, G. 1974. Toxicity of grammoxone. Rec. Vet. 150:337.
16. Morrison, M.L. and E.C. Meslow. 1983. Impacts of forest herbicides on wildlife:
toxicity and habitat alteration. Trans. North Am. Wildl. and Nat. Resour. Conf.
30:336-348.
17. Mumford, R.E. and J.O. Whitaker, Jr. 1982. Mammals of Indiana. Indiana
University Press, Bloomington. 537 p.
18. Nicolau, G.Y. and E. Socoliuc. 1980. Effects of atrazine on circadian RNA,
DNA and total protein rhythms in the thyroid and adrenal. Endrocinologie
18:161-166.
380 Indiana Academy of Science Vol. 94 (1985)
19. Pesticide Analytical Manual Vol. II. 1969. The determination of
2-chloro-2,6-diethyl-N-(methoxymethyl) acetanilide metabolites containing
2,6-diethylamine moiety. Pest. Reg. Sect. 180.249.
20. Plewa, M.J. and J.M. Gentile. 1976. Mutagenicity of atrazine. A maize-microbe
bioassay. Mutat. Res. 38:287-292.
21. Robel, R.J., CD. Stalling, M.F. Westfahl, and A.M. Kadoum. 1972. Effects
of insecticides on populations of rodents in Kansas 1965-69. Pesticides Monit.
J. 6:115-121.
22. Rodgers, R.D. and J.B. Wooley. 1983. Conservation tillage impacts on wildlife.
J. Soil and Water Conserv. 38:212-213.
23. Shin, K.H. and J.K. Moon. 1979. Analysis of atrazine herbicide residue by thin-
layer chromatography. Punsok Hwahak 4:30-32.
24. Smith, P. and D. Heath. 1976. Paraquat. Crit. Rev. Toxicol. 4:411-445.
25. Warburton, D.B. and W.D. Klimstra. 1984. Wildlife use of no-till and con-
ventionally tilled corn fields. J. Soil and Water Conserv. 39:327-330.
Acid Rain: A Synopsis
Ronald J. Galloy
Indiana Air Pollution Control Division
Indiana State Board of Health
Indianapolis, Indiana 46206
Introduction
Acid rain is a simple term used to describe both dry and wet forms of acid deposi-
tion. These depositions originate from naturally occurring and anthropogenic (man-
made) sources. At this point there is a lack of scientific understanding regarding how
much man-made pollutants contribute to this complex issue.
Research programs conducted by the federal governments Interagency Task Force
on Acid Precipitation are giving us greater understanding of acid rain. This research
continues and programs are expanding but acid rain appears to be an issue that will
require legislative action before there is full scientific conclusion on cause and effect.
Historic Note
In 1857 Robert Angus Smith, an English chemist, presented the first detailed
analysis documenting polluted precipitation and some of its harmful effects. Twenty
years later Smith authored a comprehensive precipitation chemistry study which coined
the phrase "Acid Rain."
Chemistry of Acid Rain
The acidity-alkalinity of a water solution is measured by its pH. The pH scale
ranges from 0 (extreme acidity) to 14 (extreme alkalinity) with the value of 7 being
neutral. Pure water has a pH of 7 since the dissociation of water molecules into hydrogen
(acid) and hydroxyl (alkaline) ions is very small. The scale is logarithmic and each
pH unit represents a ten-fold change in the hydrogen/hydroxyl ion concentration.
What is acid rain? Acid rain defined is rain with a pH value lower than 5.6.
The reason 5.6 is set as the determining pH is because carbon dioxide in balance with
atmospheric moisture creates a carbonic acid solution with this value. Although rain
is defined as acid below this level, naturally occurring rain may range down to 4.9
and up to 6.5. These values allow for additional acidity resulting from lightning or
alkalinity resulting from atmospheric dust.2'5 Figure 1 shows the pH scale with a list
of values for commonly found substances.
pH Scale1'2
Extreme Alkalinity 14
13 13.0 Lye
12 12.0 Household ammonia
11
10
9 8 to 9 Soap
8 8.2 Baking soda
Neutral (Pure Water) 1_ 7.4 Human blood
6 6.4 Milk
5 5.0 Carrots
4 4.6 Bananas
3 3.0 Apples
2 2.2 Vinegar
1 1.1 Stomach digestive acids
Extreme Acidity 0
Figure 1. pH scale with a list of values for commonly found substances.
381
382 Indiana Academy of Science Vol. 94 (1985)
Rain becomes acidic in several ways. Mainly this occurs from C02, NOxand S02
gases interacting with atmospheric moisture. Carbon dioxide is a naturally occurring
gas composing .03 percent of the atmosphere and nitrogen oxides result from lightning
and combustion processes. Most sulfur oxides are emitted from fossil fuel combustion
at electric generation plants. Figure 2 shows how acids are formed from mixture of
these gases with water.
Acid Formation
A. C02 + 2 H20 - H,0 + + HCOj~
Carbon Dioxide + Water Yields Hydronium Ion and Bicarbonate
B. 2 NO + 02 - 2N02
Nitrous Oxide + Oxygen Yields Nitrogen Dioxide
3 N02 + H20 - 2 HN03 + NO
Nitrogen Dioxide + Water Yields Nitric Acid and Nitrous Oxide
HNO, + H20 - H,0+ + NO,"
Nitric Acid + Water Yields Hydronium Ion and Nitrate
C. So2 + H20 - 2H2SO,
Sulfur Dioxide + Water Yields Sulfurous Acid
2 H2 SO, + Vi 02 - 2 H2 S04
Sulfurous Acid + Oxygen Yields Sulfuric Acid
H2 S04 + H20 - H,0+ + HS04"
Sulfuric Acid + Water Yields Hydronium Ion + Bisulfate
Figure 2. Acids formed from mixture of gases with water.
Certain biological filters affect the chemistry of rainwater from its initial point
of contact near grounds surface to watershed entrance. These filters include: a) the
forest canopy, bushes, other plant leaves, and greenery which collect atmospheric dusts
and add alkalinity to the water; b) the humus layer from decaying vegetation on the
ground which adds acid concentrations; and c) the soil and rock layer containing alkaline
minerals providing further alkalinity. These filters have had a constant effect on rain-
water chemistry prior to man's influence, therefore any changes to watershed chemistry
can be attributed to anthropogenic reasons.
Major ions influencing rains pH are sulfate S04~ ~ , nitrate N03~ , chloride Cl_ ,
ammonium NH4 + , calcium Ca + + , magnesium Mg + + , and potassium K + . Exactly
how much man contributes to excessive acidity through sulfate an nitrate deposition
is yet undetermined. It is believed the contributions through power plants, industrial
processes, and transportation sources are significant.
Affects of Acid Rain
Rain is the natural cleansing agent of the atmosphere. As it forms and falls to
the earth it gathers with it various pollutants including those causing acid rain. Ultimately
it is the land and watersheds which act as final pollution collectors.
Affects from acid rain are stated to include acidification of lakes resulting in
reduced or total loss of fish population, corrosion of buildings and monuments, and
reduced seed germination resulting in cuts in crop and timber production. Studies have
been conducted showing that a pH of 5 is the level where fish life in general ceases
to exist. Acid rain is also able to leach out metals from the soil including aluminum.
Once soluble, aluminum can be toxic to aquatic wildlife by clogging the gills of fish
and to vegetative species by causing a dehydration condition.
The Congressional Research Service (CRS) has completed a study listing possible
effects from the impact of acid rain on aquatic biota. According to CRS; bacteria,
algae, vegetative, invertebrate, amphibian, and fish populations shift away from acid
sensitive species. More specifically, bacterial decomposition decreases, sensitive fish
species die or experience reproductive failure and increases in aluminum make fish
Environmental Quality 383
more susceptible to death from exposure to acid conditions.4 Ironically lakes adversely
affected by acid conditions appear crystal clear due to decreases by living biota in them.
One of the major acid rain issues is degradation of the lakes and forests in the
Adirondack Park system. This park, largest in America, covers six million acres, and
is located in upper New York state. Some high elevation lakes and ponds in this park
have acid values less than 5.0 pH2 and hence do not support most form of fish life.
Also, our northern neighbor Canada, is voicing strong concern over acid deposition
originating from sources located in the United States but falling on its land and lakes.
Recent studies, yet unconclusive, indicate that decline of some forest species from Maine
to North Carolina are a result of the acid rain phenomenon.
What Should We Achieve with an Acid Rain Regulatory Program
Rain in the northeastern part of the country has been averaging 4.2 pH. The
National Academy of Sciences states that a target level of 4.5 pH is necessary to pro-
tect sensitive aquatic ecosystems from acid rain.6 This pH level allows for natural causes
of acidification and gives allowance for the fact that removal of all S02 and NOx
emissions from man-made emission sources would be economically impossible.
The U.S. EPA has estimated that at the start of the 1970's, about 26 million
tons of S02 and 17 million tons of NOx were emitted annually into the atmosphere
of the U.S. Of these totals, about 16 million tons of the S02 (62 percent) and 5 million
tons of the NOx (29 percent) were exhausted by fossil fuel burning electric generation
plants.2 In EPA's nationwide emissions report for 1982 power plant S02 and NOx
emissions amounted to 17.5 million tons and 7.5 million tons respectively.10
To achieve the necessary reductions of acid rain it is suggested that anywhere
from 3 to 12 million tons of S02 be removed yearly from power plant exhaust gases.
A lesser degree of NOx removal is also suggested.
Control Methods
Controlling acid rain means controlling S02 and NO, x emissions. Controlling S02
emissions can include the following:
1. Coal washing — Sulfur in coal occurs primarily in two forms, organic and
inorganic (pyritic). Organic sulfur, chemically bound to the coal, cannot be removed
by physical cleaning. Pyritic sulfur composing up to 45 percent of the sulfur in
coal is bound to iron and occurs as a separate particle. Up to 90 percent of the
pyritic sulfur can be removed by washing thus yielding reductions of 10 to 40
percent of the total sulfur content.8
2. Using low sulfur coal for combustion— Coal generally ranges from 0.5 per-
cent to 5 percent sulfur with western coal having a lower average sulfur content
than midwestern coal. By midwestern standards locally mined coal is considered
low sulfur when it is 2 percent or less. The lower the coals sulfur content the
lower the generated emission level of S02.
3. Exhaust gas scrubbers— Scrubbers are very expensive to install and operate
but are effective for controlling S02. All new coal-fired power plants are required
to remove 90 percent of the S02 gases or control it to a level of 1.2 pounds
per million Btu, whichever is stricter. Usually this means scrubber installation.
Controlling NO,x can be through use of low NO,x burners in power plants and in-
dustrial boilers, and through vehicle emission reductions which are now occurring from
currently implemented programs.
384 Indiana Academy of Science Vol. 94 (1985)
Liming lakes has also been suggested as a control supplement and is reasonably
cost-effective for regulating the pH of lakes.
Control Proposals
Several legislative bills have been submitted for action on acid rain. The legisla-
tion ranges from a) reductions of 3 million tons of S02 from a 10-state area, b) 10
million tons of S02 from 31 states east of and touching the Mississippi with additional
reductions of NO,x, and c) 12 million tons of S02 from the 48 contiguous states with
additional NO,x reductions. Who funds the equipment needed to yield these reductions
range from each state paying for its required equipment and associated reduction to
a national tax on most forms of electric generation applied where needed for the program.
Expense
S02 reductions are occurring as New Source Performance Standards for power
plants take effect. NO,x reductions are occurring from NSPS also, and as emission reduc-
tions from the newer auto fleet are realized. These reductions however are not happen-
ing quickly enough to abate the acid rain problem. Current lack of an effective plan
for acid rain is due to: a) the lack of conclusive knowledge about environmental effects
coupled with; b) the great expense involved to retrofit controls onto existing utility boilers.
To gain scientific understanding of this issue federal expenditures for acid rain
research in fiscal year 1985 will double to $55.5 million from the 1984 level. The
Environmental Protection Agency will receive the bulk of this with a 124 percent fun-
ding increase to $34.3 million.
Regarding expense of a control program the State of Indiana is used as an exam-
ple for a cost estimate. In 1982 Indiana's total S02 emissions from stationary sources
amounted to 1,694,000 tons with about 88 percent or 1,490,000 tons coming from
power plants.3 From an EPA survey of all coal burning public utilities in Indiana it
was determined that retrofitting scrubbers to control S02 emissions would cost 1.85
billion dollars. Operation and maintenance costs for these would amount to another
355 million dollars yearly.9 With this data an estimated expenditure of $1,241 per ton
of emissions would be required to retrofit scrubbers with an additional expenditure
of $238 per ton for yearly operation and maintenance.
To conform with acid rain control strategy, Robert McKnight, Chief Environmental
Engineer at Indianapolis Power and Light, states local utilities electric rates could be
up to 31 percent higher from the costs of controlling S02 emissions. This figure applies
to legislation such as the Stafford bill which requires each state to pay for their own
contributing share of emissions.7 Another study conducted by the Congressional Of-
fice of Technology Assessment estimates utility rate increases ranging from 5 to 19
percent as applied to various utilities in affected states. Other studies show as low
as a 2 percent increase in rates to customers based on a national tax to fund this program.
It should be understood that implementation of necessary acid rain control legisla-
tion for environmental protection could result in associated social problems in the form
of: a) displacing jobs in the coal mining industry; and b) increased utility expenses
to be shared by the poor and elderly. To minimize social disruption reasonable legisla-
tion must also account for job displacement protection and provide assistance to those
less fortunate and unable to burden the extra expense of control. If these problems
are dealt with fairly, society will surely gain from the benefits of protecting our buildings,
monuments, lakes, forests, and aquatic wildlife.
Conclusion
Our environment has improved since the institution of federal, state, and local
Environmental Quality 385
environmental management programs, however, some problems remain. Acid rain is
one of these. Environmentalists, industry, and the public all agree a solution is necessary.
Perspectives on the solution vary widely but the differences are healthy for from these
varied views a balanced effective management program will develop. For now acid
rain is long from solved but as research continues and comprehensive management
programs evolve our society will come to benefit from protection against acid rain.
Literature Cited
1. American Chemical Society, Acid Rain Information, Washington, D.C., October
1982, 8 pp.
2. Edison Electric Institute, An Updated Perspective on Acid Rain, Washington,
D.C., November 1981, 44 pp.
3. Indiana, Air Pollution Control Division, Emission Inventory Subsystem, 1982.
4. Inside EPA, Weekly Report, Washington, D.C., October 28, 1983, p. 10.
5. National Research Council, Acid Deposition; Atmospheric Processes in Eastern
North America, National Academy Press, Washington, D.C., 1983, 375 pp.
6. National Research Council, Atmosphere-Biosphere Interactions, National Academy
Press, Washington, D.C., 1982, 263 pp.
7. Stated by Robert McKnight, Chief Environmental Engineer, Indianapolis Power
and Light, in a Telephone Interview with R.J. Galloy on January 12, 1984.
8. U.S. EPA, Control Technique for Sulfur Oxide Emissions from Stationary Sources,
Research Triangle Park, April 1981, p. 4.2-10.
9. U.S. EPA, Document 600/7-8 1-0 12a, Utility FGD Survey Oct. -Dec, Research
Triangle Park, 1980, pp. A-7, 8.
10. U.S. EPA, National Emissions Data System, Nationwide Emissions Report,
Research Triangle Park, December, 1983.
GEOLOGY AND GEOGRAPHY
Chairperson: Edward Lyon
Department of Geography
Ball State University
Muncie, Indiana 47306
(317)285-1761
Chairperson-Elect: John Cleveland
Department of Geology/Geography
Indiana State University
Terre Haute, Indiana 47809
(812)749-2833
ABSTRACTS
Landfills in Marion County — A Revisit. Konrad J. Banaszak and Theodore K.
Greeman, U.S. Geological Survey, 6023 Guion Road, Indianapolis, Indiana 46254.
Seven landfills studied in the early 1970s were revisited in the fall of 1983. Four
of the fills are in coarse sediments of the White River glaciofluvial channel and
three are on the Tipton Till Plain. A map of lineaments was prepared from aerial
photographs. There is no apparent relation between those features and the hydrology
of the fills, probably because many wells are lost and four fills are in coarse material.
Of the 82 wells drilled to study the three fills in till, 38 remain. Ground-water mounds
were present at all three. At one fill, the specific conductance of water in most shallow
wells ranged from 1,200 to 10,000 micromhos per centimeter (umhos/cm), and in a
deep (162-foot) well specific conductance increased from an average of 760 umhos/cm
in the 1970s to 4,450 umhos/cm in 1983. Of the 93 wells drilled to study the four
fills in glaciofluvial material, 57 remain. In 1983, no data could be collected at one
fill. Of the remaining fills, one had no gradient change; flow was toward the river
with extremely slight vertical gradients. The second fill had no gradient change; flow
was away from the river with downward vertical gradients. The gradient at the third
fill had great change. In the 1970s, the shallow system flowed toward the river with
a horizontal gradient of 0.001 and deep system was almost flat. In 1983, the direction
of shallow and deep flow was away from the river with a horizontal gradient of 0.0025.
These results confirm the advantages of continuous monitoring and the upredictability
of changes in flow direction and gradient.
Compression Strength Testing of the Springfield Coal, Coal V, Pike County, Indiana.
K.C. Kuo and T.R. West, Department of Geosciences, Purdue University, West
Lafayette, Indiana 47907. Coal pillars are left intact in underground mines to
support the opening. Typically square or rectangular in shape, their purpose is to pro-
vide safety and continued mining while preventing surface subsidence. The optimum
design maximizes coal extraction as well.
Coal strength can be determined by in-situ tests, (time consuming and expensive)
or through laboratory testing. In the lab, different sized, cube-shaped specimens are
tested in uniaxial compression. Research on Appalachian coals has shown that strength
of cubes decreases with increasing size until a value equal to the pillar strength is obtained.
In this research, specimens of the Springfield Coal (Coal V) were collected from
an operating open pit mine, Pike County, Indiana. Cut from the working face im-
mediately behind the loading shovel, they were stored in sealed styrofoam coolers to
387
388 Indiana Academy of Science Vol. 94 (1985)
prevent moisture loss. Cracks occur in coal specimens during drying. Storage is in
a humidity chamber prior to sample preparation and for prepared samples until testing.
Specimens are cut dry using a horizontal band saw with a tungsten carbide blade.
After rough cutting, cubes are ground smooth using sand paper and a surface grinder
to assure the loading surfaces are parallel. Cubes are prepared so that loading will
be perpendicular to the bedding planes. Coal strength data for the Illinois Basin coals
will be provided in this research.
Interpretation of Glacial Geology and Groundwater Problems in East-central Indiana
using Improved Compilations of Water Well Driller's Records. Alan C. Samuelson,
Department of Geology, Ball State University, Muncie, Indiana 47306. Recently
published USGS compilations of water well driller's records in East Central Indiana
have proven to be superior to previously published general compilations. The data
were compiled for computer simulations of regional groundwater conditions, but have
been valuable in interpretation of landuse, site specific groundwater, and geologic pro-
blems involving glacial stratigraphy. The new compilations show depth and lateral ex-
tent of sand and gravel horizons. The improved maps display four to six sand and
gravel horizons per county and show distribution by elevation and thickness of each
horizon. A number of examples are presented to demonstrate data reliability as con-
firmed by subsequent tests and the resulting evaluations of geologic, engineering, and
groundwater resource problems. Specific aquifer horizons have been correlated with
outcrop and out wash soil exposures. Locations of important groundwater seepage into
stream baseflow can be identified.
Three-dimensional Patterns of Biotite Composition within the Cloudy Pass Batholith,
Washington. J.R. Sans and CD. Potter, Department of Geology, Ball State Univer-
sity, Muncie, Indiana 47306. The Cloudy Pass batholith is a small epizonal pluton
of Miocene age. Since the batholith straddles the Cascade Crest, it has been deeply
dissected by glacial erosion so that specimens could be collected over an area 14.88
by 15.26 kilometers with a vertical range of 1.54 kilometers.
The ten chemical elements most abundant in biotite (Na, Mg, Al, Si, CI, K, Ca,
Ti, Mn, total Fe) were determined by electron microprobe. Ferrous iron was deter-
mined by decomposition in a teflon bomb followed by titration of excess standard
potassium dichromate with standard ferrous ammonium sulfate.
The compositional variations of biotite were studied on the following five dif-
ferent scales extending over nine orders of magnitude (micrometers to kilometers): (1)
within a single biotite grain, (2) between grains in a single thin section, (3) between
sections from the same rock specimen, (4) between specimens from the same outcrop
and, (5) over the entire accessible volume of the batholith (about 350 cubic kilometers).
At the scale of a single biotite grain, three cations (Na, K and Mn) exhibit essen-
tially no zoning, five cations (Mg, Al, Si, Ca, and Fe) show weak zoning, and one
cation (Ti) shows strong zoning. At the three intermediate scales, specimens from the
center of the pluton show a significant range of biotite composition, especially in the
Fe/(Fe + Mg) cation ratio. Specimens from the margins and roof show a peculiar bimodal
distribution of biotite compositions. On the scale of the entire batholith, Fe/(Fe + Mg),
Mn, total Fe, and ferrous Fe decrease with elevation, whereas Mg, CI, and ferric iron
increase. All the above features of biotite are interpreted as due to subtle resetting
of composition by hydrothermal activity during the cooling history.
Geology and Geomorphic History of the Garrison Chapel Cave System, Monroe County,
Indiana. William L. Wilson and Donald W. Ash, Department of Geography and
Geology and Geography 389
Geology, Indiana State University, Terre Haute, Indiana 47809. The Garrison
Chapel Cave System, in western Monroe County, Indiana, is composed of three
hydrologically connected caves named Grotto, Shaft and Salamander. All three convey
the same drainage westward from portions of the karsted Cave Creek and Garrison
Chapel Valley watersheds. The cave stream resurges along the eastern side of Coon
Hollow and is tributary to Richland Creek via Little Richland Creek. Up to four cavern
levels are present in some portions of the system. Similar size, elevation, and fluvial
sediments have led some authors to suggest that the Main Passage in Salamander Cave,
the Big Room in Shaft, and the Main Passage in Grotto Cave were at one time in-
tegrated parts of the same large truck drainage net. Recent stratigraphic measurements
and level surveys show that the passages are not related. Upper levels are accordant
with bedding, are generally strike-oriented, have low gradients, and have sequences
of mostly silty fluvial sediment that rise to, or near to, the passage ceiling, except
where re-excavated by free surface streams. The lowest level contains an active stream,
is dip-oriented, has a gradient steeper than the local dip, consequently downcutting
at least 35 feet through the stratigraphic section. The relationship between cave passages
and their geologic setting suggests a history of initial progressively westward and
stratigraphically lower development of strike-oriented, phreatic passages that occurred
perhaps in response to base level lowering. Meander scars that rise along the cave
wall while passing downstream, indicate conduits may have developed by upcutting
to reach equilibrium with base level (paragenesis). At some places, the older, upper
levels have collapsed into the stream (lowest) level. Some cave streams appear to have
fortuitously intersected older passages and now follow the passages along certain reaches
of the stream. Thick, paragenetic sediment has been partially excavated by modern
streams that may be downcutting to reach equilibrium with base levels that were greatly
lowered by deep stage entrenchment of surface streams associated with drainage rear-
rangements of the Teays and Ohio rivers during Pleistocene glaciation.
Evidence of Algal Source of Micrite in a Saluda
Coral Zone in Southeastern Indiana
Will H. Black well
Departments of Botany and Geology
Miami University, Oxford, Ohio 45056
Introduction
The Saluda Formation has received considerable study (1, 2, 3, 4, 6, 9), as indeed
is the case with other Cincinnatian (Upper Ordovician) lithostratigraphic units. Com-
pared to other Cincinnatian (particularly Richmondian) formations, however, the Saluda
is lithologically distinct, being typically dolomitic and poorly fossiliferous. The prevalent
lithology of the Saluda is either calcitic dolomite or dolomitic micritic (micro-to cryp-
tocrystalline calcium carbonate) limestone (3). Developmentally, the Saluda again pro-
vides contrast to other Cincinnatian strata in that it is in all probability the product
of a lagoonal setting. Specifically, the Saluda is considered to have originated from
a shallow, penesaline, atoll lagoon (3, 4, 9); associated tectonism perhaps represented
the inception of the Cincinnati Arch (9). The contour of the Saluda Formation, biconvex
and lens-like (3), reflects this ontogeny.
The only really characteristic fossils of the Saluda are the compound corals,
Favistella alveolata and Tetradium approximation (4). These two corals, singly or
together, tend to form a biostromal zone (or zones), especially in Lower Saluda rocks
(3, 4, 9). The coralline zone is of considerable paleoecological significance in that it
represents the remains of a low, but broad, wave-resistant bank of corals (and other
organisms) which essentially circumscribed the Saluda lagoon (3). Circumscription by
this coral shoal produced a barrier which significantly altered depositional environ-
ment, restricting conditions lagoonward as compared with the surrounding epeiric sea.
Environmental restrictions of the shallow lagoon eventually led to increased evapora-
tion rates, salinity, and dolomitization (perhaps penecontemporaneous) within the lagoon
(3, 9).
The encircling coralline zone per se is not so highly dolomitic, and contains an
abundance of micrite. The often massive coral colonies are in some cases haphazardly
oriented, indicating at least sporadic turbulent conditions of the surrounding sea (3).
This relatively high energy coralline zone more or less effectively delimited the low
energy (quiet water) lagoon from the moderate energy epeiric sea (9). The term "reef"
is not applied to the coralline zone because of the lack of consistent structural con-
solidation (3).
As discussed by Van Hart (9) an apparent textural anomaly exists between evidence
of a coral bank reflecting turbulent conditions and the presence (in association with
the corals) of substantial amounts of ooze (micrite), which would presumably have
been winnowed away by the turbulence. Van Hart speculated that the coralline zone
might in fact represent a coral/algal complex, and that algal mats could have been
the source of the persistent micrite which can be seen in some cases to connect and
even surround the coral colonies. Although entirely logical, this idea has remained
as speculation. Direct evidence of algae or of definitive algal micrite in the Saluda
coralline zone has not been satisfactorily demonstrated. It is to this end that this
investigation was directed.
Materials and Methods
Samples were collected at two exposures of the Saluda Formation in eastern Indiana.
The first locality is on a roadcut along Highway 101, approximately 5 miles north
391
392 Indiana Academy of Science Vol. 94 (1985)
of Brookville (4.5 miles north of the Brookville Lake Flood Control Station). This
is the locality designated as "Brookville North" by Hay (5). The thin exposure of
the Saluda at this locality consists primarily of Tet radium colonies. Hay considered
this exposure of the Saluda to represent the "feather edge" of the formation (personal
communication). The second locality is 1.1 miles northwest of Versailles on Highway
421 (north), 0.2 miles north of the divergence of Highways 421 and 50. Specifically,
the locality occurs several hundred yards east of the road in the middle of the "west
branch" of Cedar Creek; here a zone of Tetradium occurs within the confines of eight
to nine feet of exposed Lower Saluda sediments (4, 9). In collecting specimens, care
was taken to sample both colonial coral {Tetradium) material and, as well, intercalary
micritic limestone areas. More than 60 thin sections were prepared, by standard techni-
ques, divided equally between the two localities discussed. Slides and samples are
deposited in the paleobotanical collection associated with the Herbarium at Miami
University (MU).
Results and Discussion
Thin sections prepared of samples taken from Tetradium colonies often revealed
associated micrite. Conversely, sectioned intercalary limestone samples frequently con-
tained Tetradium fragments. A clear association is thus apparent between the colonial
corals, or their fragments, and probable in situ micrite. Invertebrate fossils (other than
corals) found in the micrite of the coralline zone are reasonably abundant and usually
fragmentary (3), with Ostracodes perhaps most commonly observed. Such fragmen-
tary constituents, trapped in the micrite, are doubtless allochthonous with respect to
the fundamentally autochthonous coralline zone. Monticuliporid bryozoans are occa-
sionally layered external to the surface of Tetradium colonies, and possibly constituted
a minor in situ component of the coral bank.
Microscopic examination of micritic regions in thin section generally supports
the hypothesis (9) of a predominantly algal source of micrite in the coralline zone.
The visible evidence is admittedly variable, however. A sliding scale exists between
areas of pure opaque micrite and those exhibiting more or less distinct calcareous algal
tubes. In either "extreme," or examples in between, an intimate relationship of algal
micrites with surfaces of the Tetradium colonies may be observed. In clearest examples
algae appear to have grown as encrusting masses directly upon Tetradium (Figure).
Based on tube diameter, morphology, and irregularity, these fossil algae bear a greater
resemblance to cyanophytes than to rhodophytes or chlorophytes (12).
Microscopic observations made on micrites of the Saluda coralline zone are con-
sistent with those of Wolf (1 1) on certain Australian Devonian and Recent algal deposits.
In both Holocene and Paleozoic examples, Wolf observed the product of an apparently
gradational grain diminution of calcareous algal cells and filaments to cryptocrystalline
calcium carbonate. Wolf considered this "decrease in detail" to be an early diagenetic
phenomenon. He discussed the possibility that algal tissue perhaps served bacteria nutri-
tionally, and that subsequent to bacterial delay, the calcareous algal remains may have
become reduced to detrital micrite and then lithified. Wolf (11) pointed to the need
for experimentation to substantiate bacterial decay as a cause of algal micritization,
as opposed to disintegration solely by mechanical abrasion (10). Regardless, Wolf con-
cluded that a great deal of enigmatic biohermal or knoll reef micrite may be explained
by grain diminution of algal colonies.
My observations thus correspond to Wolf's (11) on textural alteration, and also
support Van Hart's (9) hypothesis of the importance of algae in the development of
the Saluda Tetradium zone. In more general terms these observations are consistent
with the belief in the significant contribution of algae to many limestones and lime
Geology and Geography
393
Figure. Encrusting algal mat material (left) in direct, perpendicular contact with large
(by comparison) tubes of Tetradium (right). In the algal material, note apparent degenera-
tion of irregular, tube-like structures to micrite. X75.
sediments (7, 8, 10). With specific reference to the Saluda coralline zone, it appears
that algae played an important role (co-significance along with corals) in its structural
establishment, in its persistence as an entity in the face of turbulence, and in genesis
of the observed high percentage of contained micrite. The unexpected abundance of
micrite in the coralline zone thus relates directly to the importance of algae in con-
struction of the zone.
Conclusions
An unexpectedly large amount of what is apparently autochthonous micrite occurs
within the coralline {Tetradium) biostrome of the Saluda Formation. Evidence accrued
in this investigation supports the hypothesis that this in situ micrite was derived in
the main from algal mats, through a process of grain diminution of calcareous algal
tubes. Rather than simply representing a coral rubble shoal, this biostrome is the rem-
nant of a coral/algal complex within which fragments of other types of fossils (e.g.,
Ostracodes) were frequently trapped. Encrusting, trepostomous Bryozoa perhaps con-
stituted a minor component of the biostrome.
Literature Cited
1. Browne, R.G. 1964. The coral horizons and stratigraphy of the Upper Rich-
mond group in Kentucky west of the Cincinnati Arch. J. Paleontology 38:385-392.
2. Foerste, A.F. 1903. The Richmond Group along the western side of the Cincin-
nati anticline in Indiana and Kentucky. Amer. Geol. 31:333-361.
3. Hatfield, C.B. 1968. Stratigraphy and paleoecology of the Saluda Formation
(Cincinnatian) in Indiana, Ohio, and Kentucky. Geol. Soc. Amer., Special Paper
95. 34 p.
394 Indiana Academy of Science Vol. 94 (1985)
4. Hattin, D.E. 1961. Notes on Richmondian stratigraphy in Southeastern Indiana,
p. 328-337. In Guidebook for Field Trips, Cincinnati Meeting, Geol. Soc. Amer.
5. Hay. H.R. 1977. Field trip No. 1— Cincinnatian stratigraphy from Richmond
to Aurora, Indiana, p. 1-1 to 1-33. In J.K. Pope and W.D. Martin (eds),
Biostratigraphy and paleoenvironments of the Cincinnatian Series, southeastern
Indiana. Guidebook, 7th Ann. Field Conference, Great Lakes Section, Soc. Econ.
Paleontologists and Mineralogists.
6. Martin, W.D. 1975. The petrology of a composite vertical section of Cincinna-
tian Series limestones (Upper Ordovician) of southwestern Ohio, southeastern
Indiana, and northern Kentucky. J. Sed. Pet. 45:907-925.
7. Pettijohn, F.J. 1975. Sedimentary Rocks (third ed.). Harper & Row Publ., New
York, Evanston, San Francisco, and London. 628 p.
8. Stockman, K.W., Ginsburg, R.N. and Shinn, E.A. 1967. The production of
lime mud by algae in South Florida. J. Sed. Pet. 37:633-648.
9. Van Hart, D. 1966. The Physical Stratigraphy of the Saluda and Whitewater
Formations (Cincinnatian Series), Southeastern Indiana. M.S. Thesis, Miami Univ.,
Oxford, OH. 142 p.
10. Wolf, K.H. 1965a. Gradational sedimentary products of calcareous algae. Sedimen-
tology 5:1-37.
11. Wolf, K.H. 1965b. "Grain-diminution" of algal colonies to micrite. J. Sed. Pet.
35:420-427.
12. Wray, J.L. 1977. Calcareous Algae. Elsevier Scientific Publ. Co., Amsterdam,
Oxford, New York. 185. p.
HISTORY OF SCIENCE
Chairperson: Gene Kritsky
Department of Biology
College of Mount St. Joseph
Mount St. Joseph, Ohio 45051
(513)244-4401
Chairperson-Elect: Gerald Seeley
Department of Civil Engineering
Valparaiso University
Valparaiso, Indiana 46883
(219)464-5120
The Rich and Varied Past of the History of Science Section
Barbara A. Seeley
805 Hastings Terrace
Valparaiso, Indiana 46383
and
Gerald R. Seeley
Department of Civil Engineering
Valparaiso University
Valparaiso, Indiana 46383
This year celebrates the centennial meeting of the Indiana Academy of Science.
In addition, this is the 40th anniversary of the first meeting of the History of Science
Section. Naturally such events call for reflection.
The History of Science is a fairly new section among the many which comprise
the Indiana Academy of Science (IAS). It is natural that histories are not attempted
until after a rich tradition already has taken root. As with any "infant," the formative
years are especially important if that "infant" is to grow into a productive "adult."
It is in this light that we look upon the early history of the section.
The IAS was already 59 years old when the minutes of the October 28, 1943
Executive Committee meeting stated the following (5):
"A recommendation was made that a chairman of a committee be appointed
for the consideration of plans for a History of Science in Indiana, including the
biographies of Indiana scientists. Said committee is to consist of a chairman,
and a member from each of the sections of the Academy. The most feasible plan
is to be presented at the next meeting of the Academy." "Professor CO. Lee
(Purdue) was elected to solicit papers concerning the history of the different fields
of science, which are to be presented at the 1944 Academy meeting in a section
on the History of Science."
At the 1944 meeting, held appropriately at Butler University, W.E. Edington
(DePauw University) presented plans for the History of Science in Indiana, including
biographies of Indiana scientists (6). John S. Wright was elected Section Chairman
for 1945.
Looking at the papers presented at the first session in 1944, one sees that the
section made a fine start with addresses to the general assembly entitled "A Historian
Views Science," by Louis Sears and "A Critique of Science" by Carroll Hildebrand.
395
396 Indiana Academy of Science Vol. 94 (1985)
The papers presented at the section include three authors whose names quickly
become familiar as one views the early years of the History of Science Section. Those
first three authors were B. Elwood Montgomery, Paul Weatherwax, and John S. Wright.
The paper by John Wright is entitled "Men of Science in Indiana, Past and Pre-
sent." It is very appropriate that we meet John Wright in this fashion since the previous
year he was given tribute for being a member of IAS for 50 years with active interest
in the Academy throughout the entire period. John Wright joined the Academy in
1893, one year after receiving his BS degree from Purdue and joining Eli Lilly as a
botanist. His interests included medical botany, histology of drugs and food, phar-
macology of plant drugs, and in later years, conservation and forestry. He was secretary
of the Academy from 1895-1904, becoming President in 1905. The 50th anniversary
of the Academy saw his active participation leading to a continued interest in the History
of Science. In his retirement from active service at Eli Lilly, he was able to pursue
this interest by advocating the publication of a historical Directory of Science for Indiana
which eventually culminated in the publication of the volume Indiana Scientists by
the IAS. As stated in the IAS tribute to him at the time of his death in 1951, (4)
"he represented the tie with the 'Giants of Other Days' for he knew them all and
he had actively served the Academy longer than any other member in its history." "He
has left an imperishable mark on the IAS and he will henceforth take his rightful
place as one of the 'Giants of Other Days."'
Another author of 1944 was Paul Weatherwax, a graduate and Professor of Botany
at Indiana University. His contributions to the History of Science Section included
several articles relating to his prime interest in the history of Indian domesticated corn.
A worldwide authority in this area, he traveled widely seeking the wild ancestor of
Indian domesticated corn. Concluding that the original ancestor was extinct, he served
on a committee of the National Research Council to preserve extant varieties contain-
ing primitive characteristics which might be needed to re-develop resistant corn varieties
for future needs. Dr. Weatherwax was a member of IAS for 63 years serving as Presi-
dent of the Academy in 1941 and as Chairman of the History of Science Section in
1949 (1). In 1966 he gave the invited paper, "Indiana Botany in Retrospect" as part
of the Academy's Symposium celebrating the Indiana State Sesquicentennial. As noted
by the editor of the symposium (2), "it is a signal honor and a mark of respect and
confidence for these men to have been chosen to write the history of their own fields
in Indiana. The collected papers published herein comprise a unique contribution to
the history of science in Indiana by those who know it best and who have helped
to make some of that history as well as write it."
Our third author of 1944, B. Elwood Montgomery (Purdue) who was elected
Fellow in 1929 contributed papers to the section over the longest time span, that being
from 1944 through 1981. His contributions covered odonatology in Indiana and America,
the domestication of bumblebees, Thomas Say Entomologist, Linnean "Elements"
in Indiana fauna and flora, the Cumberland Road, the origin and derivation of insect
names and entomological terms, and a Bicentennial Study of Indiana fauna. He served
as Chairman of the Section in 1955 and 1969.
The meeting of 1945 introduces us to the most prolific author of the History
of Science Section, Stephen S. Visher of Indiana University. Dr. Visher contributed
14 articles from 1945-1966. Being one of the nations outstanding geographers, he con-
tributed several articles in this area, including his invited paper, "A Brief History of
Geography in Indiana" for the Indiana Sesquicentennial celebration in 1966. However,
his major contribution to the History of Science included many articles on the con-
tributions and achievements of scientists in Indiana, chronicling Indiana Nobelists and
National Academy members, and searching for a key to the success of outstanding
persons with regard to their environment and geographical origins. He was editor of
History of Science 397
Indiana Scientists, a biographical directory and analysis which was published by IAS
in 1951. He served as Section Chairman in 1948 and 1959. Dr. Visher's Presidential
address of 1950 (7) contained conclusions regarding the production of this valuable
resource known as scientific leadership, conclusions which are important for us to
recall today. He viewed encouragement by one's family as highly significant in early
years. He emphasized that no scientist is self-made and that personal encouragement
by enthusiastic, stimulating teachers is deeply significant in the development of scien-
tific leaders. As evidence of this connection he noted the large number of respected
scientists trained by such great Indiana teachers as zoologist David Starr Jordan and
botanist John M. Coulter. He encouraged his colleagues and each of us today to "be
generous in encouraging our more promising students and young friends. A few
appreciative words may alter their life!"
Another long-term participant in the History of Science Section was William E.
Edington, head of Mathematics and Astronomy at DePauw University. In 1944 as
chairman of the committee, he presented plans to the Executive Committee of IAS
for the History of Science in Indiana, including biographies of Indiana scientists. He
was President of the Academy in 1937 and Chairman of the Section in 1946. He presented
papers at the section from 1948 to 1973 on topics as diverse as The Wabash Academy
of Science, the Terre Haute Scientific Society, The History of Science at DePauw,
David Starr Jordan, John P.D. John, William Ephraim Heal, and Indiana Women
in Mathematics. He was invited to join the ranks of the illustrious scientists participating
in the 1966 Indiana Sesquicentennial Symposium with his paper entitled "Mathematics
in Indiana 1816-1966, From the Rule of Three to Electronic Computers."
However, Dr. Edington's greatest single contribution to the History of Science
in Indiana was made long before the creation of the History of Science Section. He
presented a paper (3) at the 50th meeting of the IAS in 1934 honoring the founders
and charter members of the IAS entitled "There Were Giants in Those Days." This
fascinating history chronicles the influence of four distinct factors on the foundation
of the IAS: 1) the influence of the New Harmony scientific community, 2) the develop-
ment of geological investigation, 3) the influence of Louis Agassiz, and 4) the inspira-
tion derived from the American Association for the Advancement of Science. Dr.
Edington's discussion of the founding members of IAS in this paper certainly was
an appropriate beginning to his work in compiling the achievements of the contributors
to science in Indiana. Over the next years Dr. Edington contributed papers on charter
members of IAS, supplied considerable material for the book Indiana Scientists (Visher
1951) and wrote memorials for the IAS for 35 years. By so doing, in his 53 years
as a member of IAS, he had probably written more than anyone else on the history
of the Academy.
In addition to the four "giants" mentioned previously, the section continued to
attract talented leaders. These men not only served as section officers and/or Academy
officers, they have had a long association with the Academy and were prolific in the
number of papers presented at History of Science Section meetings.
The 1946 meeting brought two new contributors to the section, Charles A. Behrens
and Raymond E. Girton of Purdue. Dr. Behrens gave us The History of the First
Five Years of The IAS, the Purdue Biological Society, development of medical
bacteriology, and the landmarks in chemotherapy. He served as Academy President
in 1923 and Section Chairman in 1947.
In 1946, Raymond Girton began a 29 year tradition of contributions including
articles on developments in plant physiology, early studies in protoplasm, Joseph Priestly,
17th Century microscopists, plant physiology at Purdue in the 19th Century, 3/4 Cen-
tury of Biology at Purdue, and a Look at Academy Presidential Addresses. Prof. Gir-
ton served as Academy President in 1956 and as Section Chairman in 1951 and 1952.
398 Indiana Academy of Science Vol. 94 (1985)
CO. Lee was chairman at the first meeting of the History of Science Section
in 1944. Over the next 10 years he presented papers on the history of the School of
Pharmacy at Purdue and on the American Pharmaceutical Association form 1852-1952.
C.L. Porter presented papers from 1947-1952 including the topics of Botanists
of Purdue, Johnny Appleseed, the history and economic importance of Mentha piperita
(mint), and the history of fungus antibiosis. A 43 year member, Dr. Porter served
as President of the Academy in 1949 and Section Chairman in 1953.
In 1947 the name of William J. Tinkle appears which is to span 25 years in the
section from 1947-1973. His papers discuss various aspects of Darwinism, conserva-
tion of germ plasm, natural selection, creationism, and a profile of J. Henri Fabre.
He served as Section Chairman in 1956.
Another long term contributor was M.S. Markle of Earlham who presented papers
from 1953-1966 on the History of Science at Earlham, Dr. John T. Plummer, the
Joseph Moore Museum at Earlham, and the influence of Quakers on Science in Indiana.
He gave an invited paper at the Sesquicentennial symposium entitled "The History
of Plant Taxonomy and Ecology in Indiana." During his 58 year membership in IAS
he served as President of the Academy in 1945 and Section Chairman in 1954.
Daniel DenUyl, a contributor of more recent years, presented papers from
1953-1958 on the Civilians Conservation Corps, Charles C. Deam, forest conservation
in Indiana, and the forests of the Lower Wabash bottomlands. He served as Section
Chairman in 1957.
We have still as members today four contributors who have been members of
the Academy for 50 years or more who have also served us well in the History of
Science Section. All from Purdue, they are H.H. Michaud, M.G. Mellon, Arthur T.
Guard, and Raymond E. Girton whom we discussed earlier.
Prof. Michaud has presented papers concerning conservation of natural resources,
conservation of recreation and scenic resources, history of game regulations, and the
history of science education in Indiana high schools. He served as Academy President
in 1963 and as Section Chairman in 1958.
M.G. Mellon contributed an article on developments in the analytic balance and
was invited to present a paper on "Chemistry in Indiana at the States Sesquicenten-
nial." As President of the Academy in 1942 his address was entitled "Science, Scien-
tists, and Society." In addition he served the Section as Chairman in 1950.
Arthur T. Guard has served us as Section Chairman in 1963 and 1964 and as
Academy President in 1960. His papers include his Presidential address on "Recent
Approaches to the Study of Plant Structure" and his section presentations on "Early
Field Trips of the Indiana Academy of Science" and "John and William Bartram —
Botanists at the Time of The Nation's Birth."
The important task of recording the lives of the men and women who have shaped
the Academy's past has been ably assumed by Fay K. Daily (Butler University), our
Academy Necrologist. She has served as Chairman of the Section in 1960 and has
presented papers on Botanists of Butler University 1920-1955, some scientific expedi-
tions in the SE US taken by David Starr Jordan and an address at the 75th anniversary
of the IAS entitled "The Academy from Horse and Buggy to Jet." Most recently
she coauthored the History of the Indiana Academy of Science 1885-1984, A Centennial
Volume.
In reading the minutes of meetings, papers presented, and memorials to those
who have preceded us, it is striking to see the depth of contributions made by these
individuals. The words "friend and benefactor of the Indiana Academy of Science"
certainly apply.
In closing, we would like to turn again to words of William E. Edington, a great
chronicler of the Academy. He concluded his address to the 50th meeting of the Academy
History of Science 399
entitled, "There Were Giants in Those Days," as follows (3): "And so I come to
the conclusion. I hope this recital of illustrious names of those who have done so
much for science in Indiana and our nation, names of scientists who were once active
in our Academy as we are active today, will inspire the younger scientists of Indiana
to attempt to follow in their footsteps. Indiana produced giants in those days. It is
my hope that when the centennial meeting of our Academy is celebrated in 1984, someone
speaking in authority may say there were giants in our days."
Literature Cited
1. Daily, F.K. 1977. Necrology for Paul Weatherwax. Proc. I.A.S. 86:63-65.
2. Eberly, W.R. 1967. The History of Indiana Science. Proc. I.A.S. 76:64.
3. Edington, W.E. 1935. There were giants in those days. Proc. I.A.S. 44:22-38.
4. 1952. Necrology for John Shepard Wright. Proc. I.A.S. 61:30-32.
5. Indiana Academy of Science 1944. Minutes of the Executive Committee, Oct.
28, 1943. Proc. I.A.S. 53:XI-XII.
6. 1945. Minutes of the Executive Committee, Nov. 10, 1944. Proc. I.A.S.
54: XI.
7. Visher, S.S. 1951. Indiana Scientists. Proc. I.A.S. 60:29-36.
MICROBIOLOGY AND MOLECULAR BIOLOGY
Chairperson: J.R. Garcia
Department of Biology
Ball State University
Muncie, Indiana 47306
(317)284-4045
Chairperson-Elect: Mary Lee Richeson
Department of Biological Sciences
Indiana University-Purdue University at Fort Wayne
2101 Coliseum Boulevard East
Fort Wayne, Indiana 46805
(219)482-5546
ABSTRACTS
Effect of Cyclosporine A on Leishmania tropica. Nancy C. Behforouz, Department
of Biology, Ball State University, Muncie, Indiana 47306. The effect of Cyclosporine
A, a new immunosuppressive and antiparasitic drug was tested, both in vivo and in
vitro, on Leishmania tropica. In vitro, the drug inhibited growth of the parasite and
decreased the infectivity of the organism. Although this drug appeared to have little
or no therapeutic effect for susceptible, infected mice at the doses tested, it had a
significant, dose-dependent prophylactic effect when used two days prior and five days
following infection.
The Regulation of S-Adenosylmethionine Synthetase in Candida albicans. Richard
H. Lambert, Eli Lilly and Company, Indianapolis, Indiana 46285 and J.R. Garcia,
Ball State University, Muncie, Indiana 47306. S-Adenosylmethionine (SAM) syn-
thetase from yeast and hyphal-phase cells of the dimorphic fungus C. albicans was
characterized by kinetic analysis and response to inhibitors. SAM Synthetase is the
enzyme responsible for the synthesis of S-Adenosylmethionine (SAM), the compound
which serves as the major methyl-group donor in the methylation of macromolecules
such as DNA, RNA, and proteins. The enzyme from yeast-phase cells has a km of
0.17 mM for methionine, 0.14 mM for ATP, and is inhibited (in vitro) by dimethyl-
sulfoxide, methionine sulfone and methionine sulfoxide. They hyphal-phase SAM syn-
thetase has a km of 0.056 mM for methionine, 0.02 mM for ATP, and its activity
(in vitro) is enhanced by the inhibitors used with the yeast-phase enzyme. This preliminary
data strongly suggests that isozymes of SAM Synthetase are present in C. albicans
and possibly that the isozymes are morphology-specific.
The in vivo studies revealed that the enzyme's synthesis is repressed by the addi-
tion of methionine and that the specific activity increases during a temperature-induced
shift in morphology. In addition, it was shown that the increase in specific activity
(seen during a yeast — hyphae shift and/or when yeast cells, grown in a methionine-
supplemented medium, are transferred to a methionine-free medium) involves de novo
protein synthesis.
A Case of Tuberculosis in the University Setting. M. Langona, Department of
Epidemiology, Ball Memorial Hospital, Muncie, Indiana 47303. Since the 1970s
the United States Public Health Service has worked extremely hard in preventing the
401
402 Indiana Academy of Science Vol. 94 (1985)
transmission of communicable and infectious diseases within this country by Asian
refugees. Mandatory health screening tests for tuberculosis, leprosy, venereal disease
and other medical conditions have been provided while the refugee is still abroad,
and then again upon arrival at various U.S. ports of entry. Unfortunately, Asians
who are not refugees may immigrate into this country without appropriate healths testing
and may represent a public health problem.
This presentation will describe a case of pulmonary and extrapulmonary tuber-
culosis diagnosed in a young, pregnant Korean who recently arrived in Indiana with
her spouse who is a foreign-exchange university student. Unfortunately, the univer-
sity's health policy only required tuberculosis skin testing of the enrolled student and
not the spouse. Information will be provided about the diagnosis, epidemiologic workup,
hospitalization of the tuberculosis patient, and the dichotomy of the public health
regulations.
Scabies: A Nosocomial Outbreak. M. Langona, S. Bossung, and M. Orr, Depart-
ment of Epidemiology, Ball Memorial Hospital, Muncie, Indiana 47303. Sarcoptes
scabiei (var. hominis) an obligate ectoparasitic mite of humans continues to present
itself as a health problem within the United States. Although scabies is a non-reportable
disease and reliable data on its incidence is limited, several investigators as well as
the Centers for Disease Control report that the United States is experiencing the most
significant increase in scabietic infestations since the epidemics of World War II.
This presentation will describe a 1984 epidemic of Norwegian (crusted) Scabies
which involved the admission of a nursing home patient into a community-teaching
hospital and the subsequent nosocomial scabies outbreak of 15 hospital personnel and
their families.
The suspicion of scabies with supportive clinical and laboratory findings war-
rants control measures, and dependent on the form of scabies present, the immediate
and efficacious epidemiologic investigation within the hospital setting.
Although the 20th century clinician possesses a simple and effective cure for scabies
infestations, it is indeed disheartening that we lack the ability to eradicate this nuisance
mite.
Three Plasmid Cloning Vectors for Mammalian Cells. Steven H. Larsen and Joann
Hoskins, Department of Microbiology and Immunology, School of Medicine, Indiana
University-Purdue University at Indianapolis, Indianapolis, Indiana 46223. Plasmid
vectors based upon selection of the dominant phenotype of resistance to the G418
antibiotic have been developed. To provide this resistance, the coding sequence for
the Tn5-derived aminoglycoside phosphotransferase activity were sandwiched between
the promotor and polyadenylation signals of the thymidine kinase gene from herpes
simplex virus. This construct was placed into ampicillin or ampicillin-tetracycline resis-
tant derivatives of pBR322. One such construct, pSL72, can be stably selected in mouse
L293 cells at an efficiency equal to any previously known system (greater than 0.1%
of the cells). This plasmid appears to be selectable at a single copy per ceil. A second
vector, pSL71, is quite similar except that the copy number can be increased to about
100 genomes per animal cell. The third plasmid includes mouse cell DNA sequences
which provide the plasmid with the ability to be maintained extrachromosomally and
hence recovered again from the animal cell into bacteria.
Banking DNA for Future Diagnosis of Hereditary Diseases. Linda Madisen and M.E.
Hodes, Indiana University School of Medicine, Indiana University-Purdue University
at Indianapolis, Indianapolis, Indiana 46223. Recombinant DNA methodology
Microbiology and Molecular Biology 403
is becoming increasingly important for the detection of the carrier state of a number
of genetic diseases. After generating a series of restriction fragment length polymor-
phisms closely linked to a gene causing a disease, it is possible to predict whether
an individual has inherited the haplotype associated with the deleterious gene. Such
studies require DNA from informative relatives as well as from affecteds and so will
require the long term storage of highly polymerized DNA, a relatively new procedure
whose limitations are still being investigated.
By storing DNA at temperatures above 4°, one may cause accelerated aging and
thus mimic long term storage. We found that DNA stored in solution at -70°, -20°,
4°, 25° and 37°C for two months remains high molecular weight. Early results indicate
these different storage temperatures have no effect on restriction enzyme banding pat-
terns for Xbal, Hindlll and EcoRI. Similar incubation of the DNA at 65°C resulted
in extreme degradation. Furthermore, blood stored at -70°C for two months prior
to extraction generally yielded a quantity of high molecular weight DNA comparable
to fresh samples. Occasional frozen samples, however, yielded considerably lower DNA
quantities, all of which were high molecular weight.
An Examination of 495 Splice Junction Sequences. F.H. Norris, Eli Lilly and Com-
pany, Indianapolis, Indiana 46285, and M.E. Hodes, Indiana University School of
Medicine, Indiana University-Purdue University at Indianapolis, Indianapolis, Indiana
46223. We have performed a computer aided examination of 495 of the exon-
intron junctions reported in the June, 1984, Genetic Sequences Databank (GenBank).
We examined the junction data as a pool and also segregated according to organisms
in which they occur. The consensus sequence we found, 5'-(AC)AG/GT(AG)AGT,
is the same as that reported by Mount and others. We also find that, except for the
A or G at position +3, conservation of the sequence is highest near the splice point
and drifts with distance, with the bases on the intron side of the junction being more
highly conserved. The nine bases indicated by the consensus sequence seem to define
the junction, since we find no conserved bases within 60 bases of the splice site. Beyond
the -3 and +6 boundaries, the bases are randomly distributed.
The frequencies of occurrence of the junction sequences were tabulated. We find
no differences between the human-ape sequence frequencies and the frequencies of
the other mammalian sequences. Striking differences appear as one compares sequences
from higher and lower organisms. Over 60% of the human, but less than 40% of
the non-mammalian vertebrate sequences, are of the form /GT(AG)AG. More than
a third of the lower vertebrate junction sequences occurred one time. Perhaps because
of a host-virus relationship, we see fewer differences between human and viral than
between viral and lower vertebrate sequences.
Transcriptional Regulation of the Sporulation-specific Glucoamylase of Saccharomyces
cerevisiae. Tom Pugh and Mary Clancy, Department of Microbiology, University
of Notre Dame, Notre Dame, Indiana 46556. Sporulating cells of the yeast, Sac-
charomyces cerevisiae contain a glucoamylase activity (SAG) which is distinct from
similar enzymes found in vegetative cells. The enzyme is a glycoprotein and is capable
of releasing free glucose from maltotriose, maltodextins, amylose and glycogen, but
maltose is hydrolyzed slowly, if at all. The time of appearance of SAG activity during
sporulation corresponds to the onset of glycogen degradation, and immediately pro-
ceeds spore formation.
We have been interested in differential gene expression in sporulating yeast and
would like to know the level at which regulation of SAG activity occurs. SAG expres-
sion is prevented if cycloheximide is added to sporulating cells at any time before full
404 Indiana Academy of Science Vol. 94 (1985)
levels are attained. Antibody prepared against 400-fold purified enzyme specifically
precipitates a protein of 68K daltous from extracts of sporulating cells which have
been pulse-labelled in vivo 35S-methionine. This band is not detected at early times
in sporulation or in non-sporulating cells. This shows that the regulation of SAG activity
is not post-translational and suggests that control may be transcriptional.
We have constructed a library of S. cerevisiae DNA in the expression vector,
pBD6, and are screening for the SAG gene, using a plate assay and antibody techniques.
Development of a Model System for the Study of Murine Leukocyte Chemiluminescence.
James L. Shellhaas, Butler University, Indianapolis, Indiana 46208. A model
system was developed for the determination of the activation kinetics of murine peripheral
blood polymorphonuclear neutrophils (PMN's). Utilizing discontinous density gradient
centrifugation and dextran sedimentation, populations of PMN's were prepared of
98% purity. These cell populations were then examined for their ability to respond
with luminol-dependent chemiluminescence upon co-cultivation with the chemotactic
peptide N-formylmethionine-leucine-phenylalanine (Fmet), the tumor promotor phorbol
myristic acetate (PMA), and opsonized zymosan. Purified populations of PMN's were
also examined for their responsiveness in chemotactic assays to each of the stimulation
agents. Significant differences between the chemiluminescent kinetics of murine cells
and the published kinetics of human cells were observed. Chemotactic responsiveness
also differed in murine cells from that observed in human PMN cell populations.
Relationship between Symptomatic Resistance and Virus Production in Barley Cultivars
Inoculated with Barley Yellow Dwarf Virus. M. Skaria, J.E. Foster and R.M. Lister.
Departments of Botany and Plant Pathology and the U.S. Department of Agriculture
(Foster, Purdue University, West Lafayette, Indiana 47907. Resistance to barley
yellow dwarf virus (BYDV) disease has been identified in some Ethiopian barleys. A
genetic factor, the "Yd2" gene associated with symptomatic resistance has been trans-
ferred to several barley cultivars. Few such barleys are available as near-isogenic pair
with the only difference in presence or absence of the Yd2 gene. We investigated the
effect of the Yd2 gene on virus synthesis in three near-isogenic barley pairs. One week
old plants of California Mariout (Yd2 - ) barley and the near-isogenic CM 67 (YD2 + )
were inoculated with PAV, MAV, or RPV isolates of BYDV (i.e. transmitted by
Rhopalosiphum padi L. and Sitibion avenae (Fabr.; by S. avenae; or by R padi, respec-
tively). Inoculated plants were grown in a growth chamber at 20 ± 1°C. The virus
content of shoots and roots was assessed at six day intervals for one month by enzyme-
linked immunosorbent assay (ELISA). With PAV, overall significantly less virus was
detected in CM 67 than in California Mariout, but with MAV and RPV there were
no such differences. In other experiments PAV production behaved similarly in Prato
(Yd2 + ) barley and the near-isogenic Briggs (Yd2 - ), and in Atlas 68 (Yd2 + ) barley
and the near isogenic Atlas 57 (Yd2-). Thus, symptomatic resistance to BYDV in
barley correlates with reduced virus synthesis.
Serum Hormone Levels in Germfree and Conventional Rats: Effect of Dietary Restric-
tion. David L. Snyder and Bernard S. Wostmann, Lobund Laboratory, University
of Notre Dame, Notre Dame, Indiana 46556. Germfree* rats were used to obtain
background information on the relationship between aging, hormone levels, and restricted
dietary intake. Blood samples were obtained by heart puncture from 14 conventional
"Actinomyces sp. had previously contaminated the isolators of these GF rats. However, fecal smears did
not indicate growth of these organisms in the intestinal tract.
Microbiology and Molecular Biology 405
(CV), 27 germfree (GF), and 12 germfree but restricted (GR) Lobund-Wistar rats. Intake
for the restricted rats was 70% of ad lib. intake. All rats were males, 8 to 12 months
old, and fed natural ingredient diet L485. Samples were collected between 10 A.M.
and 12 P.M., under halothane anesthesia, and after an overnight fast. GF rats had
slightly lower serum insulin than CV rats (52.9 vs. 62.6 uU/ml) but GR were significantly
(P < 0.01) lower than GF (52.9 vs. 35.2 uU/ml). Serum glucose levels paralleled in-
sulin levels (CV:140; GF:114; GR:98 mg/dl). No significant differences were found
in total thyroxine (T4) levels (CV:6.2; GF:5.5; GR:5.5 ug/dl) and in total triiodothyronine
(T3) levels (CV:115; GF:134; GR:133 ng/dl). Significant differences were found among
the testosterone (T) levels. GR rats had higher (P < .02) T levels than Cv rats (3.4
vs. 2.1 ng/ml). GR rats had higher (P < .01) T levels than GF rats (7.8 vs. 3.4 ng/ml).
The reduction in insulin levels in GR rats may be a response to lower caloric intake
and an effort to maintain glucose levels through gluconeogenesis. Other possible fac-
tors affecting insulin and glucose levels are the lower metabolic rate of GF animals,
changes in thyroid hormones, and the anabolic effects of testosterone. Though body
weights of GR rats were only 72% of GF rats, GR rats maintained testes sizes similar
to GF rats. However, this could account only in part for the higher serum T concen-
trations of GR rats. Our findings suggest that dietary restriction of even 30% is enough
to modify hormone patterns which in turn may lead to the extended lifespan observed
in these animals.
Control of Cell Growth by Transplasmalemma Redox: Stimulation of HeLa
Cell Growth by Impermeable Oxidants
I.L. Sun, J.E. Putnam, and F.L. Crane
Department of Biological Sciences
Purdue University
West Lafayette, Indiana 47907
Introduction
Very little attention has been paid to energy rich redox agents, such as NADH,
in contact with the interior of the plasma membrane. The energy source for plasma
membrane functions such as vesicle formation and movement or transport, is always
considered to be ATP derived from mitochondria or cytoplasmic glycolysis. However,
the presence of high and low redox-potential compounds at the plasma membrane
means that energy should be available.
A transplasma membrane redox enzyme which transfers electrons from reducing
agents in the cytoplasm to external impermeable oxidants, such as ferricyanide, is pre-
sent in all cells which have been tested (1, 4, 25, 26). This redox activity has been
found to be related to several vital functions which include control and stimulation
of cell growth (7), facilitation of iron uptake (3, 18, 20, 28) and defense against bacteria
(14). In addition there is also good evidence that this redox enzyme is hormone sen-
sitive (2, 6, 11, 13), driving amino acid transport (9), including proton release (13),
and controlling adenylate cyclase activity (12). These are indications that this enzyme
has an important role in the control of cellular functions.
In this study we present evidence that impermeable electron acceptors for the
transplasma membrane redox system stimulate the growth of HeLa cells in a serum
free medium. Insulin (30/ig/ml) enhances this growth stimulation and increases the
rate of oxidant reduction by cells. Impermeable oxidants, which do not interact with
the electron transport system, do not stimulate growth. The coupling of proton release
to this electron transport indicates that local membrane energization is affected by
transmembrane electron flow and that intracellular pH may change. We propose that
such activation and the increase of cytoplasmic pH can be very important in cell growth.
Materials and Methods
HeLa cells were grown in flasks with Eagle's medium containing 10% fetal calf
serum, 100 u. of penicillin and 170 ug streptomycin per ml at pH 7.4 and maintained
in a similar medium containing 2% fetal calf serum. Cells were prepared for study
by pelleting the trypsinized suspension cultures at 27,000 g. The pellet was diluted
with TD-Tris buffer (NaCl 8g/l, KC1 0.34g/l, Na2HP04 0.1 g/1 and Trisma base 3g/l,
pH 7.5) to a final concentration of 0.1 gm cells/ml.
Growth of HeLa cells with supplements in serum free media was carried out with
cells harvested during the exponential growth phase. Insulin, ferricyanide or other ox-
idants can replace fetal calf serum as a growth factor for the replication of HeLa
cells. Cells were grown in a serum free medium. A final concentration of 0.01-1.0
mM of ferricyanide or other oxidants and 30 /xg/ml of insulin were used as supplements
for cell growth. After 2 days of incubation at 37°C, cells were harvested and a cell
survival count was taken immediately. Survival was determined by the eosin Y exclu-
sion method as described by Mighell and Shrigi (15). The colorless viable cells were
counted. Cell number was determined by counting with a hemacytometer. Cell counts
were obtained in duplicate with a cell counter after trypsinization.
407
408
Indiana Academy of Science
Vol. 94 (1985)
The rate of ferricyanide reduction by HeLa cells was determined in an Aminco
DW-2a dual beam spectrophotometer with a linear recorder, a cuvette stirrer, and a
37° temperature controlled cuvette chamber. The assay of ferricyanide reduction was
performed as described previously (5), except TD-Tris buffer instead of 0.05 M sodium
phosphate buffer, pH 7.0, was used. Absorbance changes were measured with the dual
beam at 420 nm minus 500 nm. The extinction coefficient for ferricyanide reduction
AA420 equals 1.0 mM-,.cm-1.
The reduction rate of other oxidants were measured as described above except
bathophenanthroline sulfonate (BPS) (3.3 /*M) and ferric chloride (0.33 fiM) were add-
ed into the assay mixture. Absorbance changes were measured with the dual beam
at 535 nm minus 600 nm. The extinction coefficient for oxidant reduction was based
on the formation of ferrous-BPS at AA535 which equaled 17.6 mM-lcm_l.
Oxygen uptake was measured with an oxygen electrode in 1.3 ml TD-Tris buffer
with 1 mM potassium cyanide. 0.35 fiM NADH and 0.05 gm wet weight of cells were
added to start the reaction.
Results
The impermeable electron acceptor, potassium ferricyanide, stimulates the growth
of HeLa cells under the conditions of serum deprivation as shown in Figure 1. At
concentrations 0.033 to 0.1 mM ferricyanide gives an optimum stimulation of growth,
which shows a 2-3 fold increase in cell count over the control. At concentrations above
12 r
10
X
CO
Q
E
I-
Ll)
a.
CO
-J
_l
LU
O
8 -
6 -
2 -
0
•7
-//-
0.01 0.033 0.1 0.33
LOG FERRICYANIDE (mM)
1.0
Figure 1. Stimulation of the growth of HeLa cells by ferricyanide. Cells were grown
in a serum free medium. Cell count was determined after cells grown for 48 hr. at 37°C.
Microbiology and Molecular Biology
409
0.1 mM, ferricyanide becomes less effective. Cytotoxicity is found at ferricyanide con-
centrations over 1 mM which inhibits cell growth. Additive growth effects are seen
with limiting levels of serum and ferricyanide up to the maximum growth with 10%
of fetal calf serum. In general, lower ferricyanide (0.01 mM) requires higher serum
(8°7o) and higher ferricyanide requires lower serum (4%) to reach maximum growth
(Figure 2).
0 2 4 6 8 10
CONCENTRATION OF FETAL CALF SERUM (%)
Figure 2. Dose-response growth curve to HeLa cells to serum supplemented with
ferricyanide. O O O, with ferricyanide 0.033 mM; • • • with ferri-
cyanide 0.01 mM; A A A, with ferricyanide 0.1 mM and □ □, with
ferricyanide 0.33 mM. 48 hr. culture.
Sodium ferricyanide stimulates growth and attachment of serum deficient HeLa
cells as well as potassium ferricyanide (Figure 3). However, potassium ferrocyanide,
the reduced form of ferricyanide, does not promote growth (Figure 4). The internal
oxidant pyruvate does not stimulate growth either (Table 1). These results indicate
that transmembrane electron flow must be involved in providing energy for cell func-
tion, since ferricyanide is extracellular and cannot itself provide nutrients for the cell.
410
Indiana Academy of Science
Vol. 94 (1985)
7r
O 5
x
<
E
o
m
cvi 3
if)
-j
-i
LlI
O
1 -
0
c\
7A
0 0.0033 0.01 0.033
0.1
0.33
1.0
LOG CONCENTRATION (mM)
Figure 3. Dose-response growth curve of HeLa cells to serum free medium sup-
plemented with potassium ferricyanide or sodium ferricyanide. O O O, sodium
ferricyanide and • • •, potassium ferricyanide. 48 hr. culture.
Microbiology and Molecular Biology
411
4r
10
i
O
if)
<
CO
E
o
lO
C\J
\
if)
UJ
o
VA
0 0.001 0.0033 0.01 0.033 0.1
LOG CONCENTRATION (mM)
0.33
Figure 4. Dose-response growth curve of HeLa cells to serum free medium sup-
plemented with potassium ferricyanide or potassium ferrocyanide. O O O,
potassium ferricyanide and • • • potassium ferrocyanide. 48 hr. culture.
412 Indiana Academy of Science Vol. 94 (1985)
Table 1. Effect of pyruvate and ferricyanide on the growth of HeLa cells.
Addition No. cells/25 cm2 flask
(X io"5)
Control 2.0
Pyruvate (0.01 mM) 2.1
Pyruvate (0.1 mM) 2.3
Pyruvate (1 mM) 1.4
Ferricyanide (0.033 mM) + pyruvate (0.01 mM) 3.3
Besides ferricyanide, other impermeable oxidants, such as hexaamine-ruthenium
III chloride and indigotetrasulfonate, which increase oxygen uptake, also stimulate
cell growth. Inactive oxidants, such as cytochrome c, do not promote cell replication
(Table 2). The growth response seems to be specific for active impermeable oxidants.
Table 2. Effects of other impermeable oxidants on the growth and transmembrance
redox system of HeLa cells.
Addition
No. cells/25cm2
flask
0^ upta
ke
Reduction rate
(nmoles 0,/min
/g.w.w.)*
(nmoles ferrous
-BPS/min/g.w.w.)
Control
1.74 x 105
138
0
Hexaamine-ruthenium 111
chloride (0.33mM)
4.76 x 10'
208
9.6
Hexaamine-ruthenium III
chloride (O.lmM)
16.1
Indigotetrasulfonate (0.01 mM)
8.62 x 10'
173
11.8
Indigotetrasulfonate (O.lmM)
15
Cytochrome c (l.OmM)
1.65 x 10'
0
Cytochrome c (3.0mM)
1.50 x 105
_
*g.w.w. indicates wet weight of cells
The application of insulin dramatically enhances the stimulating effect of ferri-
cyanide as indicated in Figure 5. Insulin is a well known growth stimulator. At the
optimum concentration (30 jig/ml), which stimulates growth, insulin also greatly in-
creases the rate of ferricyanide reduction by HeLa cells (Table 3). Both the initial fast
rate and long term slow rate of ferricyanide reduction are stimulated (27). There is
a close correlation between insulin and increase in transmembrane redox enzyme activity
and insulin induction of cell proliferation, as shown in Figure 6. The actual mechanism
of insulin action as a growth promoter is not clear. Our results, however, indicate
that electron flow through the transplamsa membrane electron transport system stimulates
growth and that insulin acts to increase that flow.
Discussion
The results of Ellem and Kay for melanoma cell growth on limiting amounts
of serum supplemented with ferricyanide (7) were similar to what we report here for
HeLa cells. Furthermore, Mishra and Passow (16) found that reduction of extracellular
ferricyanide by human erthyrocytes was accompanied by ATP formation, presumably
accomplished as a result of transmembrane electron flow. Recent evidence also sug-
Microbiology and Molecular Biology
413
UJ
o
6 -
O 5
(0
<
-J
Li.
CM
E
o
lO
CVJ
\
CO
3 -
O
-1—/A I l I I I
0 0.0033 0.01 0.033 0.1 0.33
LOG FERRICYANIDE (mM)
Figure 5. Dose response curve to HeLa cells to serum free medium supplemented
with ferricyanide and insulin. ▲ A ▲, without insulin, A A A
with insulin (30 ug/ml). 48 hr. culture.
gests that the electron flow is more important for cell transition from G, to S phase
than is the production of ATP therefrom (19). Therefore, there is considerable evidence
414 Indiana Academy of Science Vol. 94 (1985)
Table 3. The effect of insulin on ferricyanide reduction by HeLa cells.
Concentration of ferricyanide
Specific activity
(nmoles/min/g.w.w.)
0 mM
0 mM + I
0.0033 mM
0.0033 mM +
0.01 mM
0.01 mM + I
0.033 mM
0.033 mM +
0.1 mM
0.1 mM + I
0.33 mM
0.33 mM + 1
Fast rate
Slow rate
0
0
0
0
101
52
208
134
145
40
221
154
214
87
314
158
259
94
334
175
278
118
475
250
I indicates insulin (30 jig/ml)
that the transmembrane electron flow plays an important role in the control of cellular
function. The fact that four of the most used anticancer drugs can inhibit transmem-
brane redox enzyme activities (21-24) further supports this idea.
It has been shown in several cell types that increase of cytoplasmic pH (alkaliniza-
tion) is associated with cell division (8-17). We have previously shown that ferricyanide
induced proton release from HeLa cells (25) in concentrations that coincide with the
concentration, which gives the maximum growth stimulation. The basis for redox stimula-
tion of growth is not quite clear yet. However, it is possible that ferricyanide induced
proton release across the membrane would increase the pH of the cytoplasm and thus
increase cellular mitosis.
Transferrin can act as an electron acceptor for the transmembrane redox system
(Sun and Crane, unpublished). Part of the growth stimulatory effects of transferrin
may be based on an oxidant effect at the cell surface. However, stimulation of growth
by an oxidant is not limited to iron compounds such as ferricyanide or transferrin.
The growth of HeLa cells is also stimulated by hexaamine-ruthenium III chloride, a
trivalent cation and by indigotetrasulfonate (Table 2). The use of a series impermeable
indigo sulfonates with different redox potential shows that extracellular oxidants with
a redox potential E '7 o above -125 mV can stimulate growth (27).
The mode of action for insulin as a growth stimulator is unknown. Our results
that the insulin stimulation of ferricyanide reduction correlates with its promotion of
cell growth plus the evidence that insulin increases proton release from the cell induced
by ferricyanide (Sun and Crane unpublished) suggest that the activation of the redox
system and a stimulation of a redox driven proton pump would be a basis for insulin
action as a growth factor.
Literature Cited
1. Clark, M.G., E.J. Patrick, G.S. Patten, F.L. Crane and G. Grebing. 1981. Evidence
for the extracellular reduction of ferricyanide by rat liver: A transmembrane redox
system. Biochem. J. 200:565-572.
2. Clark, M.G., E.J. Patrick and FL. Crane. 1982. Properties and regulation of
a transplasma membrane redox system in rat liver. Biochem. J. 204:795-801.
3. Cole, E.S. and J. Glass. 1983. Transferrin binding and iron uptake in mouse
hepatocytes. Biochim. Biophys. Acta 762:102-110.
Microbiology and Molecular Biology
415
1800
1600 -
3:
1400
\
c
E
N
1200
o
E
c
w
1000
>%
^—
>
^_
o
<
800
o
H-
o
(1)
600
Q.
if)
400 -
200 -
0.1
0.3 1.0 3
Insulin (pg/ml)
10
30
Figure 6. The correlation between insulin promotion of cell growth and insulin stimula-
tion of transmembrane redox enzyme activities. O O O, slow rate of HeLa
cells ferricyanide reduction; • • • fast rate of HeLa cells ferricyanide reduc-
tion; A A A, cell growth. Left ordinate indicates the specific activity of ferri-
cyanide reduction by HeLa cells. Right ordinate indicates the stimulation of cell growth
under the condition shown in the figure.
4. Craig, T.A. and F.L. Crane. 1981. Evidence for a transplasma membrane elec-
tron transport system in plant cells. Proceed. Indiana Acad. Sci. 90:150-155.
5. Crane, F.L. and H.Low. 1976. NADH oxidation in liver and fat cell plasma mem-
brane. FEBS Lett. 68:153-156.
6. Crane, F.L., H.E. Crane, I.L. Sun, W.C. MacKellar, G. Grebing and H. Low.
1982. Insulin control of a transplasma membrane NADH dehydrogenase in
erythrocyte membranes. J. Bioenerg. Biomemb. 14:425-433.
7. Ellem, K.A.O. and G.F. Kay. 1983. Ferricyanide can replace pyruvate to stimulate
416 Indiana Academy of Science Vol. 94 (1985)
growth and attachment of serum restricted human melanoma cells. Biochem.
Biophys. Res. Communs. 112:183-190.
8. Frelin, C, P. Vigne and M. Lazounski. 1983. The amiloride-sensitive Na + /H~l~
anitport in 3T3 fibroblasts. J. Biol. Chem. 258:6272-6280.
9. Garcia-Sancho, J., A. Sanchez, M.E. Handlogten and H.N. Christensen. 1977.
Unexpected additional mode of energization of amino acid transport in Ehrlich
cells. Proc. Natl. Acad. Sci. USA 74:1488-1491.
10. Gerson, D.F., H. Kiefer and W. Grufe. 1982. Intracellular pH of nitrogen-
stimulated lymphocytes. Science 216:1009-1010.
11. Goldenberg, H. 1982. Plasma membrane redox activities. Biochem. Biophys. Acta
694:203-223.
12. Low, H. and S. Werner. 1976. Effects of reducing and oxidizing agents on the
adenylate cyclase activity in adipocyte plasme membranes. FEBS Let. 65:96-98.
13. Low, H., F.L. Crane, G. Grebing, K. Hall and M. Tally. 1978. Metabolic milieu
and insulin action in diabetes. W.K. Waldhausl ed., Excerpta Medica, Amster-
dam pp. 209-213.
14. McLoughlin, P., I.L. Sun and F.L. Crane. 1982. Membrane redox systems in
porcine neutrophils. Proceed. Indiana Acad. Sci. 91:333-339.
15. Mishell, B.B. and S.M. Shrigi. 1980. In Selected Methods in Cellular Immunology.
W.H. Freeman, Co., San Francisco, pp. 17-18.
16. Mishra, R.K. and H. Passow. 1969. Induction of intracellular ATP synthesis by
extracellular ferricyanide in human red blood cells. J. Memb. Biol. 1:214-224.
17. Moolenaar, W.H., R.Y. Tsien, D.T. VanderSaag and S.W. DeLaat. 1983.
Na + /H + exchange and cytoplasmic pH in the action of growth factors in human
fibroblasts. Nature 304:645-648.
18. Morgan, E.H. 1983. Chelator-mediated iron efflux in reticulocytes. Biochem.
Biophys. Acta 733:39-50.
19. Olivotto, M. and F. Paoletti. 1981. The role of respiration in tumor cell transi-
tion from the noncycling to the cycling states. J. Cell Physiol. 107:243-249.
20. Sijmons, P.C. and H.F. Bienfait. 1984. Mechanism of iron reduction by roots
of Phaseolus vulgaris L. J. Plant Nutr. 7:687-693.
21. Sun, I.L. and F.L. Crane. 1981. Transplasmalemma NADH dehydrogenase is
inhibited by actinomycin D. Biochem. Biophys. Res. Commun. 101:68-75.
22. Sun, I.L., F.L. Crane, H.Low and C. Grebing. 1984. Properties of a transplasma
membrane electron transport system in cultured HeLa cells. J. Bioenerg. Biomemb.
16:209-221.
23. Sun, I.L. and F.L. Crane. 1984. The antitumor drug cis diamminedichloroplatinum
inhibits transplasamalemma electron transport in HeLa cells. Biochem. Internat.
9:299-306.
24. Sun, I.L. and F.L. Crane. 1984. Bleomycin control of transplasma membrane
redox activity and proton movement in HeLa cells. Biochem. Pharmacol., in press.
25. Sun, I.L., F.L. Crane, G. Grebing and H. Low. 1984. Properties of a transplasma
membrane electron transport system in HeLa cells. J. Bioenerg. Biomemb.
16:583-595.
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impermeable oxidants through plasma membrane redox. J. Cell Biol. 99:293a-293a.
28. Thorstensen, K. and I. Romslo. 1984. Uptake of iron from transferrin by isolated
hepatocytes. Biochem. Biophys. Acta 804:200-208.
PHYSICS AND ASTRONOMY
Chairperson: Vincent A. DiNoto, Jr.
Department of Physics
Indiana University Southeast
New Albany, Indiana 47150
(812) 945-2731 ext 390
Chairperson-Elect: Ruth Howes
Department of Physics and Astronomy
Ball State University
Muncie, Indiana 47306
(317)285-5494 or 285-6268 - Dept.
ABSTRACTS
The Dynamics of the Population of the United States. Albert A. Bartlett, Depart-
ment of Physics, University of Colorado, Boulder, Colorado 80309 and Richard L.
Conklin, Hanover College, Hanover, Indiana 47243. The simple concepts of
kinematics and dynamics can be used to calculate values of interesting demographic
parameters. Date from standard statistical sources allow one to estimate that the average
speed of Americans is 0.7 m/s. Each decade the U.S. Census Bureau publishes the
coordinates of the Center of Population of the U.S. Identifying this with the center
of mass (CM) one can use these coordinates to determine the average displacement
and hence the average velocity of the CM in the decade. From the differences in velocities
of consecutive decades one can calculate the average accelerations of the CM. Vector
diagrams representing the velocity and acceleration are historically interesting, and when
the various values are used to calculate the kinetic energy, momentum, and accelerating
force the results are quite surprising.
On the Measurement of Thermal Diffusivities with Bryngdahl Interferometry. Mar-
shall P. Cady, Jr., Department of Natural Sciences, Indiana University Southeast,
New Albany, Indiana 47150. Bryngdahl shearing interferometry is a relatively in-
expensive but highly accurate method for the measurement of both temperature
derivatives of the refractive index and thermal diffusion coefficients of liquid mix-
tures. This paper reports the results of a numerical analysis into the question: Can
Bryngdahl interferometry be used to simultaneously measure thermal diffusivities? The
thermal diffusivity is important because it is proportional to the thermal conductivity
and it is the coefficient that determines the rate at which temperature changes after
boundary temperature conditions are altered.
The numerical analysis uses a model sandwich-cell experiment in which upper
and lower boundary temperatures change exponentially with a 5 minute relaxation time
and a final steady state gradient equal to 0.7°C/cm is established. To generate data,
it is assumed that the true liquid thermal diffusivity equals 0.05214 cmVmin and that
the interferometer shear parameter(2D) divided by the image height(H) equals 0.2648.
The Crank-Nicholson finite difference method is then used to solve the Fourier
temperature equation on a spatial grid of 30 points/cm every 0. 1 minutes subject to
the boundary conditions(BC). The interferometer fringe count as a function of time
is calculated from the resulting temperature data. It represents the experimental fringe
data — a quantity that is highly accurate.
To simulate an experimental effort to determine the thermal diffusivity, random
417
418 Indiana Academy of Science Vol. 94 (1985)
Gaussian noise is then placed into the BC and the thermal diffusivity in the Fourier
temperature equation is systematically adjusted until the experimental fringe data is
most nearly reproduced in a least squares sense. This computation is repeated 15 times
with the same noise band whereupon the average thermal diffusivity and its standard
deviation are computed. The noise band width represents experimental uncertainty in
BC knowledge; the standard deviation of the thermal diffusivity represents its subse-
quent uncertainty.
It is found that thermal diffusivities can be simultaneously measured to within
4% if BC are measured to within 0.006°C and to within 2% if BC are measured to
within 0.003°C. This requires a highly automated data acquisition system capable of
sampling rates greater than 50 per relaxation experiment.
The Physics of the Grist-mill. Vincent A. DiNoto, Jr., Department of Physics, Indiana
University Southeast, New Albany, Indiana 47150. Taking a step back in time
to the time of the founding of the Indiana Academy of Science 100 years ago, we
will observe some of the applied physics of this day and time. One of the major needs
of the people was the grinding of corn. First animals were used for this purpose and
then later water power. The mills were very inefficient and required a fairly large supply
of water which must have either a large vertical drop or a swift current or both. Pro-
bably more than 500 mills were built in the 1800s in Indiana with few still remaining
and even fewer still operational.
The National Optical Astronomy Observatories. Frank K. Edmondson, Department
of Astronomy, Indiana University, Bloomington, Indiana 47405. The newly
established National Optical Astronomy Observatories combines three observatories
funded by the National Science Foundation and operated under contract by the Associa-
tion of Universities for Research in Astronomy, Inc. (AURA). They are: The Kitt
Peak National Observatory (KPNO), the National Solar Observatory (NSO) and the
Cerro Tololo Inter-American Observatory (CTIO). AURA also operates the Space
Telescope Science Institute (STScI) under contract with the National Aeronautics &
Space Administration.
Indiana University was one of the seven founding members of AURA. The paper
will review the history of AURA and the three observatories. The rationale of the
new organization will be discussed.
The Manchester Interface Adapter for Commodore and Apple Microcomputers. L.
Dwight Farringer, Department of Physics, Manchester College, North Manchester,
Indiana 46962. An inexpensive interface adapter has been designed for use with
the "game" control ports of microcomputers such as the Apple II, II + , and He and
the Commodore 64 and VIC-20. It provides a convenient and safe way to utilize the
digital inputs, digital outputs, and resistive analog inputs which are accessible at those
control ports.
The digital inputs are buffered by Schmitt triggers which "clean up" certain kinds
of noisy signals and also protect the computer from damage by possible wrong con-
nections to the outside world. The digital outputs are buffered by transistors which
can drive external loads to about 50 mA. and 4 volts, and relays can be added for
controlling external loads which require external power sources. The analog inputs
are provided with switchable capacitors for adjusting the full-scale range of resistance
which the computer can read.
Use of this interface adapter with accessories such as optical sensors, thermistors,
and various kinds of sensing switches makes possible quite a variety of experiments
Physics and Astronomy 419
using the microcomputer as a laboratory instrument.
The low cost, ease of assembly, versatility, and protection of the computer which
are afforded by such an interface adapter are features which recommend its use in
many high school and college teaching laboratories.
Software for Astronomical Photometry. Jodi Hamilton and Thomas H. Robertson,
Department of Physics and Astronomy, Ball State University, Muncie, Indiana
47306. Computer programs have been developed to support and facilitate observing
projects and data reduction for astronomical photometry. These programs are designed
for observational support, data acquisition and data analysis. A primary objective for
the development of this software was to make the execution of observing programs
in observational astronomy more tractable for students with very limited experience.
Some system limitations and plans for future program development are discussed.
Licensing and Certification of Physics Teachers by Examination: What are the Dangers?
Lawrence E. Poorman, Department of Physics, Indiana State University, Terre Haute,
Indiana 47809. The Indiana General Assembly enacted legislation last spring (1984)
mandating competency testing of all prospective teachers for licensing and certifica-
tion starting July 1985. The State Licensing and Certification Commission, now defunct,
recommended the use of national teacher examinations available through Educational
Testing Service.
In August, 1984, panels were called to convene at North Central High School,
Indianapolis, to validate and determine minimum competency levels for certification
purposes. This author served on the panel to evaluate chemistry, physics and general
science examinations.
There are many concerns. As members of faculties preparing persons for teaching;
all should be very concerned with the procedural selection and administration of any
certification examination.
A System for Astronomical Photometry. Thomas H. Robertson and Jodi Hamilton,
Department of Physics and Astronomy, Ball State University, Muncie, Indiana
47306. A photometric system has been developed to serve both instructional and
research programs. The system consists of a Pacific Precision Instruments model 2426-1
photometer with a model 401 telescope coupler, an Altair computer and a twelve-inch
Tinsley cassegrain reflector. The system is capable of both DC and pulse counting
modes of operation. Limiting magnitudes observable are currently determined by night
sky brightness and the mechanical and electrical imperfections of the telesccope drive
system. Preliminary site condition tests and future plans for system upgrading are
discussed.
The Great Southern U.S. Geologic Uplift Observed in the Early Months of 1984. Gerald
J. Shea, Terre Haute, Indiana 47801. A land mass bubble of enormous propor-
tions was detected and observed using horizontal pendulum instruments of high sen-
sitivity from January to May of 1984. Its boundaries were estimated as covering an
area which included eleven states. The maximum rise appeared to be near Nashville,
Tennessee and was computed as being 5 inches. The observations were carried on using
instruments located at three different locations along the edge of the bubble.
The significance of the observation is a possible earthquake being due in the affected
area which includes the New Madrid Fault zone and the Wabash Valley Fault zone
which have been responsible for large earth displacements in the past.
The significance also may be meterological in origin being due to the intense dry
420 Indiana Academy of Science Vol. 94 (1985)
hot summer of 1983 which may have disturbed the underlying geologic formations
resulting the uplift.
One thing known for sure is no such observations of tilt have been detected
previously over the 35 years that the Terre Haute Seismological Station has been in
operation.
Astrophotography Using Celestron Telescopes. F.R. Steldt, Department of Physics,
Indiana University at Kokomo, Kokomo, Indiana 46902. A series of slides have
been taken of various heavenly bodies using Celestron telescopes. These telescopes in-
cluded the C-90, C-8, and C-14 models and all utilized portable tripod mounts. Kodak
ASA 400 high speed Ecktachrome slide film was used for the majority of the photographs
and the film was processed by a local firm using the standard procedure.
Exposure times ranged from 1/500 of a second to ten seconds for the major solar
system objects. Constellation and deep space subjects required time exposures from
one minute up to twenty minutes. The time exposures required constant manually adjusted
guiding to correct the errors in the clock drive in order to keep the subject at the same
position on the film.
The 35mm single lens reflex camera was positioned at the prime focus of the
telescope for the deep space objects and low magnification photographs of the moon
and the sun. The high magnification photographs of the moon and the planets were
taken through one of the oculars attached to the telescope. Constellation photographs
used the telescope as a guide for the camera riding piggy-back.
Using Toys to Teach Physics to Middle School Students. Nancy Watson, Burris
Laboratory School and James Watson, Jr., Department of Physics and Astronomy,
Ball State University, Muncie, Indiana 47306. Physics of toys explores the science
concepts that are used in various toys. Toys can be used to demonstrate scientific con-
cepts at all levels, kindergarten through college. Hot wheels, cereal box toys, and other
common toys are examined as examples of scientific concepts. Several areas of physics
are explored including mechanics, heat, optics, sound, and energy. The student will
learn science by "playing." The principles on which toys work are also the principles
that most objects that are used daily also work. A correlation between toys and every-
day objects will be emphasized. Students will learn observation and deduction skills
as they "play with the toys." Once the basic physics concept is discovered, the student
will use the toy to take data in an experimental setting. This data will then be used
to confirm the laws of physics.
Integer-valued Equivalent Resistances
Samir I. Sayegh and Joseph D. Lawrence
Department of Physics
Indiana University-Purdue University at Fort Wayne
Fort Wayne, Indiana 46805
Introductory physics course usually covers simple circuit theory and hence the stu-
dent faces problems of finding the equivalent resistance of two resistors placed in parallel.
The relationship is:
1/Rpn = 1/R, + 1/R2 Eq.l
The instructor's task is to find 'nice' values for R,, R2 and Req such that the student
doesn't need to waste time punching calculator keys. If the instructor restricts R,,
R2 and R„n such that they are elements of the natural numbers, then the calculation
is much simpler for the student. By restricting the resistor values in this way, we find
that we have a diophantine equation. The problem of finding a general solution to
the diophantine equation boils down to determining what conditions R, and R2 must
meet in order for R„n to be a natural number. We have found a general solution
cq
to this problem.
Our solution can benefit the physics educator as well as the mathematician. For
the introductory physics educator, our solution allows him to design assignment and
test problems which illustrate the physics concepts with a minimum of calculation time.
A quick search of introductory physics texts shows that the same 'nice' values are
abused in example after example. Our solution allows an instructor to generate un-
familiar values which are still very 'nice.' To the math educator, our equation and
solution offer a new problem in diophantine equations. In mathematics, classic diophan-
tine problems stem from geometry. An example is the problem of finding integer valued
solutions to the Pythagorean equation. Our equation offers the student a more tangible
problem. Also, many of the key techniques in number theory are illustrated by our
method of solution. With these benefits in mind, we turn to the solution.
We start by rewriting the equation form as such:
Req = R,R2/(R, + R2) Eq.2
Next, let D be the greatest common factor between R, and R2. We can write:
R, = DM R2 = DN Eq.3
where M and N are the remaining factors of R, and R2 respectively. We can see that
M and N must be relatively prime, (greatest common factor equals 1) for otherwise,
we could extract the common factor from M and N and make a further contribution
to D. We substitute these expressions for R, and R2 into Eq.2.
R-n = DMN/(M + N) Eq.4
cq
For R to be a natural number, then sum (M + N) must evenly divide some combina-
tion of D, M, and N. Since we cannot extract a common factor from M and N, then
421
422 Indiana Academy of Science Vol. 94 (1985)
the sum (M + N) cannot divide any combination which includes M or N. Therefore,
the sum (M + N) must divide D. So we can write:
D = k(M + N) Eq.5
where k is some natural number. Now we substitute Eq.5 into Eq.3. After doing this
we have:
R, = kM (M + N)
R2 = kN (M + N) Eq.6
Req = kMN
This is the general solution to our diophantine equation.
For any values of M and N such that M and N are relatively prime natural numbers,
we can generate a solution to our equation. We now define a solution, where k = 1,
to be a primitive solution. Of course all positive integer multiples of a primitive solu-
tion are also solutions to the equation. We can show that each integer valued solution
of Eq.l can be generated by Eq.6 from exactly one combination of M and N.
PROOF:
Let M and N be relatively prime natural numbers. Likewise for M' and N'.
R, = kM(M + N)R,' = k'M'(M' + N') Eq.7
R2 = kN(M + N) R2' = k'N'(M'+N')
If R, = R, ' and R2 = R2 ' then,
kM(M + N) = k'M'(M' + N') Eq.8
kN(M + N) = k'N'(M' + N')
Dividing equations, we get:
M/N = M'/N' Eq.9
For this to be true we must have
M' = cM and N' = cN Eq.120
where c is a non-zero integer. Since M ' and N ' are relatively prime, then c = 1. Hence:
M = M' and N = N' Eq.ll
which means, for a given R, and R2, only one combination of M and N will generate
Rl and R2.
From the form of our solution, M and N are indistinguishable and so we always
choose M < N. M can equal N only if the case when M = N = 1. We extended
our method of solution to an arbitrary number of parallel resistors.
An example of how to apply our method should make the solution clearer. Suppose
the instructor wants to design a problem such that R„n = 30 ohms. To start with,
we find the prime factorization of 30.
Physics and Astronomy 423
30 = 2 x 3 x 5
Next, we start choosing values for k, M, and N from the prime factors. We don't
neglect the possibility that k, M, or N could equal 1. Generally we start by choosing
values for k first and then we sort through the remaining factors for values of M
and N. Choices for M and N must meet the criterion that M and N be relatively prime
and M < N, except for M = N = 1. We go through the choosing process systematically
until we exhaust all combinations which meet the criteria. As an illustration of the
method, the combinations for R£Q = 30 ohms are presented in Table 1. By following
the procedure outlined above, the instructor can generate solutions for any value of Req.
Table 1.
Combinations for R
eq
= 30 Ohms
k
M
N
R,
R:
1
1
30
31
930
1
2
15
34
255
1
3
10
39
130
1
5
6
55
66
2
1
15
32
480
2
3
5
48
80
3
1
10
33
330
3
2
5
42
105
5
1
6
35
210
5
2
3
50
75
6
1
5
36
ISO
10
1
3
40
120
15
1
2
45
90
30
1
1
60
60
So far, our formulation has been restricted to positive integers, since ordinary
resistances are never negative. However, with minor adjustments, we can extend our
formulation to include negative integers, then we can look at another important topic
covered in introductory physics courses, namely thin lenses. The thin lens equation is:
1/f = 1/i + l/o Eq.12
Here, the values for the image and object distances may also assume negative values
by sign convention. It is easy to show that as soon as one chooses values for M and
N when the image and object distances, the focal length, and the magnification have
been determined. Namely:
o = kM(M + N)
i = kN(M + N) Eq.13
f = kMN
Magnification = -M/N
The fact that so many problem parameters are determined by choosing M and N points
to the very nice feature of design simplicity offered by our method.
To conclude, our method succeeds in simplifying the task of designing introduc-
tory physics problems. Using the formula forms:
424 Indiana Academy of Science Vol. 94 (1985)
X = kM(M + N)
Y = kN(M + N) Eq.14
Z = kMN
where k, M, and N are non-zero integers and M and N are relatively prime, we can
generate values of X, Y, and Z such that they will be non-zero integers. With this
solution, the physics educator can design problems involving parallel resistors, series
capacitors and thin lenses for introductory courses. To the math educator, our solu-
tion presents a fresh problem in diophantine equations which illustrates basic techni-
ques of number theory analysis.
PLANT TAXONOMY
Chairperson: Marion T. Jackson
Department of Life Sciences
Indiana State University
Terre Haute, Indiana 47809
(812)232-6311
Chairperson-Elect: Victor Riemenschneider
Department of Biology
Indiana University at South Bend
South Bend, Indiana 46615
(219)272-8262
ABSTRACTS
The Discovery of Native Rare Vascular Plants in Northern Indiana. James R. Aldrich,
Lee A. Casebere, Division of Nature Preserves, Indiana Department of Natural
Resources, Indianapolis, Indiana 46204 and Helene Starcs, 4250 Crittenden Avenue,
Indianapolis, Indiana 46205. Active field surveys during recent years have greatly
influenced our knowledge of the endangered and threatened flora of northern Indiana.
This report includes several new county records and the rediscovery of two species,
dragon's mouth orchid — Arethusa bulbosa (Orchidaceae) and bluebead lily — Clintonia
borealis (Liliaceae), thought to be extirpated in Indiana. A native species new to the
Indiana flora, bog valerian — Valeriana uliginosa (Valerianaceae), is also discussed.
A Preliminary Survey of Phenolic Compounds in Sympatric Populations of Quercus
shumardii and Q. rubra in Northern Indiana. Roxane A. Dupuis and Richard J.
Jensen, Department of Biology, St. Mary's College, Notre Dame, Indiana
46556. Thin layer chromatography was used to investigate phenolic profiles in
sympatric populations of Q. shumardii Buckl. and Q. rubra L. The populations sampled
were chosen because of apparent hybridization between these two taxa. Four sites in
northern Indiana were sampled in June, 1984. Several twigs were taken from each
tree and leaves were air dried in standard plant presses. Methanolic extracts were prepared
and spotted on acetate sheets coated with polyamide. Two-dimensional thin layer
chromatogrphy was conducted with several solvent systems. The patterns observed
were compared with those from trees of the same species from outside the study area.
The results suggest that both species contain "unique" compounds. Further, there is
evidence of hybridization in that several trees yield patterns that appear to be additive
with respect to the species' patterns.
Rust Species Diversity in Temperate and Tropical Regions of the Americas. J.F. Hen-
nen, R.M. Lopez-F. and M.M. Hennen, Department of Botany and Plant Pathology,
Purdue University, West Lafayette, Indiana 47907. Nineteen of the 103, (18%),
are new species. We estimate the number of vascular plant species to be 300. Indiana
is about 232,262 times larger than our Brazilian study area but has only about one
and three-fourths times as many species of rusts. In Indiana about 1 out of 14 (7.36%)
vascular plant species has a rust; while in Mogi-mirim, about 1 out of 3 (33.3%) vascular
plant species has a rust.
In our recent work on the currently known rust fungi of Brazil (Hennen et al.,
1982) we reported 687 species in 54 genera. In our general collecting work in Brazil
425
426 Indiana Academy of Science Vol. 94 (1985)
we found about one new species for each 70 collections. Except for our studies, very
little surveying for rusts in Brazil has been done. Therefore, considering the thousands
of vascular plant species known for Brazil that could serve as hosts for rusts, we predict
that when Brazil is more thoroughly studied at least 3,000 species of rusts will be found.
Literature Cited
1. Hennen, J.F., M.M. Hennen and M.B. Figueiredo. 1982 (1984). "Indice das
ferrugens do Brasil." Arq. Inst. Biol., Sao Paulo 49, supl. 1:1-201).
2. Jackson, H.S. 1921. The Uredinales of Indiana III. Proc. Indiana Acad. Sci.
1920 (1921): 165-182.
3. McCain, J.W. and J.F. Hennen. 1982. Notes on Biogeography and New Records
of Rust Fungi in the Great Lakes region. Proc. Indiana Acad. Sci. 1981 (1982):
504-514.
4. Viegas, A. P. 1943. "Alguns fungos do cerrado." Bragantia 3:49-72.
Additions to the Flora of Indiana: II. Michael A. Homoya, Division of Nature
Preserves, Indiana Department of Natural Resources, Indianapolis, Indiana
46204. Vascular plant species new to Indiana, and several that have been infre-
quently collected in southern Indiana, were discovered during the 1983-84 field seasons
by members of the Division of Nature Preserves. A partial list of species includes
blackstem spleenwort (Asplenium resiliens Kunze), sedge (Carex atlantica subsp. atlantica
L.H. Bailey), Fairy- wand (Chamaelirium luteum (L.) Gray), American pennywort
(Hydrocotyle americana L.), oval ladies'-tresses (Spiranthes ovalis Lindley), and barren
strawberry {Waldsteinia fragarioides (Michx.) Tratt.).
Assessing Variation in Mixed Oak Communities: Evaluation of Multivariate Analyses
of Morphological Data. Richard J. Jensen and Roxanne A. Dupuis, Department
of Biology, St. Mary's College, Notre Dame, Indiana 46556. Morphological varia-
tion in fruit and leaf characters was studied in a community containing three taxa
of Quercus subg. Erythrobalanus: Q. palustris Muenchh., Q. rubra L., and Q. velutina
Lam. Data were collected for sixteen individual trees. Discriminant analyses of sets
of leaves from each tree revealed that each tree represents a reasonably well-defined
multivariate entity. However, one tree tentatively identified as Q. velutina appears mor-
phologically more similar to Q. palustris. Additional analyses of both leaf and fruit
data support the hypothesis that this tree may be a hybrid between these two species.
Comparison of relationships among the trees, conducted by employing Mantel's test
of similarity of distance matrices, reveals that leaf and fruit data provide significantly
different patterns of between tree taxonomic distances. Mantel's test is demonstrated
by way of an MBASIC computer program written by RJJ.
Linear Differentiation of Allium cepa, Lens culinaris and Vicia faba Chromosomes.
R.C. Mehra, D. Fisher, S. Brekrus, S. Alwine and J. Palbykin, Indiana Universi-
ty at South Bend, South Bend, Indiana 46634 and M.G. Butler, Department of Medical
Genetics, Indiana University, School of Medicine, Indianapolis, Indiana
46223. Recently several techniques have been developed to produce bands along
the length of plant and animal chromosomes. This linear differentiation has been of
tremendous help in chromosome analysis and thus has greatly advanced human genetics
in the last fifteen years. The techniques which have had some success in linear dif-
ferentiation of plant chromosomes are: C, N, Q and silver staining. We have attemp-
ted some of these techniques on a few plant taxa. In Allium cepa, Lens culinaris and
Plant Taxonomy 427
Vicia faba, through a silver staining procedure, we have been able to localize nucleolus
organizing regions (NORS) in their chromosomes and found that a polymorphism exists
with respect to this chromosomal phenotype. We have N-banded, both L. culinaris
and V. faba and found that, whereas in L. culinaris, N-bands are mainly confined
to the centromeric region and NORS, in V. faba they are also present in the interstitial
areas of its chromosomes. On the basis of N-banding and chromosomal measurements,
we have developed an N-banded karyotype and an idiogram for L. culinaris. With
the help of a modified C-banding procedure, we have been able to localize constitutive
heterochromatin in different areas of L. culinaris and V. faba chromosomes. A com-
parison of C and N bands in these taxa show that both procedures produce bands
in same areas of the chromosomes. Evidence will be presented. In conclusion, if plant
chromosomes can be banded with the same ease as the mammalian chromosomes,
then chromosome banding will become a very powerful tool in plant biosystematics.
Vascular Flora of Grant County, Indiana: Additions and Comments. Paul E.
Rothrock, Department of Biology, Taylor University, Upland, Indiana
46989. Based upon BSC-FLIP data, 132 species of vascular plants are reported
for the first time in Grant County. This increases the county total to 568 species. Among
the additions are 45 introduced species, one species new to Indiana {Vicia dasycarpa
Tenore), and Carex woodii Dewey, an endangered species in this state. Species near
the edge of their range included Chelone obliqua L., Heracleum lanatum Michx., and
Luzula multiflora (Retz.) Lej. Two introduced species have become locally common
since Deam's flora of 1940: Rosa multiflora Thunb. ex. Murr. and Torilis japonica
(Houtt.) DC. Voucher specimens of these collections are being deposited in the Her-
barium of Indiana University.
Pre-burning Floral Inventory of Little Bluestem Prairie, Vigo County, Indiana. Rebecca
A. Strait and Marion T. Jackson, Department of Life Science, Indiana State Univer-
sity, Terre Haute, Indiana 47809. Nearly 13% of Indiana was once covered by
prairie; however, this community type is now very rare. Indiana's only known rem-
nant of sandhill prairie is Little Bluestem Prairie located in Vigo County, Indiana.
The prairie is the site for an ongoing study to assess the effect of burning on prairie
flora and fauna.
The purpose of this study was to quantitatively sample the plant communities
represented and to conduct a flora inventory prior to pre-vernal burning. Density and
cover data were taken for all plant species in 20 1 x 1 meter stratified random sample
plots in late May, mid July and late August of the 1984 growth season. Follow-up
surveys will be conducted at the same sample locations and at similar times following
a burn scheduled for early Spring 1985.
Four generalized community types have been recognized: dry sand prairie, moist
slope prairie with Equisetum, woody ravines and black locust invasion areas. Twenty-
three families and 41 genera, and at least 55 species of vascular plants were recorded
in the prairie plots. Commonly represented species include: Andropogon scoparius,
Crysopsis mariana, Euphorbia corollata, Guara biennis, Lespedeza capatitata,
Petalostemum villosum, Sorghastrum nutans and Lithospermum canescens.
An effort is being made to control black locust invasion by tree cutting followed
by basal herbicide application and burning.
The Red and Black Oaks of Indiana
Richard J. Jensen
Department of Biology
St. Mary's College
Notre Dame, Indiana 46556
Introduction
The number of species of red and black oaks (Quercus subgenus Erythrobalanus
(Spach) Oersted) in Indiana has been reported variously as nine or ten. A simple count
does not suffice because all authors do not recognize the same species. Specifically,
many authors treat the cherrybark oak as Q. falcata Michaux var. pagodaefolia Ell.
while others consider it a distinct species, Q. pagoda Ashe. If however, allowance is
made for such taxonomic problems, there is still a slight discrepancy. That is, there
has been some disagreement about which species actually occur in the state.
Deam (5) in a comprehensive treatment of the trees of Indiana, reported nine
species. Eight years later, Deam (6) reported the same nine species but the recorded
distributions of several of these had been markedly expanded. Other authors apparently
have based their reports on Deam's (6) maps. Little (16, 17) presented maps, for the
same taxa, that are virtual duplicates of Deam's (6) maps. Elias (7) followed suit,
but apparently relied on Little (16, 17) as his sources. Preston (21) reported one species,
Q. nigra L., not included by other authors. Furthermore, he indicated that Q. marilandica
Muenchh. occurs throughout Indiana while Deam (6), Little (16), and Elias (7) depicted
this species as occurring only in the southwest and southcentral portions of the state.
In a more recent edition, Preston (22) adopted Little's (16, 17) maps and no longer
included Q. nigra among the species found in Indiana. Thus, by virtue of Deam's
(6) revision, followed by gradual adoption of his maps by others, there is now an
apparent agreement on the number of species of red and black oaks in Indiana as
well as on the distribution of these species. Of course, this agreement is not a reflec-
tion of congruence between the research findings of different authors. Rather, it is
a function of later authors relying on Deam's (5, 6) work as a source for their own
reports.
A second aspect of the recorded occurrence of red and black oaks in Indiana
is the small number of hybrids that have been reported. Hybrids, or at least trees
thought to be hybrids, are encountered commonly in mixed oak forests. The existence
of hybrid individuals has been confirmed by morphological studies (12, 13, 14) and
by chemical studies (15). All species of Erythrobalanus native to Indiana have been
identified as progenitors of hybrid trees (20), yet very few hybrids have been reported
from Indiana. Deam (5) reported only two hybrids among the many specimens he
examined: one specimen of X Q. exacta Trel. from Posey County and two specimens
of X Q. leana Nutt., pro sp., one each from Lawrence and Lake Counties. Both of
these hybrids involve Q. imbricaria Michaux as one parent, with Q. palustris Muenchh.
and Q. velutina Lam., respectively, being the second parents. In 1940, Deam (6) added
one more hybrid to this list, X Q. bushii Sarg. (Q. marilandica x Q. velutina), based
on a single specimen from Knox County. While hybrids between morphologically distinct
species such as the above-mentioned are rather easily identified, it is not surprising
that Deam (5, 6) reported so few hybrids. After all, most hybrids that could be ex-
pected to occur in Indiana would involve parent species that are morphologically very
similar, thus making the hybrid difficult to detect.
Palmer (20), in a comprehensive list of hybrid oaks found in North America,
reported only two additional hybrids from Indiana: X Q. paleolithicola Trel. (Q. ellip-
429
430 Indiana Academy of Science Vol. 94 (1985)
soidalis E.J. Hill x Q. velutina) and X Q. runcinata (A. DC.) Englem. (Q. imbricaria
x Q. borealis Michaux f.). Palmer's (20) paper is the most thorough survey of oak
hybrids published to date. Since its publication, there has been no attempt to update
the literature. A number of new hybrids have been described, but none has been noted
specifically as occurring in Indiana. Nor has there been any general survey of addi-
tional hybrid reports from various parts of the country. Therefore, our current knowledge
of hybrid oaks that may occur in Indiana is essentially the same as it was thirty-four
years ago.
The research reported here had two primary goals. First, the distribution of each
species of red or black oak native to Indiana was to be brought up to date and an
attempt was to be made to verify the county records reported in Deam (5, 6). Second,
an annotated list of red and black oak hybrids found in Indiana was to be prepared
and, again, Deam's (5, 6) reports were to be checked.
Materials and Methods
The distributions of red and black oaks in Indiana were determined primarily by
examination of specimens on file at various herbaria in Indiana, Illinois, and Ohio.
The herbaria visited were those at Ball State University (BSU), Butler University (BU),
DePauw University (DPU), Earlham College (EC), the Field Museum (F), Indiana
University(IND), Miami University (MU), Purdue University (PUL), St. Mary's Col-
lege (SMC), the University of Illinois (ILL), the University of Notre Dame (ND), and
Wabash College (WAB). As specimens were examined, a record of county occurrences
was made by entering label information into a data file. The file was prepared using
the MicroLibrarian© program with an OSBORNE-1 portable microcomputer.
A data entry was not made for every specimen examined. Rather, an entry was
made for the first specimen of each taxon which could be verified for each county.
Once a county record was entered, no additional entries for that taxon in that county
were made. Thus, the list of verified county records is top-heavy with specimens from
the first several herbaria visited. On the other hand, all specimens labeled or verified
as hybrids were recorded. In addition, several reports are based on my own specimens
collected during September, 1983. These specimens eventually will be filed in the In-
diana University Herbarium (IND).
The files generated could be scanned by taxon and by county to prepare distribu-
tion maps. These maps were prepared to reflect (1) the distribution based on my data
files, (2) the distribution reported by Deam (5, 6), (3) new records since Deam's last
treatment (6), and (4) records reported by Deam (6) that could not be verified by cor-
rectly identified herbarium specimens.
The complete data file is housed on three floppy disks and comprises over six
hundred entries. As such, it is too large for inclusion here. If anyone desires a copy
of the file, sequenced either by taxon or by county, the author will provide same upon
receipt of a written request.
Results and Discussion
The results are presented in alphabetical order of the species epithets. When her-
barium specimens are cited, the format is to present the herbarium abbreviation (see
above) and accession number followed by the collector's name and collection number.
I. Quercus borealis Michx.
This is one of the more commonly encountered oak species in Indiana and, while
the maps in Deam (6; map 790) and Figure 1 indicate many gaps, probably can be
found in every county. Although I was able to verify most of Deam's records and
added eighteen additional records, including two collected by me (Whitley Co., Jensen
Plant Taxonomy
Figure 1. Quercus borealis in Indiana. In this, and all other figures, • = mapped
in Deam (1940) and verified by existing specimens; * = not mapped by Deam, but
verified by existing specimens; a letter signifies that that county was mapped in Deam,
but could not be verified and the letter reflects the herbarium Deam cited, B = Butler;
D = Q. borealis var. borealis.
432 Indiana Academy of Science Vol. 94 (1985)
83-40; LaPorte Co., Jensen 83-18), there were 10 county records on Deam's map which
could not be verified. All 10 of these were, according to Deam, based on specimens
housed at Butler University. Those specimens, if at Butler, are not housed in the her-
barium with the other oaks. In addition, there are literature reports for Q. borealis
in twelve other counties: Clay (28), Daviess (19), Fayette (25), Fulton and Grant (24),
Greene (19), Howard (8, 24), Owen (29), Switzerland (9), Union and Vermillion (26),
and White (24).
Many of the specimens examined did not have fruits, therefore it is difficult to
assess the range of Q. borealis var. borealis, the smaller fruited and less common variety
of this species. As indicated in Figure 1, only four specimens of this variety were found,
all others being Q. borealis var. maxima (Marsh.) Ashe or, for lack of fruits, were
merely identified to the species.
Even though Deam (6) stated that this species "may be entirely absent from Ben-
ton, Newton, and possibly Lake Counties . . ," his map (790) shows it in Newton
County. While, as shown in Figure 1, it also occurs in Benton and Lake Counties,
there is still validity to his comment that it "is rare or absent in the lower Wabash
Valley."
II. Quercus coccinea Muenchh.
Deam (6) reported this species from only 13 counties (Figure 2) in Indiana, re-
marking that "... is local and, no doubt, has a wider range than the map indicates."
As Figure 2 illustrates, Deam was right. Although I added only 19 counties to those
mapped by Deam (6), the distribution suggests that Q. coccinea may be expected to
occur throughout the state. I was unable to verify two of Deam's (6) county records
(Floyd and Sullivan Counties) and found a discrepancy in his text. Quercus coccinea
var. tuberculata Sarg. is reported as occurring in Vanderburgh County, yet this is not
indicated on the map (Deam's [6] map 795). As shown in Figure 2, I was able to
verify the occurrence of var. tuberculata in 13 counties and its distribution suggests
that both varieties of Q. coccinea are probably common in the state. Two other coun-
ties, not marked in Figure 2, perhaps should be included. Hale (9) reported Q. coc-
cinea from Switzerland County and Underwood (27) reported collecting Phyllactinia
suffulta from Q. coccinea in Johnson County.
III. Quercus ellipsoidalis E.J. Hill
This species is confused easily with Q. coccinea and in northern Indiana, where
their ranges overlap, it is difficult to distinguish the two. There were a number of
sterile specimens, of one of these two, which I was not able positively to identify to
species. Generally, these two species can be separated by the shape of the nut, which
tends to be elliptic in Q. ellipsoidalis and sub-globose in Q. coccinea. However, both
taxa are variable in this respect. Another character which may be used is the presence
of rings of minute pits around the apex of the nut. This feature is more common
in Q. coccinea, although it occasionally may be expressed in Q. ellipsoidalis. Sterile
specimens, especially in northern Indiana, are very difficult to identify, although careful
multivariate analyses indicate that this can be done (Jensen, unpublished).
I have no evidence that Q. ellipsoidalis occurs south of a line extending roughly
from the Benton-Warren County line in the west to Adams County in the east (Figure
3). Deam (6) stated, for Q. ellipsoidalis, that its "distribution ... in Indiana is not
known" and he reported it from only three counties. I have verified the identity of
specimens from these three counties as well as an additional sixteen counties in Northern
Indiana. As Deam (6) noted, Andrews (1) reported Q. ellipsoidalis from Monroe County,
a report probably based on misidentification of either Q. coccinea or Q. palustris.
IV. Quercus falcata Michx.
This complex assemblage consists of many named variants, of which Deam (6)
Plant Taxonomy
433
Figure 2. Q. coccinea in Indiana. □ = var. tuberculata. D = Deam's Herbarium
(now at IND).
recognized four: falcata var . falcata, falcata f. triloba (Michx.) Palmer and Steyerm.,
434
Indiana Academy of Science
L*
:t""~c
Vol. 94 (1985)
•
* /._.
"1 1 *
; r t l ;
- 1
■ 1
L_L
,_i
H
* ■*
r
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Figure 3. Q. ellipsoidalis in Indiana.
falcata var. leucophylla (Ashe) Palmer and Steyerm., and falcata var. pagodaefolia
Ell. Yet in his mapping of distributions (map 796), he lumped all under falcata sensu
Plant Taxonomy 435
lato. Thus, the distribution of each variety or form can be deduced only from the
text;/, triloba apparently occurs throughout the area mapped, variety leucophylla was
reported only from Posey County, and variety pagodaefolia was reported only from
Posey and Gibson Counties.
I have chosen to follow Ware (30), Jensen (11), and Hicks and Burch (10) by
viewing this complex as comprising two species: Q. falcata and Q. pagoda Ashe. The
former includes forma triloba and the latter includes variety leucophylla. Figures 4
and 5 present the distributions of these two species based on specimens I have examined.
For Q. falcata, my map is almost identical to Deam's (6), although I have added one
county (Greene) and was unable to verify its occurrence in two others (Knox and Jenn-
ings). The Knox Co. record was based on an IND specimen (IND 3336 = Welch and
Pocket 4996), annotated as Q. falcata by Deam, which is from Q. pagoda. No specimen
of any form or variety of this complex was found for Jennings County. In addition
to the five counties indicated in Figure 5, Q. pagoda has also been reported from
Franklin County (4), although no specimen was found to verify this report.
V. Quercus imbricaria Michx.
This species, easily identified because it is the only entire-leaved oak in Indiana,
was reported from forty counties throughout the state by Deam (6). Each of Deam's
reports could be verified along with an additional thirteen records not included in Deam's
work. One of these, for Wabash County, was made by me {Jensen 83-30) during
September, 1983. Based on the distribution shown in Figure 6, it is probably safe to
assume that this species occurs in many of those counties lacking records. Andrews
(1) did report Q. imbricaria from Monroe County.
VI. Quercus marilandica Muenchh.
Deam (6) reported this species to be "local and infrequent, mostly in the
southwestern part of the state. ..." Figure 7 is very similar to Deam's map 797;
no new records have been added, but two could not be verified. I was unable to find
a specimen from Jackson County and the two specimens at IND from Lawrence County
were both misidentified. Cain 100 (IND 3382) is a specimen of X Q. leana Nutt. In
his 1931 report on the flora of Spring Mill State park, Cain (2) included Q. marilan-
dica, but noted that the identification was based on a single questionable specimen.
Wible 234 (IND 3381), while not Q. marilandica, appears to be X Q. bushii Sarg.,
a hybrid having Q. marilandica as one of its parents. Thus, Q. marilandica must be
in Lawrence County, it is just that no confirmed specimen is on file.
VII. Quercus palustris Muenchh.
This is a very common species throughout Indiana as indicated by Figure 8 and
by Deam's (6) map 792. I was able to verify every county record reported by Deam
as well as collections for an additional 27 counties. Deam (6) commented that this
species "may be absent from Benton County," but a PUL specimen (6958 = R. Kriebel
5147) from Benton Co. is certainly this species. Deam (6) also noted that there appear
to be two forms of this species differing in nut size, with the more common form
having larger nuts. However, Deam did not specify what the sizes of the nuts are.
In my experience, the common form in Indiana is the typical small-fruited pin oak
found throughout the mid-eastern United States. I encountered a single tree in Hun-
tington County {Jensen 83-31) which had fairly large nuts, measuring almost 2.0 cm
in length and diameter. Otherwise, the specimens I have collected and examined had
nuts generally less than 1.5 cm in length and diameter. This species probably occurs
also in Shelby County for which Underwood (27) reported having found Phyllactinia
suffulta growing on pin oak. However, there appear to be no specimens on file.
436
Indiana Academy of Science
Figure 4. Q. falcata in Indiana. I = Indiana University; M = Herbarium of Scott
McCoy.
Plant Taxonomy
437
Figure 5. Q. pagoda in Indiana.
VIII. Quercus shumardii Buckl.
The distribution of this species probably has been better documented than that
438
Indiana Academy of Science
Vol. 94 (1985)
r:;
r — i — :.~it_^ •■+-
.H
■H
rvi i-
--4
"l
Figure 6. Q. imbricaria in Indiana.
of any other oak. Kriebel conducted an extensive search for it during the late 1930s
and many herbaria, particularly BU, IND, DEP, and PUL, contain duplicates of his
Plant Taxonomy
439
V
/._
l._
"I- — i
Y>~-4— /
1
I"
L
Figure 7. Q. marilandica in Indiana. I = Indiana University; K = Herbarium of
Ralph Kriebel.
collections. Most of the reports illustrated in Figure 9 are based on specimens collected
440
Indiana Academy of
Science
<
•H
Vol. 94 (1985)
,./•!* i. — v
' r > ,
i r\ _ r 1 - '
r -+ -i • r
|.j I r!_.| J j J---;
, U-j •>-
•i
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-i — <
i^-
Figure 8. Q. palustris in Indiana.
by Kriebel. As can be seen by comparison with Deam's (6) map 793, my map does
not represent a significant change. Two of the county reports by Deam, for Carroll
Plant Taxonomy
441
Figure 9. Q. shumardii in Indiana. □ = var. schneckii. K = Herbarium of Ralph
Kriebel.
and Cass Counties, could not be verified, while two others, Fountain and Whitley
Counties, have been added. The last is based on several specimens I collected in the
442 Indiana Academy of Science Vol. 94 (1985)
south-central part of Whitley County. As noted by Deam (6), Kriebel did not find
Q. shwnardii in Benton County or in the northwest part of the state. None of the
specimens I examined came from that part of the state. In addition to the counties
shown in Figure 9, this species was reported from Vermillion County by Coulter (4),
but no specimen was found to verify this report.
As does Q. borealis, Q. shwnardii has two varieties differentiated primarily by
acorn size. Deam (6) did not provide any details on the distribution of the two varieties
beyond noting that var. schneckii (Britt.) Sarg. ranges "northw. (sic) in the Mississippi
Valley to Wells County, Indiana." Whenever possible, I noted the variety for each
specimen examined and I have indicated their joint distributions in Figure 9. There
is no apparent difference in the distributions of the two varieties, although there are
two counties, Jefferson and Hancock, for which only var. schneckii has been reported.
IX. Quercus velutina Lam.
This is probably the most common oak in Indiana and Deam (6) stated that it
is "without doubt found in every county of the state." Figure 10 shows it to be recorded
from 66 counties. Deam also reported Q. velutina from Crawford, Parke, Perry, and
Pike Counties. No specimens were found for the first three of these and the single
specimen found from Pike County (IND 39321 = Deam 18353) was misidentified.
Two of the records shown in Figure 10, those for Huntington and Whitley Counties,
are based on specimens {Jensen 83-33 and 83-44) I collected during September, 1983.
In addition to the records illustrated in Figure 10, there are literature references
indicating that Q. velutina also may be found in six other counties: Clay (28), Jenn-
ings (19), Martin (23), Orange (24), Switzerland (9), and Wayne (18).
X. Hybrids
The most frequently recorded hybrid is X Q. leana Nutt., which has been found
in 13 counties in Indiana (Figure 11). Deam (6) reported this hybrid from only three
counties, but the tree on which his Lake County report (IND 18546, 18707, 70361
= Deam 18088) was based actually belongs in X Q. runcinata. Deam's report of X
Q. exacta from Posey County (IND 18560, PUL 6701 = Deam 29116) was based on
another tree which also appears to belong in X Q. runcinata. Thus, the occurrence
of X Q. exacta has not been verified for Indiana. Deam's third hybrid report could
be verified. Besides occurring in Knox County, X Q. bushii also occurs in Spencer
and, possibly (as noted under discussion of Q. marilandica), Lawrence Counties.
There are two specimens of X Q. benderi Baenitz (Q. borealis X Q. coccinea),
annotated as such by E.J. Palmer, on file: IND 70236 (Deam 62074) and PUL 7282
(Kriebel 10053). These specimens apparently came from the same tree; they were col-
lected on consecutive days (October 5 and 6, 1942) in the southeast quarter of section
24 of Wells County. The specimens are, however, not of a hybrid but rather, are
from a tree of Q. shumardii. There is another specimen which may be this hybrid.
Collected in Montgomery County, this specimen (WAB 5761 = Bed tel s.n.) was iden-
tified as Q. coccinea, but probably belongs in X Q. benderi.
Other hybrids, and the counties for which records have been verified, are given
below:
X Q. runcinata — Adams, Delaware, Gibson, Lake, Lawrence, Porter, St. Joseph,
Wells, and possibly Kosciusko, Morgan and Tipton;
X Q. paleolithicola — Elkhart, Lagrange, and possibly Kosciusko;
X Q. hawkinsii Sudw. — Lake, Laporte, Porter, Vigo;
X Q. tridentata (A. DC.) Engelm.— Crawford.
Plant Taxonomy
443
V
L-^-i---
,V
.L4
1-i ■
— 1
I.-.v-a r— 1 h..
H i. Wn^
M I
— i
Figure 10. Q. velutina in Indiana. B = Butler University; D = Deam's Herbarium.
Two other putative hybrids, X Q. mutabilis Steyerm. (Q. palustris x Q. shumardii)
and X Q. vaga Palmer & Steyerm. (Q. palustris x Q. velutina) may occur in Gibson
444
Indiana Academy of Science
Vol. 94 (1985)
Figure 11. X Q. leana in Indiana. D = Deam's Herbarium.
and Porter Counties, respectively. The specimens examined give the appearance of
being these hybrids, but the identification is tentative. Recently, one of my students
Plant Taxonomy 445
and I have conducted both morphological and chemical studies indicating that X Q.
vaga occurs in St. Joseph County. Research is being continued in order to verify its
occurrence. A complete list of specimens on which these hybrid reports are based is
available from the author on request.
XI. Excluded Taxa
As noted in the introduction, Q. nigra L. was shown to be in Indiana by Preston
(21). I found a single specimen (WAB s.n. = Clapp s.n.), collected near New Albany
in 1836, identified as Q. nigra. Deam had correctly annotated this specimen as Q.
marilandica. Interestingly, Coulter (3) cited a Deam specimen from Crawford County
in support of his claim that Q. nigra is fairly well distributed throughout the state,
near streams and swamps, sometimes in upland regions. I don't know which species
Coulter was describing, because there is no other taxon of Quercus, found in such
habitats in Indiana, that even remotely resembles Q. nigra. The confusion regarding
this taxon is exacerbated by Andrews' (1) report that it is also found in Monroe County.
The only specimen of Q. nigra that I have located, from Delaware County (BSU 1273
= Hughes s.n.), was originally identified as Q. nuttallii Palmer. Undoubtedly, this
specimen was taken from a tree planted somewhere near the Ball State University campus.
Quercus phellos L. also should be excluded. Coulter (3) noted that reports of
this species probably were based on misidentification of a narrow-leaved tree of Q.
imbricaria. I came across two specimens of Q. phellos (BU 96603 = Friesner 24804;
BSU 1278 = Olsen s.n.), both of which must have come from planted trees. The
former was collected in Versailles State Park, Ripley County, and the latter in Muncie,
Delaware County.
Acknowledgments
This research was supported by grants from Saint Mary's College and the Indiana
Academy of Science. I would like to thank Willard Yates, Richard Mayes, William
Burger, Lewis Johnson, Gene Williamson, John McCain, Almut Jones, and Robert
Petty for their assistance with this research. Almut Jones provided many helpful com-
ments in a critical reading of an earlier version of this manuscript.
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30. Ware, S. 1967. The morphological varieties of southern red oak. J. Tenn. Acad.
Sci. 42:29-36.
A Preliminary Review and Multiple-entry Key to the Rust Fungi on
Cyperaceae and Juncaceae in Indiana
John W. McCain
Department of Botany and Plant Pathology
Purdue University, West Lafayette, Indiana 47907
The Manual of the Rusts in United States and Canada (1), the crucial reference
for identification of this highly important group of plant parasitic fungi in North
America, was published in 1934, the 50th year of the Indiana Academy of Science.
As a pilot project to develop techniques for revising the manual, including computer-
aided compilations of distributions and generation of new keys, a subset of the North
American rust fungi, the Indiana species, was chosen for review in 1984, the 100th
year of this academy. The most recent compilation of the Indiana rust fungi was in
1916-1921 (3).
Because the 165 species listed by Jackson (3) still make a large group, the
focus of this paper was further narrowed. Rust fungi attack hosts in many families
of flowering plants (about 100 in North America — 1), but certain families are especial-
ly burdened with these pathogens. The rust fungi on nearly all of these major host
groups (Gramineae, Leguminosae, etc.) have been re-studied since 1934, except for
those of Cyperaceae. In fact, Savile (10) called for a "moratorium" on the publication
of records on Carex rusts until a thorough study could be done. This paper is a first
step in such a study. The rushes (Juncaceae) often occur together with the Cyperaceae
in nature. Their rust pathogens should be studied along with those on the sedges, for
they may be related as their hosts are (9). Aecia of the Indiana sedge/rush pathogens
are on hosts representing nine other families of flowering plants.
Specimen label information and annotation notes, especially including camera
lucida drawings and spore measurements, were collated for the 650 specimens of In-
diana rust fungi on sedges and rushes in the Arthur Herbarium (PUR), Purdue Univer-
sity, West Lafayette, IN. In addition, forty previously overlooked collections were found
on phanaerogamic specimens in the Kriebel Herbarium of Purdue University and twenty
new collections were made in the field.
Of the Arthur Herbarium specimens, 90% are at least 60 years old. Knowledge
of the hosts and rusts has increased over the last six decades, the distribution of these
taxa has probably changed, and the dried specimens no longer contain viable spores
for re-verification of the life cycle studies performed by Arthur (1}. In addition, the
distribution data are skewed towards Tippecanoe County (57% of the PUR specimens)
and away from eastern and southern Indiana: no specimens are available for any species
from 44 of the 92 Indiana counties, including Allen, Delaware, Floyd, Vanderburgh,
and Wayne. This is an example of the distribution of collectors, not of the taxa col-
lected (5). Clearly, systematic state-wide collecting of fresh specimens is necessary for
valid biogeographical or phenological hypotheses. A series of such trips is being planned
for 1985.
Fifteen species of rust fungi are known on Cyperaceae in Indiana, but only three
on Juncaceae. Only two of the more than 100 genera of rust fungi are represented
here: Puccinia and Uromyces. A taxonomic "splitter" might add at least six other
sedge rust species for the state, and five more occur nearby in Wisconsin or Michigan.
Although one would expect Indiana to have been well-surveyed, the complete life cycles
of four (U. junci-effusi Syd., U. minutus Diet., U. rhynchosporae Ell., and U. valens
Kern) of these 18 rust species are still unknown. The other species are heteroecious
and macrocylic (1). Fifty-six of the 217 species of Cyperaceae in Indiana (2) have been
447
448 Indiana Academy of Science Vol. 94 (1985)
collected with rust on them (26%), but only four of the 26 species of Juncaceae in
this state (15%) are known hosts.
Four of the Indiana rust species are known from only one county (P. obscura
Schroet. ex Pass., U. americanus Speg., U. junci-effusi, and U. valens); three from
two counties (P. minutissima Arth., U. minutus, and U. rhynchosporae), and five
others from three to six counties (P. cyperi Arth., P. eleocharidis Arth., P. obtecta
Peck, U. lineolatus (Desm.) Schroet. in Rabh., and U. perigynius Halst.). Puccinia
canaliculata (Schw.) Lagh. has been found in nine counties, P. angustata Peck in 10,
U. silphii Arth. in 19, P. bolleyana Sacc. in 22, P. caricina in 23 (including P. caricina
var. limosae (P. Magn.) Jorst. from one county), and P. dioicae P. Magn. in 31 coun-
ties. None of these are as widespread as their hosts. For example, Scirpus cyperinus
(L.) Kunth. is known from at least 50 counties (2), or five times as many counties
as its rust pathogen, P. angustata. Maps of all rust species collections were prepared,
but none showed any coherent geographic trends.
One-third of the Indiana sedge or rush collections bearing rust fungi could be
assigned to P. caricina, one-third to P. dioicae, and the remaining third to all the
other 16 species combined. Puccinia caricina is now known on 17 species of Carex
in Indiana and on 110 species when all U.S. and Canadian records are counted. Puc-
cinia dioicae has 26 Indiana Carex hosts and 137 total north of Mexico. Verifying
the identity of the host fragments in many of the old PUR collections is probably
impossible, but a large number of the hosts were originally named by Dr. K.K. Mackenzie
of the New York Botanical Garden, one of the foremost Carex scholars ever, so they
probably can be accepted as correct. When all the North American collections are
tabulated, a tendency appears for these two rust species to favor certain of the sections
of the genus Carex (7). For example, 36 species of the section Ovales harbor infections
of P. dioicae, but P. caricina is found in the PUR collections on only one species
of Ovales, Carex multicostata Mkze. from California. Section Acutae (especially C.
aquatilis Wahl. and C. stricta Lam.) consistently includes hosts for P. caricina (16
to 2 for P. dioicae). Some other sections for which the rust species show preferences
(based on PUR records) include Bracteosae (3 species are hosts for P. caricina, 11
for P. dioicae), Atratae (6, 1), Cryptocarpae (4, 0), Laxiflorae (6, 1), Limosae (5,
0), Montanae (1, 9), Multiflorae (0, 6), Pseudocypereae (4, 1), Stellulatai (0, 9), Sylvaticae
(= Debiles, 5, 2), and Triquetrae (3, 1). Puccinia dioicae hosts are more common
in Carex subgenus Vignea, P. caricina hosts in subgenus Eucarex. A few sections in-
clude species susceptible to both rust species: Divisae (3, 3), Heleonastes (= Canescentes,
5, 3), Hirtae (2, 4), Lupulinae (2, 3), Paludosae (3, 4), Phyllostachyeae (2, 2), Virescentes
(4, 2), and Vesicariae (= Physocarpae, 3, 3). Nevertheless, these trends may be useful
as predictors. The first rust collection on Carex davisii Schw. & Torr. was predicted
to be P. caricina because section Gracillimae included three sedge species attacked
by P. caricina but only one by P. dioicae and, in fact, the collection did key to P.
caricina (5). In the Gramineae, such fidelity of rust fungi to certain host tribes has
been used to show that a grass genus with the "wrong" rust pathogen should be re-
assigned to a different tribe (4). The section preferences of the rust fungi should now
be reported to Carex specialists who might use them to spot similar host taxonomy
corrections.
These two common rust species, P. caricina and P. dioicae, may actually be species
complexes, with arrays of aecial hosts in different families. Therefore, each may be
groups of sibling species or races isolated reproductively by their different aecial hosts,
yet scarcely or not at all separable by fungal morphology. In Europe, inoculation studies
have shown consistent separation of populations, which are recognized as distinct species
(1), and Savile has used those segregate names for his North American sedge rust col-
lections (10). The detailed life cycle studies on North American collections reported
Plant Taxonomy 449
by Arthur (1) suggested a trend to host specialization, but not strong enough to sup-
port delimitation of species at this time. Varieties or formae speciales (f. sp.) may
be justifiable. The populations of spores collected from the various aecial hosts as
yet show no readily identifiable patterns of favoring certain species or sections of the
telial host genus Carex and no immediately obvious geographic trends on the broad
scale. Perhaps fine-grained ecological data, such as whether the /?/6es-infecting isolates
of P. caricina come from drier or higher ground than the aecial collections from Ur-
tica hosts, may yield some guidance (8). No experiments ever have tested whether these
rust fungi are merely opportunistic, producing pathotypes that may utilize either aecial
host species if it is present and environmental conditions are right.
There is a slight morphological trend in the urediniospores of P. caricina collec-
tions. If the ranges of urediniospore length and width are plotted on a graph (Figure
1), the collections associated with aecia on Urtica spp. have the largest urediniospores
(data from 1), and those from sedge infections following aecia on Ribes spp. are smaller,
with a narrower length range. Most P. caricina urediniospores have 3 (or 4) equatorial
germination pores. A variant form with one pore near the hilum of the spore occurs
on Ribes and is intermediate in size between the two former groups. No patterns of
urediniospore morphology, host supraspecific taxa fidelity, or geography are evident
at this time to sort out aecial populations of P. dioicae on Onagraceae, Phrymaceae,
Valerianaceae, Thymeleaceae (these collections have been challenged as actually belonging
to a grass rust — 10), or several tribes of the Compositae. The reported variations in
aeciospores (10) have not vet been reviewed in this study.
Finally, a preliminary multiple-entry key was developed as a working tool, based
on published descriptions of these taxa (1). Because this type of key requires that all
potentially useful character states be scored for all taxa, it is useful for finding gaps
or inconsistencies in the data and, thus, show which taxa need further review. For
example, in its current form, some leads show that the published literature contains
synonymous or inconsistently applied terminology. Others, such as color characters,
are probably too subjective to prove useful. Instructions for the use of this type of
key are given by Pitt (6). Subsequently, the characters for each taxon can be listed
as formulae (part IV of the key) and be readily translatable to a computerized key.
Comparisons of the formulae reveal that some taxa are not clearly separable by the
key, unless the host has been identified first. Measurements of a sample of spore lengths
and widths are also necessary for identification of uredinial states of five of the rust
taxa on Carex, for telial specimens of three of these, and for uredinial material of
the two Puccinia rusts on Scirpus. Given an allowance for variation in some characters,
such as spore shape, the other taxa can be keyed out by qualitative features once the
host is known.
In summary, as a sample subset and first step towards a revision of the present
manual of North American rust fungi (1), the species were reviewed that infect members
of the Cyperaceae and Juncaceae in Indiana. The incomplete distribution records and
the need for further life cycle studies indicate that only collecting and testing of new
specimens, supported by computerized data management, will provide complete, cor-
rect information from which to produce an up-to-date manual.
J.F. Hennen offered critical advice on the ms.
Literature Cited
1. Arthur, J.C. 1934. Manual of the rusts in United States and Canada. Purdue
Research Foundation, West Lafayette, IN, 438 p.
2. Deam, C.C. 1940. Flora of Indiana. Dept. of Conservation, Div. of Forestry,
State of Indiana, Indianapolis, 1236 p.
450
40
Indiana Academy of Science
Vol. 94 (1985)
30
urn
20
10
r ■
!U
N
i
D
i L
!R
10
um
20
30
Figure 1 . Urediniospore size ranges in Puccinia cahcina and P. dioicae on Carex spp.
N — from uredinal infections of P. caricina following aecia on Urtica spp.; R — from P.
cahcina uredinia following aecia on Ribes spp.; U — one-pored urediniospore variants
of P. caricina following aecia on Ribes spp.; D — urediniospores of P. dioicae. Based
on data from Arthur (1).
3. Jackson, H.S. 1916-1921. The Uredinales of Indiana. Proc. Indiana Acad. Sci.
25: 429-475 (1916); II. 27: 133-137 (1918); III. 30: 165-182 (1921).
4. McCain, J.W., and J.F. Hennen. 1982. Is the taxonomy of Berberis and Mahonia
(Berberidaceae) supported by their rust pathogens Cumminsiella santa sp. nov.
and other Cumminsiella species (Uredinales)? Syst. Bot. 7: 48-59.
5. McCain, J.W., and J.F. Hennen. 1982. Notes on biogeography and new records
Plant Taxonomy
451
of rust fungi in the Great Lakes region. Proc. Indiana. Acad. Sci. 91: 504-514.
6. Pitt, I.J. 1974. A synoptic key to the genus Eupenicillium and to sclerotigenic
Penicillium species. Canad. J. Bot. 52: 2231-2236.
7. Savile, D.B.O. 1954. The fungi as aids in the taxonomy of flowering plants. Science
120 (3210): 583-585.
8. Savile, D.B.O. 1965. Puccinia karelica and species delimination in the Uredinales.
Canad. J. Bot. 43: 231-238.
9. Savile, D.B.O. 1971. Co-ordinated studies of parasitic fungi and flowering plants.
Naturaliste Canada 98: 535-552.
10. Savile, D.B.O. 1973. Aeciospore types in Puccinia and Uromyces attacking
Cyperaceae, Juncaceae and Poaceae. Rept. Tottori Mycol. Inst. (Japan) 10:
225-241.
Multiple-Entry Key to Species of Puccinia and Uromyces on Indiana
Cyperaceae and Juncaceae
Fungus Taxa.
1. P. angustata s. 1.
2. P. bolleyana
3. P. canaliculata
4. P. caricina s. 1.
4a. P. caricina var. limosae
5. P. cyperi
6. P. dioicae s. 1.
7. P. eleocharidis
8. P. minutissima
9. P. obscura
Key to Host Taxa. (Bold face type indicates rust taxa appearing under more
than one lead).
1. Carex—2, 4, 4a, 6, 8, 14, 15, 18
2. Cyperus — 3, 5 6.
3. Dulichium — 6 7.
4. Eleocharis — 7 8.
5. Eriophorum — 1 9.
10. P. obtecta
11. U. americanus
12. U. junci-effusi
13. U. lineolatus
14. U. minutus
15. U. perigynius
16. U. rhychosporae
17. U. silphii
18. U. valens
J uncus — 12, 17
Luzula — 9
Rhynchospora — 1 6
Scirpus—l, 10, 11, 13
II. Key to Uredinial Material.
A. Number of Germination Pores per Spore.
1 . One— 4
2. Two— 1, 2, 3, 4, 5, 6, 8, 9, 10, 14, 15, 16, 17
3. Three— 4, 4a, 5, 7, 12, 13, 14
4. Four— 4, 4a, 5, 7, 11, 12, 13, 18
5. Five— 7, 11
6. Six— 11
B. Spore Germination Pore Position.
1. Superequatorial to Apical— 1, 2, 6, 8, 9, 15, 17
2. Equatorial— 1, 3, 4, 4a, 5, 7, 10, 11, 12, 13, 14, 16, 18
3. Near Hilum — 4
C. Spore Length.
1. > 36 /im— 7,11
2. 25-35 urn— 1, 2, 3, 4, 5, 6, 7, 9, 10, 11, 12, 13, 15, 16, 18
3. < 25 urn— 1, 2, 3, 4, 4a, 5, 6, 8, 9, 12, 13, 14, 15, 16, 17, 18
452 Indiana Academy of Science Vol. 94 (1985)
D. Spore Shape.
1. Ellipsoid— 1, 2, 3, 4, 4a, 5, 6, 7, 8, 9, 10, 11, 12, 13, 14, 15, 16 17,
18
2. Obovoid— 1, 2, 3, 4, 4a, 5, 6, 7, 9, 13, 14, 15, 16, 17
3. Globoid— 6, 8, 9, 15
4. Oblong— 11
E. Spore Wall Color.
1. Cinnamon-brown— 1, 3, 4, 4a, 5, 6, 7, 10, 11, 12, 13, 14, 15, 17, 18
2. Chestnut-brown — 2, 4, 14, 18
3. Golden- or yellowish-brown — 3, 8, 9, 16
4. Golden-yellow — 17
F. Sorus Color.
1. Cinnamon-brown— 1, 3, 4, 4a, 5, 6, 7, 9, 10, 11, 12, 13, 14, 15, 17,
18
2. Chestnut-brown — 2, 8, 16
3. Golden-brown — 9
G. Sorus Emergence.
1. Sorus Erumpent, Pulverulent — 1, 2, 4, 4a, 6, 8, 9, 10, 12, 13, 14,
15, 18
2. Sorus Tardily Naked or Opening Only by Slits — 3, 5, 7, 11, 16, 17
H. Sorus Position.
1. Hypophyllous— 1, 2, 3, 4, 4a, 5, 6, 8, 9, 10, 12, 13, 14, 15, 16, 17,
18
2. Epiphyllous— 9, 10, 12, 13, 17
3. Culmicolous— 7, 9, 11, 17
III. Key to Telial Material.
A. Number of Cells per Teliospore.
1. Two— 1, 2, 3, 4, 4a, 5, 6, 7, 8, 9, 10
2. One— 11, 12, 13, 14, 15, 16, 17, 18
B. Pedicel Color.
1. Colored— 1, 3, 5, 7, 8, 9, 10, 11, 12, 13, 17
2. Colorless— 2, 4, 4a, 6, 11, 14, 15, 16, 18
C. Pedicel Length.
1. Less than length of spore— 3, 4, 4a, 5, 6, 7, 8, 9, 10, 11, 13, 14, 15
2. About length of Spore— 1, 2, 6, 9, 12, 15, 16, 17
3. Longer than Spore — 17, 18
D. Sorus with Paraphyses.
1. No— 1, 2, 4, 4a, 5, 6, 7, 8, 9, 12, 13, 14, 15, 16, 17, 18
2. Yes— 3, 6, 10, 11
E. Sorus Emergence.
1. Erumpent Early, Pulverulent— 1, 2, 4, 4a, 6, 7, 8, 9, 12, 13, 14, 15,
16, 18
2. Remaining Covered or Loculate — 3, 5, 6, 10, 11, 17
F. Spore Wall Thickness at Apex.
1. > 10 /Am— 1, 2, 4, 5, 6, 8, 9, 10, 13, 15
2. 5-10 iim— 1, 2, 3, 4, 4a, 5, 6, 7, 8, 9, 10, 11, 12, 13, 14, 15, 16, 17,
18
3. < 5 /mi— 1, 3, 4, 7, 11, 16
G. Spore Length.
1. > 50 /mi— 1, 2, 3, 4, 5, 6, 7, 8, 10, 11, 13
2. > 25-20 /mi— 1, 2, 3, 4, 4a, 5, 6, 7, 8, 9, 11, 12, 13, 15, 16, 17, 18
3. < 25 fim— 12, 14, 15, 16
Plant Taxonomy 453
H. Spore Wall Color.
1. Cinnamon-brown — 1, 3,
2. Chestnut-brown— 1, 2, 4, 4a, 5, 6, 7, 8, 9, 10, 12, 13, 14, 15, 16,
17, 18
3. Yellow or Golden— 9, 11
4. Colorless — 11
5. Paler in Lower Part of Spore — 3, 5, 7, 16
I. Teliospore Shape.
1. Clavate— 1, 2, 3, 4, 5, 6, 3. Obovoid— 12, 14, 15, 17,
7, 9, 10, 16 18
2. Oblong— 2, 3, 4a, 5, 6, 4. Ellipsoid— 9, 12, 13, 15,
7, 8, 11 16
5. Cylindric— 1, 10, 11
6. Cuneiform — 16
J. Teliospore Apex.
1. Rounded or Obtuse— 1, 2, 3, 4, 4a, 5, 6, 7, 8, 9, 10, 13, 14, 15, 16,
17, 18
2. Truncate— 4a, 5, 6, 7, 9, 11, 12, 14, 15, 16, 17
3. Acute, Acuminate, or Pointed — 1, 3, 4, 10, 12, 15, 16, 17, 18
IV. Species Formulas (compiled by listing each lead in the key under which the species
occurs). Uredinial states are listed first.
1. 2, 12, 23, 12, 1, 1, 1, 1; 1, 1, 2, 1, 1, 123, 12, 12, 15, 13.
2. 2, 1, 23, 12, 2, 2, 1, 1; 1, 2, 2, 1, 1, 12, 12, 2, 12, 1.
3. 2, 2, 23, 12, 13, 1, 2, 1; 1, 1, 1, 2, 2, 23, 12, 15, 12, 13.
4. 1234, 23, 23, 12, 12, 1, 1, 1; 1, 2, 1, 1, 1, 123, 12, 1, 1, 14.
4a. 34, 2, 3, 1, 1, 1, 1, 1; 1, 2, 1, 1, 1, 1, 2, 2, 2, 12.
5. 234, 2, 23, 12, 1, 1, 2, 1; 1, 1, 1, 1, 2, 12, 12, 25, 12, 12.
6. 2, 1, 23, 12, 2, 2, 1, 1; 1, 2, 12, 12, 12, 12, 12, 2, 12, 12.
7. 345, 2, 12, 12, 1, 1, 2, 3; 1, 1, 1, 1, 1, 23, 12, 25, 12, 12.
8. 2, 1, 3, 13, 3,2, 1, 1; 1, 1, 1, 1, 1, 12, 12,2,2, 1.
9. 2, 1, 23, 123, 3, 13, 1, 123; 1, 1, 12, 1, 1, 12, 2, 23, 14, 12.
10. 2, 1, 2, 1, 1, 1, 1, 12; 1, 1, 1, 2, 2, 12, 1, 2, 15, 13.
11. 456, 2, 12, 14, 1, 1, 2, 3; 2, 12, 1, 2, 2, 23, 12, 34, 25, 2.
12. 34, 2, 23, 1, 1, 1, 1, 12; 2, 1, 2, 1, 1, 2, 23, 2, 34, 23.
13. 34, 2, 23, 12, 1, 1, 1, 12; 2, 1, 1, 1, 1, 12, 12, 2, 4, 1.
14. 23, 2, 3, 12, 12, 1, 1, 1; 2, 2, 1, 1, 1, 2, 3, 2, 3, 12.
15. 2, 1, 23, 123, 1, 1, 1, 1; 2, 2, 12, 1, 1, 12, 12, 2, 34, 123.
16. 2, 2, 23, 12, 3, 2, 2, 1; 2, 2, 2, 1, 1, 23, 23, 25, 146, 23.
17. 2, 1, 3, 12, 14, 1, 2, 123; 2, 1, 23, 1, 2, 2, 2, 2, 3, 123.
18. 4, 2, 23, 1, 12, 1, 1, 1; 2, 2, 3, 1, 1, 2, 2, 2, 3, 13.
Additions to the Flora of Pike and Gibson Counties, Indiana
Thomas W. Post
Department of Natural Resources
Division of Reclamation
Indianapolis, Indiana 46204
Over a three year period several field trips were made to Pike and Gibson Coun-
ties, Indiana, to inventory potential natural areas of county and statewide significance.
The habitats investigated included upland forest, lowland forest, sloughs and seasonally
flooded bottomland areas. Since the primary reason for the field work was to look
for areas with natural integrity left to them, roadsides and fallow fields were not
investigated.
This inventory resulted in the collection of 2 vascular plant species not previously
reported for Gibson County by Deam (2) or later updates to the flora
(3,6,8,9,10,12,13,14,15). With the 2 species reported here the vascular flora recorded
for Gibson County totals 620 species: 18 fern or fern allies, 1 gymnosperm, 167
monocots, and 434 dicots.
This inventory also resulted in the collection of 10 vascular plant species not
previously reported for Pike County by Deam (2) or later updates to the flora
(4,5,6,7,12). With the ten species reported here, the vascular flora recorded for Pike
County totals 303 species: 9 fern or fern allies, 88 monocots, and 206 dicots. Of special
interest are Carex louisianica and Ilex decidua, both considered rare in Indiana by
Bacone et al (1). The Carex was collected on the shaded shore of a shallow pond
and the Ilex was collected in lowlands woods along the Patoka River.
The following list records the new record by county, plant name, and author's
collection number. The nomenclature follows Fernald (11). All specimens are deposited
in the Deam Herbarium of Indiana University.
Acknowledgments
The author would like to thank Jim Aldrich, Brian Abrell, Mike Homoya, and
Helene Stares for help in collection and identification of specimens and the Division
of Reclamation for support of this work.
Gibson County
Orchidaceae
Aplectrum hyemale (Muhl.) Torr.; #127
Cruciferae
Amoracia aquatica (Eat.) Wieg; #154
Pike County
Polypodiaceae
Cystopteris fragilis (L.) Bernh.; #118
Cyperaceae
Carex granulans Muhl; #161
Carex louisianica Bailey; #171
Liliaceae
Camassia scilloides (Raf.) Cory; #150
455
456 Indiana Academy of Science Vol. 94 (1985)
Corylaceae
Alnus serrulata (Ait.) Willd.; #138
Cruciferae
lodanthus pinnatifidus (Michx.) Stewd.; #125
Crassulaceae
Sedum ternatum Michx., #149
Oxalidaceae
Oxalis grandis Small; #121
Aquifoliaceae
Ilex decidua Walt.; #137
Compositae
Eupatorium hyssopifolium L.: #147
Literature Cited
1. Bacone, J. A. and C. Hedge. 1980. A Preliminary List of Endangered and
Threatened Vascular Plants in Indiana. Proc. Indiana Acad. Sci. 89:359-371.
2. Deam, C.C. 1940. Flora of Indiana. Indiana Department of Conservation, Divi-
sion of Forestry. Indianapolis. 1236 p.
3. Deam, C.C, R.C. Friesner, R. Kriebel, and T.G. Yuncker. 1940. Indiana Plant
Distribution Records. Proc. Indiana Acad. Sci. 50: 72-78.
4. Deam, C.C, R.C. Friesner, R. Kriebel, and T.G. Yuncker. 1941. Indiana Plant
Distribution Records, II. Proc. Indiana Acad. Sci. 51:120-129.
5. Deam, C.C, R.C. Friesner, R. Kriebel, and T.G. Yuncker. 1942. Indiana Plant
Distribution Records, III. Proc. Indiana Acad. Sci. 52:97-108.
6. Deam, C.C, R.C Friesner, R. Kriebel, and T.G. Yuncker. 1944. Indiana Plant
Distribution Records, V. Proc. Indiana Acad. Sci. 54:91-99.
7. Deam, C.C, R.C. Friesner, R. Kriebel, and T.G. Yuncker. 1945. Indiana Plant
Distribution Records, VI. Proc. Indiana Acad. Sci. 55:50-64.
8. Deam, C.C, R.C. Friesner, R. Kriebel, and T.G. Yuncker. 1947. Indiana Plant
Distribution Records, VIII. Proc. Indiana Acad. Sci. 57:81-86.
9. Deam, C.C, R.C. Friesner, R. Kriebel, and T.G. Yuncker. 1949. Indiana Plant
Distribution Records, X. Proc. Indiana Acad. Sci. 59:48-52.
10. Deam, C.C, R.C. Friesner, R. Kriebel, and T.G. Yuncker. 1951. Indiana Plant
Distribution Records, XI. Proc. Indiana Acad. Sci. 61:72-77.
11. Fernald, M.L. 1950. Gray's Manual of Botany, eighth edition, American Book
Company, New York, New York. 1632 pp.
12. Humbles, J. 1965. Indiana Plant Distribution Records, XIX. Proc. Indiana Acad.
Sci. 75:231-232.
13. Humbles, J. 1970. Indiana Plant Distribution Records, XXI. Proc. Indiana Acad.
Sci. 80:413.
14. Kay, S.B., and J. Humbles, 1974. Indiana Plant Distribution Records, XXII.
Proc. Indiana Acad. Sci. 84:428.
15. McClain, W.E. 1980. Additions to the Flora of Gibson County, Indiana. Proc.
Indiana Acad. Sci. 90:395-397.
Gravel Hill Prairies of Indiana
Thomas W. Post, John A. Bacone and James R. Aldrich
Division of Nature Preserves
Indiana Department of Natural Resources
Indianapolis, Indiana 46204
Introduction
Gravel hill prairies are native grasslands occurring on gravel deposits. They are
termed hill prairies because they occur on relatively steep south, southwest, or west
facing slopes (9). This xeric community is previously undescribed in studies of Indiana's
plant communities (12) and contain a flora quite different from the characteristic "tall
grass prairie" of northwest Indiana (5, 12). In this paper we discuss gravel hill prairies
recently located in Tippecanoe County and the methods used to locate and describe
them. We also compare them with hill prairies in other states. A preliminary species
list is presented, including information concerning some very rare floristic elements (2).
Methods
A systematic search for remaining gravel hill prairies was made in 1979 and 1980.
Past evidence of the existence of this type of prairie included early plant collection
records (8) and a report by Betz (5). The methods used were similar to those used
to find glades in Harrison County, Indiana (1). The 1971 aerial photographs were
examined at the Agriculture Stabilization and Conservation Service office, in con-
junction with the 7.5 minute United States Geological Survey topographic quadrangle
map, and the soil survey of Tippecanoe County (20). Openings along the gravel bluffs
of the Wabash River and tributaries, primarily Wea Creek, were examined in an effort
to locate natural communities. Eight sites were found to have potential. These sites
were then checked during an aerial survey in a high-winged aircraft, from an elevation
of 1000 feet.
During the ground survey three significant prairie remnants were identified. Several
other more disturbed sites, containing some prairie vegetation, were also located. The
three prairie remnants were visited throughout the past several growing seasons. A
species list was compiled for each site, and rare species were located and mapped.
Nomenclature follows Gleason and Cronquist (11).
The Study Area
All three remaining gravel hill prairies are located in central Tippecanoe County.
Historically this area was a mix of oak woods and prairies as recorded by the early
land surveyors (22). Gravel hill prairies in presettlement times covered a small percen-
tage of the total land area of TippecanocCounty, while 35% of the county was covered
by tall grass prairie typically found on silt loam soils in west-central Indiana. Today
it is difficult to find any prairie in this region, as the area has been intensively altered
by agricultural and industrial uses.
Topographic and edaphic conditions form a narrow continuum from dry to dry
mesic in the 3 sites which are located on a steep south or west facing bluff between
an elevation of 570 to 600 feet. The soil type under each prairie is a Rodman gravelly
loam (20). This soil is found on steep slopes of gravel terraces, kames and eskers,
usually along the Wabash and Tippecanoe Rivers and a few of their major tributary
streams in Carroll, Tippecanoe, Fountain, Parke and Vermillion Counties (4, 15, 18,
20, 21). Characteristics of this soil types are a dark brown gravelly loam, underlain
457
458 Indiana Academy of Science Vol. 94 (1985)
by stratified calcareous sand and gravel, mildly alkaline to alkaline in pH with high
permeability, rapid runoff and excessive drainage.
All three prairies are found within a mile of each other on the bluffs overlooking
Wea Creek. Names given to each site are Lookout Point, Wabash Breaks, and Wea
Creek respectively. Wea Creek is the largest of the three although each hill prairie
exists today as a small (less than one hectare) opening in a forest composed of Quercus
macrocarpa, Q. imbricaria, Carya ovata, Prunus serotina, and Cercis canadensis. It
is likely that each was larger in the past but has shrunk in size due to a number of
disturbances. These disturbances include horses causing erosion, brush invasion due
to lack of fire (5) and mining of the gravel resulting in the outright destruction of
part of the Wea Creek prairie.
Results and Discussion
These gravel hill prairies still exist today in Indiana due to soil and topography.
Environmental conditions are more xeric in hill prairies than in surrounding forests
due to higher light intensities, higher wind velocity, higher soil temperatures, higher
daily air temperatures and higher evaporation rates (9, 14). No hill prairies or poten-
tial hill prairies have been found on north or east facing slopes probably due to more
mesic conditions.
As a result of xeric conditions a unique assemblage of plants occurs in these gravel
hill prairies (8). This assemblage is a combination of typical prairie plants found in
Indiana and prairie plants considered rare in Indiana but common in mid-grass prairies
farther west. Prairie plants found on these sites but rarely found elsewhere in Indiana
include: Androsace occidentalis, Arenaria patula, Aster oblongifolius, Besseya bullii,
Erysimum asperum, Lithospermum incisum, and Muhlenbergia cuspidata. Astragalus
tennesseenis, Psoralea tenuiflora, Onosmodium molle var. hispidissimum, and Linum
sulcatum occurred in the Wea Creek vicinity historically. The Astragalus and Psoralea
are now considered extirpated in the state and the others endangered and threatened (2).
To date, 154 vascular plant species representing 54 families have been identified
at the three prairies. At Lookout Point 80 taxa including one rare species and 5 state
endangered species were identified. At Wabash Breaks 101 taxa were found including
one rare species and three state endangered species. At Wea Creek 86 taxa were found
to occur including one rare species and seven state endangered species (Table 1).
The visual aspect is that of mid-grass prairie, i.e. vegetation height typically 2
feet high. The common grasses found on Indiana's gravel hill prairies are Bouteloua
curtipendula, Andropogon scoparius, and Stipa spartea. Other associates included Allium
cernuum, Amorpha canescens, Aster oblongifolius, Petalostemum purpurem, Erysimum
asperum and Kuhnia eupatorioides (Table 1). These species, in conjunction with the
dominance of little bluestem and side-oats grama grass, are more typical of mid-grass
prairies.
The Illinois Natural Areas Inventory recognized both gravel hill prairie and loess
hill prairie (23). There is a close similarity in species composition between Indiana
and Illinois gravel hill prairies (9, 23). Species in common include Bouteloua curtipen-
dula, Andropogon scoparious, A. gerardi, Sporobulus heterolepis, Sorghastrum nutans,
Androsace occidentalis, Aster oblongifolius, Besseya bullii and Lithospermum incisum.
In a more recent Illinois study, 51% of the species including Lithospermum incisum,
Muhlenbergia cuspidata, Onosmodium hispidissimum and Besseya bullii reported from
the Tazewell Gravel Terrace Prairie (23) occur on Indiana gravel hill prairies. Another
of Indiana's gravel hill prairie components, Astragalus tennesseensis also occurs at this
Illinois site.
The Illinois loess hill prairies were found to occur on upper slopes of southwest
Plant Taxonomy
459
Table 1. Vascular Flora of Three Tippecanoe County Gravel Hill Prairies
Taxa
Lookout
Wabash
Wea
Point
Breaks
Creek
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
# Achillea millefolium
# Alliaria officinalis
Allium cernuum
# Amaranthus sp.
Ambrosia artemisiifolia
Ambrosia trifida
Amelanchier arborea
Amorpha canescens
Andropogon gerardi
Andropogon scoparius
# Androsace occidentals
Anemone cylindrica
Anemone quinquefolia
Antennaria plantaginifolia
Apocynum androsaemifolium
Aquilegia canadensis
# Arenaria patula
# Artemisia sp.
Asclepias syriaca
Asclepias tuberosa
Asclepias verticillata
Asclepias viridiflora
Aster ericoides
# Aster oblongifolius
# Avena fatus
# Besseya bullii
Bidens sp.
Bouteloua curtipendula
# Bromus inermis
Carex pensylvanica
Carya ovata
Cassia fasciculata
# Catalpa speciosa
Celastrus scandens
Celtis occidentalis
Cercis canadensis
Claytonia virginica
Comandra umbellata
Convolvulus sepium
Coreopsis palmata
Coreopsis tripteris
Cornus racemosa
Crataegus sp.
Cyperus sp.
U Daucus carota
Dodecatheon meadia
U Draba verna
Elymus canadensis
Equisetum arvense
Erigeron sp.
# Erysimum asperum
Euonymus atropurpureus
Eupatorium altissimum
Eupatorium rugosum
Eupatorium serotinum
Euphorbia corollata
# Euphorbia sp.
Fragaria virginiana
Fraxinus americana
Galium aparine
460
Indiana Academy of Science
Vol. 94 (1985)
Table 1.— Continued
Taxa
Lookout
Wabash
Wea
Point
Breaks
Creek
X
X
X
X
X
X
X
X
X
X
X
X
X
X
Galium circaezans
Helianthus divaricatus
Helianthus grosseserratus
Helianthus occidentalis
Heuchera richardsonii
Hypericum sphaerocarpum
Hypericum spp.
Hypoxis hirsuta
Hystrix patula
Kuhnia eupatorioides
Lactuca sp.
# Lepidium sp.
Lespedeza capitata
Lithospermum canescens
# Lithospermum incisum
Lonicera sp.
# Medicago sativa
# Melilotus alba
Mirabilis nyctaginea
Monarda fistulosa
ft Morus alba
# Muhlenbergia cuspidata
Muhlenbergia racemosa
Oenothera biennis
Opuntia humifusa
Osmorhiza longistylis
Oxalis sp.
Panicum virgatum
Panicum sp.
Parthenocissus quinquefolia
Penstemon hirsuta
Petalostemum purpureum
Phlox bifida
Physalis sp.
Physostegia virginiana
Phytolaca americana
H Plantago aristata
Platanus occidentalis
ft Poa pratensis
Polanisia graveolens
Polygonatum biflorum
# Potentilla recta
Prunus serotina
Prunus virginiana
Ptelea trifoliata
Pycnanthemum sp.
Quercus imbricaria
Quercus macrocarpa
Quercus muhlenbergia
Quercus velutina
Ratibida pinnata
Rhus aromatica
Rhus glabra
Rhus radicans
Ribes sp.
ft Robinia pseudoacacia
Rosa Carolina
Rubus flagellars
Rubus occidentalis
Plant Taxonomy
461
Table 1. — Continued
Taxa
Lookout
Wabash
Wea
Point
Breaks
Creek
X
X
X
X
X
X
X
X
X
X
X
X
X
Rudbeckia hirta
Ruellia humilis
Ruellia strepens
Sanguinaria canadensis
tt Saponaria officinalis
Scutelaria elliptica
Scutelaria parvula
tt Setaria sp.
Silphium integrifolium
Silphium terebinthinaceum
Smilax sp.
tt Solanum sp.
Solidago altissima
Solidago nemoralis
Solidago rigida
Solidago ulmifolia
Sorghastrum nutans
Specularia perfoliata
Sporobolus clandestinus
Sporobolus heterolepis
Sporobolus vaginiflorus
Taenidia integerrima
tt Taraxacum officinale
Tilia americana
Tradescantia ohiensis
Tradescantia virginiana
tt Tragopogon pratensis
* Trichostema dichotomum
tt Tridens flava
tt Trifolium repens
Ulmus americana
tt Verbascum thapsus
Veronica sp.
Verbena stricta
Vitis riparia
= on "Preliminary list of endangered and threatened vascular plants of Indiana" (2)
= Non-native Species
and west facing bluffs along major Illinois streams particularly along the Mississippi
River (9). The major difference between these prairies and the Indiana gravel hill prairies
is their soil, which is composed of loess, a windblown accumulation of silt with subor-
dinate clay and minor amounts of fine sand. However, the vegetation of these prairies
appears to be similar to the Indiana gravel hill prairies. Characteristic species of Illinois
loess hill prairies are Bouteloua curtipendula, Psoralea tenuiflora, Petalostemum can-
didum, Linum sulcatum and Lithospermum incisum. Common species included: An-
dropogon scoparius, A. geradri, Aster oblongifolius, Cassia fasciculata, Erigeron
strigosus, Euphorbia corollata, Eupatorium altissimum, Lespedesza capitata, Linum
sulcatum, Lithospermum incisum, Petalostemum purpureum, Psoralea tenuiflora, Ruellia
humilis and Sorghastrum nutans (9). All of the aforementioned species with the excep-
tion of Psoralea tenuiflora, Petalostemum candidum, and Linum sulcatum are found
on Indiana gravel hill prairies.
In Wisconsin, a similar type of hill prairie occurs on steep southwest facing slopes
on thin soil over limestock bedrock (7). A number of prairie species reported from
462 Indiana Academy of Science Vol. 94 (1985)
these areas also occur on the Indiana gravel hill prairies, including: Amorpha canescens,
Andropogon gerardi, A. scoparius, Aster oblongifolius, Bouteloua curtipendula, Com-
andra umbellata, Coreopsis palmata, Euphorbia corollata, Kuhnia eupatorioides,
Lithospermum incisum, Muhlenbergia cuspidata, Sporobulus sp., and Stipa spartea.
Curtis lists 47 prevalent species of dry prairie, 33% of which also occur on Indiana's
gravel hill prairies (7).
In Missouri, loess hill prairies, also known as "Till Slope Hill Prairies" (19) have
species such as Muhlenbergia cuspidata, Bouteloua curtipendula, and Andropogon
scoparius, and typical prairie forbs such as Lespedeza capitata, Lithospermum canescens,
L. incisum, and Monarda fistulosa.
Two types of hill prairie are reported from Iowa (16). One is a hill prairie on
steep southwest facing slope with limestone ledges and rubble present. Dominant grasses
are Bouteloua curtipendula, Sorghastrum nutans, and Andropogon scoparius. Forbs
include Aster oblongifolius, Anemone patens, Castilleja sessiliflora, Lithospermum in-
cisum, Coreopsis palmata, Liatris aspera, and Amorpha canescens. The other prairie
type is the loess hill type in extreme western Iowa that is apparently similar to the
Missouri loess hill prairie.
Summary
The gravel hill prairies in Indiana provide habitat for prairie species typical of
tall grass prairie such as big bluestem, Indian grass, tall coreopsis (Coreopsis tripteris),
bush clover, (Lespedeza capitata), switchgrass (Panicum virgatum), and flowering spurge
(Euphorbia corollata). More noteworthy though is the occurrence of a number of rare
species of plants at or very near the limit of their eastern range that are more typical
of prairie further west. Upon examining the general distribution of these plants one
discovers they are very rare in Indiana (2), and become common to abundant in Il-
linois (13, 17), Missouri (6) and farther west (3).
The Tippecanoe County gravel hill prairies and their associated rare flora repre-
sent a unique part of Indiana's natural heritage. One of the remaining gravel hill prairies
is a dedicated State Nature Preserve, one is owned by The Nature Conservancy and
negotiations are underway to protect the third prairie. Plans are also being drawn up
to monitor the rare plant species and determine how to best manage these areas. To
date, encroaching brush has been cut back and studies have started on several plant
species. Future plans call for prescribed burns to stimulate growth of prairie species
and to control woody brush encroachment.
Literature Cited
1. Aldrich, J. and J. Bacone. 1982. Limestone glades of Harrison County, Indiana.
Proc. Ind. Acad. Sci. 91:480-485.
2. Bacone, J. A. and C.L. Hedge. 1980. A preliminary list of endangered and
threatened vascular plants in Indiana. Proc. Ind. Acad. Sci. 89:359-371.
3. Barkley, T.M. editor. 1977. Atlas of the flora of the Great Plains. The Iowa
State University Press, Ames, Iowa. 600 p.
4. Bell, A. P. 1958. Soil Survey of Carroll County, Indiana. U.S.D.S. Soil Conser-
vation Service, Washington, D.C. 67 p.
5. Betz, R.F. 1978. The prairies of Indiana. Proceedings of the Fifth Midwest Prairie
Conference. Iowa State University Press, Ames, Iowa. 230 p.
6. Checklist of rare and endangered species of Missouri. 1984. Missouri Depart-
ment of Conservation. Jefferson City, Missouri. 16 p.
7. Curtis, J.T. 1959. The vegetation of Wisconsin. The University of Wisconsin Press,
Madison, Wisconsin. 657 p.
Plant Taxonomy 463
8. Deam, C.C. 1940. Flora of Indiana. Indiana Department of Conservation.
Indianapolis, Indiana. 1236 p.
9. Evers, R.A. 1955. Hill prairies of Illinois. 111. Lab. Nat. Hist. Bull. 26(5):367-446.
10. Fell, E.W. and G.B. Fell. 1956. The gravel hill prairies of Rock River Valley,
Illinois. 111. Acad. Sci. Trans. 49:47-62.
1 1 . Gleason, H.A. and A. Cronquist. 1963. Manual of vascular plants of northeastern
United States and Canada. D. Van Nostrand Company, New York, New York.
819 p.
12. Jackson, M.T. 1980. A classification of Indiana plant communities. Proc. Ind.
Acad. Sci. 89:159-172.
13. Mohlenbrock, R.H. and D.M. Ladd. 1978. Distribution of Illinois vascular plants.
Southern Illinois University Press, Carbondale, Illinois. 282 p.
14. Reeves, J.T., U.D. Zimmerman and J.E. Ebinger. 1978. Microclimatic and soil
differences between hill prairies in east central Illinois. 111. Acad. Sci. Trans.
71(20:156-164.
15. Robbins, J.M. Jr. 1978. Soil survey of Vermillion County, Indiana. U.S.D.A.
Soil Conservation Service, Washington, D.C. 124 p.
16. Schennum, W. 1984. personal communication.
17. Sheviak, C.J. 1981. Endangered and Threatened Plants, in M.L. Bowles, et al.,
editors, Endangered and threatened vertebrate animals and vascular plants of
Illinois. Illinois Department of Conservation, Springfield, Illinois. 214 p.
18. Sturm, R.H. 1966. Soil Survey of Fountain County, Indiana. U.S.D.A. Soil Con-
servation Service, Washington, D.C. 122 p.
19. Thorn, R.H. and J.W. Wilson, 1980. The natural divisions of Missouri. Missouri
Department of Conservation, Jefferson City, Missouri.
20. Ulrich, H.P. 1958. Soil survey of Tippecanoe County, Indiana. U.S.D.A. Soil
Conservation Service, Washington, D.C. 117 p.
21. Ulrich, H.P. 1967. Soil Survey of Parke County, Indiana. U.S.D.A. Soil Conser-
vation Service, Washington, D.C. 95 p.
22. United States Public Land Survey. Field notes north of the base line and west
of the second meridian, Indiana. Vol. 15. Indiana State Archives, Indiana State
Library, Indianapolis, Indiana.
23. White, J. 1978. Illinois natural areas inventory, technical report. Vol. 1. Univer-
sity of Illinois, Urbana-Champaign, Illinois. 436 p.
Vascular Plants of Barker Woods Nature Preserve, LaPorte County, Indiana
Victor Riemenschneider
Department of Biological Sciences
Indiana University at South Bend
South Bend, Indiana 46334
and
Patricia Wiese Reed
233 Hillcrest Road
Michigan City, Indiana 46360
Barker Woods Nature Preserve, the generous gift of Miss Margery Barker and
her mother to the Indiana Chapter of The Nature Conservancy in December 1974,
lies within the rapidly developing southeastern edge of Michigan City, Indiana (SW1/4,
NE1/4, Sec. 4, T 37 N, R 4 W, Michigan City West, U.S.G.S. 7.5' Quadrangle).
The 12 ha preserve is bounded on two sides by roads and housing developments. The
north boundary has some development at the northwest and the northeast corners while
the eastern boundary is undeveloped forest land similar to the preserve.
The preserve is located on the Calumet Lacustrine Plain about 4 km from Lake
Michigan. The plain formed during the latter part of the Pleistocene Epoch when
meltwaters of the retreating Lake Michigan lobe of the Wisconsinan ice sheet formed
a huge lake (Lake Chicago) behind the Valparaiso Moraine. Lake Chicago initially
stabilized at an elevation of 195 m above sea level (Glenwood Stage) which corresponds
to the elevation at the base the dune like ridge along the southern boundary of the
preserve. Later as the outlet channel eroded, the lake level decreased and finally stabilized
at 189 m above sea level approximately two km north of the preserve. The major
topographic features and surface materials were formed and deposited during this
interval.
A majority of the soils of the preserve are poorly drained except the well drained
Oakville fine sands on the dune ridge along the southern border and the moderately
well drained Brems fine sands of a lower parallel ridge in the northern third. Along
the northern edge is a strip of the poorly drained Newton loamy fine sands while the
large central lowland is a complex of the Saugatuck and Pipestone series. These two
soil types are unique to LaPorte County in Indiana. All of the soils are medium to
strongly acid and highly permeable in the upper layers except Saugatuck soils which
have an iron cemented layer within 60 cm of the surface (3).
The early land surveyors described the area as swampy along the western, southern
and eastern boundaries of section four and the northern boundary as mostly wet and
level. The northwest corner of the section was marked by a mound in prairie but all
other boundary locations were marked by witness trees that included beech, black ash,
elm, hickory, maple, pepperage, pine, popular and white oak. At the time of settle-
ment, the poorly drained soils of the preserve were seasonally ponded. During the
early years of settlement, the surface drainage systems were expanded through ditching
and a network of shallow ditches still exists in the preserve. Seasonal ponding has
rarely occurred during the last ten years.
The preserve area was purchased in 1833 and title transferred several times before
the land was purchased by Mr. Norton Barker in 1902. The records available recorded
little of the disturbance history except that some of the land was probably cleared
and farmed between 1866 and 1902. Shortly after they purchased the land, the Barkers
built a house and outbuildings on approximately two ha along the southern boundary
(now owned by the National Audubon Society). State Forester, Charles Deam, visited
465
466 Indiana Academy of Science Vol. 94 (1985)
the property on July 24, 1928 to determine the land's suitability for Classified Forest
designation. He described the forest as white, black and red oaks, sugar maple, hickory
and elm with some basswood, beech, aspen, sassafras and yellow birch. Based on his
recommendations the Barkers planted about two ha to spruce, white and red pine (Dr.
LaTourette Stockwell, personal communication). With the exception of one or two
fires, the forest has received little disturbance since the 1930s.
Methods
Data collection on the flora of the preserve was initiated in 1979 with periodic
visits to the preserve and the recording of species present in various locations. Later,
north-south compass traverses were made at approximately 30 pace intervals. Also,
a few east-west traverses were made at 50 pace intervals. As each new species was
encountered, the associated species were recorded as well as comments on its relative
abundance. Additional notes were made while laying out permanent plots and during
the mapping of the endangered and threatened species by Reed (6). Voucher specimens
have been collected for the Graminae and Cyperaceae and a few other species.
Results and Discussion
An alphabetical list of the species recorded for the preserve is presented in Table
1. The 161 species represent 110 genera and 46 families based on family classification
by Gleason and Cronquist (4). Twenty-three families are represented by only one species.
Eleven species are considered alien to Northern Indiana. The top five families include:
Cyperaceae with 15 species, Compositae with 12 species, Rosaceae with 11 species,
Ericaceae with 10 species and Graminae with 9 species. Members of the Ericaceae more
than any other family are responsible for giving a northern aspect to the shrub and
herb levels of the preserve.
Swink and Wilhelm (7) have assigned to each taxon in the Chicago region a
numerical value that "... expresses a taxon's relative autecological value with respect
to all other taxa in the flora." For native taxa, the values range from 0 for taxa that
are nearly ubiquitous under a broad set of synecological conditions to 10 for plants
that typify stable or near climax conditions and exhibit relatively high degrees of fidelity
to a narrow range of synecological conditions. Plants rare to the Chicago area were
given a value of 15 and plants threatened or endangered in the region were assigned
a value of 20. Introduced taxa were given ratings from - 3 to 2 with the lower rating
given those that detract from our landscape.
The rating column in Table 1 provides the Swink and Wilhelm value for each
of the taxa listed in their book. The average value for all taxa is 7.1 and 18% (27
taxa) have values of 15 or 20. A frequency plot of the number of species versus rating
values results in a bimodal distribution with peaks at values of 5 and 15. Along with
their rating system they present a formula for a natural area index. The index value
is obtained by multiplying the average taxon rating by the square root of number of
taxa. A rating of 50 or better is considered a high quality natural area. The index
value for Barker Woods is 90.8 which indicates the quality of the flora and the value
of the area as a nature preserve.
Eight of the species, Betula papyhfera, Carex arctata, C. folliculata, C. interior,
Epigaea repens, Melampyrum lineare, Pyrola rotundifolia var. americana and P. ellip-
tica, are either state threatened (ST) or state endangered (SE) (1). The first two species
are abundant in the preserve (Table 1). Epigaea repens and Pyrola elliptica are limited
to a few individuals or colonies (6). Carex interior exists as single clump in the central
lowland. Most of these populations are disjuncts with no other populations close enough
to maintain gene flow with species distribution centers.
Plant Taxonomy
467
Table 1 . Alphabetical list of vascular plants of Barker Woods Nature Preserve, LaPorte
County, Indiana including relative abundance and numerical rating for Chicago region.
Species
Rating
Abundance
7
A
2
c
5
U
8
c
4
u
6
u
5
R
8
A
15
R
5
U
10
R
5
C
4
C
10
U
6
u
-2
c
15
c
15 ST
c
2
c
15
u
15
u
15 SE
A
10
u
4
u
20
u
10
20 SE
R
4
C
10 SE
R
15
R
15
R
1
U
8
10
5
A
Id
C
5
R
8
R
20
R
6
R
0
C
15
R
10
C
5
U
1
u
2
u
5
c
5
R
10
R
5
R
15
U
6
c
15 ST
R
0
U
2
u
10
R
1
R
5
U
1. Acer rubrum L.
2. Agrimonia gryposepala Wallr.
3. Agrostis scabra Willd.
4. Amelanchier arborea (Michx. f.) Fern.
5. Amphicarpa bract eat a (L.) Fern.
6. Antennaria plantaginifolia (L.) Hook
7. Aquilegia canadensis L.
8. Aralia nudiculis L.
9. A. racemosa L.
10. Arisaema atrorubens (Ait.) Blume
1 1 . Asclepias exalt at a L.
12. Aster cordifolius L.
13. A. lateriflorus (L.) Britt.
14. A. macrophyllus L.
15. Athyrium filix-femina (L.) Roth
16. Berberis thunbergii DC.
17. Betula lutea Michx. f.
18. B. papyri/era Marsh
19. Boehmeria cylindrica (L.) Sw.
20. -Botrychium dissectum Spreng.
21. Brachyelytrum erectum (Schreb.) Beauv.
22. Carex arctata Boot.
23. C. crinita Lam.
24. C. cristate/la Britt.
25. C. digitalis Willd.
26. C. festucacea Schkuhr.
27. C. folliculata L.
28. C. hirtifolia Mackenz.
29. C. interior Bailey
30. C. intumescens Rudge.
31. C. laxiculmis Schwein.
32. C. laxiflora var. blanda (Dewey) Bott.
33. C. longii Mackenz.
34. C. mesochorea Mackenz.
35. C. pensylvanica Lam.
36. C. swanii (Fern.) Mackenz.
37. Carya ovata (Mill.) Koch
38. Chelone glabra L.
39. Chimaphila maculata (L.) Pursh.
40. Cicuta maculata L.
41. Circaea quadrisulcata (Maxim) French. & Sav. var.
canadensis (L.) Hara
42. Copt is groenlandica (Oedar) Fern.
43. Cornus florida L.
44. C. obliqua Raf.
45. C. racemosa Lam.
46. Corylus americana Walt.
47. Danthonia spicata (L.) Beauv.
48. Dentaria laciniata Muhl.
49. Desmodium nudiflorum (L.) DC.
50. D. paniculatum (L.) DC.
51. Dryopteria noveboracensis (L.) Gray
52. D. spinulosa (O.F.Muell) Watt
53. Epigaea repens L.
54. Euonymus alatus (Thunb.) Sieb.
55. Euphorbia corollata L.
56. Fagus grandifolia Ehrd.
57. Fragaria virginiana Duchesne
58. Fraxinus americana L.
468
Indiana Academy of Science
Vol. 94 (1985)
Table 1. — Continued
Species
Rating
Abundance
59.
60.
61.
62.
63.
64.
65.
66.
67.
68.
69.
70.
71.
72.
73.
74.
75.
76.
77.
78.
79.
80.
81.
82.
83.
84.
85.
86.
87.
88.
89.
90.
91.
92.
93.
94.
95.
96.
97.
99.
100.
101.
102.
103.
104.
105.
106.
107.
108.
109.
110.
111.
112.
113.
114.
115.
Galium aparine L.
G. pilosum Ait.
G. triflorum Michx.
Gaultheria procumbens L.
Gaylussacia baccata (Wang.) K. Koch
Geum canadense Jacq.
Glyceria striata (Lam.) Hitchc.
Goodyera pubescens (Willd.) R. Br.
Hamamelis virginiana L.
Helianthus divaricatus L.
Hieracium gronovii L.
H. scabrum Michx.
Hydrophyllum virginianum L.
Ilex verticillata (L.) Gray
Iris sp.
Lindera benzoin (L.) Blume
Liriodendron tulipifera L.
Lonicera japonica Thunb.
L. tatarica L.
Lycopodium lucidulum Michx.
Lycopus virginicus L.
Lysimachia cilia ta L.
L. terrestris L.
Maianthemum canadense Desf.
Medeola virginiana L.
Melampyrum lineare Desr.
var. pectinatum (Pennell) Fern.
Mitchella repens L.
Monotropa hypopithys L.
M. uni flora L.
Morus alba L.
Nyssa sylvatica Marsh.
Oenothera laciniata Hill.
Onoclea sensibilis L.
Osmorhiza claytoni (Michx.) C.B. Clarke
Osmunda cinnamomea L.
O. regalis L.
Oxalis europaea Jord.
Panicum dichotomum L.
P. lanuginosum Ell.
var. implicatum (Scribn.) Fern.
P. latifolium L.
Parthenoscissus quinquefolia (L.) Planch
Pedicularis canadensis L.
Penstemon digitalis Nutt.
Phytolacca americana L.
Pinus banksiana Lamb.
P. resinosa Ait.
P. strobus L.
P. sylvestris L.
Poa compressa L.
Podophyllum pelt a turn L.
Polygonatum pubescens (Willd.) Pursh.
Polygonum punctatum Ell.
Potentilla simplex Michx.
Prenanthes altissima L.
Prunella vulgaris L. var. lanceolata
(Bart.) Fern.
Prunus serotina Ehrh.
P. virginiana L.
1
10
5
10
9
0
4
20
8
5
6
7
5
9
7
10
-2
-1
15
6
4
8
15
10
15 ST
15
15
15
-1
3
6
8
0
20
3
7
2
10
4
2
20
20
0
5
7
6
4
10
1
Plant Taxonomy
469
Table 1. — Continued
Species
Rating
Abundance
5
C
15 ST
U
10 ST
R
7
C
4
A
8
R
8
A
7
A
6
C
0
U
-3
u
6
R
1
R
1
U
5
C
-3
U
3
u
9
A
0
R
1
U
6
A
5
R
2
C
15
A
-3
U
7
c
5
R
4
R
6
C
9
R
6
U
2
R
15
C
2
R
4
U
5
c
8
A
1
U
9
A
10
U
15
R
0
U
7
R
3
U
10
u
15
R
116.
Pteridium aquilinum (L.) Kuhn
117.
Pyrola rotundifolia L. var. americana
(Sweet) Fern.
118.
P. elliptica Nutt.
119.
Pyrus melanocarpa (Michx.) Willd
120.
Quercus alba L.
121.
Q. bicolor Willd.
122.
Q. palustris Muench
123.
Q. rubra L.
124.
Q. velutina Lam.
125.
Ranunculus abort ivus L.
126.
Rhamnus frangula L.
127.
Rhus copallina L. var. latifolia Engler
128.
R. glabra L.
129.
R. radicans L.
130.
Ribes cynosbati L.
131.
Robinia pseudo-acacia L.
132.
Rubus allegheniensis Porter
133.
Rubus hispidus L. var. obovalis
(Michx.) Fern.
134.
Rutnex obtusifolius L.
135.
Sambucus canadensis L.
136.
Sassafras albidum (Nutt.) Nees
137.
Scutellaria lateriflora L.
138.
Smilacina racemosa (L.) Desf.
139.
Smilax rotundifolia L.
140.
Solanum dulcamara L.
141.
Solidago caesia L.
142.
S. juncea Ait.
143.
S. nemoralis Ait
144.
S. rugosa Ait.
145.
Spiraea tomentosa L. var. rosea (Raf.)
Fern.
146.
Symplocarpus foetidus (L.) Nutt.
147.
Tradescantia ohiensis Raf.
148.
Trientalis borealis Raf.
149.
Triodia flava (L.) Smyth
150.
Ulmus rubra Muhl.
151.
Vaccinium angustifolium Ait.
var. laevifolium House.
152.
V. corymbosum L.
153.
Verbascum thapsus L.
154.
Viburnum acerifolium L.
155.
V. recognitum Fern.
156.
Viola pallens (Banks) Brainerd
157.
V. papilionacea Pursh.
158.
V. sagittata Ait.
159.
V. sororia Willd.
160.
Vitus aestivalis Michx.
161.
Woodwardia virginica (L.) Sm.
1 . Species nomenclature is based on Fernald (2). Rating is based on values given by Swink and Wilhelm (7). A dash
indicates no value given for that species. The state endangered and threatened species are identified by SE and ST
adjacent to rating value. Abundance values are the subjective estimates of the two authors. Letters indicate relative
abundance as follows: A — species abundant throughout the preserve. C — species is common in most of the preserve.
U — species is limited in its distribution and/or abundance. R — species is found in only one small area of preserve
or restricted to a few individuals.
470 Indiana Academy of Science Vol. 94 (1985)
One curious note is the relatively recent appearance of Chimaphila maculata in
the flora. During many visits to the preserve, we normally travel a loop path that
probably served as fire access road during the 1967 fire. On the north loop of this
trail there presently exists five plants of this species which was first recorded in 1983
as two plants present. Since this species is considered rare in the Chicago area (7),
the possible migration to and establishment in a new area gives hope for its continued
survival in the regional flora.
A majority of the alien species are limited to disturbed areas on the perimeter
of the preserve. However, three species (Lonicera japonica, L. tatarica and Euonymus
alatus) appear to be increasing numbers and distribution within the preserve and should
be monitored to determine their impact on native taxa.
Conclusion
Barker Woods has proven to be an excellent addition to the State Nature Preserve
System and exemplifies how much present and future Hoosier generations owe a debt
of gratitude to those who protected remnants of our natural heritage.
The preserve protects eight state threatened and endangered species and many
other species considered rate in the Chicago Region. In addition, one of the soil mapping
units is uncommon in the state and much of the mapping unit is threatened by
urbanization. The unique combination of plants, soils and lake affected climate pro-
vides an excellent opportunity for gaining a better understanding of our Indiana flora.
We thank Helen Stare and Jim Aldridge for their assistance in identifying the
sedges. A special note of thanks to the Barker Woods Preserve Management Commit-
tee and The Nature Conservancy for encouraging scientific study of the preserve and
to the Barker family for preserving the area.
1. Bacone, J. A. and C.L. Hedge. 1980. A preliminary list of endangered and
threatened vascular plants in Indiana. Proc. Ind. Acad. Sci. 89:359-371.
2. Fernald, M.L. 1950. Gray's manual of botany, 8th ed. Amer. Book Co., New
York. 1632 p.
3. Furr, G.F. 1982. Soil survey of LaPorte County, Indiana. U.S. Department of
Agriculture, Washington. 162 p. + maps.
4. Gleason, H.A. and A. Cronquist. 1963. Manual of vascular plants of northeastern
United States and adjacent Canada. Van Nostrand, New York, 810 p.
5. Hill, J.R., D.D. Carr, E.J. Hartke and C.B Axelroad. Geology as a contribution
to land use planning in LaPorte County, Indiana. Indiana Geol. Surv. Spec. Rept.
14, Bloomington. 28 p.
6. Reed, P.W. 1985. Population studies of threatened and endangered plants of Barker
Woods Nature Preserve, LaPorte County, Indiana. Proc. Ind. Acad. Sci. 94:
(in press).
7. Swink, F. and G. Wilhelm. 1979. Plants of the Chicago region. Morton Arboretum,
Lisle, IL. 922 p.
PSYCHOLOGY
Chairperson: Robert Fischer
Department of Psychological Science
Ball State University
Muncie, Indiana 47306
(317) 285-1713
ABSTRACTS
Marking in Submissive Male Gerbils after Contact with a Dominant Male and His
Odors. A.M. Fullenkamp, Kim Duffy, Robert A. Vance and Robert Fischer, Depart-
ment of Psychological Science, Ball State University, Muncie, Indiana 47306. The
marking behavior of dominant and submissive male Mongolian gerbils (Meriones
unguiculatus) was observed in two experiments within an open field. In the first
experiment, the 60 x 80 cm field was divided in half by a wooden board, each half
contained an elevated square. After a half hour of separation, the board was removed,
and marking behavior and time spent in the area were recorded. The next day the
male judged to be dominant was again placed in the divided field and allowed to mark
the area for a half hour. When the male and the board had been removed, the sub-
missive was given free run of the open field. There were no significant differences
in the submissive male's tendency to mark or spend time in either area.
In the second experiment, the submissive male was presented with the same open
field, this time studded with four squares. One square was clean, one was marked
by a novel submissive, one was marked by the familiar dominant male and one had
been marked by the subject. Others have hypothesized that the submissive would avoid
or be intimidated by the odors of the dominant male. Our results indicate that the
submissive marked the clean square the most, but also tended to mark the dominant
square more than his own.
Heterosexual Social Interactions in the Syrian Hamster. Bonnie Gray, Robert Fischer
and Gary Meunier, Ball State University, Muncie, Indiana 47306. The female
hamster modifies her social interactions with males as a function of her estrous cycle.
In order to describe these changes, twenty-four naturally cycling females were observed
interacting with restrained male pairs in a Y-shaped choice apparatus on each of the
four cycle days. Males differed in terms of dominance status. Variables measured in-
cluded the number of female approaches, time spent in proximity, sniffing, and vaginal
marking. Behavioral changes which varied significantly over the four day cycle were
the amount of marking and the number of sniffs exhibited by the females. Marking
increased throughout the estrous cycle, reaching a peak on the day prior to estrous.
No effect for male status was found. Sniffing varied both as a function of cycle state
and male status with the dominant male receiving the greater number of sniffs. No significant
effects were found for approaches or for time spent in proximity. It is suggested that
female marking probably subserves a general advertisment function, while sniffing is
more discriminating and could reflect female proceptivity. Such olfactory investigation
likely influences the probability of the occurence of other behaviors and would form
the basis of female choice.
The Several Themes of Adolescence. Barbara Kane, Indiana State University, Terre
Haute, Indiana 47809. This author finds several themes manifested in adolescent
471
472 Indiana Academy of Science Vol. 94 (1985)
life: autonomy, control self-absorption, intensity, definition, extremism, and sexuality.
A theme is a central idea, an underlying issue, or a repetitive pattern. Other authors,
notably Bios, Elkind, Erikson, A. Freud, Hall, Marcia, and Sullivan, have addressed
the topic of adolescence, but none has seen this number or variety of specific major
concerns.
Autonomy is the adolescent's drive to separate and counterdrive to cling. Con-
trol includes relinquishing control, resisting control, achieving control, and pro-social
control. Self-absorption is the egocentrism and narcissism of adolescence. Intensity
refers to adolescents' urgency and passion, and their need for immediacy. Definition
is the adolescent's striving to replace confusion and ambivalence with identity and
commitment. Extremism refers to intrapersonal and interpersonal polarities that are
seen in adolescents' needs, interests, values, and activities. Sexuality includes the lust
dynamism and the need for intimacy.
At no other time in the life span are the themes more prominently displayed.
They become significant in the lives of pre-adolescents and become increasingly important
with the youngsters' emergent adolescence. They begin to fade in late adolescence,
and in adulthood they become integrated into the personality and lifestyle. Their moment
is over; their impact on adolescence, however, influences adulthood and beyond.
Psychovector Love Scale and its Differentiability. Oliver C.S. Tzeng and Roberta
Schliessmann, Department of Psychology, Indiana University-Purdue University at
Indianapolis, Indianapolis, Indiana 46223. —The concept of love has been the cen-
tral issue in human communications and interpersonal dating behaviors. However, as
to its nature and determinants, our society still has little insight beyond what have
been written by poets and novelists. Although many attempts have been made to pur-
sue scientific knowledge of love in the research community, there still exists numerous
questions unanswered, and controversies unresolved.
This study conducts a comprehensive evaluation of the issues in the theories and
measurements of love relationships. As a result, a process-oriented measurement scale,
called Psychovector Love Scale, was developed. Based on the combinations of three
behavioral signals (retreat — approach — attack) and four primordial dispositional pro-
cesses (mobilization, unification, affinity and variety), twelve basic emotions were
constructed to measure diverse situational encounters between two people. Empirical
data were collected from over 600 adults at five different levels of interrelationships.
Inter-group comparisons resulted in significant patterns of interactive dynamics and
prospects of future behaviors. The utilities of this development were discussed in the
realm of various counseling and psychotherapeutic purposes.
Orwell's 1984, Skinner's Walden III, Marx' Classless Society and other Utopias: An Ex-
ploration of Human Expectation and the Psychological Factors in a "Perfect Society."
John M. Vayhinger, 1235 Favorite Street, Anderson, Indiana 46013. The
infantile expectation in the early infant of control over one's environment and
the hope a a human Eden or heaven on earth or a perfect political society is
wide spread among socieites and writers. With the growing possibility of a control
of behavioral environment (family structure, educational in-put, media and literature
manipulation), of chemical control of emotions, and the political control of groups,
cultures are experimenting with various forms of "Utopias," and control over individual's
lives from 'womb to tomb.'
From Walden II to Beyond Freedom and Dignity, B.F. Skinner proposed schemes
to implement a society without hunger, oppression, poverty, competition, frustration
or uneven distribution of the products of civilization and community.
Psychology 473
In the Marxist-Leninist countries, classical conditioning has been applied through
Pavlovian Institutes, political and economic control with various effectiveness in making
"The New Soviet Man" and the "New Chinese Man."
In literature Orwell warned (or predicted) a totalitarian society for England where
Big Brother Is Watching You through telescreens, where 'war is peace,' 'freedom is
slavery,' 'ignorance is strength,' and where Newspeak has eliminated the possibility
of even thinking independently and individual action, especially what is known in the
scientific community as accuracy in research.
This presentation will explore the use of classical and operant conditioning in
these three Utopias, (1) positive fictional (Walden II), (2) politically repressive (Marxist-
Leninist theory in Russia and China), and (3) negative fictional ("1984").
Personality Types and Perceptual-motor Performance. Roger Ware and Charles
Yokomoto, School of Science and Engineering, Indiana University-Purdue University
at Indianapolis, Indianapolis, Indiana 46223. Voluntary student participants were
administered the Myers-Briggs Type Indicator to assess their personality type, and subse-
quently performed the Mirror Tracing Task over three consecutive trials. Previous results
using only one trial were not supported, but new findings emerged. Individuals with
a sensing (S) preference have significantly longer overall performance time than in-
dividuals with intuitive (N) preference. Individuals with a thinking preference (T) have
shorter performance times on the first trial but slightly longer performance times on
the second and third trials than individuals with a feeling preference (F). Similar results
occurred with errors and error time. Previous research suggesting that individuals with
an NT preference are somewhat brighter and quicker is supported, but the present
results also suggest that individuals with SF preferences appear to catch up over time,
at least on learning a new perceptual-motor task.
Munro's Doctrines: A Forgotten Pioneer in Holism and Hypnosis
Walter Hartmann
Department of Behavioral Sciences
Purdue University Calumet
Hammond, Indiana 46323
One of the characteristics of this post-Freudian age is the fast developing holistic
approach to the person. Side by side with the explosive growth in microbiology,
neurology, and all the "hard" sciences, goes the growing recognition that the human
being is a totally reacting organism. "Mind" does not control body, or body "mind" —
they function as a unit.
Among the phenomena illustrating the development of holism are psychologists
being appointed to teach medical students (e.g., 33); a prestigious medical journal
publishing a layman's self-cure, largely by positive emotions (3), and this layman subse-
quently becoming a member of a medical faculty; and a Nobel Prize winning im-
munologist listing hypnosis as one established, indirect immunodepressor (12).
Hypnosis, the most obvious area where psyche and soma, mind and body, meet
and interact, is also in the midst of a research explosion and of accepted and respected
practice. One major trend here is also toward demythologizing, generalization, and
synthesis. The conceptual baggage — like trance, surreal capabilities, addressing the
subconscious — is being lightened.
Of course, this new direction is not the only one; argument is carried on. However,
statements like the following are today common.
The far reaching influence of suggestion or the personal influence of the physi-
cian . . . (is) not even faintly appreciated by the profession . . .
Physiology, psychology, and biology are on friendly terms.
Education is another form of suggestion
We now realize that . . . both mind and body constitute a manifestation of the
real self in action.
I use the terms "hypnotism" and "suggestion" as synonymous terms.
Our therapeutic measures must be in accordance with an individual's preconceiv-
ed beliefs.
But these are not contemporary quotations. They are culled, almost randomly,
from A Handbook of Suggestive Therapeutics, Applied Hypnotism, Psychic Science
by Henry S. Munro, M.D., second edition— 1908 (14, pp. 5, 6, 11, 14, 19, 31).
Seemingly a historical curiosity, the book does contain Edwardian prose,
philosophical speculation, purely anecdotal evidence. To a contemporary scientist, it
is in many ways naive.
However, with and underlying all this, Munro presents startlingly modern and
challenging ideas about hypnosis and suggestion, and propounds sophisticated holistic
approaches generally.
Yet, Munro is unknown. The Handbook is out of print. Altogether only four
references to Munro were found (4, 8, 9, 32). No mention of him occurred in any
standard treatise. No worker in the field was found who knew about Munro.
Henry S. Munro is a forgotten pioneer.
475
476 Indiana Academy of Science Vol. 94 (1985)
The Handbook indicates little more than that he was a downstate Georgia physi-
cian at the turn of this century. Lengthy investigation and the cooperation of many
resulted in considerable information on Munro's work and life. An outline follows.
Munro's Handbook . . . went through four editions (13, 14, 15, 16). The first
and second editions, 1907 and 1908 (13, 14), are identical; the third and fourth, 1911
and 1917 (15, 16), revised and enlarged, differ in that the fourth edition has two addi-
tional chapters.
The two later editions differ interestingly from the earlier two. The later ones
are much more like modern publications, in style, make-up, index and references.
The third and fourth editions have a chapter on psychoanalysis. Munro treats
Freud with respect, but critically. Among other matters, Munro discusses Freudians'
rejection of hypnosis, quoting Freud himself (6) as not really negative toward it.
Ten papers by Munro were found (17 through 26), one published twice, in dif-
ferent journals (26). Two further papers are referred to, but could not be traced. In
addition, there are four pamphlets Munro issued (27, 28, 29, 30); here too he ag-
gressively propagates his teachings, but also sells and advertises in questionable taste.
The following themes are prominent in Munro's work.
Munro teaches transcendence of the mind-body dichotomy. He stresses and deals
with attitudes and expectations as involved in the etiology of health and disease. He
recognizes the social environment. He insists that the physiological machinery is basic,
but is affected by "psychotherapy" or "suggestive therapeutics."
Munro defines these as "mental influences in the treatment of disease . . . with
the definite understanding that any influence . . . exerted in any way by the personali-
ty of the physician . . . come(s) under the broad domain of Suggestive Therapeutics"
(14, p. 16); or "Suggestive Therapeutics is the sum total of the influence exerted by
the physician ... to help toward recovery, or for relief of mental or physical symp-
toms. This is always accomplished through the normal physiological processes" (14,
p. 19).
Munro teaches that the organism has as a whole and in each of its parts and
systems — cell, nerve, muscle, gland, immune machinery, etc. — a capacity to organize
itself for growth and well-being. This capacity depends greatly on attitudes, expecta-
tions, and conscious and unconscious goals. These in turn depend on the environment,
social and personal.
Attitudes and expectations, then, largely determine mental and physical health and
hygiene; and they are in very large measure a function of education, widely understood —
church, body politic, family interactions, among others.
Regarding hypnosis, Munro writes, "Hypnotism is the art of persuading an in-
dividual to act upon or execute an idea or series of ideas, either consciously or sub-
consciously. The condition is brought about by suggestion" (14, p. 11); and "hyp-
notism (is) the induction of a mental and physical condition in which the subject is
more amenable or susceptible to suggestion" (14, p. 18).
Always stressing the inter-relatedness of mind and body, Munro deals with hyp-
nosis and suggestion in connection with childbirth, anesthesia and pain control generally,
placebos, sex, religion, personality formation, education, physical and mental hygiene,
among others.
With and underlying all the physical means directed toward the organism's
machinery, the healer, then, has at his disposal the powerful and widely applicable
tools of "suggestive therapeutics" or "psychotherapy" — of suggestion.
Munro writes of societal factors in the development of attitudes hindering or
furthering mental and physical health and hygiene in an almost Marxist manner:
"... environment has been the actual creator of man"; "capital(ists) . . . disorganize
. . . equilibrium (to) better control the ignorant laboring classes. . . " (26).
Psychology 477
Munro explicitly addresses himself to the ordinary general health provider. In-
deed, he censures authorities like Janet, Binet, Dubois, Prince, Putnam, "and many
others," because they "apparently would limit the field of psychotherapy to neurology
and psychiatry when it is equally applicable ... in all classes of professional work"
(15, p. 28). Munro contrasts the restrictedness of Freud's theory and method, quoting
Freud (6), with the wide applicability of Munro's teachings.
The healer always affects patients through suggestion by his personality and de-
meanor. He should learn about this tool and use it deliberately and efficiently. Munro
stresses that psychotherapy (as he uses the term) is one, albeit powerful, tool, not
a system of therapy.
Munro writes in so many words that he hopes to get the readers to act on his
ideas (14, p. 12), i.e., by his definition, to hypnotize them. He tries to influence his
audience also by suggesting — again, in so many words — that the money going into
charlatans' pockets could and would go into theirs, if they followed his advice (16,
p. 162).
Munro writes he had read the old writers on hypnotism. It seems obvious, already
from his book's title, that he was familiar with Bernheim (e.g., 2). Munro at times
still writes of hypnosis as a kind of sleep, following Bernheim (2) and others; but
Munro uses this quite perfunctorily. And Munro stresses even more strongly than Bern-
heim that hypnosis is itself also due to suggestion. He insists it is a function of the
subject, not of the hypnotist. Munro displays a strongly patient-centered attitude, though
his technique is fast and brisk.
Munro clearly was a highly skilled hypnotist; according to one report, he once
convinced a skeptical audience by taking dozens of people randomly off the street
and hypnotizing all of them in short order. An eye witness reports seeing Dr. Munro
hypnotizing a male subject rapidly and fully, in 1918; throughout, Munro used nothing
but a firm, soothing voice, some arm and hand motions, and eye contact.
Apparently Munro acquired his skills from entertainers; an old photograph has
been described (by people who saw it) of Munro with a comfortable and unconcerned
looking youngster with many pins stuck into him. However, while reporting that he
had used hypnosis for entertainment, Munro repeatedly and strongly declares such
use of hypnosis absolutely unacceptable.
Originally, Munro's technique was in line with Kroger's statement that the technique
rests to quite an extent on misdirection (10, p. 7); but Munro wrote later (15, p. 72),
"... the methods which are here described are not those I am presently employing
in my practice, for in no case now do I use the least bit of deception."
Munro anticipated modern thoughts and findings in many ways. For example,
Glass and Barber (7) report that a placebo can be as effective as traditional trance
induction — Munro uses placebos routinely in rapid procedures (14, pp. 23, 217, and
passim); Barber and Calverley (1) find that the operator's tone of voice can affect
suggestibility — Munro suggests similarly (14, pp. 161, 257).
Kroger writes:
. . . hypnotic responses . . . are due to subjective mechanisms inherently present
in all individuals. ... It is indeed a wise hypnotist who knows who is hypnotizing
whom! (10, p. 8);
and
. . . the patient actually induces the hypnosis through his own convictions (10,
p. 31).
Over fifty years before this, Munro wrote:
How frequent it is that the operator becomes hypnotized instead of the subject,
478
Indiana Academy of Science
Vol. 94 (1985)
thinking that it was some power he was exerting over the hypnotized individual,
rather than the use of an inherent quality . . . within the individual himself (14,
p. 38).
Also, he wrote of the necessity to "secure the accord of the patient" (14, p. 131).
Further, "Hypnotism is a self-induced psychological condition. You do not hypnotize
an individual — you simply get him to do it himself" (15, p. 162).
■"■~';.v-.-;.;V''::- ".;'•;_,'
Henry Sumner Munro
Psychology 479
Henry Sumner Munro was born April 19, 1869, on his family's 3000 acre planta-
tion at Putnam, Georgia, a small place seven miles from both Buena Vista and Ellaville,
small communities not far from Americus, Georgia. Putnam served this plantation,
Springdale Farm, and a similar, neighboring one, Peachtree, belonging to the family
of Munro's mother. Today Putnam is just a name. Munro was the fifth of six surviv-
ing children.
Munro's family originated in Scotland, France, and England. They were unconven-
tional, moving about a great deal; usually they exhibited intellectual aspirations and
capacities. Among Munro's many interesting ancestors were Tories and his great-
grandfather, Dr. Joseph deLespine, French army surgeon, who came here in 1778 with
a French fleet supporting the American Revolution.
Munro's father, George Washington Coe Munro, self-educated, explosive, was
a progressive farmer, a kind of pioneer of the New South; his mother, of the promi-
nent Stevens family, a gentle but powerful woman.
Munro began his education in a one-room schoolhouse built on the plantation,
where his eldest sister taught her siblings, other relatives, and neighborhood children.
This school soon moved to larger church premises.
Munro attended what was then Emory College, 1885-7, where he successfully
engaged in public debating. He wrote later that he did not finish for lack of funds;
that back home he "studied medicine" by poring over the Pharmacoepedia in the
Ellaville drugstore.
Still, in 1891 Munro graduated from the University of Maryland School of Medicine
and College of Physicians and Surgeons. In that same year, he married the daughter
of a prominent local physician. He practiced with him for a while, then in his own
practice in Ellaville.
Early in 1895, Munro undertook three months of post-graduate study in New York.
At some times, Munro practiced in Americus, probably around 1896 and around
1906. Mostly his practice was in Ellaville 1891-1909, where by 1907 he had had quite
an imposing home and office built, apparently with help from his father-in-law. Known
as the Gingerbread House, it stands today, with a caduceus prominently chiseled over
the entrance.
Munro became "impressed with . . . the psychic factor in therapeutics" about
1892 (14, p. 5). In 1899, he started lecturing to physicians' groups all over the country
(ibid.). For example, in 1907 Munro lectured in Rochester, Minnesota, where he met
the Mayos' anesthetists; probably in Fargo, North Dakota; in St. Louis, Missouri,
to "75-100 leading physicians"; in Chicago. His Handbook . . . (13, 14, 15, 16) is
an outcome of these lectures.
Munro clearly was away from Ellaville a great deal. Even so, his practice became
known as a "hypnosis clinic," where he treated quite a number of patients from far
away, apparently referred by physicians who had heard him. He called his Ellaville
practice "Poplar Grove Sanatorium," himself its Superintendent (see 18). The telephone
number is said to have been "1." Poplars (actually, sycamores) still stand around
that house.
In April of 1909, Munro and his family, including now six children, moved to
Omaha, Nebraska. Munro leaving the family, Mrs. Munro and the children, aided
by her family, soon returned. Divorce proceedings, instigated by Munro and decided
firmly against him, followed in 1915.
From 1909 to 1916, Munro lived and practiced in Omaha; he had offices in the
Brandeis Theater Building.
By 1916, Munro's specialty is listed as psychiatry in the American Medical Dic-
tionary. Among other things, he served as medico-legal expert; he appeared in a number
of cases for elderly persons whose families were trying to have them declared incompe-
480 Indiana Academy of Science Vol. 94 (1985)
tent. He used hypnotic techniques to demonstrate their competence — at least, physical —
quite dramatically in court. He reports receiving $500 for one such case.
In 1917, Munro married his second wife. In 1918, the Munros moved to Lincoln,
Nebraska.
For the birth of the first daughter of this second marriage, 1918, Dr. and Mrs.
Munro returned to Putnam, where his eldest sister now ran what was left of the
plantation. Munro attended his wife's confinement; in line with his — and present —
teachings, Mrs. Munro was up and about three days after giving birth, to the surprise,
even scandal, of local opinion.
From 1919 to 1942, Munro lived in Portland, Oregon. In 1921, his last child
was born. In 1923, he worked on a new, fifth, edition of his Handbook; he wrote
later that, feeling provoked and disadvantaged by the publisher, he aborted this effort.
Though listed many years as a physician in the City Directory, Munro did not
take out medical license in Oregon, feeling at odds with the medical establishment
by this time. He continued lecturing, probably until about 1927. He produced literature
to educate the public to his ideas of health and hygiene (e.g., 30). Around 1935 he
produced a tonic in his home, the "Eutrophic Research Laboratory." Indeed, in 1932
he was convicted of practicing medicine without a license; however, neither the judge
in the case nor Munro seem to have taken the conviction very seriously. At any
rate, Munro carried on as before.
The depression his the Munros very hard. He lost his home. In 1936, he was
working in a quarry, almost certainly on a WPA project. Relations within the family
were very difficult.
In 1942, Munro moved by himself to Hood River, Oregon; received welfare. Old
age assistance began in 1945. By 1945, Munro resided in a Hood River hotel. In 1953,
divorce occurred again.
On March 1, 1958, Henry Sumner Munro, M.D., died, aged almost 89, in a Hood
River nursing home.
Obviously, Munro's life was characterized by much strain and difficulty, due con-
siderably to his own unconventional and awkward personality. Even so, into advanced
age Munro was vitally interested, physically and mentally vigorous, and proud of it,
and angry.
Munro's teachings about holistic health care and about hypnosis and suggestion
were controversial, but for a considerable time not out of the main stream.
Munro received a hearing. His wide lecturing provided a reasonable income. His
Handbook went through four editions. It is probable, though unconfirmed, that the
Handbook was used at Tulane University. Munro had the approval of many colleagues,
quite a few prominent. For example, in 1914 he presented a paper (25) to the national
meeting of Alienists and Neurologists, apparently at the request of the president of
the Chicago Medical Society; Munro was appointed to a committee of five to draft
recommendations, endorsed unanimously, for the prevention of insanity in the United
States. Munro was not alone; see for example Magaw (11), one of the Mayos'
anesthetists, whom Munro quotes — she quite unconcernedly stresses suggestion in
anesthesia.
At the same time, Munro was fighting opposition and disapproval. He attacks
medical education, over-specialization, professional narrow-mindedness. After about
1919, Munro wrote later, the medical school — presumably in Omaha — tried to quell him.
Munro's holism became less and less part of the mainstream. The knowledge
of the machinery of the body and of technology improved; the healer was seen, and
saw himself, largely as mechanic. Hypnosis and suggestion, too, was handled mainly
in the laboratory and in elaborate theory building, if it was dealt with at all by respec-
table scientists.
Psychology 481
At the same time, Munro's writings and his life indicate he was difficult, offen-
sive, often quite shrill, perhaps a bit paranoid. Observers of the family agree the Munro
men were intellectual, but "mean," "lacked common sense."
At any rate, Munro became more and more isolated, disappointed and disgruntled.
However, this development also reflects what went on generally, as becomes clear,
for example, fromOberndorf(31)andHale(9). Indeed, it seems a reflection in microcosm
of the tensions and conflicts pervading the field (see, e.g., 5); and, of course, they
still go on (e.g., 34). They may be adumbrated by the tension between generalist and
specialist, synthesis and analysis, mental and physical, holism and reductionism.
Here, then, is a physician outside the prestige and power centers of medicine
and psychology. Whatever his weaknesses, he used an original and powerful mind,
brought a messianic fervor to bear, exhibited courage and strength.
He began over eighty years ago to bring one approach to the person to the general
worker in health and hygiene, and continued this for over twenty years, achieving
and propounding important insights.
In the end, the mainstream left his current, and he was forgotten. Recently, the
field he cultivated strenuously has been worked again.
So, Dr. Henry S. Munro, pioneer in holism and in hypnosis and suggestion, can
be appreciated again.
Acknowledgments
This presentation is the result of astonishingly generous and effective coopera-
tion and support by colleagues and librarians at Purdue; by workers at many institu-
tions; by acquaintances of, and especially by members of the families of Dr. Munro.
This paper is dedicated to one of them: the late F.X. Dever, M.D., of New York,
who had himself begun an investigation of this subject, and who made all his materials,
thoughts, and counsel available.
The project was supported also by a semester's sabbatical leave and by Depart-
mental funds at Purdue University Calumet.
Literature Cited
1. Barber, T.X., and D.S. Calverly, 1964. Effect of E's tone of voice on "hypnotic-
like" suggestibility. Psychol. Reports. 15:139-144.
2. Bernheim, H. 1957. Suggestive therapeutics. Associated Booksellers, Westport,
Conn. 420 pp. (Originally published 1888).
3. Cousins, N. 1976. Anatomy of an illness (as perceived by the patient). New England
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4. Elman, D. 1968. Hypnotherapy. Westwood, Los Angeles, Calif. 336 pp. (Originally:
1964, Findings in hypnosis).
5. Esper, E.A. 1964. A history of psychology. Saunders, Philadelphia, Pa. 368 pp.
6. Freud, S. 1909. Selected papers on hysteria and the psychoneuroses. In Monographs
on nervous and mental disease, published by Jeliffe, S.E., and W.A. White (eds.),
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7. Glass, L.B. and T.X. Barber, 1961. A note on hypnotic behavior, the definition
of the situation and the placebo effect. J. of nervous and mental disease.
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8. Grinstein, A. 1958. Index of psychoanalytic writings. Internat'l University Press,
New York, N.Y. 6 volumes.
9. Hale, N.G., Jr. 1971. Freud and the Americans. Oxford University Press, New
York, N.Y. 574 pp.
482 Indiana Academy of Science Vol. 94 (1985)
10. Kroger, W.S. 1963. Clinical and experimental hypnosis. Lippincott, Philadelphia,
Pa. 361 pp.
11. Magaw, A. 1906. A review of over fourteen thousand surgical anasthesias. J.
of surgery, gynecology, obstetrics. Dec.:795-799.
12. Medawar, P.B. 1968. Genetics and the future of man — Recent advances in the
immunology of transplantation. Graham Memorial Lectures, Washington Univer-
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13. Munro, H.S. 1907. A Handbook of suggestive therapeutics, applied hypnotism,
psychic science. C.V. Mosby, St. Louis, Mo. 353 pp.
14. 1908 2nd edition, 359 pp.
15. 1911. 3rd edition, revised and enlarged. 409 pp.
16. 1917. 4th edition, revised and enlarged. 481 pp.
17. 1903. Psychic force, a therapeutic power. Medical Index — Lancet, Kansas
City, Mo. (Read before Jackson County Med. Soc, Kansas City, Mo., 9/10/ '03).
18. 1908. The influence of suggestion as an adjunct in the administration of
anesthetics. St. Louis Med. Review. 57:383-391.
19. 1909. The general practician in the realm of psychotherapy. Amer. J. of
Clin. Medicine, Chicago, 111. 16:747-752.
20. 1909. Psychotherapy in relation to the expectant mother. Annals of
Gynecology and Pediatry, Boston, Mass. 22:1-6.
21. 1909. Personality — its significance in therapeutics. Annals of Gynecology
and Pediatry, Boston, Mass. 22:65-74.
22. 1910. Psychotherapy in relation to the general practice of medicine and
surgery. Medical Herald, St. Joseph, Mo. 24:271-286.
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The Amer. Practitioner, New York, N.Y. 47:578-591, 619-633.
24. 1914. Instinct, intellect, and the game. J. of the Amer. Institute of Homeopathy,
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of Alienists and Neurologists of the U.S., Chicago, 111., 1914). Also 1918 in Western
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29. 1917. An open question and answer. Pamphlet.
30. 1937. Why not wake up and live? Sales pamphlet.
31. Oberndorf, C.P. 1964. A history of psychoanalysis in America. Harper Torch-
books, New York, N.Y. 280 pp. (Originally published by Grune and Stratton, 1953).
32. Stanton, H.E. 1979. Short-term treatment of enuresis. Amer. J. of Clinical Hyp-
nosis. 22:103-107.
33. Thompson, R.J., Jr., and J.D. Matarazzo. 1984. Psychology in US medical schools:
1983. American Psychologist. 39:988-995.
34. Wertheimer, M. 1972. Fundamental issues in psychology. Holt-Rinehart-Winston,
New York, N.Y. 278 pp.
SCIENCE EDUCATION
Chairperson: Linda Hamrick
The Canterbury School
5601 Covington Road
Fort Wayne, Indiana 46804
(219)432-7776
Chairperson-Elect: Gary Dolph
Indiana University at Kokomo
Kokomo, Indiana 46902
(317)453-2000
ABSTRACTS
Ideas Concerning the Use of Computer Data Acquisition Systems to Improve Teaching
Effectiveness within the Laboratory. Marshall P. Cady, Jr., Department of Natural
Sciences, Indiana University Southeast, New Albany, Indiana 47150. The tradi-
tional molecular weight determination by freezing point experiment has been modified
into an extremely short experiment and coupled with an HP-85 computer/HP-3438A
digital multimeter data acquisition system. Educational benefits that result from this
condensation of time and technology are discussed in this paper along with some aspects
of laboratory philosophy. The benefits may be applicable in many different lab situa-
tions and to many different disciplines. They include (a) freeing the student from the
time consuming tasks which do not develop skills, (b) clearly emphasizing skills which
are to be developed, (c) forcing the student to focus on interpretation of data, and
(d) making it possible to encourage students to plan the redesign of an experiment.
A New and Challenging Science Program from AAAS for Grades 7 and 8. Walter
Cory, Coordinator for School Sciences, Indiana University, Bloomington, Indiana
47405. A new program, Science Resources for Schools, is being distributed to
schools in nine states and the District of Columbia by the American Association for
the Advancement of Science. The development of the materials is supported by a grant
of 3 million dollars from the Standard Oil Company of Ohio (SOHIO).
Packets mailed to the schools include copies of Science '84, Science Books and
Films, a bulletin for the principal, pamphlets and fact sheets on science and science
careers, posters, teaching notes and a project newsletter.
Titles of the teaching guides include:
Bubbles,
Maps and Mapping,
Image Markers,
Fluid Patterns,
Cooking and Science.
There are hopes that this program will be expanded into additional states in 1985-86,
including Indiana; but that will depend upon the availability of new funding. Examina-
tion of some of these activities for students indicate that they would help students
to understand science better, and to enjoy science. If you are a teacher of grade 7
or 8, you should write to AAAS Office of Science and Technology Education, 1776
Massachusetts Avenue, NW, Washington, DC 20036 to let them know you wish infor-
mation on the Science Resources Program for Grades 7 and 8.
483
484 Indiana Academy of Science Vol. 94 (1985)
Determining Needs: First Step for Improving Science and Mathematics Instruction in
Rural High Schools in Northwestern Indiana. G. Earle Francq and Jerry M. Col-
glazier, Indiana Department of Education, Indianapolis, Indiana 46204. Fostering
scientific and mathematical literacy for Indiana students is considered an essential goal
for assuring economic and technological prosperity. However, adequate opportunities
may not exist in many rural schools to develop these proficiencies because of unique
conditions associated with the rural environment. Rural areas have financial and popula-
tion characteristics which not only differ from urban communities, but also vary in
comparison across other rural regions. Consequently, one must first identify the uni-
que circumstances associated with a particular rural region before any procedures can
be implemented to improve educational settings.
This report examines evidence pertaining to science and mathematical educational
needs as indicated for selected rural high schools of northwestern Indiana.
Superintendents, high school principals, and high school physics, chemistry, and calculus
teachers responded to both a Science/Mathematics Needs Assessment and a Stage of
Concerns Questionnaire. Results revealed that (1) available funds were inadequate for
modernizing or replacing laboratory equipment to keep pace with current technology,
(2) availability of qualified applicants who could satisfy the multi-subject needs of
the schools was inadequate for filling physics, chemistry, and calculus teaching vacan-
cies, and (3) teachers were not provided adequate inservice to keep abreast of instruc-
tional development in physics, chemistry, and calculus.
Educational institutions and agencies can help to resolve the equipment, teacher
licensing, and inservice factors which are barriers affecting implementation of science
and mathematics instruction. These efforts would certainly aid future rural citizens
to develop their abilities to apply scientific and mathematical skills for personal, social,
and economic advantages.
The Layered Classifier: A More Effective Method for Studying Seasonal Changes in
Forest Cover Types Using Satellite Data. D. FABian Lozano-Garcia and Roger M.
Hoffer, Department of Forestry and Natural Resources and Laboratory for Applica-
tions of Remote Sensing, Purdue University, West Lafayette, Indiana
47907. Considerable interest has been generated in recent years concerning the
destruction of forests in many countries of the world. There has been speculation con-
cerning possible relationships between the increased level of carbon dioxide in the
earth's atmosphere. Landsat multispectral scanner data provides a unique opportunity
to study the earth's surface and the extent and condition of various cover types, in-
cluding forest cover. Many different computer-aided analysis techniques have been
developed to classify Landsat multispectral scanner (MSS) data. However, most of
these techniques are designed to utilize a single data set. To determine changes in the
extent or condition of the forest canopy that may occur over time requires overlaying
multiple data sets and a different approach to the analysis of such a multi-temporal
data set.
This research examines the effectiveness o f the Layered Classifier as a possible
technique for analyzing multi-temporal Landsat data. The test site was near the Monroe
Reservoir in the Hoosier National Forest. Landsat satellite data, obtained on four dates
throughout the year were digitally registered and analyzed. The results show that the
Layered Classification technique enable more accurate classification results to be
obtained, and at far less cost (in terms of computer time needed) than were obtained
by simply combining the data from two dates and applying a standard maximum
classification algorithm. These results provide significant insights into effective techni-
ques for using satellite data to monitor changes in forest canopy conditions or areal
extent.
Science Education 485
Synthesis Experiments for High School Chemistry. James George, Department of
Chemistry, DePauw University, Greencastle, Indiana 46135. The preparation of
compounds is an aspect of chemistry which can readily be incorporated into a high
school chemistry laboratory program. While working with high school aged students
during the past few years, I have had the opportunity to develop several such experiments.
They can be performed quickly and inexpensively with simple laboratory equipment.
They also require relatively little preparation time for the teacher. Included are the
synthesis of potassium aluminum sulfate, aspirin and salicylic acid, potassium acid
tartrate, potassium trioxalatoferrate, potassium chlorate, and cuprous chloride. Copies
of the experiments will be available for high school teachers and students.
The International Challenge: A Comparison of Science Education Models from Four
Nations. Linda Hamrick, The Canterbury School, Fort Wayne, Indiana 46804 and
Harold Harty, Department of Science and Environmental Education, Indiana Univer-
sity, Bloomington, Indiana 47405. Within the curricular formats of the United
States, it becomes relatively easy to lose perspective concerning standards, requirements
and course options relative to other nations. This presentation will highlight the major
underlying themes, the curricular requirements and program structures regarding science
education in Hawaii (as a cultural sub-group/The Kamehameha Schools), Great Britain,
Saudi Arabia and China. The conspectues will follow the results into each country's
mainstream of life as their science education practices affect life-long approaches to
education in general, the sciences specifically, and the society which they seek to educate.
Summative statistics regarding course requirements contrasted with the United States
will attempt to establish a perspective of our own practices in an international con-
figuration. The viewpoint emerges highlighting the difficulty of managing to support
a technological society in this country when our public secondary institutions generally
require only about one third the number of science courses compared to other in-
dustrialized nations. Implications for changing the number and type of required courses
are discussed.
A New Approach to Fostering Scientific Literacy among Indiana's Secondary School
Students. Susan M. Johnson, Department of Biology, Ball State University, Muncie,
Indiana 47306. During the past summer millions of Americans sat before their
televisions enraptured as Olympic athletes explored the limits of human physical strength
and agility. The same sort of delight in viewing athletic competitions results in legions
of avid sports fans who energetically support their favorite teams and encourage their
children to develop athletic skills. Competitive sporting events have enormous popular
appeal. For science educators who watch students and parents come alive at a basket-
ball game but turn somnolent at the mention of science, or who watch school boards
loosen the purse strings for athletic programs and close them abruptly when support
for science programs is sought, the question arises: Is there a way to generate popular
enthusiasm for science that approximates even a fraction of the excitement, support,
and prestige associated with sports?
In answer to this question, leaders in Delaware and Michigan have established
statewide Science Olympiads, which encourage students to explore the limits of their
mental strength and agility. This paper describes the Science Olympiad concept and
the effects of the Olympiad on school science programs in Delaware and Michigan.
The paper also discusses the possibilities and practicalities of implementing a Hoosier
Science Olympiad.
486 Indiana Academy of Science Vol. 94 (1985)
Science Training for the Industrial Environment (STIE). Paul B. Kissinger, Depart-
ment of Physics and Astronomy and John A. Ricketts, Department of Chemistry,
DePauw University, Greencastle, Indiana 46135. This paper describes STIE, a
new DePauw program developed specifically for science-oriented students who plan
to have significant, future managerial or sales responsibilities. STIE is based on the
premise that our society is growing more complex with greater technological orienta-
tion; consequently, managers who can "speak the language" of research scientists and
understand the patterns, problems and time frames associated with industrial research
and development should be extremely valuable in helping to direct and focus an in-
stitution's resources. Accordingly, broadly prepared individuals whose backgrounds
include both strong scientific and managerial training will be increasingly in demand
by basic industries, high-tech companies, government agencies and the like. The rela-
tionship of TIE to the DePauw Management Fellows Program will be outlined; in
addition, various course syllabi, laboratory exercises and the role of off-campus in-
ternships will be stressed. A specially developed seminar to explore the interrelation-
ships between society, science and technology also will be discussed. Although STIE
has significant professional orientation because it is designed to give students specific
preparation for the early, crucial stages of their business careers, it will be shown that
STIE students will have no difficulty satisfying DePauw's broad, traditional, liberal
education requirements.
Field Biology: A Blow to Provincialism. Rosalie Kramer, Department of Biology,
Indiana University East, Richmond, Indiana 47374. Students at small, nonresidential
campuses tend to be older and more closely tied to their community than do tradi-
tional students. Often they lack experience with environments and cultures other than
the ones on their immediate area. The field biology course at Indiana University East
has provided the faculty with the means to expose students to environments that are
totally new.
Through the vehicle of the course, students become acquainted with:
(1) new biological organisms and their environments,
(2) new subcultures and lifestyles, and,
(3) often new interpersonal interactions.
Students learn to work together, share living space, and take responsibility for
themselves and others. All the students were amazed at the amount of biology learned
in a two-week period as well as personal growth achieved.
Indiana University East's field biology course was conducted in the Florida
Everglades and Keys.
Speaking of Sex — A Presentation on Terminology for Students in Reproductive Biology
Classes. John Richard Schrock. Association of Systematics Collections, Museum
of Natural History, University of Kansas, Lawrence, Kansas 66045. Just as dif-
ferent political groups use different vocabularies, groups of people use different
vocabularies concerning sexuality. This brief classroom presentation describes the five
general attitudes toward sexuality and accompanying vocabularies and explains the
necessity of using detailed rationalistic terms in the explanation of current knowledge
in reproductive biology and in future scientific research. This presentation has been
found in reducing the conflict between students and parents on sex education.
Improving the Results of Molecular Mass Determination Experiments by Using a
Microelectronic Thermistor Device. Richard E. Schuley, Seymour, Indiana 47274
Science Education 487
and Marshall P. Cady, Jr., Department of Natural Sciences, Indiana University
Southeast, New Albany, Indiana 47150. An inexpensive, easy to use microelec-
tronic thermistor was developed for use in introductory chemistry labs. The specific
purpose of the device is to collect data for use in freezing point depression studies
of solutions. The device consists of an Omega 44006 thermistor interfaced with an
Intersil 7106 evaluation microprocessor kit and an LCD display. Modification of the
kit was necessary to alter the original function, the measurement of voltage, to display
resistance. Over small temperature ranges, it was found that the resistance displayed
by the LCD is related to the Kelvin temperature by the formula 1/T = A + B InR
where T is the temperature in Kelvin units, R is the resistance in kilo-ohms and A
and B are constants that vary with each thermistor. Each thermistor must be calibrated
separately. In this case, the development of the values of the constants was done by
computer, but can be done by simple substitution of variables when two temperatures
and their respectively resistances are known. The device was found to be accurate to
within 0.03 Kelvin units. Using the device, error in molecular mass experiments, which
is normally high, was reduced to a value of under two percent. As of the middle of
1984, the cost of this device, which must be assembled by the operator, was under
fifty dollars.
An Introductory Titration for First Year Chemistry Students: A Comparison of Antacid
Effectiveness. Katharine Sessions, The Canterbury School, Fort Wayne, Indiana
46804. High school students appear to learn abstract concepts more readily when
those concepts are presented via demonstrations using familiar materials. The effec-
tiveness of various antacids against stomach acid can be demonstrated with a simple
titration. This experiment is effective in introducing the physical techniques of titra-
tion, as well as providing an opportunity to discuss proper choice of an indicator,
selection of normality range for the standard solution, and potential difficulties en-
countered in titrations due to extraneous materials such as fillers found in antacid
tablets. The experiment, as well as teaching basic techniques, also satisfies student
curiosity concerning common advertising claims made by competing drug companies.
Using the Microcomputer to Teach Science in the Elementary Classroom. Stanley
S. Shimer, Science Teaching Center, Indiana State University, Terre Haute, Indiana
47809. In the third and fourth grade classes the unit on planets has been a typical
one for many years. The presentation will demonstrate how the microcomputer with
software can stimulate interest and motivate students to ask many questions. The soft-
ware can provide students data to be collected on solar distances, rates of travel by
various modes and their weight on the selected heavenly body. The program titled
"Solar Distances" also helps students to learn the sequence of the planets from the
Sun to Pluto. The worksheet was designed to be used with the software from Min-
nesota Education Computer Consortium (MECC) listed as Elementary Volume 4, Ver-
sion 1 1983.
Computer Aided Classroom Presentations in Chemistry. James T. Streator, Depart-
ment of Chemistry, Manchester College, North Manchester, Indiana 46962. Using
a computer in the classroom frequently requires a lot of out-of-class preparation time.
To minimize preparation time and to allow fairly spontaneous presentations in class,
input to the game port of a Commodore 64 has been used to generate multicolor displays
in class. The Koala Pad has been used to make presentations in beginning chemistry
classes as well as in Analytical and Physical Chemistry. Topics in bonding and struc-
ture and the drawing of titration curves and phase diagrams have been enhanced using
488 Indiana Academy of Science Vol. 94 (1985)
this device. A simple analog interface developed at Manchester College has also been
used in classroom demonstrations to show phenomena such as phase changes and oscilla-
tions. Both of these devices will be demonstrated and the impact of using such devices
will be discussed.
Color Vision: A Lecture Demonstration of Afterimages. Albert A. Williams, Depart-
ment of Biology, Manchester College, North Manchester, Indiana 46962. To the
majority of students, at any level of biology education, the areas of directly perceivable
human physiology have always been the most interesting. How they see, hear, balance,
feel pain, hunger, thirst, etc. captures the attention of students far more easily than
the details of meiosis or the Krebs cycle. Instructors can capitalize on this inherent
interest in order to lead the student deeper into the subject matter.
The area of visual perception lends itself exceptionally well to graphic demonstra-
tions that illustrate some very basic physiological concepts. By sequentially demonstrating
the nature of colored light and our ability to respond to, and differentiate between,
a broad spectrum of observable colors we can lead a class into a discussion of how
our retina/cortex complex recognizes individual colors. Generating a series of simple
and complex afterimage patterns for the entire class clearly demonstrates two major
physiological concepts: 1) that our visual sensory receptors are subject to exhaustion
and 2) that our color perception depends on a mental interpretation of the stimuli
rather than a direct stimulus/receptor relationship. These demonstrations can then lead
into a discussion and subsequent verification of the cellular basis and tri-color theory
of color perception at a level appropriate for the specific course.
A simple system for generating afterimages for an entire class will be demonstrated.
CLIMATE: A Microcomputer Program Allowing Student Preparation of
Climatic Maps for Indiana
Gary E. Dolph
Department of Botany
Indiana University at Kokomo
Kokomo, Indiana 46902
When studying the natural history or ecology of Indiana, one topic that the students
should master is climate, because of its influence on the distribution of plants and
animals in the State. However, effective class discussion of a series of climatic maps
can be very difficult. To increase student understanding of climatic variation in In-
diana, I prepared a computer program, called CLIMATE, for use on the Apple He.
Using CLIMATE, the students can generate their own climatic maps of Indiana, and
they can compare variation in climate with variation in plant distribution and plant
morphology. CLIMATE'S data base contains information on temperature and precipita-
tion from 91 weather stations in Indiana and the adjacent states of Michigan, Ohio,
Kentucky, and Illinois. Information on seven different climatic variables is available:
temperature, range in temperature, precipitation, biotemperature, potential
evapotranspiration ratio, effective temperature, and equability. Copies of CLIMATE
as well as the other support programs mentioned in the text may be obtained by send-
ing a blank disk to the author.
Weather Stations
Figure 1 shows the locations of the 91 weather stations used as data sources in
Indiana, Michigan, Ohio, Kentucky, and Illinois. The monthly normals of temperature
and precipitation for the period from 1941 to 1970 are recorded for each weather sta-
tion in the data base (11-15). More recent information is available from the National
Climatic Center in Asheville, North Carolina, but it was not used for two reasons.
First, the measurements of temperature and precipitation for the period from 1941
to 1970 are as accurate as for any subsequent period. Second, the older data were
used to minimize the ever increasing effects of industrialization and urbanization follow-
ing World War II (10). However, if more recent information is desired, a totally new
data base may be constructed using the programs (MTEMP and APREC) discussed
below.
Temperature data are available from 65 weather stations in Indiana; and precipita-
tion data are available from 84. Because information about both temperature and
precipitation is required to calculate the potential evapotranspiration ratio, only infor-
mation from weather stations recording both temperature and precipitation is included
in the data base. This requirement limited the number of usable weather stations in
Indiana to 65. The remaining 26 weather stations in the data base are from the sur-
rounding states of Michigan, Ohio, Kentucky, and Illinois. Inclusion of these 26 weather
stations minimizes the distortion of the climatic contours at Indiana's borders.
Programs for Modifying and Collecting Climatic Data
Four programs (Mean TEMPerature, (MTEMP), MODify TEMPerature
(MODTEMP), Average PRECipitation (APREC), and MODify PRECipitation
(MODPREC)) were used to collect and correct the initial climatic data. These four
programs do not require the user to specify how much data the programs will be handl-
ing. Therefore, these programs can be used to collect and correct variable amounts
489
490
Indiana Academy of Science
Vol. 94 (1985)
• 3
Figure 1. Locations of the 91 weather stations for which climatic data are available
when using CLIMATE. The names of the weather stations are given in Table 1.
of climatic data for any state or combination of states. Except for prompts specifying
the type of data to enter (e.g., precipitation or temperature), MTEMP and APREC func-
tion similarly as do MODTEMP and MODPREC. Simplified flow charts for these
two sets of programs are given in Figure 2. MTEMP and APREC are diagrammed on
the left; and MODTEMP and MODPREC are diagrammed on the right.
MTEMP and APREC are used to collect data. First, each program dimensions
an array to hold either temperature or precipitation data. Then, APREC or MTEMP
Science Education 491
Table 1. Key to the weather stations referenced in Figure 1. Unless otherwise stated,
the weather stations are in Indiana.
Number Weather Station
1 Eau Claire, Michigan
2 Three Rivers, Michigan
3 Coldwater State School, Michigan
4 Chicago University, Illinois
5 Gary
6 Hobart
7 Valparaiso Waterworks
8 LaPorte
9 South Bend WSO
10 Goshen College
1 1 Angola
12 Montpelier, Ohio
13 Wheatfield
14 Plymouth Power Substation
15 Warsaw
16 Albion
17 Waterloo
18 Kankakee, Illinois
19 St. Joseph's College, Collegeville
20 Winamac
21 Rochester
22 Columbia City
23 Fort Wayne WSO
24 Paulding, Ohio
25 Kentland
26 Fowler
27 Delphi
28 Wabash
29 Marion
30 Huntington
31 Berne
32 Van Wert, Ohio
33 Hoopeston, Illinois
34 Frankfort Disposal Plant
35 Kokomo
36 Salamonia
37 Danville, Illinois
38 Crawfordsville Power Plant
39 Whitestown
40 Anderson Sewage Plant
41 Winchester Airport
42 Greenville Water Plant, Ohio
43 Paris Waterworks, Illinois
44 Rockville
45 Greencastle
46 Indianapolis WSO
47 Greenfield
48 Cambridge City
49 Richmond Waterworks
50 Terre Haute
51 Franklin
52 Shelbyville Sewage Plant
53 Greensburg
54 Brookville
55 Hamilton/Fairfield, Ohio
56 Palestine, Illinois
57 Elliston
58 Indiana University, Bloomington
59 Columbus
492 Indiana Academy of Science Vol. 94 (1985)
Table 1. — Continued
Number Weather Station
60
Olney, Illinois
61
Vincennes
62
Edwardsport Power Plant
63
Washington
64
Crane Naval Depot
65
Highway 50 Bridge, Shoals
66
Purdue Experimental Farm, Oolitic
67
Bedford
68
Seymour
69
North Vernon
70
Covington WSA, Kentucky
71
Abbe Observatory, Cincinnati, Ohio
72
Fairfield Radio, WF1W, Illinois
73
Johnson Experimental Farm
74
Princeton
75
Paoli Radio, WVAK
76
Salem
77
Scottsburg
78
Henryville State Forest
79
Madison Sewage Plant
80
Vevay
81
Williamstown, Kentucky
82
McLeansboro, Illinois
83
Mt. Vernon
84
Evansville WSO
85
Tell City Power Plant
86
Jeffersonville
87
Louisville NSO, Kentucky
88
Anchorage, Kentucky
89
Henderson, Kentucky
90
Owensboro, Kentucky
91
Irvington, Kentucky
sequentially requests the following information: 1) the number of the weather station,
2) the mean temperature or average precipitation for each month beginning with January,
and 3) the mean annual temperature or average annual precipitation over the thirty
year period from 1941 to 1970. When these operations are completed, the information
for that weather station (one record) is transferred to the disk. The data base which
is ultimately constructed represents a random access data file. Each record contains
only one complete data set (i.e., temperature or precipitation data for a single weather
station). This structure allows for fast access to the data, whether the required infor-
mation is located in the first or last record in the file, through the use of a record
number, the weather station number.
MODTEMP and MODPREC are used to make additions or corrections to the
data base constructed using MTEMP and APREC. First, MODTEMP or MODPREC
requests the number of the weather station (record number) for which incorrect infor-
mation has been recorded. This entire record will be printed on the monitor. A correc-
tion is made by specifying the number of the month having incorrect data (1 for January
up to 13 for the thirty-year mean). After any corrections have been made, the new
record replaces the old on the disk.
Using these programs, two data bases were constructed for use with CLIMATE:
TEMPDAT (TEMPerature DATa), containing temperature data; and PRECDAT
(PRECipitation DATa), containing precipitation data. The data were recorded in the
Science Education
493
( START J
DIMENSION
VARIABLES
I
r INPUT WEATHER
STATION NUMBER
T
INPUT TEMPERATURE
OR PRECIPITATION DATA
I
INPUT MEAN ANNUAL
TEMPERATURE OR AVERAGE
ANNUAL PRECIPITATION
WRITE THE RECORD
IN A DATA FILE
ADD
ANOTHER
RECORD
NO
YES
( STOP J
( START J
DIMENSION
VARIABLES
REQUEST A
WEATHER STATION
DISPLAY THE DATA
T
)
r SPECIFY FIELD TO
BE CORRECTED
["ENTER CORRECTION
REPLACE RECORD
IN THE DATA FILE
X
MODIFY
ANOTHER
RECORD
NO
YES
( ST0P )
Figure 2. Simplified flow charts illustrating the functioning of MTEMP and APREC
(left) as well as MODTEMP and MODPREC (right).
English system. This mode of data storage was adopted for two reasons. First, the
data may still be compared directly to the information in the original data sources
(11-15). Second, although CLIMATE provides information to the students in the metric
system, the program can be modified to work with information in the English system.
Climate
The flow chart in Figure 3 illustrates how CLIMATE functions. At the start of
execution, CLIMATE constructs two arrays, one for temperature data and one for
precipitation data, based on the two data bases, TEMPDAT and PRECDAT. Then,
CLIMATE sequentially requests the following information: 1) the type of climatic
parameter to analyze; 2) the time period over which to calculate the climatic parameter;
and 3) the number of the weather station at which to calculate the climatic parameter.
Seven different climatic parameters may be studied (see below). Values for these climatic
parameters may be calculated for the whole year or for selected portions of the year,
494
Indiana Academy of Science
Vol. 94 (1985)
( START )
DIMENSION VARIABLES
INPUT CLIMATIC DATA FROM DISC
I
CONVERT DATA TO METRIC SYSTEM
CHOOSE CLIMATIC
PARAMETER
T
INPUT TIME PERIOD
T
^CHOOSE WEATHER
STATION
CALCULATE RESULTS
<
k.
DISPLAY RES
ULTS J
X
ANOTHER YES
ANALYSIS
NO
( ST0P )
Figure 3. Simplified flow chart showing the overall operation of CLIMATE.
Science Education 495
such as the growing season. Any of the 91 weather stations may be chosen as a data
source. After the calculations are completed, the results are displayed on the monitor.
After each calculation, the student can decide whether or not to continue with the
analysis.
Climatic Parameters
Seven climatic parameters can be studied using CLIMATE: temperature, range
in temperature, precipitation, biotemperature, potential evapotranspiration ratio, ef-
fective temperature, and equability. In addition to being unique expressions of various
aspects of the environment, these seven climatic parameters represent three different
methods of correlating climate with plant form. First, several attempts have been made
to relate mean annual temperature, mean annual range in temperature, and/or total
annual precipitation to variation in vegetational cover or to variation in plant form
(e.g., 9, 16-20). Second, Holdridge (7, 8) uses mean annual biotemperature, average
annual precipitation, and the potential evapotranspiration ratio to define the life zone,
the basis of his vegetational classification. Finally, Bailey (1-4) believes that effective
temperature (how frost free a given locality is) and equability (the freedom of a locality
from extreme heat and cold) can be used along with mean annual temperature and
mean annual rainfall to delineate the distribution of modern plant communities.
Average temperature is calculated by summing the mean temperature for a specific
number of months and then dividing by the total number of months in that time period.
The whole year or only a portion of the year may be selected. When values for the
whole year are requested, the mean annual temperature is calculated. If only a portion
of the year, such as the growing season, is desired, two numbers ranging from 1 (January)
to 12 (December) are used to indicate the starting and stopping months for the
calculation.
The range in temperature is calculated by subtracting the lowest mean monthly
temperature in the selected time period from the highest. When values for the whole
year are used, the mean annual range in temperature is calculated. The range in
temperature for less than a full year is requested by selecting the appropriate starting
and stopping months.
Summing the average monthly precipitation over a specified time period yields
the total precipitation. If values for all twelve months are requested, the average an-
nual precipitation is calculated. The total precipitation for periods of less than a year
is calculated by selecting the appropriate starting and stopping months.
Biotemperature is the average temperature between 0°C and 30°C at which vegeta-
tional growth takes place (7, 8). The average biotemperature is the sum of the mean
monthly temperatures between 0°C and 30°C over a specific time period divided by
the total number of months in that time period. Because months having a mean value
above 30°C or below 0°C are not used to calculate biotemperature, values for average
biotemperature of less than 0°C or greater than 30°C are not possible. When data
for all twelve months are used, the mean annual biotemperature is calculated. If values
above 30°C or below 0°C are not encountered within the specified time interval, the
calculated values of average biotemperature and mean temperature are equal. The
calculated values of average biotemperature and mean temperature are equal in warm
temperate states such as North and South Carolina (6), but they are not equal in cold
temperate states such as Michigan.
The potential evapotranspiration ratio is a climatic index calculated using both
biotemperature data and precipitation data (7, 8). First, potential evapotranspiration,
a hypothetical number, is calculated by multiplying the biotemperature for a specific
time interval by 58.93. Then, the potential evapotranspiration ratio is calculated by
496 Indiana Academy of Science Vol. 94 (1985)
dividing the potential evapotranspiration by the average precipitation for the same time
interval:
PER = (58.93 x T<bioVP,
where the potential evapotranspiration ration (PER) is a dimensionless number, T> °'
is the biotemperature given in °C, and P is the total precipitation given in mm. A
value of 1.00 indicates that precipitation balances potential evapotranspiration over
the time period selected. The most favorable habitats for plant growth and human
activity cluster about a potential evapotranspiration ratio of 1.00. As the potential
evapotranspiration ratio increases, the precipitation is less than needed to balance poten-
tial evapotranspiration and the habitat becomes more arid. As the potential
evapotranspiration ratio decreases, the precipitation is more than is needed to balance
potential evapotranspiration and the habitat becomes more moist. The potential
evapotranspiration ratio ranges from a high of 32.0 in the super-arid humidity pro-
vince to a low of 0.125 in the super-humid humidity province. The potential
evapotranspiration ratio may be calculated for the whole year or for a portion of the
year by selecting the appropriate starting and stopping months.
Effective temperature is the temperature, expressed in °C, at the beginning and
end of a warm period in which vegetational growth occurs (1-4). This warm period
will be largely free from frost. Effective temperature (ET) is calculated as:
ET = (8T + 14A) / (8 + A),
where T is the average temperature in °C and A is the range in temperature in °C
for a specific time period. The change from cold climates to warm climates is indicated
by an increase in the value of effective temperature. Polar climates have an effective
temperature of less than 10°C; cool midlatitude climates of between 10°C and 14°C;
warm midlatitude climates of between 14°C and 18°C and tropical climates of more
than 18°C. Each of these categories can be further subdivided into smaller climatic units.
Equability measures the freedom of a specific locality from extremes of heat or
cold regardless of whether or not the extremes are perennial or seasonal (1-4). Equability
(M) is calculated using the following formula:
M = 109.0 — 30 log ((T— 14)2 + (1.46 + 0.366A)2),
where T is the average temperature in °C and A is the range in temperature in °C
for a specific time period. Equability is a dimensionless index, ranging from 0 to 100.
The lower the equability, the more extreme the climate. A climate which is totally
free from extremes of heat or cold has an equability of 100. The outer limit for temperate
climates is an equability of 55.
Classroom Uses
Rather than discussing what must appear to be an endless series of climatic maps
in the classroom, CLIMATE allows two different types of analyses to be carried out.
First, after analyzing the data and drawing the appropriate climatic maps, the students'
maps can be compared with published climatic maps for Indiana (5, 10). Normally,
the students' maps will lack the smooth contours characteristic of the published maps.
Factors that can influence a climatic measurement, such as elevation, exposure, or
the time of day when the measurement was taken, require that a climatologist treat
Science Education 497
these measurements as relative and not absolute values. Therefore, there is a tendency
for the more divergent measurements to be rejected when publishing a climatic map.
The rejection of some data results in a climatic map which is more aesthetically pleas-
ing even if it is not absolutely accurate. By comparing the students' maps with the
published maps, the role judgment plays when producing climatic maps can be discussed.
Second, when used in conjunction with vegetation maps, the relationship between vegeta-
tion and climate as well as the relative merits of the three methods of climatically
classifying vegetation may be assessed.
Evaluation
The teaching effectiveness of CLIMATE and two other simulations (work in pro-
gress) was evaluated using a pretest-posttest design with experimental and control groups.
Both groups were given the same 50 question pretest and posttest. Based on the results
of these tests, the mean gain scores registered for each group were calculated. Two
different tests (a separate variance estimate of the difference in mean gain scores and
an analysis of covariance for the posttest scores of the experimental and control groups)
were run to see if the gain in the experimental group was significantly greater than
the gain in the control group. In each case, the difference was found to be significant
at a 5°Io significance level.
Outcomes
A number of important student outcomes result from using CLIMATE:
1) By calculating the positions of their own contour lines, the students gain
valuable insights into how maps, such as weather maps, geologic maps, or vegeta-
tion maps, are constructed.
2) After discussing trends in climatic variables, the students should find it easier
to identify trends in other mappable variables, such as vegetation characteristics.
3) By being actively involved in constructing and analyzing climatic maps, the
students gain greater appreciation for the variation in Indiana's ciimate than could
be obtained by passively attending lecture.
4) By comparing their maps with published climatic maps for Indiana, the students
can see how the acceptance or rejection of data from various weather stations
can influence the final form of their maps.
5) By comparing the ability of different climatic parameters to explain the varia-
tion in vegetation patterns in Indiana, the students can begin to explore the rela-
tionship between vegetation and climate.
Acknowledgments
This material is based upon work supported by the National Science Foundation
under Grant No. SER 8004789. The author would like to express his thanks to Dr.
Lian-Hwang Chiu for assisting with the statistical analysis and to Dr. Charles R. Bar-
man for critically reviewing the manuscript.
Literature Cited
1. Axelrod, D.I. and H.P. Bailey. 1969. Paleotemperature analysis of Tertiary floras.
Palaegeogr., Palaeclimatol., Palaeoecol. 6:163-195.
498 Indiana Academy of Science Vol. 94 (1985)
2. Bailey, H.P. 1960. A method of determining the warmth and temperateness of
climate. Geograf. Ann. 42:1-16.
3. . 1945. Toward a unified concept of temperate climate. Geograph. Rev.
54:516-545.
4. 1966. The mean annual range and standard deviation as measures of disper-
sion of temperature around the annual mean. Geograf. Ann. 48:183-194.
5. Dolph, G.E. 1984. Leaf form of the woody plants of Indiana as related to en-
vironment. In: N.S. Margaris, M. Arianoustou-Farragitaki, and W.C. Oechel
(Eds.), Being Alive on Land, pp. 51-61, Tasks for Vegetational Science, Vol. 13,
Dr. W. Junk Publishers, The Hague, 322 pp.
6. and D.L. Dilcher. 1979. Foliar physiognomy as an aid in determining
paleoclimate. Palaeontographica, Abt. B, 170:151-172.
7. Holdridge, L.R. 1967. Life zone ecology. Tropical Science Center, San Jose, Costa
Rica, 206 pp.
8. , W.C. Grenke, W.H. Hatheway, T. Laing, and J. A. Tosi, Jr. 1971. Forest
environments in tropical life zones: A pilot study. Pergamon, New York. 747 pp.
9. Raunkiaer, C. 1934. The life-forms of plants and statistical plant geography. Ox-
ford University Press, Oxford, 632 pp.
10. Schaal, L.A. 1966. Climate. In: A. A. Lindsey (Ed.), Natural Features of Indiana,
pp. 156-170, Indiana Academy of Science, Indianapolis, 597 pp.
1 1. U.S. Department of Commerce Environmental Data Service. 1973a. Climatology
of the United States, No. 81 (Illinois). Monthly normals of temperature, precipita-
tion, and heating and cooling degree days 1941-1970. Asheville, North Carolina.
12. . 1973b. Climatology of the United States, No. 81 (Indiana). Monthly nor-
mals of temperature, precipitation, and heating and cooling degree days 1941-1970.
Asheville, North Carolina.
13. 1973c. Climatology of the United States, No. 81 (Kentucky). Monthly nor-
mals of temperature, precipitation, and heating and cooling degree days 1941-1970.
Asheville, North Carolina.
14. . 1973d. Climatology of the United States, No. 81 (Michigan). Monthly nor-
mals of temperature, precipitation, and heating and cooling degree days 1941-1970.
Asheville, North Carolina.
15. . 1973e. Climatology of the United States, No. 81 (Ohio). Monthly normals
of temperature, precipitation, and heating and cooling degree days 1941-1970.
Asheville, North Carolina.
16. Wolfe, J. A. 1969. Paleogene floras from the Gulf of Alaska region. U.S. Geol.
Surv. Open-file Rep. 114 pp.
17. . 1971. Tertiary climatic fluctuations and methods of analysis of Tertiary
floras. Palaeogeogr., Palaeoclimatol., Palaeoecol. 9:27-57.
18. . 1978. A paleobotanical interpretation of Tertiary climates in the Northern
Hemisphere. Amer. Sci. 66:694-703.
19. . 1981. Paleoclimatic significance of the Oligocene and Neogene floras of
the northwestern United States. In: K.J. Niklas (Ed.), Paleobotany, Pa/eoecology,
and Evolution, Volume 2, pp. 79-101, Praeger, New York, 269 pp.
20. and D.M. Hopkins. 1967. Climate changes recorded by land floras in nor-
thwestern North American. In: K. Hatai (Ed.), Tertiary correlations and climatic
changes in the Pacific, pp. 67-76, Symp. Pacific Sci. Congr., Tokyo.
A Summer Institute in Microcomputer Applications
for Secondary School Science Teachers
L. Dwight Farringer
Department of Physics, James T. Streator,
Department of Chemistry, and
Albert A. Williams,
Department of Biology
Manchester College
North Manchester, Indiana 46962
Introduction
Last winter, the Indiana Consortium for Computer and High Technology Educa-
tion announced its support of 1984 summer institutes for Indiana teachers. The in-
stitutes were to provide computer-related training opportunities related to the teachers'
professional interests and local curriculum requirements. Teacher training institutions
in the state were invited to submit proposals for such institutes.
The Consortium approved Manchester College's proposal for a two-week institute
in microcomputer applications and interfacing for secondary school science teachers.
This institute was to be aimed at teachers who already had some programming ability
and who desired to increase their capabilities for utilizing the microcomputer as a science
teaching tool. Participants could earn three semester-hours of graduate credit for the
concentrated two-week experience.
Sixteen participants were selected, with the requirement that they already have
at least a year of experience in using and programming microcomputers in science
teaching. Teaching areas of the participants included physics, chemistry, biology, and
computer science. Geographically, their schools were distributed over the northern part
of the state, as far south as Indianapolis.
Nature of the Institute
It was planned that participants would work on projects which would be related
to their own teaching areas for the ensuing school year and which would run on
microcomputers available for use in their local schools.
Because interfacing and graphical procedures are quite hardware-dependent, it
was important that participants work on computers of types available to them in their
schools. To supplement the microcomputers available at Manchester College, participants
were invited (but not required) to bring microcomputers with them to the institute.
It turned out that 12 of the 16 participants brought microcomputers; some were from
their schools, and some were the individuals' own computers. A few of the participants
brought two different types of computers, as they expected to utilize them for dif-
ferent purposes in their schools.
It was made clear to the applicants that the hardware-dependent parts of the
work would be planned for Apple and Commodore 64 and VIC-20 computers. It was
explained that some users of other types of computers would be accepted as participants
in the institute, if they were willing to devote some of their efforts to making the
necessary adaptations to their computers. Several participants did at least part of their
work with Radio Shack and Atari computers, and they were generally able to make
desired adaptations to those computers.
Though the participants all had prior experience with microcomputers, there was
considerable variation in their areas of competence. Some were not very confident
of their abilities for ordinary programming in BASIC — such things as using data arrays,
499
500 Indiana Academy of Science Vol. 94 (1985)
getting data in and out of disk files, and performing string operations. On the other
hand, several of the participants were able to write machine-language subroutines to
perform such things as high-speed data moving operations.
The teaching staff consisted of the three authors of this paper, representing the
disciplines of biology, chemistry and physics. Each of us has worked for several years
on computer-related projects in our disciplines. We believe that the interdisciplinary
composition of the institute— both the participants and the teaching staff— was an im-
portant factor in the value of the experience. In most high schools, the science teaching
and learning are not highly specialized along the lines of traditional science disciplines.
Many problems encountered by science teachers in learning how to use the microcom-
puter are quite similar, regardless of which science discipline is involved. Throughout
the institute, it was clearly a meeting of science teachers, not of computer game players.
Topics and Projects
The institute emphasized use of the microcomputer as:
1) A computational tool for problem-solving, laboratory data analysis, simulation,
and modeling.
2) A graphical medium for plotting functions and displaying spatial relationships.
3) A laboratory instrument for interfacing with experimental apparatus.
The daily schedule of the institute involved morning, afternoon, and evening sessions,
with the evenings somewhat optional, depending on interest and stamina. Evening par-
ticipation was often about 80 percent.
Approximately one-third of the institute time was devoted to lecture/demonstra-
tion presentations by the teaching staff and two-thirds to work on individual projects.
Presentations by the instructional staff centered on:
1 ) Programming techniques for science teaching applications — e.g. com-
putation, graphing, sorting, storing and retrieving data.
2) Interfacing of microcomputers with external apparatus.
3 ) Transducers for physical measurements.
4) Demonstration of applications developed by the instructional staff.
5 ) Demonstration of software from various sources.
Hands-on experiences of all the participants included:
1 ) Assembling and testing of an interface board (the MIA, or Manchester
Interface Adapter) designed for use with Apple or Commodore
computers.
2 ) Assembling and testing of accessories for use with the MIA — pushbutton
switches, thermistors, light sensors, and connecting cords.
3 ) Sharing of science teaching ideas and software and hardware ideas among
the participants.
4) Trying out and adapting ideas from various sources — from the instruc-
tors, other participants, reprints of published articles, and commercial
software.
5 ) Becoming acquainted with the various types of microcomputers
represented in the institute.
Beyond these common experiences, there was much variation of individual
experiences. Some of the participants worked on improving their general programm-
ing skills in areas related to science teaching. Others used their previously acquired
Science Education 501
skills to develop particular applications in their teaching areas. Some worked mostly
on software development; others also did considerable work on apparatus for par-
ticular experiments.
The following summary gives an idea of what was done:
7 participants centered their efforts primarily on physics-related projects,
3 on chemistry-related projects, and 3 on biology-related projects.
All 16 participants successfully completed the assembly and testing of the Manchester
Interface Adapter and its accessories for timing and for measurement of temperature
and light intensity.
7 worked on temperature calibration and measurement procedures for
use with thermistor temperature probes.
3 worked on applications of light intensity measurement with photosen-
sors, e.g. observing the progress of chemical or biological processes
in terms of the light transmission through a test tube in which the pro-
cess is occurring.
6 worked on various motion timing experiments — e.g. timing of pendulums
and freely falling objects.
5 worked on other timing experiments — e.g. pulse-rate timing, animal ac-
tivity, strobe light calibration, and stop-watch type of computer
applications.
3 used the computer to control external devices — e.g. controlling a strobe
light, or controlling an electromagnet in a release mechanism for mo-
tion experiments.
5 developed instructional programs other than direct laboratory measure-
ment programs — e.g. simulations and demonstrations.
6 developed various utility programs for doing useful tasks related to
science teaching — e.g. graphic routines, data collecting and analysis,
record keeping and grading.
8 worked on various kinds of hardware development — devising apparatus
for experiments, adapting the MIA to other computers than the Apple
and Commodore for which it was designed, making a voltage analog-
to-digital converter, etching circuit boards for additional MIA's, etc.
4 did some work which involved creating machine-language subroutines
to achieve precision of timing or increased speed of certain computer
operations.
Evaluation
Evaluations were obtained from the participants on the appropriateness and quality
of the instruction, the value of the projects on which work was being done, and the
degree to which individual needs were being met. Predominantly good-to-excellent ratings
were given in all of these aspects of the institute.
Because the institute was aimed at the needs of science teachers who had already
achieved "computer literacy" and wanted to increase their ability to program and use
microcomputers as regular teaching tools, we were very interested in gaining ideas about
factors in the secondary school teaching environment which would favor the effective
utilization of microcomputers in science teaching.
One factor which came through very strongly was that integrating the powers
of microcomputers into science teaching depends strongly upon having the computers
available in science classrooms and laboratories for everyday use — not just occasionally
getting the use of a computer borrowed from a central computer room. Many impor-
tant uses of computers in science teaching depend upon the creativeness of teachers
502 Indiana Academy of Science Vol. 94 (1985)
and students to grasp the opportune occasions for applying the computer to those
tasks for which it is especially suited.
The institute participants quickly grasped the values of laboratory interfacing to
achieve microcomputer-based systems for experiments, demonstrations, and simula-
tions. The direct hands-on student involvement in such activities requires ready avail-
ability of multiple microcomputers.
Many of the institute participants have had experience with various under-$300
microcomputers — e.g. several models of Commodore, Radio Shack, and Atari
computers — which have really fine computational, graphical, and interfacing capabilities.
It seems likely that this can be an important key to making it feasible for science
teachers to have multiple computers for laboratory use — if school systems do not in-
sist that more expensive computer systems are the only ones which are worth buying.
If we have an opportunity to run a similar institute again next summer, the most
likely improvements to be achieved would be: 1) more effective adaptation of interfac-
ing techniques to several different types of inexpensive microcomputers, and 2) more
effectively air-conditioned working space for the institute.
Use of a Microcomputer to Enhance the Coin Flip Probability
Exercise in the General Biology Laboratory
K. Michael Foos
Department of Biology
Indiana University East
Richmond, Indiana 47374
Simple, but effective, software should be available for microcomputer use in the
classroom. I have heard over and over that there is not adequate software support
for biology, and I have come to believe that this is, in fact, the situation. I believe
that one way to combat this lack of available software is to produce software ourselves.
Having viewed several biology software packages, I am certain that if we, as teachers,
write our own software it will be as good as that on the market. And, there is a distinct
advantage to writing one's own software. When you write your own software you
can tailor it to your unique situation. Further, by developing a simple software package
one is often stimulated to think of another, more complex use for the microcomputer
in the classroom.
It is an attempt to encourage biology teachers to develop their own microcom-
puter software that is the primary purpose for this paper. The program described in
this paper is not particularly sophisticated, or complex. It is not difficult, or fancy.
It was designed to perform a particular task in my biology classroom, and it does
that one simple task rather well. I share it with you for your use, and perhaps more
importantly, to stimulate you to develop a better program, one that can do the same
task better or one that can expand the focus of this program.
It is common to introduce the topic of genetics with a discussion of probability
(1-3). Without a basic understanding of probability it is difficult or impossible to fully
discuss the concepts of Mendelian genetics. In the laboratory, probability can be
demonstrated easily using coins. However, there is a major limitation to flipping coins.
Students can be asked to do a limited number of coin flips before they become weary
and their flipping fingers get sore. But, it can be valuable for students to do fifty
or one hundred flips, as many laboratory exercises recommend.
After flipping their coins, students can be asked to calculate their ratios of results.
But, with such a small sample the likelihood of obtaining highly accurate results is
small. And, obtaining results far from the theoretical expected results can cause more
questions than it answers. One partial solution to this problem is to add all of the
individual sets of data to produce a larger class sample. This almost always gives a
sample result that more closely approximates the theoretical than the individual ob-
tains (except in those rare cases where an individual happens to hit the theoretical result).
The concept of increased accuracy with increased sample size is all too often ig-
nored. Using small numbers that can be obtained by manipulating coins provide a
very small sample size and thus not very accurate results. Further, the time required
to gather data from coin flips would be very substantial if there were a large enough
number of flips to be statistically significant.
The computer program that I am about to describe is a simple one devised to
show both the coin flip probability and the increased accuracy obtained when using
an increased sample size. This program was originally written in Applesoft BASIC
and will run on any Apple II or Apple He. There are no sophisticated programming
techniques that restrict the use of this program to the Apple. It could be run on almost
any computer with very slight modification. The program described here was not designed
to stretch the limits of the computer. It was designed to provide data for students
503
504 Indiana Academy of Science Vol. 94 (1985)
to analyze. And, the program is simple enough that almost anyone with any computer
experience can design such a program.
A listing of the program is included in Table 1.
Table 1 . Listing of the Probability Coin Flip Program for One Coin.
100 REM **COIN FLIP PROGRAM**
110 REM **M. FOOS, 11/12/81**
120 REM **TO DO SIMPLE PROBABILITY**
130 HOME: PRINT
140 PRINT "CLASSICALLY STUDENTS HAVE STUDIED"
150 PRINT "PROBABILITY BY FLIPPING COINS."
160 PRINT: PRINT "AS OUR ECONOMY IS CHANGING TO BECOME"
170 PRINT "MORE AUTOMATED, IT WOULD SEEM THAT"
180 PRINT "FLIPPING COINS WOULD ALSO BECOME"
190 PRINT "AUTOMATED."
200 PRINT: PRINT "THIS PROGRAM IS DESIGNED TO FLIP COINS"
210 PRINT "ELECTRONICALLY AND TO PRESENT THE"
220 PRINT "RESULTS OF THE FLIPS ON THIS SCREEN."
230 PRINT: PRINT "BECAUSE THE COMPUTER FLIPS THE COINS SO"
240 PRINT "RAPIDLY IT IS DIFFICULT TO SEE, IT WILL"
250 PRINT "ALSO COUNT THE NUMBER OF FLIPS IT DOES."
260 PRINT: PRINT "TO USE THIS PROGRAM ALL YOU HAVE TO DO"
270 PRINT "IS SELECT THE TOTAL NUMBER OF FLIPS YOU"
280 PRINT "WANT THE COMPUTER TO DO."
300 GOSUB 1000
600 HOME: N = 0:H = 0:T = 0:PRINT: PRINT
606 PRINT "HOW MANY COIN FLIPS DO YOU WANT THE"
610 PRINT "COMPUTER TO DO? ENTER YOUR NUMBER AND"
612 INPUT "PRESS RETURN.";N
620 HOME: PRINT: PRINT
630 X = INT(RND(1)*2)
640 IF X=l THEN PRINT "H";:H = H+1
650 IF X = 0 THEN PRINT "T";:T = T+1
660 IF H + T = N THEN GOTO 680
670 GOTO 630
680 PRINT: PRINT CHR$(7): PRINT " HEADS = "H" TAILS = "T
690 TH=TH+H:TT = TT + T
700 GET X$
710 IF X$ = "X" THEN 730
720 GOTO 800
730 HOME:PRINT:PRINT:PRINT
740 PRINT "THE TOTAL NUMBER OF COIN FLIPS DONE IN"
750 PRINT "THIS LAB WAS ";TT + TH
760 PRINT:PRINT:PRINT "THE RESULTING DISTRIBUTION IS "
770 PRINT "LISTED BELOW:"
780 PRINT:PRINT " HEADS = "TH" TAILS = "TT
790 END
800 HOME
810 FLASH
820 PRINT "HHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHH"
821 PRINT "HHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHH"
822 PRINT "HHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHH"
823 PRINT "HHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHH"
824 PRINT "HHHHHTTTTTTTTHTHHHHHHHHTTTHHTTTTTTTHHHH"
825 PRINT "HHHHHTHHHHHHTHTHHHHHHHHHTHHHTHHHHHHTHHH"
826 PRINT "HHHHHTHHHHHHHHTHHHHHHHHHTHHHTHHHHHHTHHH"
827 PRINT "HHHHHTHHHHHHHHTHHHHHHHHHTHHHHHHHHHHTHHH"
828 PRINT "HHHHHTHHHHHHHHTHHHHHHHHHTHHHHHHHHHHTHHH"
829 PRINT "HHHHHTHHHHHHHHTHHHHHHHHHTHHHHHHHHHHTHHH"
830 PRINT "HHHHHTHHHHHHHHTHHHHHHHHHTHHHHHHHHHHTHHH"
831 PRINT "HHHHHTHHHHHHHHTHHHHHHHHHTHHHHHHHHHHTHHH"
Science Education 505
Table 1. — Continued
832 PRINT "HHHHHTTTTTHHHHTHHHHHHHHHTHHHTTTTTTTHHHH"
833 PRINT "HHHHHTHHHHHHHHTHHHHHHHHHTHHHTHHHHHHHHHH"
834 PRINT "HHHHHTHHHHHHHHTHHHHHHHHHTHHHTHHHHHHHHHH"
835 PRINT "HHHHHTHHHHHHHHTHHHHHHHHHTHHHTHHHHHHHHHH"
836 PRINT "HHHHHTHHHHHHHHTHHHHHHHHHTHHHTHHHHHHHHHH"
837 PRINT "HHHHHTHHHHHHHHTHHHHHHHHHTHHHTHHHHHHHHHH"
838 PRINT "HHHHHTHHHHHHHHTHHHHHHTHHTHHHTHHHHHHHHHH"
839 PRINT "HHHHTTTHHHHHHHTTTTTTTTHTTTHTTTHHHHHHHHH"
840 PRINT "HHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHH"
841 PRINT "HHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHH"
842 PRINT "HHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHH"
843 PRINT "HHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHHH"
844 GOSUB 1000
860 GOTO 130
1000 PRINT:INVERSE:PRINT" < PRESS RETURN >
1001 NORMAL
1002 GET A$
1003 IF A$ = CHR$(13) THEN RETURN
1004 PRINT CHR$(7): GOTO 1000
The program can be broken into three parts. The first part consisting of lines
100-300 is an introduction to the program. This kind of an introduction is especially
important for students who have little or no working knowledge of the computer.
It is designed to put them at ease and give them a starting point for using the com-
puter. It helps them feel that they know what to do.
The second part of the program included in lines 600-790 actually makes the pro-
gram work. In looking at this section it is obvious that several different things happen.
In line 600 the screen of the computer's monitor is cleared and all of the counters
are set to zero. The student is then asked to enter a number of coin flips to be done
by the computer. Line 630 is the heart of this program. By using the computer's random
number generator, a number is selected. This instruction designates that the random
number selected be any integer less than two. Thus, the computer's random number
must be either a 0 or a 1.
Lines 640 and 650 instruct the computer to print the letter 'H' if the computer
selects a 1 and to print a letter 'T' if the computer selects a 0. These lines also add
the number of times a 'H' and a 'T' have been printed.
Lines 660 and 670 instruct the program to continue printing 'H' and 'T' until
the total number of letters printed equals the number requested by the student in the
beginning. When the correct total numbers of letters has been printed, the computer
prints the totals at the bottom of the screen as directed by line 680. Line 690 keeps
a running total of all of the counts in the session.
The 'GET' command used in line 700 accepts a single character from the keyboard
without using the < RETURN >. If a capital 'X' is pressed at this time, the computer
will print the aggregate total of all of the runs in the session as dictated in lines 730-780,
and will end the execution of the program (line 790).
The instructor will normally press this key after all of the students have received
their data. It is possible that a random stroke could cause this to happen out of turn,
but there is only a small probability.
Striking any key but the capital 'X' will cause the screen to show a flashing display.
This third part of the program is listed in lines 800-843. This display is nothing more
than a device to show that the computer is turned on and working. It lets the student
506 Indiana Academy of Science Vol. 94 (1985)
know that no one is currently using the computer, and it is free for their use. It flashes
on the Apple and could be enhanced to be colorful. It does not use either of the graphics
modes of the Apple and is therefore directly adaptable to another brand of computer.
However, the 'FLASH' command is a uniquely Apple command.
The last few lines 1000-1004 help to 'goof -proof the program. These lines pro-
hibit the input of any character other than the < RETURN > key when that response
is requested. The buzzer also sounds if any character other than the < RETURN >
is pressed.
By altering the lines 600-790 it is easy to modify this program to simulate flipping
two or three coins simultaneously. These modifications, listed in Table 2 and Table
3 expand the usefulness of the program from the simple probability relationships one
finds with a single variable to the consideration of two or three variables. This leads
to the logical discussion of dihybrid and trihybrid crosses as well as the typical
monohybrid cross. This simple modification expands the one program into three pro-
grams and thus increases its effectiveness.
Table 2. Modification for the Coin Flip Program for Two Coins.
600 HOME:N=0:HH=0:HT = 0:TH=0:TT = 0:PRINT:PRINT
606 PRINT "HOW MANY COIN FLIPS DO YOU WANT THE"
610 PRINT "COMPUTER TO DO? ENTER YOUR NUMBER AND"
612 INPUT "PRESS RETURN.";N
630 X= 1NT(RND(1)*4)
635 IF X=0 THEN PRINT "HH ";:HH = HH + 1
640 IF X = l THEN PRINT "HT ";:HT = HT+1
645 IF X = 2 THEN PRINT "TH ";:TH=TH + 1
650 IF X =3 THEN PRINT "TT ";:TT = TT+1
660 IF HH + HT + TH+TT = N THEN 680
670 GOTO 630
680 PRINT:PRINT CHR$(7): PRINT "HH = "HH
682 PRINT "HT = "HT
684 PRINT "TH = "TH
686 PRINT "TT = "TT
690 A = A + HH:B = B + HT:C = C + TH:D = D + TT
700 GET X$
710 IF X$ = "X" THEN 730
720 GOTO 800
730 HOME:PRINT:PRINT:PRINT
740 PRINT "THE TOTAL NUMBER OF COIN FLIPS DONE IN"
750 PRINT "THIS LAB WAS ";A + B + C + D
760 PRINT:PRINT:PRINT "THE RESULTING DISTRIBUTION IS"
770 PRINT "LISTED BELOW:"
779 PRINT
780 PRINT "HEADS • HEADS = "A
781 PRINT "HEADS • TAILS = "B
782 PRINT "TAILS • HEADS = "C
783 PRINT "TAILS • TAILS = "D
790 END
Table 3. Modification for the Coin Flip Program for Three Coins.
600 HOME:N=0:A = 0:B = 0:C = 0:D = 0:E = 0:F = 0:G = 0:H = 0
602 PRINT:PRINT
606 PRINT "HOW MANY COIN FLIPS DO YOU WANT THE"
610 PRINT "COMPUTER TO DO? ENTER YOUR NUMBER AND"
612 INPUT "PRESS RETURN. ";N
620 HOME:PRINT:PRINT
630 X = INT(RND(1)*8)
Science Education 507
Table 3. — Continued
635 IF X=0 THEN PRINT "HHH ";:A = A+1
637 IF X=l THEN PRINT "HHT ";:B = B+1
639 IF X=2 THEN PRINT "HTH ";:C = C+1
640 IF X = 3 THEN PRINT "HTT ";:D = D+ 1
643 IF X = 4 THEN PRINT "THH ";:E = E+ 1
645 IF X = 5 THEN PRINT "THT ";:F = F+1
647 IF X = 6 THEN PRINT "TTH ";:G = G+1
650 IF X = 7 THEN PRINT "TTT ";:H = H + 1
660 IFA + B + C + D + E + F + G + H = N THEN GOTO 680
670 GOTO 630
680 PRINT:PRINT CHR$(7)
681 PRINT "HHH= "A
682 PRINT "HHT = "B
683 PRINT "HTH = "C
684 PRINT "HTT = "D
685 PRINT "THH = "E
686 PRINT "THT = "F
687 PRINT "TTH = "G
688 PRINT "TTT = "H
690 TA = TA + A
691 TB = TB + B
692 TC = TC + C
693 TD = TD + D
694 TE = TE + E
695 TF = TF + F
696 TG = TG + G
697 TH = TH + H
700 GET X$
710 IF X$ = "X" THEN 730
720 GOTO 800
730 HOME:PRINT:PRINT:PRINT
740 PRINT "THE TOTAL NUMBER OF COIN FLIPS DONE IN"
750 PRINT "THIS LAB WAS ";TA + TB + TC + TD+TE + TF + TG + TH
760 PRINT:PRINT:PRINT "THE RESULTING DISTRIBUTION IS"
770 PRINT "LISTED BELOW:"
779 PRINT
780 PRINT "HEADS • HEADS • HEADS = "TA
781 PRINT "HEADS • HEADS • TAILS = "TB
782 PRINT "HEADS • TAILS • HEADS = "TC
783 PRINT "HEADS • TAILS • TAILS = "TD
784 PRINT "TAILS • HEADS • HEADS = "TE
785 PRINT "TAILS • HEADS • TAILS = "TF
786 PRINT "TAILS • TAILS • HEADS = "TG
787 PRINT "TAILS • TAILS • TAILS = "TH
790 END
The simple computer program described here works well in a classroom with a
discussion of probability and was designed primarily to be used in a first semester
college biology course. It works well to provide a large number of coin flips very rapidly.
This program will generate 100 coin flips in about three seconds; five thousand coin
flips can be generated in two minutes and ten seconds. Larger samples can be generated
by the computer if needed. Students may also compare their own actual coin flips
to the computer. When students do smaller numbers of coin flips and compare their
data with a larger number that the computer has generated, they almost always become
aware of the greater accuracy of the larger sample.
This program makes it possible for an instructor to dwell on the increased accuracy
of large sample sizes to provide accurate data approaching the theoretical results one
508 Indiana Academy of Science Vol. 94 (1985)
would anticipate. Without such a laboratory example students often do not fully realize
this relationship between the increased accuracy and the increased sample size. It is
possible with this program to compare relatively large sample sizes for accuracy. In-
dividuals could compare 1000 coin flips with 10,000 coin flips to determine the amount
of increased accuracy with that amount of increase in sample size.
Also, it is often interesting for students to see the total results in the laboratory.
Just the difference between one individual's sample size and the sample size of the
entire class is often enough to be striking.
This is just one simple example of the use of a microcomputer to enhance a
laboratory exercise commonly used in biology. There are without a doubt many other
instances in which computer enhancement would lead to a better understanding of
biological principles. I would like to challenge you to develop computer software that
works for you, and then tell the rest of us.
Literature Cited
1. Baker, Jeffrey, J.W. and Garland E. Allen. 1982. The study of biology. Fourth
edition. Addison-Wesley Publishing Co. Reading, MA. 971 p.
2. Curtis, Helena and N. Sue Barnes. 1981. Invitation to biology. Worth Publishers.
New York. 696 p.
3. Purves, William K. and Gordon H. Orians. 1983. Life: the science of biology.
Sinauer Associates. Boston. 1182 p.
Two-year College Biology Instructors' Perceptions
about their Role Expectations
Lawrence Scharmann* and Harold Harty
Science Education Program Area
Indiana University
Bloomington, Indiana 47405
Perspective
Two-year colleges, during their early years, were in search of professional identity
and recognition. While striving to appease both scientific and educational facets of
the higher education establishment, two-year colleges sought to maintain their innovative
instructional image while enhancing their professional status. To attempt this, they
raised their standards and applied more rigor for hiring faculty by requiring (in most
disciplines) a masters degree in the subject field for which the candidate was seeking
employment. Prior to these new standards, they were hiring instructors with a variety
of experiences and a diversity of backgrounds represented by business, industry, former
high school teachers and dissatisfied university professors. Past history (Palinchak,
1973; Monroe, 1976) depicts these early two-year college instructors as being expected
primarily to fulfill a set of role expectations very similar to those of secondary school
instructors. The more recently hired instructors, however, have a better command and
depth in their academic disciplines; they have been hired as subject specialists. Albeit
a discussion of detailed past role expectations for two-year college instructors, have
the new hiring criteria promoted a similar rigor in present expectations? And if so,
are our present graduate training programs adequate to meet both the hiring criteria
as well as the presently perceived role expectations? This study attempts, in part, to
respond to these questions.
Much literature (Rosen, 1976; Edwards, 1977; Chiapetta and Collette, 1978; Horak
and Lunetta, 1979) exists regarding the expectations of the secondary school biology
teacher; however, very little literature has been generated with respect to the expecta-
tions of two-year college science instructors (Cohen and Brawler, 1980; 1983). Butzow
and Quereshi (1978) were concerned about the validty of expectations and noted that
an expectation needs to be defined in terms of its demonstrability by science teachers
or instructors and its ability to be observed. When generating statements of expecta-
tions, it appeared helpful to synthesize broad categories of skills or competency areas
into which specific expectations can be grouped. Simpson and Brown (1977) developed
seven general and fundamental competency areas for science instructors; these were
(1) professional knowledge, (2) knowledge of science, (3) planning skills, (4) evalua-
tion skills, (5) instructional skills, (6) management skills, and (7) human relation skills.
Unfortunately, present hiring criteria only account for a candidate's qualifications in
the area of knowledge of science. The subject specialist's degree program often precludes
skill training in planning, evaluation, instruction, management, or human relations.
Two-year colleges have thus been left to hire new faculty that need to be exten-
sively prepared by way of inservice programs to remediate inadequate preservice univer-
sity graduate program deficiencies. Dean (1970) remarked that most of the prepara-
tion for biology instructors has been through a teaching assistant program whereby
the training had been less than adequate. Bleyer (1979), nearly a decade later, also
reported that university teaching assistant programs for preparing two-year college in-
structors appears to be grossly inadequate where the teachers tend to be either narrow
* Current address: Department of Biology, Indiana University, Indiana, Pennsylvania.
509
510 Indiana Academy of Science Vol. 94 (1985)
subject matter specialists or secondary school-oriented education majors.
When considering adequate or enhanced preparation, Palinchak (1973) and Roueche
(1983) concluded that two-year college instructors need to be trained in programs that
not only provide subject area expertise but also a background in learning theory, pro-
gram planning, curriculum design, instructional strategies and evaluation techniques.
Related to this stance, Roueche and Hurlburt (1968) found that two-year college students
in terms of their overall impression rated educationally-trained biology instructors
significantly higher than scientifically/research-trained biology instructors; students also
claimed that they learned more from educationally-trained biology instructors. The
relative importance of educationally-related role expectations and the ability or inability
of our present graduate training program to meet these needs is being discussed by
both university science educators and biology professors (Dowling and Roland, 1982;
Coleman and Selby, 1983). Discussion is not sufficient, however, to document the
need for enhanced preparatory programs to remediate the perceived deficiencies of
current two-year college instructors.
Methodology
The purpose of this exploratory survey was to examine the perceptions and obser-
vations of practicing two-year college biology instructors about their role expectations,
and to document a testable role state-of-the-art which hopefully might be pursued more
rigorously from a research framework and more indepth from a qualitative perspec-
tive. To collect data, a survey instrument, "Two-Year College Biology Instructors'
Role Expectation Inventory," was developed and validated.
The survey instrument is composed of 15 Likert items which examine two-year
college biology instructors' role expectations. Abbreviated phrasing of the items can
be found on the left hand side of Table 1. The respondents are asked to rate the
items on a scale of 5 (a must), 4 (very desirable), 3 (desirable), 2 (some importance),
and 1 (unimportant). A sixteenth item is also provided for the respondent to write
in a role expectation not included in the survey instrument. In addition, ample space
is provided for respondents to write down, "a final thought" on the improvement
of biology instruction at their institution. Self administration time was field tested
to be about 15 to 20 minutes.
Face validity of the items and content validity of the instrument were established.
Face validity, a measurement of item relevance, was determined by 2 science educators
with the assistance of a community college biology instructor who were involved with
the generation and refinement of the items. During the item refinement process em-
phasis was given to the relevancy of the substance of the items, and the degree to
which the items purported to measure the role expectations. Content validity, a most
basic validation process, to determine representativeness, was assessed for the "Two-
Year College Biology Instructors' Role Expectation Inventory" by a panel of 3 "experts-
judges" who provided reaction, input and evaluation. The judges were 2 regional junior
college and 1 community college professors of biology. The judges were requested
to mark-up, make marginal notes or comments on, and rewrite, eliminate or add items
to a preliminary draft (22 items) of the survey instrument. The validation dimensions
reacted to by the judges were (1) representativeness of the items from a total pool
or universe of items dealing with the role expectations, (2) relevance (how pertinent)
of the items to the need to conduct a survey of this nature, (3) clarity and under-
standing of the items by the target population, and (4) utility/usefulness of the knowledge
production resulting from the collected information. The constructive suggestions of
the judges resulted in discarding some items, rephrasing of some items and eliminating
redundant verbiage. Several items were eliminated because of lack of relevance, perceived
respondent lack of interest, and perceived respondent attention time.
Science Education
511
Table 1. Summary of Findings on Role Expectations of Two-year College Biology
Teachers
____^ Measures
Skill-
Mean
Rank
Competency
Expectations — — _______^
Rating
Order
Category
1.
Possess Secondary Schools Teaching Experience
2.3
14
Instructional
2.
Teach Courses Other than Biology
1.9
15
Knowledge of
Science
3.
Design/Implement Laboratory Exercises Supplemental
to Content Learning
4.1
4-5
Planning
4.
Design a Variety of Experiences that Maximize Student
Learning
4.0
6-7
Instructional
5.
Measure and Evaluate Student
Learning
4.4
2
Evaluation
6.
Be Cognizant of Theories and Principles of
Learning
2.9
12
Professional
Knowledge
7.
Willing/Able to Experiment with and Evaluate Different
Methods of Instruction
3.5
9
Instructional
8.
Strive to Assist Each Student to Achieve Success
4.5
1
Human Relations
9.
Encourage Students to Develop their Own Values Rather
than Imposing Values
3.6
8
Human Relations
10.
Point Out to Students the Implications Biology Has on
Everyday Life
4.2
3
Human Relations
11.
Advise Students Whose Careers Involve Required Study
in the Sciences
4.0
6-7
Human Relations
12.
Demonstrate the Ability to Carry Out Scientific Research
2.6
13
Knowledge of
Science
13.
Keep Abreast of New Scientific Theories and
Discoveries
4.1
4-5
Professional
Knowledge
14.
Work with Other Faculty in an Interdisciplinary Course
of Study
3.3
10
Instructional
15.
Participate in Educational and Community Service
3.1
11
Human Relations
A total of 240 two-year colleges were selected from a possible of 1,169 (21%)
offering general biology using a table of 5,000 random numbers. In order to obtain
a nationwide sample of present biology instructors, a directory of institutions published
by the American Association of Community, Junior and Technical Colleges was used
for selection purposes. The validated instrument was mailed to the 240 institutions
with directions in a cover letter to forward it to at least one biology instructor or
science department chairman who could best represent the following demographic criteria:
(1) at least 3 years of instructional experience; (2) at least a masters degree in either
biology or science education; and (3) should reflect the institution's role expectations.
Because of the high cost factor, no follow-up mailings were conducted, nor were duplicate
responses accepted from any single institution to avoid unrepresentative bias from those
institutions or states.
The instrument was responded to by biology instructors or science department
chairs representing 126 institutions (53 % response rate) geographically located in 38
states. Although the 53% response rate might be questionable in some circles, the respec-
tability of the response rate can be enhanced by its representing an 11% nationwide
sample of all two-year college institutions that offer biology. The expectation statements
have been rank-ordered in Table 1 by the arithmetic mean. The best possible rating would
be a mean value of 5.0, conversely the lowest would be a mean of 1.0. The statements
have been listed in the order they appeared on the survey. The fifteen items have,
in addition, been classified as to skill-competency areas (Simpson and Brown, 1977;
Butzow, 1978; Chiapetta and Collette, 1978; Horak and Lunetta, 1979).
512 Indiana Academy of Science Vol. 94 (1985)
Findings
The 15 expectations found in Table 1 lack a statement of two-year college biology
instructors' content proficiency or effectiveness in teaching the subject matter. Items
of this nature were excluded because practitioners might have construed it as a state-
ment of their lack of expertise; these items were eliminated from the original list of
22 expectations at the recommendations of the judges serving as content validators.
If the respondent considered these two expectations "a must," then they could be
listed and rated under the "write in" expectations. Their absence from the survey in-
strument does not preclude their importance, but implies that they exist inherently
in the other expectations and not as separate entities.
The expectation receiving the highest rating (Table 1) was "striving to assist each
student to achieve success," and running a close second was "measuring and evaluating
student learning." The expectation viewed with the least favor was "teaching courses
other than biology." Another expectation receiving a somewhat low rating was "possess-
ing secondary school teaching experience." There were three frequently occurring "write
in" expectations; these were "exhibiting a dedication to teaching" (N=14; M=4.4),
"participating in faculty governance" (N=9; M =4.3), and "demonstrating total pro-
fessionalism" (N= 11; M=4.1). Many of the write-in's were very similar to or exten-
sions of the listed 15 expectations.
While individual item-analysis, statement by statement, is in itself interesting, this
analysis does not determine the general trend of perceived role expectations deemed
as fundamental competency areas for science instructors (Simpson and Brown, 1977).
Therefore, a "lumped" or grouped-analysis was performed on these identified skill
areas by determining an overall arithmetic mean for all items/statements belonging
to a particular category. When considering the general skill-competency areas, excluding
planning and evaluation which contained only one expectation respectively, human
relations was deemed the most important (M = 3.9). The remaining areas in rank order
were professional knowledge (M = 3.5), instructional (M = 3.3), and knowledge of science
(M = 2.3). However, when bringing the one-item areas of planning and evaluation under
the instructional category, a higher mean of 3.6 surfaces.
The final discussion or "final thought" item allowing for divergent commentary
tended to "converge" on the preparation of two-year college biology instructors as
the major undergirding factor that would most contribute to the improvement of the
biology programs at their respective institutions (N = 54; M = 3.7). These comments,
at times, were very specific, even getting into what courses need to be taken at the
preservice or inservice levels. Some respondents even sketched out a complete preser-
vice course of study and/or inservice course sequence. Many of the respondents also
proposed a dual masters degree program, one in biology and one in science education
with an emphasis on learning theory and evaluation.
Discussion
The "ideal" (Hammons, 1979) two-year college biology instructor probably needs
appropriate training in "broad-based" biology, teaching methods, and the
philosophies/practices of two-year colleges. Learning psychology and evalua-
tion/measurement might be included under teaching methods. In any instance, it appears
that meeting the "ideal" set of role expectations at the two-year college level requires
a more indepth scientific background than programs for secondary school biology in-
structors, yet it also demands a more professional educational background than pro-
grams for university professors. One way of meeting this requirement would be for
two-year colleges to demand a dual or double masters degree. It is simply unrealistic,
however, to suggest this amount of preservice preparation! Some universities presently
Science Education 513
suggest a masters degree in the natural sciences coupled with a few basic educational
training courses in teaching methods, philosophy and evaluation. But this approach
to preparation, as evidenced by the respondents, appears too non-integrated to be useful
in meeting the "ideal" for two reasons: (1) A masters degree in the natural sciences
is too research oriented to allow flexibility in establishing a broad-base of science con-
tent necessary for effective two-year college instruction, and (2) the educational com-
ponent is too heavily oriented toward secondary instruction to be relevant to the two-
year college setting. In other words, simply patching together two unrelated programs
is not an effective solution (Hansen and Rhodes, 1982).
One potential solution would be for universities to continue research for establish-
ment of an integrated independent masters degree in biological education. This degree
would allow candidates for two-year college instructional positions to establish a broad-
base scientific background (without requiring specialized research orientation) while
integrating professional education coursework specifically designed to meet the unique
individual needs of the two-year college setting. Indeed, the results of this survey sug-
gest that the two programs of study need not be mutually exclusive, but mutually
beneficial and complementary.
No attempt is made, however, to delineate a specific program of coursework that
might be more consistent with the perceived role expectations than may presently exist
for two-year college biology instructors. The collected data are limited and respondents
greatly differed on the specific biological science content that should be included in
an "ideal" program.
Implications for Practice
Two questions were stated in the opening paragraph that have a direct bearing
on both the findings of this study as well as the implications for practice. First, stan-
dardization of hiring criteria has allowed two-year college administrators the oppor-
tunity to expect a minimal level of competence in biological science content attained
by prospective faculty candidates. This unfortunately, precludes the expectations of
instructional capability, course development skills, and evaluate competencies deemed
necessary by the population of practicing two-year college biology instructors. The
results of this study suggest, therefore, that while both hiring criteria and role expecta-
tions have become more rigorous, these two factors are inconsistent with one another.
Second, the discrepent inconsistency of these factors is indirectly attributable to the
inadequacy of graduate training programs to supply the appropriate experiences re-
quired by a two-year college faculty candidate's opportunity to become proficient in
both biological content and instructional competence.
In terms of recommendations for change, the findings of this study speak to both
prospective faculty candidates and administrative practitioners. First, prospective two-
year college biology instructors need to ensure that their program of study allow suffi-
cient flexibility and/or opportunities to gain experience in the application of biological
content through instructional techniques and practice. This might be accomplished as
part of a cooperative internship program between two-year colleges and university
graduate schools. Second, two-year college administrators need to consider a prospec-
tive faculty candidate in terms of demonstrable instructional competence and not simply
in terms of content proficiency. If an internship program is not feasible, then renewed
cooperation between two-year colleges and university graduate institutions needs to
occur in order to establish a degree program more suited to the needs of the two-year
college setting. In summary, perhaps alternatives to the MS/Ph.D. degree structures
need to be reexamined (Hansen and Rhodes, 1982).
514 Indiana Academy of Science Vol. 94 (1985)
Suggestions for Further Study
Based on the quantitative and qualitative findings of this survey effort, the following
hypotheses have been generated for testing in future inquiry endeavors:
Two-year college biology instructors see themselves as teachers rather than as
scientists.
A primary self-assessed role expectation is being able to relate to students as
individuals.
The development of biological applications and human values takes precedent
over student content mastery.
The individualizing of instruction in two-year college biology is made available
through evaluative techniques, various teaching methods, and a variety of learn-
ing experiences.
Two-year college biology instructors view scientific research as a relatively low
priority expectation.
Two-year college biology instructors are not expected to teach courses other than
biology
Experience as a secondary school science teacher is not deemed a necessary prere-
quisite for successful two-year college biology teaching.
Knowledge of theories and principles of learning are low priority expectations
for two-year college biology instructors.
Literature Cited
1. Bleyer, D. Higher education's omission: The preparation of community college
teachers. Community College Review, 1979, 6, 46-51.
2. Butzow, J.W., and Quereshi, Z. Science teachers' competencies: A practical ap-
proach. Science Education, 1978, 62, 59-66.
3. Chiappetta, E., and Collette, A. Secondary science teacher skills identified by
secondary science supervisors. Science Education 1978, 62, 67-71.
4. Cohen, A.M., and Brawler, F.B. New directions for community colleges (no.
41): evaluating faculty and staff. San Francisco, California: Jossey-Bass, Inc.,
Publishers, 1983.
5. Cohen, A.M., and Brawler, F.B. New directions for community colleges (no.
31): teaching the sciences. San Franciso, California: Jossey-Bass, Inc., Publishers.
1980.
6. Coleman, W.T., and Selby, C.C. Educating Americans for the 21st century.
Washington, DC: The National Science Board Commission on Precollege Educa-
tion in Mathematics, Science, and Technology, 1983.
7. Dean, D.S. Pre-service preparation of college biology teachers: A search for a
better way. Washington, D.C.: Commission on Undergraduate Education in the
Biological Sciences, American Association for the Advancement of Science, 1970.
8. Dowling, N.G., and Roland, H. Institutional and faculty life cycle changes. Com-
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10. Hammons, J.O. The multi-faceted role of an 'ideal' community college faculty
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to the Ph.D. Community College Review, 1982, 10, 52-58.
Science Education 515
12. Horak, W., and Lunetta, V. Science teacher types: A study of beliefs about the
importance of specific teaching behaviors. Journal of Research in Science Teaching,
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13. Monroe, C.R. Profile of the community college. San Francisco, California: Jossey-
Bass Publishers, 1976.
14. Palinchak, R. Evolution of the community college. Metuchen, New Jersey:
Scarecrow Press, Inc., 1973.
15. Rosen, S. Can science education mass produce superteachers? Science Education,
1976, 60, 53-60.
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College Research Review, 1968, 2, 1-3.
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SOIL AND ATMOSPHERIC SCIENCES
Chairperson: David R. Smith
Department of Geosciences
Purdue University
West Lafayette, Indiana 47907
(317) 494-3285
Chairperson-Elect: Charles L. Rhykerd
Department of Agronomy
Purdue University
West Lafayette, Indiana 47907
(317)494-8101
ABSTRACTS
Land Cover Classification of Rupgang Thana Dhaka, Bangladesh Using Landsat MSS
Data M.F. Baumgardner, N.N. Chaudhuri and S.J. Kristof, Purdue University,
West Lafayette, Indiana 47907. In Bangladesh, the Rupgang Thana study area
is located along the shore of the Lakhya River. It has a tropical monsoon climate.
The soils are derived mainly from alluvial deposits. During the monsoon rain season
most of the area is inundated. The major food crops are rice, wheat, and mustard;
the cash crops include jute, tea, sugar cane and tobacco. Bamboo, jackfruit, mangoes
and bananas are commonly grown on small areas in and around the villages.
The land cover of the study area was first spectrally classified by computer-
implemented pattern recognition techniques using Landsat multispectral scanner (MSS)
data collected on 3 January 1977 and 2 February 1980.
Ten spectral classes from the data of January 1977 and 12 classes from the February
1980 data were displayed in color-coded digital format on a digital image display. Dif-
ferent colors were assigned to each class. Grouping of classes was accomplished to
avoid color discrimination difficulties. The area is relatively flat, and the fields are
generally small, seldom larger than two or three hectares. Classification results of Landsat
MSS data with a resolution of 60 x 80 m (< 0.5 Ha) were limited in delineating in-
dividual fields, but were successful in delineating soils with slight differences in eleva-
tion and internal drainages. The better drained soils are highly correlated with the
cultivation of wheat and mustard; the poorly drained soils with rice.
Air Temperature Fluctuation in Alabama During the Annual Solar Eclipse on 30 May
1984. William R. Gommel, Douglas W. Poad and John W. Wicker, Department
of Earth-Space Sciences, Indiana Central University, Indianapolis, Indiana
46227. Using a sling psychrometer 7 statute miles north of Prattville, Alabama,
in front of the Robert Murphy residence (which is also 3/4 mile south of Pine Level)
on the east frontage of U.S. Highway 31, a reduction in air temperature of 5.0°F
was observed from first contact (61.1°F at 9:49 A.M. or 14:49 GMT) to annularity
(56.1°F at 11:17 CDT). The observations which were taken on the eclipse centerline
registered a somewhat smaller wet-bulb temperature decrease of 2.8°F from 49.7°F
to 46.9°F during the same period, and relative humidity increased from 42% to 48%.
By the end of the partial phases at fourth contact (12:52 P.M. CDT), the air temperature
had increased to 62.7°F, and relative humidity had decreased to 42% once again — the
same value recorded at the beginning of the partial phases.
517
518 Indiana Academy of Science Vol. 94 (1985)
The sky was clear, and horizontal visibilities were greater than 10 miles throughout
the eclipse. A brisk northerly surface wind of 15-20 knots during the early phase diminish-
ed dramatically by annularity. Electric lights suddenly lighted although it was near
mid-day, and an eerie quiet briefly fell over the earth during the awe-inspiring event.
By noon CDT, winds were increasing in velocity and once again reached earlier brisk
speeds by 12:30 P.M. when the temperature recovered to 61.6°F — the same value recorded
at first contact.
On the centerline, during this eclipse, the meteorological elements changed similarly
(but with less magnitude) to changes observed throughout the night under the synoptic
conditions. In other parts of the United States (including Indiana) where only partial
phases occurred, the observed changes were not as dramatic.
Engineering Properties of Indiana Peats and Mucks. Paul Joseph and C.W. Lovell,
School of Civil Engineering, Purdue University, West Lafayette, Indiana 47907. A
study is currently under way at Purdue to investigate the problems associated with
the use of peats and mucks as foundation materials. These materials pose problems
to the foundation engineer because of their low shear strength, high compressibility,
and time dependent behavior. The previous construction method was to excavate the
peat or muck, and replace it with a better foundation material. Nowadays, however,
due to ecological and financial reasons, this method is no longer popular. An alter-
native is to construct on the peat or muck itself. However, there appears to be some
confusion as to the definition and classification of peats and mucks. Further, the un-
disturbed sampling and testing of these materials has proved difficult, because of their
low strength and high water contents. The high variability of these materials also causes
problems.
This paper looks into the definition of peat and muck and suggests a few tests
that can be conducted in a highways research laboratory for accurate classification
purposes. An undisturbed sampling technique and procedures for undisturbed testing
are also presented.
The consolidation of peat is highly complex, in view of the large strains and time
dependent deformations. Further this creep is not linear in most cases. The results
of various tests on a typical Indiana peat and also a muck are presented.
Characterization of Indiana Soils by Porosimetry. C.W. Lovell, School of Civil
Engineering, Purdue University, West Lafayette, Indiana 47907. All engineering
characteristics and properties of soils and rocks are influenced by the distributions
and arrangements of solids within the mass, i.e., the fabric. While direct measurement
and quantification of the fabric is extremely complicated, the size distribution of the
pores between the solids can be quickly determined. The common technique is mer-
cury intrusion porosimetry, wherein the relationship between the pressure on the mer-
cury and the pore size are intruded by incrementally increasing the pressure on the
mercury and measuring the quantity intruded. Results are expressed in terms of either
cumulative or differential frequency distributions of pore sizes.
These distributions have been simply correlated to a number of behavioral responses
of soils (both sandy and clayey) and rocks. The most useful correlations are those
involving permeability. It is believed that the porosimetry technique has many applica-
tions in understanding and predicting the behavior of earthen materials.
Survey of the Mineral Composition of Forage Crops in Portugal. C.L. Rhykerd, S.E.
Fowler, Afonso de Almeida, A.M. Ferreira, Nuno Moreira, C.H. Noller and
J.L. Ahlrichs, Departments of Agronomy and Animal Sciences, Purdue University,
Soil and Atmospheric Sciences 519
West Lafayette, Indiana 47907, 1 .U .T. A. D. , Vila Real, Portugal and University of Evora,
Evora, Portugal. Limited data are presently available concerning the mineral com-
position of Portuguese forage crops. The following experiment was conducted to deter-
mine the mineral concentrations in forage samples collected in Portugal from university-
conducted experiments. Three major studies were made representing oats and vetch,
corn, and common forage crops, with each addressing the following objectives:
1) to determine the mineral composition of the forage samples
2) to relate the mineral value to sufficiency levels for the various forage species
3) to compare the mineral values to mineral nutritional requirements of beef cattle,
dairy cattle, and sheep
4) to evaluate the quality of the forage and make recommendations relative to
its (nutritional) improvement.
Mineral analysis by means of an emission spectrograph of forage samples from
Portugal revealed many low concentrations within the small grain crops, forage grasses,
and forage legumes, when harvested as forage.
Data obtained in this study emphasize the importance of legumes to Portuguese
livestock farmers. Legumes, if properly inoculated, are able to utilize atmospheric N
and thereby eliminate the need for N fertilizer. In addition, legumes are high in pro-
tein and contain considerably higher concentrations of certain minerals, especially Ca
and Mg, which are essential to proper mineral nutrition of growing and lactating rumi-
nant animals.
Soils an Important Component in a Digital Geographic Information System. C.R. Valen-
zuela, T.L. Phillips, M.F. Baumgardner, and L.A. Bartolucci, Purdue Univer-
sity, West Lafayette, Indiana 47907. There is an increasing use of Digital Geographic
Information Systems to meet the demand for specific, accurate and rapid information
of our resources. The degree of usefulness of this information depends on the accessibility
and efficiency of the methods utilized for input, storage, analysis and retrieval of
information.
The demand for accurate and rapid soil information is growing in our modern
society, thus the element soil, because of its importance, is a basic component in a
Geographic Information System.
The Indiana Soil Associations Map at a scale of 1:500,000, was manually digitized,
projected to the Albers equal-area map projection, rasterized, and stored in a Geo-
referenced Data Base created for the State of Indiana.
Using the digital soils data stored in the Geo-referenced Data Base, new sets of
data were generated by changing the coding of the soils associations or by combining
two or more of these new generated products. Among the new digital data generated
from the soils data are: Prime Agricultural Lands, General Slope Information and
Potential Erosion Data.
In addition, mapping and inventory errors were investigated in relation to the
cellularization of spatially distributed soils data set, and an attempt was made to define
an appropriate cell size, so that the mapping and inventory errors will be within the
cartographic standards.
Wet Atmospheric Deposition in Indiana
J. A. Andresen, W.W. McFee, J.L. Ahlrichs
Department of Agronomy
Purdue University, West Lafayette, Indiana 47907
and
K.T. PawU
Department of Land, Air and Water Resources
University of California-Davis
Davis, California 95616
Introduction
Atmospheric deposition of materials by precipitation has become an increasing
area of concern. Although the role of this deposition in the environment and its sources
are still largely uncertain (Miller, 1984), it is thought that precipitation chemistry,
especially acidity, can be highly influenced by anthropogenic emissions of sulfur and
nitrous oxides (Cogbill and Likens, 1974, Likens, 1976, Martin and Barber, 1977).
To monitor atmospheric deposition, several networks have been developed over
N. America and Europe. Beginning in July, 1982, weekly precipitation samples were
collected in West Lafayette, IN as a part of the National Atmospheric Deposition Pro-
gram Network (NADP), organized under the USDA Cooperative State Research Ser-
vice Interregional project IR-7. The network was created in 1978 and currently has
a total of 177 stations across the U.S. The network has excellent common procedures
for field work, standardized equipment at each site, and has all samples analyzed at
a central laboratory location, minimizing procedural biases and enhancing the capability
for regional and temporal comparisons.
Samples at the West Lafayette sites are taken weekly at the Purdue Agronomy
Farm (1 1 km NW). On-site measurements of precipitation pH and specific conductivity
are performed on a small aliquot. The remaining sample is shipped in its collection
bucket to the NADP Central Laboratory for a detailed laboratory analysis including
pH and conductivity, as well as soluble concentrations of Ca, Mg, K, Na, NH4, N03,
CI, S04, and P04 ions. In addition, detailed on-site precipitation records are main-
tained, giving a thorough history of rainfall duration as well as collector performance.
On-site pH and conductivity measurements were made only when the total precipita-
tion sample weight exceeded 70g (1.03mm liquid precipitation equivalent), otherwise
the entire sample is shipped to the Central laboratory.
Results and Discussion
The field pH measurements taken at the Agronomy Farm field site are shown
in Figure 1 for the entire two-year period ending 10 July, 1984. Eighty-eight weekly,
on site, observations were taken (out of 104 possible). In the remaining weeks there
was either no precipitation or the volume was less than 70 ml. The data points and
a moving three-week average line are shown for comparison. The scatter of the points
is relatively high with a range from 3.21 to 4.93 pH hunits. Also, a definite periodicity
can be seen these first two years, with pH maxima occurring in the winter months
and pH minima occurring in the summer. The annual mean pH based on weekly,
on-site measurements is 4.15.
The Central Laboratory data yielded similar results. At the time of writing this
presentation, laboratory analyses were available up through the end of May, 1984 (98
weeks), on 86 samples from weeks with sufficient precipitation for analysis. The mean,
volume-weighted pH for the laboratory measured samples was 4.36. Figures 2a and
521
522
Indiana Academy of Science
Vol. 94 (1985)
CD
Q.
"o3-° | ) / i ! i i ; i i i i i i i i | ii i i \ i i
2 JUL JAN JUL JAN JUL
1982
!
1983
Date
1984
Figure 1 . On-site precipitation pH measurements at West Lafayette, In for the two-
year period 13 July, 1982 - 10 July, 1984.
2b show the S04 and N03 deposition vs. time for the 98-week period. The S04 ion
also shows some periodicity, but approximately 6 months out of phase with the pH
plot, with summer maxima and winter minima. This would be "in phase" with an
H+ ion concentration plot. This seasonality (especially pH has been observed in other
parts of the NADP network (Semonin and Stensland, 1984) and may be related to
T — i — i — i — i — p — i — i — i — n — i — i — i — i — i — i f i — i — r-i — r
JUL JAN JUL JAN JUL
1982 1983 1984
Date
Z JUL JAN JUL JAN JUL
1982 1983 1984
Date
Figure 2. a) Sulfate and b) Nitrate deposition at West Lafayette, In for the 98-week
period 13 July, 1982 - 1 June, 1984.
Soil and Atmospheric Sciences
523
temperature. The NO, ion, on the other hand, shows less seasonality and appears less
random. NADP Network results for mean pH and S04 ion concentration in the sum-
mer and winter of the 1982 are shown in Figures 3 and 4. Again, the seasonal periodicity
4.4
Summer 1982 Volume Weighted Mean pH
Winter 1982 Volume Weighted Mean pH
Figure 3. NADP Network results for mean precipitation pH in a) summer and b)
winter, 1982 (NADP, 1984).
524
Indiana Academy of Science
Vol. 94 (1985)
Summer 1982 Sulfate Ion Concentration(mg/l)
1.0
Winter 1982 Sulfate Ion Concentration(mg/l)
Figure 4. NADP Network results for mean sulfate ion deposition in a) summer and
b) winter, 1982 (NADP, 1984).
of both data sets is quite distinct. Indiana lies on the western fringes of the area of
most acidic rainfall and highest S04 deposition with extends north and eastward into
the northeastern U.S. and southeastern Canada.
Soil and Atmospheric Sciences 525
Statistical Data Analysis
The laboratory results were analyzed with stepwise multivariate linear regression
with computer programs from the Statistical Package for the Social Sciences (SPSS8.5.3)
at the Purdue Univ. Computing Center. The initial results (with 86 observations) showed
great variation in all variables. Careful scrutiny of the data revealed that most of the
large concentrations of ions were obtained in small precipitation volumes. Martin and
Stensland (1984) have suggested sample contamination may be a problem in very small
samples in large containers due to chemical reactions with the material of the collec-
ting bucket itself during the storage time before lab analysis. The enhanced scavenging
effect of light showers would also contribute to high concentration in the small samples.
When data from sample volumes of less than 70 ml were removed, reducing the number
of observations to 76, the analysis gave more plausible results and drastically reduced
the standard deviation of the ion concentrations, in some cases by as much as 480%.
The correlation coefficients for various ions and processes with H ion concentra-
tion are shown in Table 1. The listed ions were chosen on the basis of concentrations
because the other measured ions were at least an order of magnitude less and were
frequently below the detection limits of the instruments. The hydrogen ion concentra-
tion was estimated using the relationship:
H = 10 exp(-pHlab).
Weekly mean temperature (TEMP) at the site was included as a variable in hopes
of seeing the effects of temperature-dependent oxidation on sulfur and nitrogen oxides.
Precipitation volume (VOL) and the difference of summed anions and summed cations
(IONDIF, in meq/1) were also used in the correlations. The IONDIF should be closely
correlated with H if no organic ions are in the system and all important inorganic
ions are being measured.
Table 1 . Correlation coefficients for ions and several processes vs. H ion concentrations.
H
so4
NO,
HN,
Ca
VOL
PHlab
TEMP
IONDIF
.596
.864
.582
.214
.205
-.216
-.718
.350
TEMP
.529
.541
.160
.243
.416
.024
.585
PHlab
-.857
-.686
-.550
-.187
-.412
.235
VOL
-.250
-.242
-.450
-.187
-.389
Ca
.510
.519
.547
.359
NH4
.084
.513
.588
NO,
.467
.575
so4
.604
The correlation coefficients show S04 to be significantly (a = .05) correlated
N03, Ca, NH4, as well as being the best predictor of H, while nitrate shows significant
correlation with S04, Ca, NH4. This is not surprising as much research has found
these two anions to be dominant in precipitation chemistry (Likens et al., 1979, Gran-
nat, 1972) and a major source of its acidity (Baker et al., 1981). The higher correla-
tions of S04 and N03 with the difference in summed anions and cations also support
this. The positive correlation between S04 and temperature and the increased summer
sulfate levels found at the site have been observed elsewhere (Musold and Lindqvist,
1983).
If the source of all S04 and N03 ions were S02 and nitrous oxides, then every
526 Indiana Academy of Science Vol. 94 (1985)
equivalent of N03 and SO„ should have an equivalent of H originally associated with
it, and as suggested by Skeffington (1984), the slope of a linear regression between
H and the oxidized form should be very close to 1. Our data yielded regressions of
H = 1.09 S04 - .004 for S04 ion in meq/1 and H = .180 N03 + .006 for NO,
ions in meq/1. The intercepts of both equations are near the origin, and in the case
of S04, the slope is not significantly different from 1.0, which should be the case
if S02 is the predominant source of S04. For N03, however, the slope is much less
than unity, and we must infer that the source of much of the N03 is not from nitrous
oxides, but from other sources. This makes sense as the site is relatively distant from
any large point sources.
A surprising result was the intercorrelation of Ca with the other ions. We expected
a negative or insignificant relation with H + , as most terrestrial dust contains Ca in
the form of CaC03, causing neutralization (Harrison and Pio, 1983). We found in-
stead a significant positive correlation. This suggests a more neutral salt of sulfate
or nitrate as the predominant Ca form. The acidic contribution (if any), however,
is not apparent.
As expected, Volume shows a negative correlation with all ion concentrations,
implying that the majority of the aerosols are scavenged during the initial phases of
precipitation events.
Overall, if we use a multiple linear model to regress the data stepwise with H
ion concentration, we obtain:
H = .013 S04 - .110 NH4 + .025 N03 + .0014 TEMP + .027 Ca - .057
where all concentrations are in mg/1 and temperature is in °F. Rejection in the step-
wise method was at the a = .05 level. The model yields an R2 value of .78, with
S04 explaining 36% of the H variance.
Impact on Indiana Soils
If we integrate the major ions (by mass), including S04, N03, and NH4, we get
average annual depositions of 8.43 kg S/ha and 6.18 kg N/ha. This corresponds well
with the 8.0 kg S/ha deposited at Champaign-Urbana, IL in 1982. The S value may
be a bit low as the averaging period did not include the last six weeks of the two-year
period, when concentrations are usually higher than the average. The acidifying effect
of these ions in precipitation is believed to have little, if any, impact on most Indiana
soils due to the large buffering capacity present (CAST, 1984). While the N deposition
represents a small fraction of total N used in agriculture, the S deposition may supply
a significant portion of most agricultural needs. Most crops in Indiana remove
approximately 18 kg S/ha annually (Terman, 1978).
Conclusions
Atmospheric deposition through precipitation has been monitored since the sum-
mer of 1982 at the Purdue Agronomy Farm as a part of the NADP network. The
mean, volume-weighted precipitation pH is 4.36, while annual S deposition in the form
of soluble S04 averaged 8.43 kg/ha, enough to meet some of the needs of growing
crops in Indiana thus reducing the need for S fertilizers. Nitrogen deposition in the
form of NH4 and N03 is on the same order of magnitude, 6.18 kg N/ha, but represents
a much smaller portion of the total plant needs.
The H and S04 ion concentrations show a definite seasonal trend, higher in the
summer and lower in the winter. Multiple correlations indicate that Ca, S04 and N03
ion concentrations in precipitation are significantly correlated with H concentrations.
Soil and Atmospheric Sciences 527
The concentrations and total deposition at this station fit the regional pattern por-
trayed by other network stations throughout the Midwest.
Literature Cited
1. Baker, M.B., D. Caniparoli and H. Harrison. 1981. An analysis of the first year
of MAP3S rain chemistry measurements. Atmos. Envir. 15:43-55.
2. CAST. 1984. Acid precipitation in relation to agriculture, forestry, and aquatic
biology. Council for Agricultural Science and Technology Report 100. Ames, Iowa.
3. Cogbill, C.V. and G.E. Likens. 1974. Acid precipitation in the northeastern United
States. Water Resour. Res. 10:1133-1137.
4. Grannat, L. 1972. On the relation between pH and the chemical composition
in atmospheric precipitation. Tellus 24:550-560.
5. Harrison, R.M. and C.A. Pio. 1983. Size differentiated composition of inorganic
atmospheric aerosols of both marine and polluted continental origin. Atmos. En-
vir. 17:1733-1738.
6. Likens, G.E. 1976. Acid Precipitation. Chem. Eng. News 54:29-44.
7. Likens, G.E., R.F. Wright, J.N. Galloway and T.J. Butler. 1979. Acidic Rain.
Scient. Am. 241:43-51.
8. Martin, A. and R.F. Barber. 1977. Some observations on acidity and sulphur
in rain water from rural sites in central England and Wales. Atmos. Envir.
12:1481-1487.
9. Martin, W. and G. Stensland. 1984. Personal communication.
10. Miller, J.M. 1984. Acid Rain. Weatherwise 37:233-239.
11. Musold, G. and O. Lindqvist. 1983. Correlations between meteorological data
and water-soluble sulphur compounds in fine aerosols. Atmos. Envir. 17:1253-1260.
12. N.A.D.P. 1984. Annual summary of precipitation chemistry for 1972. National
Atmospheric Deposition Program, Nat. Res. Lab., Colorado St. Univ., Ft. Col-
lins, Co.
13. Semonin, R.G. and G.J. Stensland. 1984. Acid Rain Trends? Weatherwise
37:250-251.
14. Skeffington, R.A. 1984. The chemistry of bulk precipitation at a site in southeast
England-II. Relationships between ions and comparison with other sites. Atmos.
Envir. 18:1695-1704.
15. Terman, G.L. 1978. Atmospheric sulphur — the agronomic aspects. Tech. Bui.
23, The Sulphur Institute. 15 p.
The National Weather Service Rainfall
Data Collection Network in Indiana
John T. Curran, Albert P. Shipe and Edward C. Yess
National Weather Service Forecast Office
Indianapolis International Airport, Indianapolis, Indiana 46251
The primary mission of the National Weather Service is to provide severe weather
warnings and flood warnings to the public. To accomplish the flood warning part
of this mission, staff members at the Indianapolis weather office collect daily reports
of precipitation measurements taken across Indiana by the official hydrologic network.
These reports represent a large part of the official National Weather Service cooperative
network in Indiana.
There are nearly 180 official National Weather Service cooperative stations in
the state. But because of monetary constraints, only about 10 percent of these observers
are asked to report on a daily basis. However, when rainfall amounts exceed one half
inch, then nearly 60 percent of the network observers report. In the Central Region,
of which Indiana is a part, it is estimated that the present average network efficiency
is only 30 to 50 percent.
Various sources in the climatological literature suggest that one observing site
for every 50 square miles is necessary to accurately describe the spatial variation of
rainfall. This is especially important where convective precipitation contributes one
third or more of the annual precipitation total. It should be obvious that with an
average of one official observer in each 205 square miles, the reporting network in
Indiana falls far short of this goal. Around 750 stations would be needed just in our
state to approximate the one observation per fifty square miles criteria.
One example of how the lack of reporting observers in the official network has
sometimes failed the public to some degree occurred in June of 1980. On this occa-
sion, data collected from the official network was sufficient to accurately forecast some
flooding along the White River in Central Indiana. The network reports also enabled
Weather Service forecasters to issue flood statements for tributaries and small streams
in Kokomo and some other areas of Central Indiana. However, when television coverage
of the resulting flood in Kokomo was reviewed, it became obvious that the predictions
for small stream flooding were too low because of grossly inadequate rainfall data.
There are always ongoing plans and goals within the Weather Service to upgrade
the present hydrologic networks. Unfortunately, this sort of change seems to take many
years and millions of dollars and these funds are rarely available. So, to try to prevent
a repeat of the Kokomo flood situation, the hydrologic unit staff at the Indianapolis
Weather Office appealed to the Indiana Amateur Radio Operators who have always
been a public spirited group. They responded in the way that the nation has come
to expect and went to work to solve the problem.
In October 1980, a small group of 40 Amateur Radio Operators organized by
Mr. Herb Clark, began collecting rainfall reports on a daily basis; forwarding them
to the Indianapolis Weather Office. Mr. Ray Fullman of Brownsburg became the local
relay for these reports. The individual check-ins were then compiled by Mr. Fullman
and called into the Indianapolis Weather Office by telephone. Under Mr. Clark's and
Mr. Fullman's guidance and determination, the Indiana "WETNET", as the network
is now called, quickly grew to 60 daily reports, then to 100 reports, and finally to
the current number of about 150. The Indianapolis Amateur Radio Network is now
approximately the same size as the official Weather Service Hydrologic Network.
Results of the Amateur Radio Rainfall Reporting Program have met or exceeded
529
530 Indiana Academy of Science Vol. 94 (1985)
all expectations. Since the Network's inception, the addition of this data has helped
the isohyetal analyses of every major rainfall or snowfall that has occurred in Indiana.
For example, in 1981, forecast of flooding along the Tippecanoe River was possible
only because of Amateur Radio Reports. Many times, excessive rainfall from isolated
thunderstorms has been verified only through these reports. However, our goal of
at least one Amateur Radio rainfall report per Indiana county has yet to be realized.
The project has not been without difficulties. Because of the time involved in
the collection and processing of so many reports by hand, the Amateur Radio operators
seemed to reach their limit with 150 daily reports. While this number has effectively
doubled the precipitation network that existed in the June 1980 flood event, it still
leaves the state far short of that optimum spacing of one observation in each fifty
square miles. Large gaps remain in the network. Some counties are still without an
observer that reports on a daily basis. Many more reports are needed but the collec-
tion and relay of these numbers of reports has created a bottleneck.
Fortunately, the home computer has appeared in the "shack" of many Amateur
Radio operators during the past several years and this will apparently provide an answer
to the data collection problem. Mr. Tom Bowen, Amateur Radio Operator and manager
of the Indiana "WETNET" has written several programs for the Commodore 64 com-
puter to monitor and reformat the reports into a message that can be used by National
Weather Service computers. Test transmissions have proven successful and further
refinements are planned in the last two months of 1984. As we go "on line" with
the data transfer computer to computer, an almost unlimited number of reports may
be processed and the existing bottleneck eliminated.
The Amateur Radio Precipitation Network is far from the precise official net-
work that has been established and is maintained by the National Weather Service.
Very few private citizens can afford a standard rain gage that cost over $100 or a
recording rain gage that can cost in excess of $3500. Although Weather Service per-
sonnel instruct the Amateur Radio operators in the proper exposure requirements for
rain gages, no staff member is able to visit and inspect all the sites. Many of the
observers use the inexpensive 6 inch Tru Check Wedge Plastic Gage that is available
in many hardware stores. A few have the more expensive 11 inch plastic gage that
has been certified by the National Weather Service for use. Still, these reports are
quite legitimate when closely compared with those of the official network.
Another difficulty is turnover of participants in the Net. When an observer quits
for any reason, there is no guarantee that a replacement will be found at or near the
previous location. During the past year, the rate of observer turnover has been nearly
15 percent.
In spite of these and some other disadvantages, the usefulness of the data received
makes the network worthwhile to maintain. Not only does the National Weather Ser-
vice use the information on a real time basis, but the Agricultural Center at Purdue
University as well as State and other Federal Agencies use the data for maintaining
climatology records, for research, and water level maintenance to name a few examples.
Local electronic archiving of the rainfall data has not been possible to date. Machine
manipulation of the data also cannot be done at the Weather Service Forecast Office
level as yet. However, we feel that this information could be of significant value in
mesoscale analysis for research and other purposes if placed in the right hands. The
National Weather Service will certainly cooperate with any such requests.
The future holds promise for an increasingly efficient and acceptable hydrologic
network. As expertise grows in the use of satellite imagery to estimate rainfall amounts
and correlation of both, Doppler and conventional radar observations to rainfall is
better understood, these networks may become redundant. Full automation of data
Soil and Atmospheric Sciences 531
collection and processing is certainly in the future, but until this becomes a reality,
we will continue to rely on the Amateur Radio Network and other hydrologic net-
works to aid in warning the public of excessive rainfall events and any resulting floods.
Soil Survey in Indiana: Past, Present and Future
D.P. Franzmeier, H.M. Galloway and J.E. Yahner
Department of Agronomy
Purdue University
West Lafayette, Indiana 47907
Soil survey activity in Indiana began in 1902 with publication of the map and
report for Posey County and continues to the most recent report, for Orange County,
in 1984. An earlier review (18) traced development of this program from 1902 through
1976. Surveys, made originally to help agriculturalists extrapolate results of field
experiments from the plots to other locales, have adapted to the changing needs in
successive periods. Since World War II rapid land use changes in housing, industry,
retail marketing and recreational developments and the expansion of agricultural and
forestry operations as well as environmental concerns of a rapidly expanding popula-
tion have all stimulated interest in using soils surveys. Equitable farm land taxation
has been another stimulus. Land qualities which soil surveys describe has made them
of inestimable value to Indiana citizens.
This paper reviews important historic phases of soil survey progress during the
past 82 years. The first phase spanned 57 years and included growth of knowledge
about soils and improvement in survey techniques. A total of 64 surveys were published,
all having colored line maps. These surveys will eventually be replaced by newer ones.
In the second phase, which spanned 24 years in the decades of the 1960s, 70s and
80s, the survey program expanded dramatically and surveys were published, or will
be published, for each county of the state. These surveys have detailed maps published
on air-photo base maps and greatly expanded reports tailored to the needs of users
in a greatly expanded audience for soil information. The third phase, beginning now,
will emphasize use of the surveys.
Phase 1. Early Growth of Indiana's Soil Survey: 1902-1959
Early soil mapping was done by the Bureau of Soils and later by the Bureau
of Chemistry and Soils of the U.S. Department of Agriculture (USDA) cooperatively
at first with the State of Indiana Department of Geology and after 1920 with the Pur-
due University Agricultural Experiment Station (AES). Later, progress continued with
Bureau of Plant Industry, Soils and Agricultural Engineering of USDA cooperating
with AES.
Field mapping was done early on plane tables at a scale of 1 inch per mile and
later 2 inches per mile. Many of the earlier workers had more training in geology
than in soils. Little was known about soil morphology but the geologic nature of soil
parent materials was better understood. Reports included only general observations
of crop growth and cropping systems followed on various soils and little reference
was made to research on crop production or how it could be adapted locally. Reports
improved greatly after 1922 when soil management chapters written by Purdue AES
workers were added to survey reports.
Two Important Advances
In 1922 Thomas M. Bushnell began his long career as head of the soil survey
activities for AES at Purdue University. His early work with the Bureau of Soils in
Lake County and elsewhere and with aerial observation and photographs in World
War I helped him to later pioneer the use of air-photos in soil mapping, which had
a very lasting effect on later survey program development.
533
534 Indiana Academy of Science Vol. 94 (1985)
Bushnell obtained air photos from the Army and from highway engineers for
experimental use in early surveys (2). He later ordered a complete set for Jennings
County and used these as field base maps for the soil survey. Jennings was the first
county to be mapped entirely on air photos (3). In 1936 he summarized the influence
of photos on the soil survey program (4). Air photos allowed better visualization of
important landscape features by soil mappers and aided them to see greater detail and
locate the observed soil features more accurately in relation to buildings, roads, streams,
cropland, pasture and wooded area boundaries.
As knowledge about soil-landscape relations increased, Bushnell and his colleagues
began to appreciate the "catena" concepts used by Milne (30) in Africa to explain
the nature of groups of soils formed from similar parent materials. He applied "catena"
ideas to Indiana soil relationships to aid development of the taxonomy of Indiana
soils in 1938 (5) and in 1939 (6) and 1942 (7). He further refined the system and put
it into a key form in the Story of Indiana Soils in 1944 (8), a work termed "a classic
of organization" by Kohnke et al. (28).
Early Improvements Set Stage for Better Surveys Later
With increasing knowledge of soils, greater accuracy in mapping with air-photos,
and increasing studies from which to draw interpretations of soil behavior under dif-
ferent uses, a slow revolution occurred in the soil survey program. Soil mapping was
done in more detail and reports became more comprehensive so that county survey
publication time increased from 1 or 2 years in early surveys to 5 years by 1930, and
to 6 years by 1940 (18).
Emphasis on making soil surveys for farm planning increased as soil conserva-
tion programs got under way with the Soil Conservation Service (SCS) in 1935. Con-
servation demonstration farms were established and the Soil Conservation District (SCD)
work with individual farmers started after 1938 and expanded rapidly. These changes
and diversion of mappers into other pursuits during World War II resulted in a drastic
increase in time between completion of the mapping of a county and publication of
the survey. The interval was 10 years in the early 1950s (18).
The SCS benefitted from knowledge gained in past surveys and developed a system
of soil conservation surveys made for individuals farms or groups of cooperating farms
as owners requested technical assistance. Beginning around 1950, symbols on soil map
units included slope range in one of seven classes, and, effects of past erosion in one
of three classes, both important in use and management (56). Such surveys preceded
any farm planning assistance. Maps enabled use of a land capability classification system
to help farmers understand the restrictions imposed by features like wetness, erodibility,
and low water holding capacity on the soils of their farms. In counties with SCDs
and active soil conservation programs, there was more interest in developing farm plans
than in concentrating on soil survey publication.
In 1951 Herbert P. Ulrich became the second soil survey leader responsible for
the operations of the soils survey for AES. Prior mapping experience in 12 or more
Indiana counties made him particularly capable of expanding knowledge about soil
parent materials (48, 49) useful to field soil scientists in succeeding years. He also
assisted in support of other studies about loess distribution and soil mineralogy begun
by White (51, 52), Bailey (1), Post (33, 34), and others. T.C. Bass became SCS State
Soil Scientist in 1945 after serving in that capacity in Wisconsin. Bass and Ulrich capably
led the survey for many years.
Ulrich assisted greatly in improving soil survey report usefulness. In the 1943
survey of Knox County, he introduced the first general soil association map of Knox
County which led to the small-scale, colored maps in the standard soil reports of 1960
and later. He first added estimated crop yields at two levels of management in the
Soil and Atmospheric Sciences 535
Vanderburgh County Survey of 1944. They provided economic background for selec-
ting farm practice combinations. These innovations were used in a number of publica-
tions delayed by World War II up to Carroll and Tippecanoe counties in 1959. In
these he introduced block diagrams relating soils and topography, tables suggesting
use and management, including rotations for soils of similar management groups, and
the first tie-in with the land capability classification system used widely by SCS in
farm planning.
Soil Survey Functions Combined Under Soil Conservation Service
In 1952 the soil survey publication functions in the USDA Bureau of Plant Industry
were combined with the soil conservation surveys made by SCS. Cooperation with
the AES at Purdue continued. Soil conservation surveys, which were available to be
used immediately in farm planting, were improved in nature and quality to make them
adaptable to later publication as part of a county soil survey.
Phase 2. Era of Standard Soil Surveys for Publication 1960-1987
Mapping for farms of SCD, later named Soil and Water Conservation District
(SWCD), cooperaiors still look priority in planning survey progress but it became
an integral part of a complete county survey. Air-photos at a scale of 4.0 inches (a
few 3.2) per mile were selected as base maps.
Standard soil survey reports included soil descriptions designed for lay people
and more technical and detailed ones for soil scientists. Reports also provided a colored
general soil map showing the broad soil associations with brief descriptions of the
soils and land uses as described by Zachary and others (57). They included a glossary
of technical terms and tabular presentations of soil properties and of interpretations
relative to using the soil map units for agricultural, forestry and engineering uses. The
first publication of this kind in Indiana was for Fayette and Union counties in 1960.
Tables of test data useful to highway engineers and a section on wildlife were
added in 1962 in Owen County; soil series classified according to the U.S. Soil Tax-
onomy (of most interest to scientists) was first added in 1967 in Parke County. Tables
of soils and outdoor recreation potential and of trees and shrubs for wildlife planting
came in 1969 in Allen County, while soil limitations for six common classes of out-
door recreation came in 1971 in Howard County. A chapter on town and country
planning rating soils for homesites, septic disposal systems, local roads and streets,
sewage lagoons, landfills and species for landscape planting emerged in 1972 in Lake
County. The reports showed a continuous improvement in meeting the needs of the user.
Other Materials Support Use of Published Soil Surveys
Management recommendations must be revised frequently. Thus, specific ones
were omitted from soil survey reports and are presented in several extension publica-
tions usually prepared by Cooperative Extension Service, AES, and SCS people. Fer-
tilization is recommended in line with soil test reports. Guides to economic productivity
levels for all soils are updated periodically (17, 50). Available soil water capacity studies
by Wiersma (53, 54) aid productivity estimates. Drainage needs refer to a farm drainage
guide (29), irrigation potential to another guide (25), and adaptation of tillage-planting
systems to yet another (20).
A highly valuable guide to conservation planning resulted from work of Wischmeier
and others (55) in developing the universal soil loss equation. With it farm planners
can predict long-time average soil losses from specific tracts of land under various
cropping and management systems. No advance has helped make soil survey maps
more useful to farm planners and landowners in agricultural programs!
For those needing to relate to soils on a broader basis than single farms other
publications are supportive. A series of general soil maps and a users guide were up-
536 Indiana Academy of Science Vol. 94 (1985)
dated in 1975 (19) and a guide to properties of soil series (24) was published in 1977.
They reflected some of Harry M. Galloway's depth of experience in helping farmers
and others use their soils while preserving them for future generations. A 1977 colored
map of soil associations (41) and a 1982 key relating soils to each other and to their
environments (14)are useful to agency program planning, planning commissions and
others. To help persons understand and judge Indiana soil properties, the high school
soil judging program manual (50) was expanded in 1978 to be used by a wider audience.
Donald P. Franzmeier became the third and present state soil survey leader respon-
sible for soil survey operations in 1970. He recognized the potential for learning more
about the soils being mapped as surveys progressed. He organized a soil characteriza-
tion laboratory at Purdue to perform analyses of soil samples taken by field soil scien-
tists of SCS and IDNR during survey operations. Results are in a series of AES station
bulletins from 1977 to 1984 (41). Field and laboratory procedures were described in
a 1977 bulletin (13).
Field soil scientists also measured water table depths and crop yields, especially
as affected by erosion. With assistance of SCS soil scientists, especially Frank Sanders,
papers were published about soil moisture regimes (12, 15), organic soils (37), history
of the Miami series in Indiana (38), and dark-colored northern Indiana soils (36).
Graduate students working with Franzmeier studied various aspects of soil genesis and
many of them worked on soil survey parties during summers to gain experience and
gather field research data. Included were studies on soil water regimes, Harlan (22);
remote-sensing, Steinhardt (44) and Cipra (9); micromorphology, Steinhardt (43);
manganese minerals in soils, Ross (35); soil formation and fragipan development in
loess-derived soils, Harlan (23), Norton (32), and Steinhardt (45, 47); eolian processes,
Franzmeier (11) and Miles (31); hydraulic conductivity and morphology of Clermont
soils, King (26, 27); and plant nutrients in trees, Crum (10). Data relating the organic
matter content and color of silt loam soils were summarized by Steinhardt (46).
County and State Sources Help Finance Surveys
In the mid 1960s, because of rapid land use changes, various counties supplied
funds to obtain soil survey information to help with land use decisions. Lake County,
in cooperation with Purdue, supported a soils extension specialist; Howard County
provided funds for extra assistance from SCS in preparing reports for its planning
commission; Elkhart and Clay counties were the first to supply funds to get their surveys
completed sooner; and in Miami, Marion, Johnson, and Kosciusko counties state and
county funds supported half the cost of field mapping.
In 1968, Ray Dideriksen became State Soil Scientist when T.C. Bass retired, and
in 1972 H. Raymond Sinclair was appointed to that position. With Franzmeier, they
directed the survey through this period of expansion.
Accelerated Soil Survey 1974-1987
The 1973 Indiana legislature passed a bill requiring that soil survey information
be used to evaluate farmland for tax assessment. At that time less than 40 of the stan-
dard surveys suitable for such use had been completed. In 1974 additional state money
was provided through the Indiana Department of Natural Resources (IDNR) to match
county and federal money to complete the field work of the standard survey in 1984.
At that time SCS employed 28 field soil scientists. To complete the survey in 10 years
it was projected that this number would remain constant and that IDNR, through
the State Soil and Water Conservation Committee, would employ a number of soil
scientists, increasing to 31 around 1980 and decreasing to none by the end of 1984.
This projected schedule for total number of soil scientists was followed fairly well
except that now, toward the end of the program, more are employed by IDNR and
fewer by SCS (Figure 1). State and county funds supported IDNR employees and federal
Soil and Atmospheric Sciences
537
60 r
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50
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o
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o
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ir\
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1970
72
74
76
82
84
86
88
78 80
YEAR
Figure 1. Number of field soil scientists in the Cooperative Soil Survey of Indiana,
their employing agency, and source of funds.
and state funds supported SCS employees. Also, three counties with surveys started
in the 40s and published in the late 50s were added, extending the program. By 1987
all field soil mapping should be completed and by 1990 all surveys should be published.
Surveys for 53 counties will be completed through this program (Table 1). In 1974
the estimated cost of the program was $15,700,000 with 48% from state funds, 31%
from federal funds, and 21% from county funds.
Table 1 . Dates or projected dates ( > 1984) of completion of field mapping and publication
of soil surveys of Indiana.
County
Mapping
Completed
Published
County
Mapping
Completed
Published
Adams*
1982
1985
Madison
1961
1967
Allen
1961
1969
Marion
1974
1978
Bartholomew
1971
1976
Marshall*
1978
1980
Benton*
1983
1986
Martin*
1982
1986
Blackford & Jay*
1982
1985
Miami
1976
1979
Boone
1970
1975
Monroe*
1977
1981
Brown*
1984
1986
Montgomery*
1982
1986
Carroll*
1986
1989
Morgan*
1978
1981
Cass*
1978
1981
Newton*
1986
1989
Clark & Floyd
1967
1974
Noble
1973
1978
Clay*
1979
1982
Ohio* (with Dearborn)
1977
1981
Clinton*
1977
1980
Orange*
1980
1984
Crawford
1968
1975
Owen
1959
1964
Daviess
1968
1974
Parke
1959
1967
Dearborn & Ohio*
1977
1981
Perry
1963
1969
Decatur*
1979
1983
Pike*
1982
1986
Dekalb*
1979
1982
Porter*
1977
1981
Delaware
1967
1972
Posey
1977
1979
538
Indiana Academy of Science
Vol. 94 (1985)
Table 1. — Continued
Dubois*
1977
1980
Pulaski
1964
1968
Elkhart
1967
1974
Putnam*
1978
1981
Fayette & Union
1952
1960
Randolph*
1981
1986
Floyd (with Clark)
1967
1974
Ripley*
1981
1985
Fountain
1961
1966
Rush*
1981
1985
Franklin*
1983
1987
Scott
1958
1962
Fulton*
1982
1986
Shelby
1967
1974
Gibson*
1984
1987
Spencer
1966
1973
Grant*
1983
1986
St. Joseph
1973
1977
Greene*
1983
1986
Starke*
1979
1982
Hamilton
1975
1979
Steuben*
1977
1981
Hancock
1974
1978
Sullivan
1962
1971
Harrison
1969
1975
Switzerland*
1983
1986
Hendricks
1970
1974
Tippecanoe*
1987
1990
Henry*
1981
1986
Tipton*
1984
1987
Howard
1965
1971
Union (with Fayette)
1952
1960
Huntington*
1979
1983
Vanderburgh
1971
1976
Jackson*
1983
1986
Vermillion
1976
1978
Jasper*
1982
1986
Vigo
1970
1974
Jay* (with Blackford)
1982
1985
Wabash*
1979
1983
Jefferson*
19g0
1984
Warren*
1985
1988
Jennings
1971
1976
Warrick
1975
1979
Johnson
1974
1979
Washington*
1983
1986
Knox*
1978
1981
Wayne*
1981
1986
Kosciusko*
1983
1987
Wells*
1986
1989
LaGrange
1977
1980
White*
1978
1982
Lake
1966
1972
Whitley*
1983
1987
LaPorte*
1977
1982
Lawrence*
1981
1984
'In accelerated soil survey program.
The employees trained during the program are benefitting Indiana in many ways.
Some continued as soil scientists or conservationists with SCS. Many continued with
IDNR in the Department of Reclamation while others took positions with agencies
like ASCS, PA, FHA or coal companies, or went on to graduate school. In all cases
their soil training is of great benefit to people in Indiana and in other states where
they are employed.
Computer Storage and Interpretation of Surveys
Joseph E. Yahner realized the advantage of storing soil maps in computers and
using the system to make various kinds of soil interpretations (42). Early systems,
using computer cards and the Purdue main computer, were tried in Elkhart, Dubois,
and Miami counties. They utilized a 2.5-acre grid-cell. Then a system was developed
for storing attributes (properties) of soil map units and soil and land owner maps by
1 1/3-acre grid-cells on the county FACTS terminals (39). This system was designed
largely to provide average productivity values for specific tracts of land. Although
initial computerization was to accommodate land evaluation for tax assessment,
experience in the pioneering counties has shown that people other than tax assessors
make extensive use of the system. For example, in Miami County, many rural appraisers
and realtors furnish a computer-printout of a soils grid-cell map and a summary of
soil productivity information to their clients (personal communication Jack Hart,
Cooperative Extension Service). The survey has been processed by computer in 19 coun-
ties to date (Figure 2).
Soil and Atmospheric Sciences
539
In progress
Field work
complete.
Not published
Published
-
/
I>
/
,
,
Published and
entered into
computer data
base.
Figure 2. Status of soil surveys and computer storage of surveys in Indiana as of
November 1984.
Phase 3. Future Plans for Soil Survey
Future Soil Survey After 1987
In the first two phases of the soil survey the goal was to produce published soil
540 Indiana Academy of Science Vol. 94 (1985)
surveys of each county of the state. After more than 80 years of making soil surveys
the procedures for describing and classifying soils and conducting surveys were described
in detail in handbooks and manuals. The process of making and publishing a soil
survey became so tightly prescribed that there was little allowance for creativity and
innovation by the soil scientists.
In the future, however, the situation will change dramatically. For the third phase
of the soil survey, which begins now as soil scientists complete their assignments in
the second phase, the goals and tasks are not as well defined as they were in the two
earlier periods. There are very few guidelines for the survey. Individual soil scientists
will have to develop new ideas. Indiana will be the first state in the midwest, and
one of the earliest in the country, to complete the standard soil survey mapping.
In proposing the goals and objectives of future phases of the soil survey we have
made two assumptions: that detailed surveys of entire counties will not be produced
in the immediate future, and that the major goal of the program will be to help people
use soil information, much of which is in published soil surveys. This will continue
and expand the survey extension education programs that began in 1958 (21). In
examining these goals we realize that some of the information needed to serve the
public is not available. Over the years the mapping goals were so demanding that soil
scientists had little time to measure the properties of the soils they were mapping.
Also, some of the field work of the "modern" detailed surveys will be 40 years old
when the last survey is published. Thus a major effort will be to collect and update
information and make it available to assist users of the survey.
Over the years the major use of soil surveys has been for planning farming opera-
tions and this will continue to be an important application of the information. In the
future, however, two new programs will be major users of soil information — using
the computer to store and interpret soil surveys, and evaluating soils for on-site home
waste disposal.
In much of Indiana a large percentage of the soils are not suited to conventional
septic-tank systems for home waste disposal because they have high water tables or
are too slowly permeable. Innovative systems can be used successfully on many of
these soils as demonstrated in the On-Site Waste Disposal Project led by J.E. Yahner
at Purdue. One is the mound system, in which effluent absorption lines are placed
in a mound of sand built on the undisturbed soil in order to create a zone of un-
saturated soil above the water table to effect purification of the effluent. Another
is the pressure distribution system, in which effluent is pumped into the drainage lines
for even distribution. Accurate soil information is essential for the successful opera-
tion of these systems. It is necessary to decide if a system can be installed and, if
it can, where it should be located and how it should be designed. Contractors must
also be taught to construct the system without damaging the soil.
With this background, we will outline the major tasks of the future of the soil
survey as we perceive them. We realize that this transition phase must remain flexible
to adjust to changing needs. Five major objectives are suggested.
Objectives of Program
Integrating and Updating Surveys
Some work is needed to bring older surveys up to the standards of the most re-
cent surveys. This is especially true for interpretations because now many more are
made than were made in earlier surveys. Also, individual county surveys were made
by different people at different stages of knowledge of the soils, so adjoining county
surveys do not match very well in some places. To help those who use surveys across
county lines we need to define mapping units state-wide and show how these units
Soil and Atmospheric Sciences 541
fit the landscape. This might be done by defining soil-landscape units and mapping
them on U.S. Geological Survey topographic maps.
So/7 Investigations
During the course of the survey practically all of the efforts were directed to
mapping soils and developing reports. In addition, laboratory characterization data
were obtained for many counties. Now we need to learn more about the properties
of the soils themselves, especially properties not measured in the laboratory. Many
of the interpretations we are now making are based largely on estimated properties
and some of these estimates were made from little factual information. In the future
we need to obtain laboratory characterization of soils in counties mapped before the
soil characterization laboratory was started. We especially need to measure field pro-
perties such as hydraulic conductivity (permeability), seasonal water table depths,
available water capacity, and bulk density (to characterize compaction) which are
necessary for designing farm drainage systems, on-site waste disposal systems, predic-
ting the water storage in soil profiles, and recommending suitable tillage systems. We
also need to measure crop yields, especially on sloping and eroded soils, to support
yield estimates made for land evaluation and tax assessments.
On-site Investigations
Soil maps can be used for many interpretations, but for some soil uses a soil
scientist must investigate specific conditions of the site. This is especially important
for on-site waste disposal — whether or not a system can be installed and, if it can,
where it should be located, how it should be designed, and when and how it should
be constructed. The nature of these investigations and the relation of soil properties
to the kinds of recommendations made are not well established. They need to be deter-
mined in conjunction with the On-Site Waste Disposal Project. In agriculture, on-site
investigations will continue to be needed for installing drainage systems, constructing
erosion control and water detention structures, and planning other farm operations.
Increasingly, soil scientists will be called on to identify soil compaction problems and
advise how to prevent compaction and improve compacted soils. These investigations
and interpretations also are not established.
Soil Mapping
Detailed soil maps and reports need to be prepared for special areas, such as
research farms, developing areas, reservoirs and other high-intensity uses. They will
draw on experience gained in the survey program.
It will be necessary to map land use and flood hazards during the growing season,
which are necessary for land assessment, and store this geographic information in the
computer. Soil scientists will be called on to assist with this mapping. Experience has
shown that soil scientists learn about soil properties and how they relate to using the
soil by mapping them. When phase 3 of the soil survey program begins a large group
of soil scientists with this background will be available, but this storehouse of knowledge
will not last forever. Some mapping programs will be necessary to provide training
and experience for future generations of soil scientists.
Education
Education is a two-way street. Soil scientists need to be educated themselves as
well as educate others. Many of the programs are new and the nature of the job will
be determined by those in the position, in contrast to the previous soil survey program
in which the nature of the job was well established before an individual soil scientist
filled a position.
In this kind of innovative program, mistakes will be made, but to learn from
them will require frequent interchanges between soil scientists and researchers, and
542 Indiana Academy of Science Vol. 94 (1985)
among the soil scientists. To be effective in their work soil scientists must have a keen
interest in maintaining and improving their skills. In the process many will obtain
advanced degrees. They will also participate in short training sessions in Indiana or
elsewhere. They must also be interested in teaching others such as conservationists,
contractors, engineers, and sanitarians. They will continue some education programs
in place since 1958 (21) and develop others as needs arise. How well these great infor-
mation resources are utilized to better the patterns of land uses by Indiana citizens
will depend on the ingenuity of workers training themselves for the future.
Summary
In 1902 the first soil survey in Indiana, for Posey County, was published. Beginning
then, we have identified three phases of the survey. In the first phase, colored line
maps were published for 64 counties between 1902 and 1959, mostly at a scale of
1 in.T mi. (1: 63, 360). Jennings County, mapped in the early 1930s was the first county
in the U.S. to be mapped entirely on air photos. Very little mapping was done during
and immediately after World War II.
Our "modern" published surveys were produced, or are being produced in the
second phase of the survey in which the mapping began about 1952 and the report
was published in 1960. The surveys were all made and published on air photo base
maps, mostly at a scale of 1:15, 840 (4 in.: 1 mi.) with some at 1:20,000 (3.2 in.:l
mi.). Now, (November, 1984) the field work is finished in 87 counties and 5 county
surveys are in progress; surveys are published for 60 counties. The field work is scheduled
to be finished in 1987.
The third phase of the survey, which begins as soil scientists complete their regular
soil mapping assignments, will emphasize learning more about the soils mapped and
helping people use soil information. Also, some "modern" surveys, made during a
35-year span, will need to be brought up-to-date and integrated with other surveys.
The challenge is to protect one of the state's most precious resources for future
generations.
Literature Cited
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unglaciated area. Proc. Ind. Acad. Sci. 68:343-48.
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37:63-72.
3. Bushnell, T.M. 1929. Aerial photographs — Jennings County. Proc. Ind. Acad.
Sci. 39:229.
4. Bushnell, T.M. 1936. Aerial photography in Indiana. Proc. Ind. Acad. Sci. 46:142.
5. Bushnell, T.M. 1938. Taxonomy of Indiana soils. Proc. Ind. Acad. Sci. 48:113.
6. Bushnell, T.M. 1939. Outline of classification of Indiana soils. Proc. Ind. Acad.
Sci. 49:151-58.
7. Bushnell, T.M. 1942. Some aspects of the soil catena concept. Soil Sci. Soc. Am.
Proc. 7:466-476.
8. Bushnell, T.M. 1944. The story of Indiana soils. Purdue Univ. Agr. Exp. Stn.
Spec. Circ. 1. 52 p.
9. Cipra, J.E., D.P. Franzmeier, M.E. Bauer, and R.K. Boyd. 1980. Comparison
of multispectral measurements from some nonvegetated soils using LANDSAT
digital data and a spectroradiometer. Soil Sci. Soc. Am. J. 44:80-84.
10. Crum, J.R. and D.P. Franzmeier. 1980. Soil properties and chemical composi-
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Soil and Atmospheric Sciences 543
11. Franzmeier, D.P. 1970. Particle size sorting of proglacial eolian materials. Soil
Sci. Soc. Am. Proc. 34:920-924.
12. Franzmeier, D.P., D. Wiersma, S. Brownfield, J. Robbins, J. Shively, and R.
Wingard. 1973. Water regimes of some Indiana soils. Purdue Univ. Agr. Exp.
Stn. Research Bull. 904, 19 pp.
13. Franzmeier, D.P., G.C. Steinhardt, L.D. Norton, and J.R. Crum. 1977. Soil
characterization in Indiana: I. Field and laboratory procedures. Purdue Univ.
Agr. Exp. Stn. Research Bull. 943, 30 p.
14. Franzmeier, D.P., and H.R. Sinclair, Jr. 1982. Key to soils of Indiana. Purdue
Univ. Coop. Ext. Ser. AY-249, 16 p.
15. Franzmeier, D.P., J.E. Yahner, G.C. Steinhardt, and H.R. Sinclair, Jr. 1983.
Color patterns and water table levels in some Indiana soils. Soil Sci. Soc. Am.
J. 47:1196-1202.
16. Franzmeier, D.P., J.E. Yahner, G.C. Steinhardt, and H.R. Sinclair, Jr. 1984.
Water table levels and water contents of some Indiana soils. Purdue Univ. Agr.
Exp. Stn. Research Bull. 967. 49 p.
17. Galloway, H.M. 1962. Establishing crop potentials for Indiana soil types. Proc.
Ind. Acad. Sci. 71:335-340.
18. Galloway, H.M. and J.E. Yahner, 1976. Indiana's soil survey adjusts to chang-
ing needs. Proc. Ind. Acad. Sci. 85:391-404.
19. Galloway, H.M., J.E. Yahner, G. Srinivasan and D.P. Franzmeier. 1976. Users
guide to general soil maps of Indiana. Purdue Univ. Coop. Ext. Ser. AY-50 series
supplement. 27 p.
20. Galloway, H.M., D.R. Griffith, and J.V. Mannering. 1977. Adaptability of various
tillage-planting systems to Indiana soils. Purdue Univ. Coop. Ext. Ser. AY-210.
21. Galloway, H.M. and J.E. Yahner. 1978. Soil survey education: the Indiana pro-
gram. Jour. Soil & Water Cons. 33 (3): 11 1-1 14.
22. Harlan, P.W., and D.P. Franzmeier. 1974. Soil-water regimes in Brookston and
Crosby soils. Soil. Sci. Soc. Am. Proc. 38:638-643.
23. Harlan, P.W., and D.P. Franzmeier. 1977. Soil formation on loess in southwestern
Indiana: II. Distribution of clay and free iron oxides and fragipan formation.
Soil Sci. Soc. Am. J. 41:99-103.
24. Indiana's soil series and their properties. 1981. Purdue Univ. Coop. Ext. Ser.
AY-212.
25. Irrigation of field crops in Indiana. 1977. Purdue Univ. Coop. Ext. Ser. ID-119.
28 p.
26. King, J.J. and D.P. Franzmeier. 1981. Estimation of saturated hydraulic con-
ductivity from soil morphological and genetic information. Soil Sci. Soc. Am.
J. 45:1153-1156.
27. King, J. J., and D.P. Franzmeier. 1981. Morphology, hydrology, and manage-
ment of Clermont soils. Proc. Ind. Acad. Sci. 90:416-422.
28. Kohnke, H. and A. Ohlrogge. 1976. Historical highlights in Indiana soil science.
Proc. Ind. Acad. Sci. 85:385-89.
29. Mannering, J.V., D.P. Franzmeier, and others. 1984. Indiana drainage guide.
Part 1. Soils drainage recommendations. Purdue Univ. Coop. Ext. Ser. ID-160.
11 p.
30. Milne, G. 1935. Composite units for the mapping of complex soil associations.
Trans. Third Int. Cong. Soil Si. Vol. 1:345-347.
31. Miles, R.J., and D.P. Franzmeier. 1981. A litho-chronosequence of soils formed
in dune sand. Soil Sci. Soc. Am. J. 45:362-367.
32. Norton, L.D., and D.P. Franzmeier. 1978. Toposequences of loess-derived soils
in southwestern Indiana. Soil Sci. Soc. Am. J. 42:622-627.
544 Indiana Academy of Science Vol. 94 (1985)
33. Post, D., A.L. Zachary and H.P. Ulrich. 1970. Characteristics of Pembroke soils
from Indiana. Proc. Ind. Acad. Sci. 79:396-404.
34. Post, D. and J.L. White. 1970. Quantitative mineralogical analysis of soil clays.
Proc. Ind. Acad. Sci. 79:405-411.
35. Ross, S.J., Jr., and D.P. Franzmeier. 1976. Mineralology and chemistry of
managanese oxides in some Indiana soils. Soil Sci. Soc. Am. J. 40:137-143.
36. Sanders, F.W., and D.P. Franzmeier. 1974. Classification of some dark-colored
northern Indiana soils. Proc. Ind. Acad. Sci. 83:433-438.
37. Sanders, F.W., and D.P. Franzmeier. 1976. Coprogenous earth in organic soils
in northern Indiana. Proc. Ind. Acad. Sci. 85:377-384.
38. Sanders, F.W., H.R. Sinclair and H.M. Galloway. 1979. History of Miami series
in Indiana. Proc. Ind. Acad. Sci. 88:405-411.
39. Santini, Judy, J.E. Yahner, and D.P. Franzmeier. 1983. Soil maps and inter-
pretations system. FACTS user guide. Purdue Univ. Coop. Ext. Ser. FX-71 (AY).
40. Soil Survey Staff. 1977. Map of the soil associations of Indiana. Purdue Univ.
Coop. Ext. Ser. AY-209.
41. Soil Survey Staff. 1977 to 1984. Soil characterization in Indiana series. Purdue
Univ. Agr. Exp. Stn. Station Bulletins 174, 175, 222, 274, 323, 360, 412, and 451.
42. Srinivasan, G., J.E. Yahner, and H.M. Galloway. 1976. Soil information for
land use decisions: a scientific approach. Proc. Ind. Acad. Sci. 85:371-375.
43. Steinhardt, G.C., P.W. Harlan, S.J. Ross, and D.P. Franzmeier. 1974. Micromor-
phological analysis of selected Indiana soils. Proc. Ind. Acad. Sci. 83:439-445.
44. Steinhardt, G.C., D.P. Franzmeier, and J.E. Cipra. 1975. Indiana soil associa-
tions compared to Earth Resources Technology Satellite imagery. Proc. Ind. Acad.
Sci. 84:463-468.
45. Steinhardt, G.C., and D.P. Franzmeier. 1979. Chemical and mineralological pro-
perties of the fragipans of the Cincinnati catena. Soil Sci. Soc. Am. J. 43:1008-1013.
46. Steinhardt, G.C., and D.P. Franzmeier. 1979. Comparison of organic matter con-
tent with color for silt loam soils of Indiana. Comm. in Soil Sci. and Plant Anal.
10(10): 1271-1277.
47. Steinhardt, G.C., D.P. Franzmeier, and L.D. Norton. 1982. Silica associated with
fragipan and non-fragipan horizons. Soil Sci. Soc. Am. J. 46:656-657.
48. Ulrich, H.P. 1937. Surface geology — Bartholomew and Brown counties. Proc.
Ind. Acad. Sci. 48:113.
49. Ulrich, H.P. 1959. Wisconsin moraines as a source of loess in Fayette and Union
counties. Proc. Ind. Acad. Sci. 68:349-53.
50. Walker, Carl F. 1976. A model to estimate corn yields for Indiana soils. Unpubl.
M.S. thesis, Agron. Dept. Purdue Univ.
51. White, J.L., G. Talvenheimo, M.G. Klages and M.M. Phillipe. 1957. Survey of
mineralogy of Indiana soils. Proc. Ind. Acad. Sci. 66:232-241.
52. White, J.L., and Maribel Cruz. 1971. Soil colloids and behavior of pesticides
in soils. Proc. Ind. Acad. Sci. 81:305.
53. Wiersma, Dan. 1962. Increasing crop potentials through water availability. Proc.
Ind. Acad. Sci. 71:347-52.
54. Wiersma, Dan et al. 1984. Soil water characteristics data for some Indiana soil
profiles. Purdue Univ. Agr. Exp. Stn. Station Bull. 452. 153 p.
55. Wischmeier, W.H. and D.D. Smith. 1978. Predicting rainfall erosion losses. USDA
Agricultural Handbook 537. SEA/USDA in Coop, with Purdue Agr. Expt. Stn.
58 p.
56. Yahner, J.D., G.C. Steinhardt, D.P. Franzmeier, H.M. Galloway, and A.L.
Zachary. 1980. Understanding and judging Indiana soils. Purdue Univ. Coop.
Ext. Ser. ID-72. 59 p. + plates.
Soil and Atmospheric Sciences 545
57. Zachary, A.L., and D.F. Post. 1962. Origin, characteristics, and management
of the soil associations of Allen County. Proc. Ind. Acad. Sci. 72:330-337.
Gust Fronts in Doppler Radar Data
Diana L. Klingle and David R. Smith
Department of Geosciences
Purdue University
West Lafayette, Indiana 47907
Introduction
A gust front is the boundary between the horizontally propagating cold air outflow
from a thunderstorm and the surrounding environmental air. The sharp changes in
wind speed and direction across a gust front can produce turbulence and wind shear
of sufficient magnitude to be hazardous to aircraft during takeoff and landing. Analyses
of aircraft accident statistics published by the National Transportation Safety Board
for the years 1976-78 indicate that one of the most significant hazards to aviation
is low altitude wind shear (12). It is in response to such hazards that research projects
such as JAWS (Joint Airport Weather Studies) have been conducted. Gust fronts,
as well as downbursts and tornadic phenomena, constitute a hazard to aviation, but
it is impossible to detect the low altitude wind shear they produce with the conven-
tional radars currently in use. Doppler radars are capable of sensing air motions and,
therefore, are useful tools in the detection of this aviation hazard. This paper examines
the use of Doppler radar in the detection of thunderstorm gust fronts and their associated
wind shear patterns.
Background on Gust Fronts
A. Gust Front Structure
A gust front is the leading edge of an outflow which is produced when the
thunderstorm downdraft reaches the ground and spreads horizontally. The passage
of the gust front is often accompanied by a sharp rise in pressure, a decrease in
temperature, and abrupt changes in wind speed and direction (4). As the cooler, denser
outflow intrudes into the warmer, less dense environmental air, the warm air is lifted
up and over the outflow boundary (Figure 1). This intrusion of colder air into warmer
has been likened to a gravity current (1, 6, 8, 16).
Studies of laboratory gravity currents have illustrated the presence of phenomena
which have counterparts in thunderstorm outflows. Fluid within the outflow moves
faster than the outflow boundary. Under the proper conditions, friction between the
HIGH TURBULENCE
GUST FRONT
BOUNDARY
wake f~ ~-^vV2C — __ LOCATION OF
COLD AIR _ 1 \\ — — — RADAR BEAM
(From Thunderstorm)
WARM
AIR
UNDERCURRENT ^ \^ H|GH
TURBULENCE
Figure 1. Schematic diagram of the vertical structure of a thunderstorm outflow
and gust front. Motion is relative to the gust front. (Adapted from Goff, 1975)
547
548 Indiana Academy of Science Vol. 94 (1985)
fluid and the surface across which it propagates causes the lowest layers of the flow
to be retarded. Some of the fluid is deflected downward, producing the "backflow."
The fluid above this friction layer moves faster and protrudes ahead of the surface
boundary. This protrusion is known as the "nose" of the gust front (Figure 1). The
advancing fluid is deflected upward at the leading edge producing a bulge known as
the "head." Studies have shown evidence that these features also exist in nature (5,
7). A turbulent "wake" region is located behind the head. The leading edge of the
outflow is not an impermeable boundary. Along with lifting, mixing of the environmental
and outflow air occurs at the outflow interface, which produces yet another turbulent
region.
B. Doppler Radar Signatures of Gust Fronts
It has been shown that Doppler radar is capable of detecting thunderstorm outflows
(2, 13, 17). The abrupt change in wind speed and direction mentioned previously can
be sensed by Doppler radar and displayed such that regions of radial shear are apparent.
Doppler radars sense the component of the wind along the radar beam; inbound (i.e.,
toward the radar) is considered negative, outbound is positive.
There are some difficulties which may prevent gust front detection by radar (18).
For example, the distance of the center of the radar beam above the surface increases
with distance from the radar due to the curvature of the Earth. A shallow outflow
at a large distance from the radar may be below the beam, and thus go undetected.
Near the radar, ground clutter contaminates the signal. Range folding (targets beyond
the unambiguous range appear to be located within the first trip) can mask the gust
frontal signature. Despite these problems, gust fronts can generally be detected in the
Doppler data at ranges up to 100 km.
1. Reflectivity
Gust fronts are often associated with "thin line" echoes in radar reflectivity fields.
Strong gradients in the refractive index at the leading edge of the outflow have been
cited as a possible explanation of this phenomena (3, 14, 15). Also, it is believed that
the thin line is caused by insects which are picked up and carried along by the outflow
and by birds that feed on these insects (9, 10). More recently it has been suggested
that the thin line is produced by the "precipitation roll," that is, by precipitation par-
ticles which are swept along with the outflow winds as they move away from the parent
storm (17). Others have suggested that this thin line is caused by the accumulation
of dust and debris particles.
2. Doppler Velocity
Gust fronts can be identified in the Doppler wind field as linear patterns of radial
shear. As an example, assume a gust front is approaching the radar from the west.
A reasonable first approximation is that winds within the outflow are oriented perpen-
dicular to the gust front and therefore have a strong radially inbound component in
regions where the gust front is perpendicular to the beam (Figure 2). Environmental
winds ahead of the gust front are typically from the southeast to southwest quadrant
and display outbound ( + ) or weak inbound (-) velocities. Moving away from the
radar toward the gust front along a radial, one finds the Doppler velocities changing
from positive (or weak negative) to negative (or more strongly negative) as the gust
front is encountered. This abrupt change in Doppler wind speed produces a linear radial
shear signature at the leading edge of the outflow.
The gust front tends to curve (Figure 2) and portions of its length may become
aligned along a radial. When this occurs, the flow is primarily across the beam and
as such is sensed as zero velocity by the Doppler radar. Identifying the radially-oriented
portions of the gust front in the radar velocity field can be difficult, yet important
in the interpretation of the strength of the outflow.
Soil and Atmospheric Sciences
549
Figure 2. Schematic diagram of the horizontal structure of a thunderstorm outflow
and gust front. Winds within the gust front tend to flow perpendicular to the gust
front. The dashed lines indicate possible locations of a radar beam which scans the
outflow.
Case Study
As an example of the ability (and difficulties) of Doppler radar to detect gust
fronts, a case study is presented. This case involves two gust fronts which were pro-
duced at different locations along the same line of storms over Oklahoma on 26 April
1984. Photographs (Figures 3-6) of the reflectivity and Doppler velocity displays from
the Doppler weather radar of the National Severe Storms Laboratory (NSSL) in Norman,
OK show the major features of this storm and accompanying gust fronts.
The line of storms displayed in Figures 3a and b was initiated by a rapidly moving
cold front, advancing toward the moist unstable air over central and eastern Oklahoma,
producing severe thunderstorms and tornadoes. At 20:21:09 CST, the first outflow
boundary produced by this line (cursor) had not separated from the parent storm to
form the thin line reflectivity signature, but roughly paralleled the 16dBZ reflectivity
contour at the leading edge of the parent line. (A value of 10 dBZ has been added
to the displayed values in order to bring weak signals above the display threshhold.)
This gust front was identified as such by the linear radial shear pattern in the velocity
display (Figure 3b). Radial wind speeds within the outflow average 19 ms "' inbound.
While this gust front was scanned, it never separated from the storm to form the thin
line signature (Figure 3a). However, the radial shear line was present in all low eleva-
tion angle scans.
In Figure 3b, all of the velocities within the outflow (west of the shear line at
the leading edge) are inbound (negative). This is no longer the case in Figure 4b. Except
for a narrow band of negative velocities behind the leading edge at the north end
of the gust front (label A), the velocities within the outflow are positive. It is believed
that the radar beam is cutting through the head of the gust front and sensing en-
vironmental winds (roughly from the southwest at 27 ms ') on either side. The loca-
tion of the radar beam illustrated in Figure 1 indicates a possible configuration for
550
Indiana Academy of Science
Vol. 94 (1985)
Figure 3(a)
Figure 3(b)
Figure 3. Plan Position Indicator displays of (a) reflectivity and (b) mean Doppler
velocity from the Norman, OK Doppler radar for 26 April 1984, 20:21:09 CST. Note
information common to all radar display photographs: The legend at the right of the
photographs indicates the date (4/26/84) and time (20:21:09 CST) of the PPI scan.
Beneath these lines are the color categories (0 through E) and their associated reflec-
tivity (in dBZ) or velocity (in ms ') values. Category F (white) is reserved for the
cursor, navigation aids, and range rings (white arcs on the displays). "R40" shows
that the range marks are separated by 40 km. The azimuth and range of the center
of the display are given by CAZ (320°) and CRG (60 km). Storm motion (SM) is the
speed and direction of the storm (00@000) that is subtracted from the velocity field
so that the displayed velocities are storm-relative. The azimuth, range and height above
the ground of the center of the cursor is shown by AZ+ (314°), RG+ (55 km) and
HT+ (0.5 km). At the time of the photograph, the radar was pointed at azimuth
AZ (269°) with an elevation angle EL (0.5°). All following radar display photographs
are interpreted similarly.
that in Figure 4b. This situation illustrates how a change in the elevation angle of
the radar can alter the appearance of the gust front on the radar displays.
The line of storms continued to propagate east-northeast and, at about 2040 CST,
it became evident that a second gust front was being produced by the cell at the south
Soil and Atmospheric Sciences
551
Figure 4(a)
Figure 4(b)
Figure 4. Plan Position Indicator displays of (a) reflectivity and (b) mean Doppler
velocity from the Norman, OK Doppler radar for 26 April 1984, 20:21:58 CST.
end of the line. Figure 5 shows the reflectivity (5a) and Doppler velocity (5b) fields
for this gust front. The outflow is defined in the reflectivity field as a thin line echo
(cursor) with an average reflectivity of 7dBZ (lOdBZ has been added to the display).
In this case there is no pronounced radial shear in the Doppler velocity field (Figure
5b) to indicate the presence of the outflow boundary. The only evidence of a gust
front is the velocity field is the slight decrease in inbound velocities from about 23
ms"' (east of the cursor) to 23 ms"' behind the boundary (cursor).
As the storms propagate to the northeast, the parent cell of the southern gust front
continues to create a boundary which moves eastward. Figure 6 shows the reflectivity
and Doppler velocity displays of the southern gust front at 22:04:53 CST after it has
moved east of the radar. The thin line echo (Figure 6a; cursor) is still evident (average
reflectivity is 21dBZ), but a change has taken place in the velocity field (Figure 6b).
In Figure 5, it was noted that there was no radial shear line associated with this gust
front. In Figure 6b, the zero velocity line separating the positive velocities near the
radar from the negative velocities of the environmental air is quite pronounced (cur-
sor). Thus, as this gust front evolved, it developed both the thin line and radial shear
signatures.
552
Indiana Academy of Science
Vol. 94 (1985)
Figure 5(a)
Figure 5(b)
Figure 5. Plan Position Indicator displays of (a) reflectivity and (b) mean Doppler
velocity from the Norman, OK Doppler radar for 26 April 1984, 20:47:23 CST.
The Doppler radar displays are very useful not only for qualitative descriptions
of phenomena such as gust fronts and their associated signatures, but also for quan-
titative measurements of outflow characteristics such as wind speed within the outflow,
peak reflectivity along the gust front, etc. Data from ten gust fronts were collected,
tabulated, and analyzed (11) to determine the expected Doppler velocities within the
outflow, presence or absence of a thin line echo or radial shear signature and value
of Doppler radial shear at the outflow leading edge. It was reported that:
A) Doppler winds within the outflow usually never exceeded 32 ms"'.
B) The Doppler radial shear was greatest in areas where the gust front was perpen-
dicular to the radar beam.
C) A thin line echo was present in seven of the ten gust fronts and in two
of the seven, the thin line developed after the radar had begun to scan the gust front.
D) In nine of the ten cases, the gust front could be identified as a line of radial
shear in the Doppler wind field.
Soil and Atmospheric Sciences
553
Figure 6(a)
Figure 6(b)
Figure 6. Plan Position Indicator displays of (a) reflectivity and (b) mean Doppler
velocity from the Norman, OK Doppler radar for 26 April 1984, 22:04:53 CST.
Conclusions
The gust front produces low altitude wind shear which can be hazardous to air-
craft, particularly during takeoff and landing. As shown here, these outflow boun-
daries can sometimes be identified as thin lines of reflectivity or lines of radial shear
or both. The thin line echo is detected only after the gust front has separated from
the parent storm whereas the radial shear line, if present, is detectable at any stage
in the gust front life cycle. Gust fronts that do not separate from the storm are not
dangerous because pilots do not usually fly into high reflectivity areas. When an outflow
boundary moves away from the storm, its reflectivity decreases and the gust front
becomes more difficult to detect. Relying on reflectivity alone as a measure of the
potential hazard is unwise because these low-reflectivity outflows can harbor signifi-
cant, possibly dangerous wind shear. Consequently, the use of Doppler velocity is essen-
tial to adequately detect the gust front outflow. The Doppler velocity field clearly displays
the zone of wind shear associated with the thunderstorm outflows, thereby providing
a distinctive signature of the gust front. Also, the ability to detect hazardous shear
in its formative stages allows one to track the shear line as its signal strength decreases.
554 Indiana Academy of Science Vol. 94 (1985)
Such a capability can provide the pilot with sufficient advanced warning to avoid the
potential hazard of low altitude wind shear associated with thunderstorms.
Literature Cited
1. Benjamin, T.B., 1968: Gravity current and related phenomena, J. Fluid Mech.,
31, pp. 209-248.
2. Brandes, E.A., 1976: Gust front evolution in severe thunderstorms: Preliminary
investigation with Doppler radar, Preprints, 7th Conf. on Aerospace and
Aeronautical Meteor., pp. 56-61.
3. Brown, H.A., 1960: Report on radar thin lines, Proc. 8th Wea. Radar Conf.,
Amer. Meteor. Soc, Boston, MA, pp. 65-72.
4. Byers, H.R. and R.R. Braham, Jr., 1948: The Thunderstorm, U.S. Govt. Print.
Off., Washington, D.C., 287 pp.
5. Charba, J., 1972: Gravity current model applied to analysis of squall-line gust
front, NOAA Tech. Memo. ERL NSSL-61, National Severe Storms Laboratory,
Norman, OK, 58 pp.
6. Charba, J. and Y. Sasaki, 1971: G-current model applied to analysis of squall
line gust front, Preprints, 7th Conf. on Severe Local Storms, pp. 277-283.
7. Goff, R.C., 1975: Thunderstorm outflow kinmatics and dynamics, NOAA Tech.
Memo., ERL NSSL-75, 63 pp.
8. Goldman, J.L. and P.W. Sloss, 1969: Structure of the leading edge of thunderstorm
cold air outflow, Preprints, 6th Conf. on Severe Local Storms, pp 75-79.
9. Harper, W.G., 1958: Detection of bird migration by centimetric radar: A cause
of radar angels, Proc. Roy. Soc, B149, pp. 484-502.
10. Harper, W.G., 1960: An unusual indicator of convection, Marine Observer,
30:36-40.
11. Klingle, D.L., 1984: A gust front case studies handbook, U.S. Dept. of Transpor-
tation, Rept. no. DOT/FAA/PM-84/15 (ATC-129), 100 pp.
12. Laird, B.G. and J.E. Evans, 1982: FAA Weather surveillance requirements in
the context of NEXRAD, U.S. Dept. of Transportation, Report no.
FAA-RD-83-111 (ATC-112), 141 pp.
13. Lee, J.T., J. Stokes, Y. Sasaki and T. Baxter, 1978: Thunderstorm gust fronts —
observations and modeling, U.S. Dept. of Transportation, FAA Rept. No.
FAA-RD-78-145, 100 pp.
14. Leach, W., 1957: Observed characteristics of convective cell bands, Proc. 6th
Wea. Radar Conf., pp. 151-156.
15. Luckenback, G., 1958: Two examples of non-precipitating echoes as observed
on AN/CPS-9 radar, Proc. 7th Wea. Radar Conf., pp. D41-D47.
16. Simpson, J.E., 1969: A comparison between laboratory and atmospheric density
currents, Quart. J. Roy. Meteor. Soc, 95, pp. 758-765.
17. Wakimoto, R.M., 1982: The life cycle of thunderstorm gust fronts as viewed
by Doppler radar and rawinsonde data, Mon. Wea. Rev. 110(8), pp. 1060-1082.
18. Zrnic, D.S. and J.T. Lee, 1983: Investigation of the detectability and lifetime
of gust fronts and other weather hazards to aviation, U.S. Dept. of Transporta-
tion, FAA Final Rept. no. DOT/FAA/PM-83/33, 58 pp.
An Analysis of the 28 March 1984 Tornado Outbreak in the Carolinas
T.E. Klingler and D.R. Smith
Department of Geosciences
Purdue University
West Lafayette, Indiana 47907
Introduction
On 28 March 1984, a fast-moving, rapidly-deepening cyclone moved across the
southeastern United States. Upon reaching the Carolinas, the system spawned 22 tor-
nadoes which claimed 57 lives and 1248 injuries.1 Extensive damage was incurred after
the northeastward sweep of the storm across northern South Carolina through the
northeastern portion of North Carolina during the period 2130Z on 28 March to 0315Z
on 29 March (Figure 1). Of the documented tornadoes, seven were intensity F4 (Fujita
REPORTED TORNADOES
MARCH 28, 1984
- ■ - ■ ■ — ,J - -
V" /\h ifT'i
v Vr
/'
•-v ■<§■& i
\
v
s
/
x >
y V./V-'
../•
"■^'r-.-v
./
/"* \
<^r"
i.
' "~ -~ 7— ——___..
.,_.._ ____
18 ^S xl
^ .. *>
f
14 x*^ V—. ■*£'* ■/
- 7""""* '-\
1 2
l, 12/
- *>> '
\> ^
z\
Ik /
>&
\
/
Vn
<r
,,..<'
y
Figure 1. PROAM analysis region displaying 22 tornado tracks of the Carolina Out-
break. Individual tracks are numbered by time from earliest to latest.
1. Storm Data, March 1984.
555
556 Indiana Academy of Science Vol. 94 (1985)
Scale) and five were F3 making this the largest tornado outbreak in terms of casualities
and damage since the "Superoutbreak" of 3-4 April, 1974.
In this paper a case study of the 28 March outbreak is presented. The synoptic
environment is examined with emphasis focused on the analysis of mesoscale features.
The Purdue Regional Objective Analysis of the Mesoscale (PROAM) is employed for
the analysis of surface meteorological variables (1). PROAM utilizes data routinely
collected and distributed via the FAA 604 teletype circuit and produces objectively
analyzed plots for several variable fields such as temperature, pressure, specific humidity,
specific-humidity convergence, vorticity, and streamlines. PROAM has proved effec-
tive for analysis of mesoscale features in earlier case studies (2) and for identifying
severe weather signatures useful in a forecast mode (3). Hourly radar summaries from
the National Meteorological Center (NMC) are extensively used for the analysis region.
This paper will examine the Carolinas outbreak to diagnose the storm as well as to
continue to study the potential of PROAM as a tool for forecasting severe weather.
Synoptic Features
The surface low-pressure system responsible for spawning the Carolina tornadoes
was located in northeastern Texas at 0000Z 28 March with minimum pressure at 989
mb. Figure 2 displays the movement of this low-pressure center with the associated
Figure 2. Diagram displaying track of the low-pressure system from 0000Z 28 March
to 0000Z 29 March in 3-hour intervals. Frontal lobes for even hours are solid and
frontal lobes for odd hours are hollow. Central pressure values are underlined and
encoded using the standard convention (i.e., "89" = 989 mb).
frontal locations through the 24-hour period prior to the outbreak. The system initially
moved quickly eastward, but slowed as it occluded over northern Alabama. By 1500Z,
the system was centered in western North Carolina, but the minimum pressure of the
storm had changed in magnitude only slightly. By 1800Z, a second low-pressure center
began to intensify over central North Carolina. As the system shifted eastward with
the progression of the low center, the warm air sector in the western portion of the
Carolinas occupied the coastal plain east of the mountains. The warm front began
to accelerate due to decreased surface friction over the more uniform terrain (4). This
occurred after 1800Z and resulted in a flattening out of the surface frontal wave struc-
Soil and Atmospheric Sciences
557
ture. Minimum pressure values began to decrease with readings of 988 mb at 1800Z,
976 mb at 2100Z, and 978 mb at 0000Z 29 March. At 0000Z, the low center was
located at the North Carolina-South Carolina border. The warm front stretched to
the northeast into southeastern Virginia and the cold front extended to the southwest
into central Georgia.
The potential for severe weather development was supported by upper-air condi-
tions at 0000Z on 29 March (not shown). Flow was southwesterly with strong currents
throughout the troposphere. For example, Charleston, South Carolina (CHS) reported
winds of 95 kts at the 500-mb level and 120 kts at the 300-mb level (Charleston did
not report 850-mb level winds, although nearby Waycross, Georgia (AYS) reported
60-kt winds). The Lifted Index at 0000Z at 29 March at CHS was -7, which indicated
a definite severe weather threat.
Examination of the NMC National Radar Summaries (not shown) for hours
preceding the Carolina storms revealed a distinct line echo as early as 1535Z (28 March).
This line was associated with the cold front and extended from northern Alabama
to southwestern Mississippi with the maximum echo top of 31,000 ft (9450 m). At
1638Z, a tornado watch (WT055) was issued for east-central Alabama and most of
the northern half of Georgia, reaching the extreme northwestern corner of South
Carolina. Maximum echo tops increased during the next few hours. At 1835Z, the
maximum reported top was 37,000 ft (11,280 m) over central Alabama and by 1935Z,
a 40,000-ft (12,195-m) top was observed over western Georgia. A second tornado watch
(WT056) was issued at 1914Z extending from northeastern Georgia across the Carolinas
to the coastal plain. Intensification of convective activity continued as demonstrated
by increased thunderstorm development (maximum echo tops of 48,000 ft [16,463 m]
over northeastern Georgia by 2135Z).
Regional Objective Analysis
Times of the individual tornadoes are charted in Figure 1. The first reported
tornado occurred at 2130Z in northwestern South Carolina (Track 1). Tomadic storms
continued eastward and entered North Carolina at 0030Z (Track 11) and by 0155Z
were raging in the extreme northeastern corner of North Carolina (Track 18).
Figure 3 displays the analyses of the NMC National Radar Summaries translated
RADAR SUMMARY CHART
28 MAR 84 - 22357.
RADAR SUMMARY CHART
\
29 MAR 84 - 0035Z
Figure 3. NMC Radar Summaries translated onto PRO AM analysis region. Con-
tours represent echo-intensity levels of 1, 3, and 5 respectively. Maximum echo tops
in hundreds of feet are underlined.
558
Indiana Academy of Science
Vol. 94 (1985)
onto the PROAM analysis region for hours 2235Z and 0035Z. During these periods,
the development of the storm can clearly be seen by the eastward progression of the
maximum echo tops evidenced by the 5 3, 000- ft (16,154-m) top in western South Carolina
at 2235Z and the pair of 45,000-ft (13,716-m) tops in southeastern North Carolina
at 0035Z. A comparison of the radar summaries with the tornado tracks in Figure
1 shows the maximum tops from 2235Z to 0135Z nearly coincide with a reported tor-
nado. Times and locations of the maximum tops for 2235Z and 0035Z correlate very
well to tornado tracks 4 and 11, respectively.
The Radar Summaries in Figure 3 show a very distinct line echo extending from
southeastern North Carolina to central Georgia. This line was propagating along the
surface frontal position, and from 2235Z to 0035Z displayed a distinctive bend associated
with the counterclockwise circulation around the low-pressure center which was at this
time located over northern South Carolina. This was also the period of the most inten-
sive tornadic activity.
Figure 4 is the objective analysis of the surface pressure field at 2300Z. The 2100Z
analysis (not shown) indicated a pair of low-pressure troughs located over southeastern
West Virginia and northeastern Georgia, respectively. The southern trough eventually
P (MB)
28 MAR 84 - 2300Z
Figure 4. Objective analysis of surface pressure. Contour intervals are 2 mb.
Soil and Atmospheric Sciences
559
deepened and, when compared with the Radar Summaries, appeared very nearly in
the same area as the maximum echo top as it progressed eastward. A mesohigh was
situated between the troughs at 2100Z in extreme western North Carolina and was
apparently due to the outflow from the main storm. This ridge is apparent at 2300Z
shown in Figure 4 and later as a bulge in the isobars just north of the southern trough
at 0000Z and 0100Z (not shown).
An analysis of the 2300Z surface relative-vorticity field is illustrated in Figure
5. An axis of positive vorticity appeared earlier over western South Carolina at 2200Z
VxV (S"1 x 107 )
28 MAR 84 - 2300Z
Figure 5. Objective analysis of relative vorticity. Solid contours represent positive
vorticity and broken lines represent negative vorticity. Contour intervals are 2x10 ~5
s_l. Values are scaled by a factor of 107.
(not shown) and extended northeastward along the storm line. The axis becomes clear-
ly evident in Figure 5. The vorticity maximum over western South Carolina at 2300Z
(1.762x10 ~4 s~ ') agrees closely with the maximum echo top on the 2235Z Radar Sum-
mary in that area (see Figure 3). Throughout the tornadic period of the storm, this
axis of maximum vorticity was coincident with the main storm line. This is important,
because areas of strong positive relative vorticity indicate low-level convergence that
560
Indiana Academy of Science
Vol. 94 (1985)
must lead to compensating upward vertical motion. Therefore, areas of strong mesoscale
relative vorticity usually signify areas of strong convection. Storm outflow areas are
again evident in Figure 5 by the large areas of negative vorticity northwest of the positive
axis.
Figures 6 and 7 display the temperature and streamline objective analyses for
2300Z, respectively. The frontal region can clearly be seen in both analyses. The strong
temperature gradient in Figure 6 through North Carolina extending southwest to north-
east delineates the frontal boundary. The streamline analysis displays the wind shift
at the frontal zone with a distinctly southerly component to the south and a more
northerly component to the north.
An interesting comparison can be made between Figures 6 and 7 and the 2235Z
Radar Summary (Figure 3). A thermal ridge existed at 2300Z in southern North Carolina
(Figure 6). The southerly winds evident from the streamline analysis at that time (Figure
7) ensured that the storm line had an abundant supply of warm, unstable air by low-
level advection. Another feature of Figure 6 is the existence of a cold-air pocket in
extreme western North Carolina. This feature was evident throughout the period 2100Z
T (DEG F)
28 MAR 84 - 2300Z
Figure 6. Objective analysis of surface temperature. Contour intervals are 5°F.
Soil and Atmospheric Sciences
561
to 0100Z. It appeared in the same area as the mesoscale pressure ridge shown in Figure
4 and is further evidence of a storm outflow region.
Objective analyses of specific humidity (not shown) indicate an abundant supply
of moisture in the warm sector of this system. A ridge of specific humidity was present
in eastern South Carolina from 2100Z through 0100Z. The specific humidity in the
ridge exceeded 15 g/kg throughout the period. This warm, moist air in this region
provided the fuel for the intensification of the storm line producing the severe weather.
Figure 8 illustrates the series of objective analyses of specific-humidity convergence
for the period 2200Z to 0100Z. This parameter, when compared to the Radar Sum-
mary Charts, provides an excellent indicator of the presence of severe weather. At
2200Z, an axis of maximum-moisture convergence appeared at the western edge of
South Carolina. This feature matches the line of maximum echoes at 2135Z (not shown).
The magnitude of the specific-humidity convergence maximum at 2200Z was 2.66x10- 3
g/kg s- '. A ridge in the moisture convergence field is very distinct at 2300Z, forming
an axis along nearly the same position as the line echo of the 2235Z Radar Summary.
The magnitude of the maximum value at 2300Z was 3.73x10 3 g/kg s '. Such a
STREAMLINES
28 MAR 84 - 2300Z
Figure 7. Objective analysis of surface streamlines.
562
Indiana Academy of Science
Vol. 94 (1985)
-V-(VQ) (G/KG S"1 x 10* ) 28 MAR 84 - 2200Z -V-(VQ) (G/KG S'1 x 10* ) 28 MAR 84 - 2300Z
-V(VQ) (G/KG S"1 x 10*) 29 MAR 84 - 0000Z -V-(VQ) (G/KG S"1 x 10*) 29 MAR 84 - 0100Z
Figure 8. Objective analysis of surface specific-humidity convergence. Solid contours
represent convergence and broken contours represent divergence. Contour intervals
are 1x10 ~4 g/kg s~ '. Values are scaled by a factor of 104. Note the progression of
maxima during the tornadic period of the storm.
significant short-term increase has been found to indicate an intensification in storm
activity (2). This case confirmed the earlier work because just after 2200Z, the magnitude
of the ridge had decreased slightly, but the maximum again appears in the region of
most intense radar echoes. An area of specific-humidity divergence was seen consistently
throughout the period in western North Carolina. Along with the evidence presented
from the temperature, pressure, and vorticity analyses, this again gives a good indica-
tion of a storm outflow region.
PROAM analyses greatly aided in the study of the Carolinas outbreak. The tem-
poral and spatial resolution of the surface-variable fields used enhanced the ability
to diagnose conditions prior to and during tornadic events on 28 March. It is hoped
that further study of this case will continue to yield more evidence to enable forecasters
to more accurately predict severe local storms.
Soil and Atmospheric Sciences 563
Summary
The 28 March Carolinas Tornado Outbreak claimed more lives than any out-
break since 1974. Seven F4 and five F3 tornadoes were generated, resulting in 57 fatalities
and over 1200 injuries. A deep, fast-moving low-pressure system provided the trigger
for the development of an intense line of storms, many of which were tornadic.
PROAM analyses were very useful in discussion of the mesoscale features of the
outbreak. Radar Summary Charts indicating precipitation-echo intensities as well as
maximum echo could be compared to objectively analyzed parameters for a complete
picture of the storm development. Specific humidity convergence was the most impor-
tant variable analyzed, because it directly indicated the presence of severe weather.
This provides further evidence for the use of PROAM, in conjunction with other data,
as a tool for short-term prediction of severe weather.
Literature Cited
1. Smith, D.R. and F.W. Leslie. 1984. Error determination of a successive correc-
tion type objective analysis scheme. J. Atmos. and Ocean. Tech. 1:120-130.
2. Smith, D.R. and S.D. McCauley. 1983. Mesoanalysis of the surface features
associated with the Shelby County, Kentucky tornado (20/21 March 1982). Preprints
of the 13th Conference on Severe Local Storms, Tulsa, OK, AMS. 308-311.
3. Snow, J.T., D.R. Smith, F.W. Leslie and R.H. Brady. 1983. Mesoanalysis of
surface variables associated with the severe weather of 9-10 July 1980. Nat. Wea.
Dig. 8: 28-39.
4. Spar, J. 1956. An analysis of a cyclone on a small synoptic scale. Mon. Wea.
Rev. 84:291-300.
Response of Forage Crops to Dolomitic Lime
Ana L. Pires, J.L. Ahlrichs and C.L. Rhykerd
Instituto Universitario de Tras-os-Montes e Alto Douro, Vila Real, Portugal
and Agronomy Department, Purdue University, West Lafayette, Indiana 47907
Introduction
Most pastures of the Tras-os-Montes region in Northern Portugal, are established
on acid soils. These soils normally have very low levels of exchangeable Ca and Mg
which impede the normal development of plants and can also cause animal health
problems. Grass tetany, a metabolic disorder in ruminants related to a low level of
Mg in forages, has been frequently observed in this region, especially in late winter
or early spring.
The objective of this study is to verify whether correcting the acidity of a represen-
tative soil from this region with dolomitic lime improves the quality and yield of ryegrass,
tall fescue, white clover and subterranean clover. The concentration of Mg in the forage
will be of special interest. A value greater than 0.20% (10) is commonly accepted as
the level where the occurrence of grass tetany is diminished. However if the forages
have a high K concentration, greater than 3%, it is suggested that the Mg level should
be higher than 0.20% (5).
Materials and Methods
The experiment was conducted in polyethylene pots containing 4 kg of air dry
soil (< 5 mm fraction) taken from a sandy loam xerofluvent whose characteristics
are found in Table 1. The forage species tested were Trifoliwn subterraneum L., cultivar
Table 1. Chemical properties of the soir1'.
PH
(H20)
Ca+ +
Mg+ +
Na +
K+ H+ +
A1++ +
CEC
Base Organic
saturation matter
P,Os
K20
_ %
- ppm -
>200
5.3
1.32
0.20
0.04
0.25 8.16
9.83
16.99 1.53
140
(1) top 20 cm; pH in 1:2.5 soil solution ratio; Exch. bases by 1 N ammonium acetate pH 7.0; Exch. H + and Al + + +
by barium chloride-triethanolamine, pH 8.0; organic matter by method of Tinsley (C x 1.724); P20, and K20 by
method of Enger-Riehm.
Clare (subterranean clover), Trifoliwn repens L., cultivar G. Huia (white clover), Festuca
arundinacea, cultivar Manade (tall fescue) and Lolium perenne x L. multiflorum, cultivar
G. Manawa (hybrid ryegrass). The forages were sown in November and each species was
thinned to 25 plants per pot. The legume seeds were inoculated with Rhizobia.
Each species were subjected to soil treatments of three levels of dolomitic lime
(63.5% CaC03 and 37.8% MgC03): 0, 15 or 30 g/pot which corresponds to 0, 7500
and 15000 kg/ha, respectively. All treatments were replicated four times. A nutrient
solution was applied before the species were sown. The quantities, added in mg/pot
were: N 840 for grasses and 112 for legumes, P 414, K 520, S 100, B 2.7, Mo 0.1,
Zn 13.2, Cu 3.2, Mn 11.0, Fe 20.0 (7).
The plants were irrigated daily with distilled water up to two-thirds of their
maximum water retention capacity.
565
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Indiana Academy of Science
Vol. 94 (1985)
The tall fescue, ryegrass and subterranean clover herbage were cut at 3 cm above
the soil in March, April and May. The white clover herbage was cut in April, May
and June. All species were flowering at the time of their last cutting.
The herbage was dried at 60°C for 24 hours for the determination of dry matter
production, and ground to pass a 1-mm screen. The ground tissue was digested using
nitric-perchloric acid and Ca and Mg determined by atomic absorption spectrophotometry
and K by flame photometer. After the final harvest, the soil was analyzed for pH
to determine if a shift in pH occurred.
Results and Discussion
Dolomitic lime significantly increased (P < 0.05) the total D.M. production of
all the species (Table 2). A positive yield response occurred for all white clover cuttings,
Table 2. Effect of species and rate of dolomitic lime on forage dry matter production
('.)
Species and
rate of
Cutting
dolomitic
lime*2'
1
2
3
Total
g/pot
Tall fescue
0 g/pot
5.40
9.91
15.65
30.96
15
4.80
11.50
20.83
37.13
30
5.12
12.20
18.61
35.93
LSD (0.05)
n.s.
1.78
4.87
5.52
Ryegrass
0 g/plot
9.38
12.58
23.55
45.51
15
9.34
13.77
27.65
50.76
30
10.29
14.13
30.56
54.98
LSD (0.05)
n.s.
1.37
3.45
4.15
Sub. clover
0 g/plot
5.42
9.12
8.15
22.69
15
6.45
12.81
14.00
33.26
30
6.43
12.84
14.97
34.24
LSD (0.05)
n.s.
3.56
3.35
4.14
White clover
0 g/pot
4.58
3.28
3.96
11.82
15
8.35
14.44
13.26
36.05
30
8.63
14.86
16.54
40.02
LSD (0.05)
1.28
3.41
3.40
6.80
(1) Values are means of 4 replications
(2) g/pot
with an increase in D.M. production from the first level of lime of 82%, 340% and
234% for the first, second and third cuttings, respectively. The other species had signifi-
cant yield responses only for the second and third cuttings. Fescue increased 15 and
33%, ryegrass 9 and 17% and subterranean clover 41 and 77%, respectively.
While all species responded well to the liming treatments the total yield of all
cuttings of tall fescue, white clover and subterranean clover did not yield significantly
higher (P < 0.05) at the 30 g than at the 15 g treatment. However ryegrass production
did respond to the application of 30 g/pot (P < 0.05). The increase obtained at the
Soil and Atmospheric Sciences 567
first level of liming was 20%, 47%, 205% and 12%, respectively for the four species.
The ryegrass had a 20% increase at the second level of liming. The average final pH
obtained with 0, 15 and 30 g of dolomitic lime per pot was 4.9, 6.0 and 6.6, respectively
(Table 3).
Table 3. Effect of species and rate of dolomitic lime on final soil pH values^1)
Treatments Fescue Ryegrass Sub. clover White clover
g/plot — - — soil pH -
0 4.7 4.8 4.7 5.2
15 6.2 6.3 5.8 6.0
30 6.7 6.6 6.6 6.7
(1) Values are means of 4 replications
The increase in D.M. production with dolomitic lime is probably due to an im-
provement of the general conditions of nutrition of which the most commonly sug-
gested are an improved assimilation of P and a decrease in the solubility of Al and
Mn (11, 16, 17). Although no data was recorded, the treatments in which the acidity
was corrected had longer roots with much more branching and root hair development
and the number of nodules on the clovers were much greater. It has been verified
that nodulation is affected not only by the acidity of the medium but also by the
lack of Ca (1).
While Ca concentration did increase with dolomitic lime application, the increase
was not significant (P < 0.05) for the two grasses tested, with the exception of the
second cutting of tall fescue (Table 4). For all treatments, the Ca concentrations that
ranged from 0.40 to 0.67% in the grasses and from 0.76 to 1.89% in the clovers are
generally considered as sufficient for a good plant growth (1, 13).
The Mg concentration was significantly improved (P < 0.05) from that obtained
in the control (Table 4) when dolomitic lime was applied, although the analytical values
were generally higher in the 30 than in the 15 g/pot treatment the differences were
not significant at the P < 0.05. The average Mg concentration of tall fescue and ryegrass
increased from 0.11 and 0.10% in the control to 0.33 and 0.24%, respectively in the
15 g lime treatment. In the white clover and subterranean clover, it increased from
0.18% in both controls to 0.42 and 0.51%, respectively in the 15 g treatment. In all
cuttings, the Mg levels obtained with the dolomitic lime treatments were greater than
0.20, the value commonly accepted as sufficient for ruminant nutrition.
The levels of Mg found in the two grasses on the unlimed soil were very low
(0.09 to 0.12%), but no Mg deficiency symptoms were observed. The symptoms usually
occur, according to Grunes et al. (5) when the percentage of Mg in the D.M. varies
between 0.05 and 0.10. Embleton (3), however, referred to somewhat higher Mg values.
The Mg level for the clovers were much higher than those of grasses on the unlimed
soil and only in the first cutting of the white clover and third cutting of the subterra-
nean clover did the Mg concentration, 0.14 and 0.11% respectively, approximate the
plant deficiency value of 0.12-0.14% (13).
The occurrence of low tissue Mg is reasonable since the level of exchangeable
Mg in the soil, 0.20 meq/100 g, is considered low (2). Hogg and Karlovsky (8) stated
that when the exchangeable Mg is less than 0.20 meq/100 g of soil, a deficiency should
be expected. Felbeck (4) suggested that application of Mg should be recommended
when the level of exchangeable Mg is less than 0.41 meq/100 g of soil. Horvath and
Todd (9) recommended that for good plant growth the exchangeable Mg level ought
568
Indiana Academy of Science
Vol. 94 (1985)
Table 4. Effect of species and rate of dolomitic lime on the concentration of Ca, Mg
and K in herbage dry matter^1)
Species and
Calcium
Magnesium
Potassium
rate of
dolomitic
lime^!'
1
Cutting
2
3o)
1
Cutting
2
3«
1
Cutting
2
3(0
0/q
Fescue
0
0.40
0.43
0.40
0.12
0.10
0.10
3.76
2.09
1.29
15
0.41
0.61
0.43
0.29
0.36
0.35
3.51
2.19
1.16
30
0.47
0.54
0.43
0.32
0.35
0.36
3.78
2.21
1.35
LSD (0.05)
n.s.
0.08
n.s.
0.06
0.02
0.02
n.s.
n.s.
n.s.
Ryegrass
0
0.43
0.67
0.42
0.09
0.11
0.09
3.69
1.86
1.14
15
0.44
0.67
0.47
0.21
0.27
0.24
3.93
2.04
1.23
30
0.46
0.67
0.44
0.23
0.30
0.25
3.67
2.28
1.34
LSD (0.05)
n.s.
n.s.
n.s.
0.02
0.03
0.04
n.s.
n.s.
0.17
Sub. clover
0
1.12
0.78
0.76
0.24
0.18
0.11
3.00
2.64
2.34
15
1.26
1.39
1.59
0.49
0.48
0.55
2.42
2.05
1.26
30
1.48
1.38
1.89
0.52
0.48
0.61
2.44
2.30
1.36
LSD (0.05)
0.13
0.15
0.32
0.05
0.05
0.07
0.41
n.s.
0.59
White clover
0
1.12
1.25
1.20
0.14
0.21
0.20
3.39
3.57
3.42
15
1.34
1.37
1.34
0.35
0.40
0.51
3.20
2.10
1.45
30
1.50
1.54
1.60
0.36
0.42
0.56
3.36
1.19
1.54
LSD (0.05)
0.28
n.s.
0.24
0.05
0.06
0.08
n.s.
0.54
0.95
(1) Values are
means
of 4 replication!
(2) All species
exhibit flowers
(3) g/pot
\^
to be at least twice the exchangeable K+ level and the Ca:Mg ration ought to be
5:1. In this soil, the Mg level was less than the K level and the Ca:Mg ratio was 6.6:1.
Conforming to what would be expected (13, 14, 18, 19), the Ca and Mg concen-
trations of the clovers were superior to those of grasses. However, the difference in
levels of Mg between the grasses and clovers is not as accentuated as in the case of
Ca, a fact already observed by Metson and Saunders (14). The fact that the legumes
have greater concentrations of Ca and Mg than the grasses points out that the risk
of grass tetany would be less in legume or grass-legume pastures.
The K concentration in the herbage was also determined (Table 4) because K
utilization could be less because of the dolomitic lime application. The K concentra-
tion of the treatments in which the acidity liad been corrected was not significantly
different (P < 0.05) than those found in the control, with exception of the second
and third cuttings of white clover which decreased 41% and 58% respectively, the
third cutting of subterranean clover which decreased 48% and the third cutting of
ryegrass which increased 8% when the percentage change is calculated from comparison
of the control with the 15 g liming treatment. The K concentration decreased in general
from the first to the final cutting. In the final cutting, the K level in the grasses was
very low, ranging from 1.14 to 1.35%, but no deficiency symptoms were observed.
These symptoms usually occur when the percentage of K is less than 2.2% (15, 22).
The K concentration in the clovers is considered medium (12, 21).
Soil and Atmospheric Sciences 569
The great decrease in the K levels in the 2nd and 3rd cuttings is due perhaps
to the fact that the K, in contrast to Ca and Mg, is a nutrient with major seasonal
variation, having marked monthly fluctuations which are in general similar in grasses
and legumes (14). Generally the levels of K in forages attains a maximum level at
the end of winter, decreasing after this until the beginning of summer at which time
it attains its lowest value (6, 14).
Conclusions
The results obtained showed that the correction of acidity by the use of dolomitic
lime (63.5% CaC03 and 37.8% MgCO,), produced a quantitative and qualitative im-
provement in production. The benefit of this action is due to the general improvement
of the nutritional status of the soil, shown by the pH (H20) rise to average values
of 6.0 (15 g/pot) and 6.6 (30 g/pot), and to an increase in the available Ca and Mg
for the plants.
The total D.M. production of tall fescue, subterranean clover and white clover
increased 20, 47 and 205%, respectively, with the addition of 15 g/pot of dolomitic
lime. Ryegrass, unlike the other forages, gave additional increase in production with
30 g of dolomitic lime per pot (P < 0.05).
The qualitative improvement of production refers essentially to the increased Mg
concentration in the D.M. to values greater than 0.20%, a value which is considered
sufficient for good plant development as well as for supplying animal needs. The average
Mg concentration of tall fescue, ryegrass, subterranean clover and white clover in-
creased to 0.33, 0.24, 0.51 and 0.42% with the application of 15 g/pot of dolomitic
lime (P < 0.05).
Pastures established on the acid soils of the Tras-os-Montes region with low levels
of exchangeable Ca and Mg should have the soil pH corrected with dolomitic lime
and not with calcitic lime. Legumes contain considerably higher concentrations of Ca
and Mg than grasses and should always be grown in pastures whenever possible in
order to reduce the risk of incidence of grass tetany.
Literature Cited
1. Chapman, H.D. 1973. Calcium. In "Diagnostic Criteria for Plants and Soils."
p. 65. Ed. H.D. Chapman, Univ. of California, Riverside, U.S.A.
2. Bolton, J. 1972. Effects of potassium, magnesium and sodium fertilizers and lime
on the yield and composition of crops in a ten year experiment at Rothamsted.
Rep. Rothamsted Exp. Stn. 2:102-110.
3. Embleton, T.W. 1973. Magnesium. In "Diagnostic Criteria for Plants and Soils."
p. 225. Ed. H.D. Chapman, Univ. of California, Riverside, U.S.A.
4. Felbeck, G.T. 1959. In "Magnesium and Agriculture." Ed. G.C. Anderson, E.M.
Jencks, and D.J. Horvath, pp. 96-105. West Virginia Univ., Morgantown, West
Virginia, U.S.A.
5. Grunes, D.L., P.R. Stout and J.R. Brownell. 1970. Grass tetany of ruminants.
Advances in Agronomy 22:331-374.
6. Hannaway, D.B., L.P. Bush and J.E. Leggett. 1980. Plant Nutrition: Magnesium
and Hypomagnesemia in Animals. Univ. of Kentucky. Agric. Exp. Station Bulletin
716.
7. Hewitt, E.J. 1966. Nutrient reagents. In "Sand and Water Culture Methods Used
in the Study of Plant Nutrition" C.A.B. p. 430 (2nd ed.) England.
8. Hogg, D.E. and J. Karlovsky. 1968. The relative effectiveness of various Mg
fertilizers of a Mg-deficient pasture. N.Z. Agric. Res. 11:171-183.
9. Horvath, D.J. and J.R. Todd. 1968. Magnesium supplements for cattle. Proc.
570 Indiana Academy of Science Vol. 94 (1985)
23 Texas Nut. Conf. Texas Agric. Exp. Stn., College Station, pp. 96-104.
10. Kemp, A. 1960. Hypomagnesaemia in milking cows: the response of serum
magnesium to alterations in herbage composition resulting from potash and nitrogen
dressing on pasture. Neth. J. Agric. Sci. 8:281-304.
11. Lopez-Hernandez, D. and C.P. Burnham. 1974. The effect of pH on phosphate
adsorption in soils. J. Soil Sci. 25:207-216.
12. Martin, W.E. and L.J. Benny. 1971. Field evaluation of CSPS on rangeland clovers.
13. McNaught, K.J. 1970. Diagnosis of mineral deficiencies in grass-legume pastures
by plant analysis. Proc. XI International Grassl. Congress, pp. 334-338.
14. Metson, A.J. and W.M.H. Saunders. 1978. Seasonal variations in chemical com-
position of pasture. Part one — Ca, Mg, K, Na and P. N. Z. J. Agric. Res.
21:341-353.
15. Reid, R.L., E.K. Odhuba, and G.A. Jung. 1967. Evaluation of tall fescue pasture
under different fertilization treatments. Agron. J. 59:256-271.
16. Santos, J.Q. 1976. Infuencia da calgem e da adubacao na cultura do bersim.
Sep. do Vol. XXXVI dos Anais do I.S.A. pp. 71-82. Lisboa.
17. Sims, J. and B.G. Ellis. 1983. Adsorption and availability of phosphorus follow-
ing the application of limestone to an acid, aluminous soil. Soil Sci. Am. J.
47:888-893.
18. Todd, J.R. 1961. Mg in forage plants. Part one — Mg contents of different species
and strains as affected by season and soil treatment. J. Agric. Sci. 56:411-415.
19. Turner, M.A., V.E. Neall and G.F. Wilson. 1978. Survey of Mg content of soils
and pastures and incidence of grass tetany in three selected areas of Taranaky.
N. Z. J. Agric. Res. 21:583-592.
20. T.V.A. 1976. Greenhouse Techniques for Soil-Plant-Fertilizer Research. Bulletin
y-104 (May), Nat. Fert. Dev. Center.
21. Ulrich, A. 1945. Critical phosphorus and potassium levels in ladino clover, Soil
Sci. Soc. Amer. Proc. 10:150-161.
22. Widdownson, F.V., A. Penny and R.J.B. Williams. 1965. An experiment measuring
effects of N, P and K contents of grass. J. Agric. Sci. 64:93-99.
A Wind Tunnel Investigation of Roughness Parameters for
Surfaces of Regularly Arrayed Roughness Elements
Wayne F. Rostek, Jr. and John T. Snow
Department of Geosciences
Purdue University
West Lafayette, Indiana 47907
Introduction
A few recent studies (5, 7) have attempted to assess the impact of surface roughness
on laboratory simulated tornado-like vortices. In these studies, the roughness proper-
ties of the surface have been described only qualitatively, or estimated by semi-empirical
methods (6, 9). The parameter most frequently cited to quantify the degree of roughness
of a surface is the roughness length, z0, because it is an invariant characteristic of a
given surface and is dependent only upon its physical properties.
Dessens (5), in a study using small pebbles to form a rough surface, applied Lettau's
(9) formula to estimate z0- Leslie (7), assumed z0 to be about 1/30 of the height of the
individual roughness elements. However, Leslie did not account for the area density
of the elements. He did recommend that it would be desirable to evaluate z0
qualitatively. This provided the motivation for the current investigation, which is the
first part of a two phase research project aimed at determining the effect of varying
degrees of surface roughness on vortices produced in the Purdue Tornado Vortex
Chamber (TVC). (The reader is referred to Church, et al. (3) for a schematic diagram
of the Purdue TVC.)
Presented here are preliminary results of an experiment to estimate the roughness
length, z0> friction velocity, V*, and zero plane displacement, D for several surfaces of
regularly arrayed roughness elements. This investigation has been performed to develop
a set of standard, reproducible rough surfaces for which the above parameters (in
particular, the roughness length) are known. These surfaces will then be used to deter-
mine the effects variations in surface roughness have on tornado-like vortices.
Simulating the Atmosphere
To study the effects of surface roughness on tornado-like vortices, the roughness
properties of the surface to be used in TVC must be properly scaled to those of sur-
faces likely to be encountered in nature. The roughness properties of a surface affect
the overlying swirling flow by modifying the properties (in particular, the depth of
5) of the boundary layer that feeds into the base of the vortex. If 5 is directly related
to z0, the desired similarity can be obtained by matching the ratio z0/6 in the TVC to
that in the atmosphere.
Zo| - _Zo I ...
6|ATM 5 |TVC. l '
For sink-type flow in the TVC, the boundary layer depth has been found by
Baker (1) to be given by
6TVC = 7.5 r Rer ~°-\ (2)
where r is the radial distance inward from the outer edge of the lower surface, and
Rer is the radial Reynolds number given by
571
572 Indiana Academy of Science Vol. 94 (1985)
Rer = U0ro/v, (3)
where U0 is the radial velocity at r0,
r0 is the radius of the updraft hole, and
v is the kinematic visosity.
By Eq. 2, using typical values for U0 and r0, the boundary layer depth in the TVC at
r0 has a range from 6 to 19 cm. Cermak (2) indicates that a typical boundary layer depth
of the atmosphere is of the order of 500 m. With these values, the ratio of the depth
of the atmosphere boundary layer to that of TVC boundary layer is between 3000
and 8000. Naturally occurring roughness lengths for the Earth's surface are nominally
less than 2 m. (It is generally assumed that a highly urbanized area has z0 ~ 2 m.)
Using this maximum value to obtain a bound for similarity in the inflow layer, Eq.
1 indicates that z0 in the TVC should be no greater than 0.08 cm.
Experimental Procedure
Since the direction of the surface inflow varies in the TVC, symmetric roughness
elements must be used. Also, distribution of the elements over the surface must be
reasonably uniform in all directions. These two conditions ensure that the surface
roughness characteristics will be independent of the inflow angle. Further, the individual
roughness elements must be reproducible and uniform in size and shape. We have
chosen to use 0.64 cm diameter cylindrical pegs mounted on commercial pegboard.
It was impractical to directly investigate the roughness properties of the roughened
disk due to its size. Instead, a test surface of similar characteristics was fabricated
by mounting pegs on a sheet of pegboard 122 by 244 cm, and having a square arary
of holes on 2.54 cm centers. The pegboard surface and the wooden pegs were given
two coats of marine varnish before assembly. This test surface was then placed in
the working section of a large wind tunnel. Once in place, the peg area density (number
of pegs per unit area) could be variable by systematically removing pegs.
Figure 1 is a view looking downstream in the wind tunnel and it shows the sur-
face completely covered with some 4600 pegs (0.155 pegs cm-2) of height 1.27 cm.
The test surface has been raised above the lower surface of the wind tunnel to escape
the surface boundary layer. Vertical profiles of the horizontal wind in the boundary
layer over the surface were measured using a hot film anemometer system. The measured
uncertainty of the velocities was ± 1 .0 cm s _ ' . The anemometer probe was rigidly
attached to a remotely controlled movable arm in the wind tunnel (Figure 2). The
movable arm was connected to a transducer which produced a voltage proportional
to the position of the arm. This system was calibrated with respect to height to deter-
mine each measurement level to within ±0.1 cm.
To obtain a velocity profile in the surface boundary layer, the probe was posi-
tioned 219 cm downstream of the leading edge of the test surface. The freestream
velocity was then set to a nominal value of 10 m s-'. This value was selected to
give a good signal throughout the boundary layer. The noted freestream velocity and
the distance of the measurement position downstream from the leading edge gives a
Reynolds number of 1.4 x 106 (the average wind tunnel temperature was 23°C). The
flow in the boundary layer over the surface was therefore very turbulent. The probe
was then moved down through the boundary layer and measurements were taken at
various levels. These levels were chosen on the basis of the "double-levels" rule described
by Lettau (8).
To obtain a good estimate of the mean velocity, the output signal of the
anemometer was first linearized, then filtered with a passive first-order filter having
Soil and Atmospheric Sciences
573
Figure 1 . Test surface completely filled with pegs looking downstream into the working
section of the wind tunnel. The wind tunnel carriage with hot-film anemometer probe
is in the background.
Figure 2. Close-up of the instrumented carriage. The sensing probe is held in the
clamp that extends to the left of the arm.
574 Indiana Academy of Science Vol. 94 (1985)
a 5 s time constant. Preliminary measurements showed that below two peg heights
above the surface, the air flow reflected details of the flow around individual pegs.
For this reason, measurements were taken only above two peg heights.
Reduction of Data
A modified form of the log-linear wind profile that is appropriate for rough sur-
faces is given by
U(z) = V* In (z - D), (4)
k z0
where U(z) is the velocity at some height z,
V* is the friction velocity,
k is von Karman's constant (= 0.4),
z0 is the roughness length, and
D is the effective obstacle height, = d _ z0 where
d is the zero plane displacement.
Figure 3 is a sketch of a typical boundary layer wind profile; superimposed is
the corresponding "best fit" log-linear profile. There are three important regions in
this idealized profile. Region I is the flow in the canopy region. It is in this region
that the wind interacts with the individual roughness elements. Region II is where the
flow reflects the integrated effects of the full rough surface. In this region, the shear
stresses are approximately constant (10). It is in this constant stress layer that the log-
linear wind profile is valid and closely coincides with the observed boundary-layer wind
profile. Region III is the freestream region of the profile where the velocity becomes
nearly independent with height.
Figure 3 also illustrates the relationships between D, z0, and d. The displace-
ment, d, is obtained by extrapolating the log-linear wind profile downward to the height
when U = Orns-1. The extrapolation is shown as the lower dashed portion of the
curve in Figure 3. The effective height D is related to the area density of the obstacles.
As more obstacles are added to a given area, the value of D approaches the height
of the obstacles. The parameter z0 is the difference between d and D.
The friction velocity, V*, is equal to (-u 'w ')0-5, where (-u 'w ') is the Reynolds
stress. Therefore, the constant stress region is also a region of constant V*. From
mixing length theory, the differential equation for the wind profile is given by:
(— )
V z-D/
du = 1_ |
dz k
Rearranging Eq. 5 to obtain an expression for V* and using a centered finite dif-
ference scheme to estimate du/dz from the measurements,
V* = (z - D)k Au. (6)
Az~
Values of V* can be computed in this manner, and a region of nearly constant friction
velocity (therefore constant stress) identified. Eq. 4 can then be fit to the portion of
the observed velocity profile that exhibits a reasonably constant V* to estimate the
roughness parameters.
Soil and Atmospheric Sciences
575
d
D
Observed Profile
Log Profile
Region of Interest
(Constant Stress)
PI
PI
u
Figure 3. Idealized boundary layer profile with a corresponding log-linear wind pro-
file. The hatched boxes represent typical roughness elements. The three regions are:
I., canopy flow; II., constant stress flow; III., freestream flow
Figure 4 is a typical wind profile for the most dense array of pegs (0.155 pegs
cm 2). Notice that the velocity at high levels is approaching a constant value in agree-
ment with Figure 3. Since the lowest measurement level is approximately two peg heights
above the surface, the flow in the canopy does not appear in this profile. Figure 5
is a profile of friction velocity computed using Eq. 6. The values were computed for
the data shown in Figure 4, and then smoothed using a five point moving average.
Figure 5 shows that a region of nearly constant V* exists between 3 and 6 cm.
The three roughness parameters were estimated by means of a non-linear least
squares fit of the log-linear profile to the data. This fitting was accomplished using
Indiana Academy of Science
Vol. 94 (1985)
10
11
Figure 4.
(peg area
6 7 3 9
UELOCITY (M/S)
Observed boundary layer wind profile for the most dense array of pegs
density = 0.155 pegs cm-2).
a numerical routine similar to that of Covey (4). This scheme first minimizes the square
error in the velocities by picking successive values of D. Once this has been done,
the values of z0 and V* are computed along with the standard errors for D, z0 and
V*. By fitting the log-linear wind profile to the observed data in Figure 4 only in
the constant stress region (Figure 5), the roughness parameters are found to be:
V* = 1.41 ± 1.43 ms ',
D = 0 ± 1.73 cm, and
z0 = 0.565 ± 0.70 cm.
The large standard errors are due in part to large turbulence at low levels. Even though
the linearized output from the anemometer was filtered, obtaining a mean velocity
value was difficult. In the future, work such as this should make use of computer
data acquisition to more objectively determine mean velocity values.
Results
Table 1 gives the results for four surfaces using 1 .27 cm high by 0.64 cm diameter
pegs, plus results for a pegboard surface and a plywood surface (the plywood surface
is the "smooth" surface which most closely resembles the normal TVC surface). These
results are as one would expect in that both V* and z0 decrease with decreasing peg
area density. The effective obstacle height is zero, even in the case of the most dense
array of pegs tested, because the maximum peg/surface interface area is still a small
fraction of the entire test surface.
Soil and Atmospheric Sciences
577
13
12
11
/sie
o 9
8
o
M 6
UJ
i 5
4
3
0
6. 155 Pegs/-=.q. cm.
2u
,:•
I | I I I I
I I I I
I I I
I I I I I
.5 1 1.5 2 2.5
FRICTION UELOCITY (M/S)
Figure 5. Friction velocity profile corresponding to velocity profile in Figure 4. Values
are obtained using Eq. 5.
Area Density
Peg cm 2
V*
m s '
D
em
z
0
cm
zQ (ATM)
m
0.155
1.44
0
0.565
33.2
0.078
1.37
0
0.460
27.0
0.039
1.17
0
0.278
16.3
0.019
0.94
0
0.144
8.5
PEGBOARD
0.50
0
2.41xl0~3
0.14
PLYWOOD
0.44
0
5.29x10" 4
0.03
Table 1. Roughness characteristics of the family of surfaces which use 1.27 cm high
by 0.64 cm diameter cylindrical pegs as roughness elements. The rightmost column
is
zQ based on TVC conditions of r(
40 cm and Rer = 5 x 10\
The last column in Table 1 shows z0 scaled up to the atmosphere for TVC condi-
tions of r0 = 40 cm and Rer = 5 x 103 giving <5TVC = 8.5 cm. Recalling from above
that for the natural surfaces, z0 < 2 m, one can see that the test surfaces considered
to this point correspond to extremely rough surfaces in the atmosphere. Because of
this, we are now examining surfaces with pegs that are only half as tall as those used
in this study. Preliminary results indicate that surfaces with these shorter pegs will
be better scaled to the atmosphere by being within the desired upper limit of z0 <^
0.08 cm.
578 Indiana Academy of Science Vol. 94 (1985)
Summary
A family of surfaces, each consisting of a field of cylindrical pegs of uniform
size arrayed in a regular pattern, has been developed to simulate the rough surface
of the earth. By measuring the velocity profile in the boundary layer above these sur-
faces when installed in a wind tunnel, then fitting a log-linear profile to the constant
stress portion of the wind tunnel boundary-layer profile, values for z0, V*, and D
have been obtained for each surface. Since z0 is dependent only upon the surface itself,
it can be used to characterize each surface.
As expected, the values of z0 decreased with decreasing peg area density. However,
the z0 values for the 1.27 cm high pegs were all "large" when scaled to the atmosphere.
Because of this, additional surfaces with pegs that are only half as tall as now being
examined. Preliminary results indicate that the roughness properties of these additional
surfaces will be better scaled to the atmosphere.
The authors would like to acknowledge Mr. R.L. Pauley for his assistance and
support. They also thank Dr. J. Katz, Dr. W.L. Wood, and Mr. D.L. Cochran for
allowing the use of the wind tunnel located in the Hydromechanics Lab of the Civil
Engineering Building at Purdue University. This work is supported by the National
Science Foundation under grant ATM 82-03757.
Literature Cited
1. Baker, G.L.: 1981: Boundary layers in laminar vortex flows, Ph.D. thesis, Purdue
University, W. Lafayette, IN, 143 pp. (Available from University Microfilms,
Inc., Ann Arbor, MI, Order Number DA 82-10153).
2. Cermak, J.E., 1971: Laboratory simulation of the atmospheric boundary Layer,
AIAA Journal 9, 1746-1754.
3. Church, C.R., J.T. Snow, G.L. Baker, and E.M. Agee, 1979: Characteristics
of tornado-like vortices as a function of swirl ratio: a laboratory investigation,
J. Atmos. ScL, 36, 1755-1776.
4. Covey, W., 1963: A method for the computation of logarithmic wind parameters
and their standard errors, Production Research Report No. 72, U.S. Dept. of
Agriculture, 28-33.
5. Dessens, J., 1972: Influence of ground roughness on tornadoes: a laboratory simula-
tion, J. Appl. Meteor., 11, 72-75.
6. Kondo, J., 1971: Relationship between the roughness coefficient and other
aerodynamic parameters, J. Meteor. Soc. Japan, 49, 121-124.
7. Leslie, F.W., 1977: Surface roughness effects on suction vortex formation: a
laboratory simulation, J. Atmos. Sci., 34, 1022-1027.
8. Lettau, H.H., 1967: Problems of micrometeoroligcal measurements (on degree
of control in out-of-doors) experiments), in The Collection and Processing of
Field Data, (E.F. Bradley and O.T. Denmead, eds.) Interscience Publishers, Div.
of J. Wiley and Sons, 597 pp.
9. , 1969: Note on aerodynamic roughness parameter estimation on the basis
of roughness-element description, /. Appl. Meteor., 84, 828-832.
10. Raupach, M.R., A.S. Thorn, and I. Edwards, 1980: A wind tunnel study of tur-
bulent flow close to regularly arrayed rough surface, Boundary-Layer Meteorol,
18, 373-397.
A Comparison of Soils on Unreclaimed 1949 Indiana Coal
Stripmine Surfaces in 1964 and 1981
John Richard Schrock
Association of Systematics Collections
University of Kansas
Lawrence, Kansas 66045
and
Jack R. Munsee
Department of Life Science
Indiana State University
Terre Haute, Indiana 47809
Introduction
In 1964, Munsee (10) studied the ecology of ants on relatively barren unreclaimed
Indiana spoil banks originally deposited in 1949-51. In 1981, Schrock (15) repeated
the methodology at the same 21 sites, now mostly revegetated. While the primary focus
of both studies was the surface active insect populations, extensive samplings of both
the soil and vegetation at each site were used in analyses to explain the distribution
of selected insects. Of these major factors studied by Munsee and Schrock — soils, vegeta-
tion, and insects — this paper summarizes the soil data.
Since there is variation in 1) the amount of sulfur compounds in the parent material,
2) the climate and particularly the available moisture, and 3) the mining techniques,
there is variation in the pH ranges found on coal stripmines. Previous studies (5, 13,
19) have all assumed that changes over time could be studied at one point in time
on a series of multi-aged spoil banks, but this approach may be limited because of
different initial pH values. The Munsee and Schrock studies, 17-years apart, constitute
the first real-time comparison of changes on humid Midwestern spoil banks over a
substantial period of time.
Methods
In 1964, Munsee selected an ecological study area in the old Sunspot Mines. The
location of the stripmines is south of Centenary in Vermillion County, Indiana, in
Township 14N, Section 24(10). An unnamed rangeline road runs north and south and
intersects State Road 163 which connects Clinton with Centenary. The old stripmines
border the rangeline road one mile south of this intersection. The study area is 0.366
km (1200') west of the rangeline road and is mapped in Figure 1.
The spoilbanks resulted from surface coal mining by Ayrshire Colleries from 1949
to 1951. Ayrshire sold about 300 acres of the spoilbanks to the Clinton Chapter of
the Isaac Walton League. The protection provided by the League and the relative in-
accessibility of the research site have prevented disturbance of the research site over
the past 41 years.
Site Descriptions
Twenty-one research sites, 19 on mined spoilbanks and two in an adjacent unmined
area, were selected by Munsee in 1964 with guidance from Dr. Leland Chandler from
Purdue University. The plots were selected to provide a similar age of spoils and an
assortment of exposures and slopes (Figure 1, Table 1). In addition each site needed
to provide sufficient area and accessibility to conduct the research.
The size and shape of plots varied due to topography but the two insect pitfall
traps placed in each plot were located along the center line in all cases (Figure 2).
579
580
Indiana Academy of Science
Vol. 94 (1985)
Figure 1. Map of physical features in research area. Contours are not surveyed but
merely represent an approximation of the topography. The age of the spoilbank ridges
is given by year mined. Average site pH for both study years is given in the box tangent
to the research site label.
Soil and Atmospheric Sciences 581
Table 1. Physical features of the twenty research sites sampled in 1964 and 1981.
Exposure, slope and plot size are taken from Munsee's 1964 measurements. Solar radia-
tion level were calculated from Bufoo et al. (8) according to the exposure and slope
of each site.
RESEARCH
AGE+
EXPOSURE
SLOPE
AREA
PLOT !
SIZE
DAILY
ANNUAL
SITE
(Years)
unmined
360 circle
155 SSE
(%)
(sq. meters)
(Feet)
35 x 35
SOLAR RAD*
799
SOLAR RAD**
W
22
113.8
233398
A
unmined
135
SE
4
452.9
75 x
65
839
207225
B
14
31
290
WNW
7
250.8
60 x
45
828
192962
C
14
31
155
SSE
10
92.2
32 x
31
833
220118
D
14
31
92
E
52
146.3
45 x
35
643
161680
E
14
31
235
SW
56
146.3
45 x
35
604
211457
F
14
31
160
SSE
59
81.3
35 x
25
535
239559
G
14
31
90
E
58
113.8
35 x
35
606
154046
H
13
30
255
WSW
61
241.6
65 x
40
591
180892
I
13
30
165
SSE
3
181.2
65 x
30
840
207398
J
14
31
350
N
9
185.8
50 x
40
818
175920
K
14
31
345
NNW
51
167.2
45 x
40
491
063947
L-0
14
31
340
NNW
2
204.4
110 x
20
838
196585
M
13
30
130
SE
7
167.2
60 x
30
836
211182
N
13
30
295
WNW
18
34.8
25 x
15
797
178027
P
12
29
210
SSW
50
69.7
30 x
25
618
233645
Q
12
29
305
NW
7
111.5
40 x
30
825
187165
R
12
29
225
SW
52
139.4
50 x
30
634
216188
S
12
29
320
NW
7
162.6
70 x
25
825
187165
T
12
29
30
NNE
35
81.3
35 x
25
659
102617
+Age of
site
in 1964 an
d 1981.
*Cal.
/cmz/day. **Cal . /
cmz/year .
Plots were often oriented to avoid stagnant ponds, ravines and unmanageable slopes.
However, in spite of avoiding problematic areas, this still sampled a variety of posi-
tions, centering the pitfall traps on tops, slopes and bases of ridges and defining the
location of soil samples. Detailed notes and a sketch map drawn by Munsee in 1964
proved vital in 1980 when Schrock proposed a follow-up study at the site. In spite
of extensive revegetation in 1981, natural landmarks permitted Munsee to locate the
pitfall trap locations and reestablish plots. The correct position, to within several feet
or closer, was confirmed for most sites by tree counts taken in 1981.
Slopes
Slope at each site was determined by an Abney level in 1964. In spite of erosion,
the topography was assumed unchanged in 17 years and slope measurements were not
repeated in 1981. Visual inspection of the sites appeared to verify this assumption.
Site Orientation
Orientation of each slope was measured by a pocket compass with magnetic north
the reference in 1964. This direction was considered unchanged in 1981. Theoretical
daily and annual solar radiation were calculated from exposure and slope according
to Buffo et al. (4). Since the research area was at 39 degrees and 40 minutes north,
the 40-degree-table data could be used with less than 0.5°7o error. Values in
Cal./sq.cm./day and Cal./sq.cm./year were extrapolated to the nearest degree slope
within the appropriate aspect (i.e., NNW, ESE, etc.). Daily solar radiation values were
calculated for each site on June 22, the approximate mid-point in the study periods.
Stage Designation
In 1964, Munsee designated his research sites by letters, beginning with "Site
A" in the south end of the area and working north along the mined ridges and west
to east when sites cut across ridges (Figure 1). Originally, "Site A" was the only un-
mined site. But several weeks into his study, Munsee added an unmined woodland
582
Indiana Academy of Science
Vol. 94 (1985)
site, "Site W," south of site A. Therefore, sites labelled by letters close together in
the alphabet are physically closer together on the research area, with the exception
of site W which is off the spoilbanks from Site A. This explains why tabular data
on sites is presented in this study in the order "W, A, B, C, D . . . T."
Soil Acidity
In 1964, Munsee collected ten soil samples from each site. With only sites W
and A extensively vegetated, the sampling procedure on bare ground posed little distur-
bance to plants or the surface-active fauna. His samples were extracted with a soil
probe to a depth of 30.5 cm. (one foot). These 2.5 cm. diameter (one inch) cores
were taken at four equally-spaced points along each diagonal of the site. Two addi-
tional samples were taken at a ". . . .convenient distance on either side of the point
of crossing of the two diagonals." The general scheme is given in Figure 2 although
1981
Figure 2. Patterns of soil sampling used in the two research years. The rectangular
sites varied in size and proportions. Although the cores on the 1964 diagonals were
equally spaced, they could not be relocated with accuracy. 1 through 10 are soil sample
cores; pitfall traps are along midline.
the distance between cores varied widely with the different-sized research sites. For
each of the ten cores, soil acidity was tested, usually on site but occasionally upon
returning to the laboratory, using Purdue University quick-test reagents and color charts.
At each core site, the surface material and material at 15.2 and 30.5 cm. (six and
12 inches) were tested. This provided values at three depths at each of ten points.
Only rarely did boulders prevent a full 30 readings per site.
In 1981, this same sampling pattern was not replicated for two reasons. First,
while the pitfall trap locations themselves were relocated with some assurance less than
a meter from their 1964 positions, the placement of the cores on unmeasured diagonals
and at a "convenient distance" along the meridian meant that the 1981 core samples
could be from one to three meters distant from the original 1964 core sites. Even more
importantly, vegetation and a thin organic layer had developed at all but site D. Sampling
ants with pitfall traps, Greenslade (7) found that such mechanical disturbances of organic
soil greatly distorted the pitfall catch of ants in the area. Likewise, Joosee (8) found
disturbances altered springtail sample numbers and traced the hyperactivity of these
insects to increased carbon dioxide resulting from any soil disturbance increasing the
Soil and Atmospheric Sciences 583
respiration by soil microorganisms. These effects would be minimal while the spoils
were generally barren but had to be considered significant in 1981. Since this research
effort was mainly directed at accurately re-sampling surface-active arthropods (11),
and since the 1964 cores were not marked for relocation, it was decided to make no
soil disturbance beyond the minimum required for setting the pitfall traps in the ground.
However, since the traps required a 20.3 cm. diameter (eight inch) core, four samples
were taken from the core, both at the surface and from the 15.2 to 30.5 cm. (six
to 12 inch) layers.
In 1981, soil samples were stored in marked plastic bags and returned to the
laboratory. Two of the four samples taken were tested at the surface and again at
the six to 12 inch layer. Soil was mixed with distilled water at a lg.:lml. ratio and
stirred with a glass rod. A Corning Digital 109 General Purpose pH Meter was used
to measure the pH, after the electrodes were allowed to equilibrate from five to 15
minutes. We recognize Smith and Sobek's (16) concern that pH readings with the elec-
trode placed in a soil suspension sediment (vs. supernatent fluid) are usually lower.
However, this would only shift the results and not change the relationships (Figure 4).
Soil Moisture and Infiltration
A field capacity test as described by Meyer and Anderson (9) was run by Munsee
in 1964. Soil from the core samples was air-dried and the large soil aggregates were
broken by a mortar and pestle. Particles larger than 0.495 mm. diameter were ex-
cluded from the test. The soil was tamped into a 500 ml. graduated cylinder. A stan-
dard quantity of water (100 ml.) was added at the top and the depth of penetration
was recorded for each of seven days. Comparable quantities of soil from 20 cylinders,
one sample from each research site, wetted to field capacity, were weighed and oven-
dried at a low setting (about 38 degrees C, 103 degrees F) for 24 hours. Percent moisture
content was calculated on an oven-dry soil basis. The same methodology was used
in 1981. In addition, in 1981 an attempt was made to layer the column with material
from a similar horizon. Since soil at each level (surface, 15.2 cm. and 30.5 cm.) had
been bagged separately, it was possible to place 30.5 cm. soil at the bottom of the
cylinder and surface soil at the top. It was found that the inner diameter of 500 ml.
graduated cylinders varied slightly. Since depth of water penetration is determined by
the soil's ability to hold water, the depth reading was adjusted to compensate for more
or less soil volume. The precise internal diameter of the 1964 cylinders is not known.
Soil Texture
The separation of soil fines into sand, silt and clay fractions was done using the
Bouyoucos (2) method for both 1964 and 1981 samples. The one inch (2.5 cm.) soil
cores did not always provide sufficient soil for both the pH and texture analyses in
1964 and additional cores were gathered. Since an organic layer was essentially absent
from the mined sites in 1964, this extensive soil sampling probably did not disturb
the strip mine biota. In 1981, the texture samples were drawn from the 15.2 and 30.5
cm (six and 12 inch) layers.
In both years, hydrogen peroxide was used to remove organic matter where it
was found in appreciable amounts. Soil samples were dried, soil aggregates broken
and all particles larger than 2 mm. were sifted out using soil sieves. Fifty grams of
soil were carefully weighed and transferred to a flask of 100 ml. of 5% sodium
hexametaphosphate (Calgon). This was shaken occasionally and allowed to stand over-
night so that all aggregates could slake.
The following day the soil suspension was poured into a modified blender and
mixed for two minutes. The suspension was transferred to a Bouyoucos cylinder and
agitated with a metal churn. The Bouyoucos hydrometer was immediately inserted and
readings were taken at 40 seconds and 6 hours 52 minutes.
584 Indiana Academy of Science Vol. 94 (1985)
Since the hydrometer reading varies with temperature, temperature readings were
made with each recording in 1981 and 0.36 units were added to or subtracted from
the hydrometer reading for each degree above or below 20 degrees C respectively. Aside
from the temperature corrections, we feel the texture tests were conducted identically.
Operator bias between 1964 and 1981 of course can never be ruled out.
Clustering Methods, Correlations and Principal Components Analyses
As a part of a broad ecological study, physical features of the stripmine sites
were defined as the pH, percent bareground, water capacity, maximum water penetra-
tion, percent silt, percent clay, plant hits and tree basal area (as architectural features),
annual and daily solar radiation, and slope.
To detect the complex patterns of similarity between sites based on physical features,
a cluster analysis was performed using each site as a case. For this task, BMDP statistical
program P2M was used as modified to run on the Kansas University Computing Center
Honeywell DPS-3/E.
The distance between two cases of data is defined as the chi square test of equality
of the two sets of frequencies. The computer program begins by comparing each pair
of cases and using this chi-square test, joins the closest two cases. When two cases
are joined, a new centroid is formed by averaging each variable. In the next round
of searching for the shortest distance, this centroid is compared with other candidates
for membership to the next larger cluster. The number of cases (or pseudo-cases) is
reduced by one at each step until all are clustered.
It is also possible to estimate the influence of these physical factors from the
combined effect of all factors using principal components analysis. For this task, BMDP
statistical program P4M was used as modified on the K.U. system mentioned above.
Table 2. Soil pH values from sample cores at various depths in both 1964 and 1981
at three sites. Values for 1964 from Munsee's unpublished field data. (*) acid slick.
S = Surface. 6" depth = 14 cm.
SITE P 1964 SITE P 1981
2.54 cm. (1") Sample Core # 20.32 cm. (8") Sample Core //
123456789 10 1 2
s
5.6
4.2
3.8
4.6
3.8
3.8
3.8 4.2 6.2
5.0
6"
6.6
4.2
4.2
3.8
3.8
3.8
3.8 3.8 4.6
4.2
12"
3.8
3.8
3.8
3.8
3.8
3.8
3.8 3.8 3.8
3.8
Veg.
N
N
N
N
N
N
N* N N
N
s
6.0 5,
.5
6.4 6.4
6"-
12"
6.4 6,
.5
7.0 7.1
Veg.
Y
Y
SITE D 1964 SITE D 1981
2.54 cm. (1") Sample Core // 20.32 cm. (8") Sample Core #
1 2
1
2
3
4
5
6
7
8
9
• 10
S
4.2
3.8
3.8
3.8
3.8
3.8
3.8
4.2
3.8
3.8
6"
4.2
3.8
3.8
3.8
3.8
4.2
3.8
4.6
3.8
4.6
12"
5.2
3.8
4.2
3.8
4.2
4.6
3.8
4.6
3.8
4.6
Veg.
N
N
N
N
N
N
N
N
N
N
S
3,
.8 3.6
3.
.9 3.8
6"-
12"
3.
3,
.4 3.6
.6
3,
4.
.8 4.4
.2
Veg.
N
N
SITE G 1964 SITE G 1981
2.54 cm. (1") Sample Core // 20.32 cm. (8") Sample Core //
123456789 10 1 2
S 6.6 7.4 6.0 5.2 6.4 7.4 7.4 7.4 3.8 7.4
6" 5.2 6.0 3.8 4.6 4.6 7.4 5.6 6.0 4.2 7.4
12" 3.8 3.8 3.8 4.2 4.2 4.6 4.6 4.2 3.8 7.4
Veg. YYYYYYNYYY
s
5.7 5.6
4.5 6.1
6"-
12"
6.2 6.2
6.4 6.5
Veg.
Y
Y
Soil and Atmospheric Sciences
585
For reasons described in Schrock (15), the PCA's were performed on log-transformed
data and 1.0 was added to all original values before transformation to avoid difficulty
with zero values.
Correlations were extracted from the correlation matrices produced as an in-
termediate stage in the construction of the PCA's.
Results
Soil Acidity
The slick acidic surfaces observed on the research sites in 1964 were absent at sites
by 1981 . From Table 2 it is obvious that site P has become less acidic and is revegetating.
Site D has remained acidic and barren. While two sample cores are less representative
than ten, they nevertheless portray a consistent pattern of neutralization of soil except
at site D.
The average pH at each site in 1964 and 1981 is placed on Figure 1 and demonstrates
the variation within both acidic and calcareous banks.
The extent to which site pH has changed over 17 years at all mined sites together
is illustrated in Figure 3. The variation in pH at each site in 1964 and 1981 is given
20%-
10%-
20%- —
r— 50%-i
1981 MINED SITES N-186
10%-
1964 MINED SITES N = 539
2=3
H \
pH (34) (3.6) 33 (4.0) 42 (44) 4.6 4.8 5<0 5.2 (54) 5£ 5.8 6.0 &2 64 6.6 (68) 7.0 (7*2) 7.4 pH
Figure 3. Frequency distribution of soil pH values at all mined sites in 1964 and
1981. Since the 1981 values were recorded to the nearest 0.1 pH unit (black) and the
1964 values were restricted to color indicators of varying resolution (white), the 1981
values are clustered into groups matching the 1964 indicator ranges for comparison (gray).
in Figure 4 by a circle with a radius of one standard deviation. These figures indicate
which sites have moved well away from values held in 1964.
The common range for mineral soils in humid regions is 5.0 to 7.2 pH (3). Of
three originally strongly acidic soils, only D has failed to move into this "normal"
range. Of six calcareous soils, all have dropped to a pH well within this common range.
Soil Moisture and Infiltration
In 1964, Munsee noted that "... soil samples through which the water per-
colated most rapidly likewise show the greatest depth of penetration during the 7-day
period." (Figure 5). The "slow" sites D, B, T, M . . . etc. were all of high clay con-
tent while the "fast" sites W, F, A, C . . . etc. were progressively higher in sand
content (Figures 6-9).
Wetted-zone depth increased, though not uniformly, in 1981. Soil moisture-holding
decreased in 1981. While water transmission is known to be low in spoils (6), this
is the first direct evidence of substantial improvement over time.
586
Indiana Academy of Science
Vol. 94 (1985)
Figure 4. Changes in soil pH between 20 research sites in 1964 (W, A, B . . . T)
and 1981 (W, A', B' . . . T'). Circles are centered on average pH value and the
radius is drawn as one standard deviation.
Soil Texture
In contrast to other mining areas, stone and gravel did not appear dominant at
these sites. No attempt was made to quantify coarse material either year, but it was
noticeable that soil fines were well in excess of the discarded coarse material in most
cases. In 1981, some larger aggregates were found to be very soft and readily crumbled
in the preparations for soil tests. Occasional chunks of soft coal were picked up that
could be crushed between fingers, and tiny black flecks of such material were evident
in some of the soil infiltration columns.
In addition to the unmined site W, sites C and F were sandy enough to be noticeably
different from other spoils in the field in 1964 and 1981. The high silt reading for
site P (Figure 9) is extreme and would be questionable if it were not for corroborating
Soil and Atmospheric Sciences
587
D B T M
Q H S
3 N
R
K
G
E
J
L-0
C
a
F
w
6
»
i
1 2.
10
i m
--
i
i
j_
i
I
j_
i
i
12
1
Li ± j
1 1
p^
i
i
-* l_J
-
i
i
IS
u
—
-
18
*****
*#
L-J
20
Water Penetration Depth (cm. in 1 & 7 days)
m
^"^
—
Ql96<
□ 1911 «--
Soil Moisture (% of dry weight)
40%
30%
M M _
"i n-i "i
-l,-
—j
-.
"1
1-1
r-Tl
_
20%
1
1
10%
f
1
~> b t ^
1 Q H S
P
M
R
K
3
E
J
L-0
c
3,
h
w
Figure 5. Changes in depth of water infiltration and changes in soil moisture at each
research site in 1964 and 1981.
90 80
30
20
70 60 50 40
% SAND <
Figure 6. Soil textures at near-level ridge-top sites (less than 10% slope) measured
in 1964 (I, J, S) and 1981 (I ', J', S ').
588
Indiana Academy of Science
Vol. 94 (1985)
90 80 70 BO 50 40 30 20 10 0
% SAND <
Figure 7. Soil textures at near-level sites (less than 10% slope) measured in 1964
(W, A, B, C, L-O) and 1981 (W\ A', B', C, L-O ').
with a third. This becomes an "apples-and-oranges" problem in which we find ourselves
asking if a similarity in vegetation between site B and C is more or less important
than a similarity in soil texture between site B and C, for instance. To ask which sites
are most similar without regard to some organism's response or other criteria, leaves
the clustering by similarity dependent on an artificial weighting of one for each factor.
The internodal distance and even the clustering order will be prejudiced by the factors
chosen and this uniform weight for each factor.
Based on slope, soil silt and clay fractions, soil pH, moisture-holding capacity
and maximum water penetration, percent bareground, exposure, slope, plant hits (out
of 100 point drops) and basal areas of trees as architectural features, and daily and
annual solar insolation, the sites do not cluster in a completely arbitrary manner in
observations: "On the occasions that the pitfall traps were flooded in Site P the liquid
contents of the trap bottles turned into a grey, viscous, colloidal suspension" (10).
Pitfall samples from site P preserved in vials from the rainy weeks in 1964 had con-
siderable silt despite several rinsings during extraction of insects. By 1981, site P has
crossed the texture chart to reside with the clay banks (Figure 9). None of the flooded
traps, including traps at site P, exhibited high amounts of silt in 1981.
General Site Characteristics
If two sites were identical in soil pH, texture, water capacity, and had similar
trees and herbacious cover, we would consider them more closely related to each other
than to a barren site with different soil properties. The task becomes more difficult
Soil and Atmospheric Sciences
589
30 SO 70 SO 50 40 30 20 10
%SAND < —
Figure 8. Soil textures at steep sites (greater than 50% slope) measured in 1964 (E,
F, G, H, K, R) and 1981 (E ', F', G', H\ K\ R ').
when the first site shares some characteristics with the second site, other characteristics
1964 and these clusters hold similar patterns in 1981 (Figure 10). Since sites W and
A cluster together, this lends some biological credence to the process. Sites C-I-M,
J-S-Q-L-L-N-B, and G-D remain associated. Sites T and H have made major changes
in associations and site K is unique in physical features in 1964 and even more so in 1981.
Correlations between selected site characteristics were remarkable for their con-
sistency in sign between 1964 and 1981 (Figure 11). Strong correlations between some
factors indicate that, while one may not directly cause changes in a second, both have
a strong relationship to a "common environmental factor."
Principal Components Analysis
Variation from all of these soil factors is arrayed for the first three principal
components (described below) in both years in Figures 12 and 13. Several characteristics
are not totally independent. Silt and clay (plus sand) total unity. The clay fraction
holds water and this water-holding ability prevents deeper soil penetration. Potential
solar radiation is determined from tables involving slope as one of three factors. Never-
theless, each of these three factors contributed sufficient unique variation to be included
in correlations with various arthropods discussed elsewhere (15).
When variation among sites' physical features is analyzed on the basis of twelve
measured variables for 1964, the sites array as shown in Figure 12 with the first three
factors explaining the greatest share of combined variation. Variation in factor 1 is
composed of water capacity, maximum water penetration, bare ground, pH, plant hits
590
Indiana Academy of Science
Vol. 94 (1985)
60
%SAND « —
Figure 9. Soil textures at remaining sites measured in 1964 (D, M, N, P, Q) and
1981 (D\ M\ N\ P', Q'). Note the drastic change at site P.
and tree basal area. Factor 2 is composed of soil texture, plant hits, tree basal area,
maximum water penetration and pH. Factor 3 is composed of variation contributed
from daily and annual insolation, slope and pH. Therefore, pH is the only characteristic
to account for major variation on all three axes. The first three factors account for
72 percent of total site variation. Sites A and W are together far in the background
while site P is near site D on factors 1 and 3 but distant on factor 2. Most of the
spoilbanks except P float slightly in front of the factor 1x3 plane. By 1981 (Figure
13), site P has returned to the fold of strip mine sites. The strip mine sites have also
tightened up, reflecting the decrease in extreme soil factor readings and fairly uniform
increases in plants and trees. Nevertheless, roughly two clusters of strip mine sites
remain: the B-M-S-Q-I-L-N-J-C tight cluster found loosely on the positive side of factor
3 in 1964, and the R-G-T-H-E-K cluster found on the negative side of factor 3 in 1964.
By 1981, the first three factors only account of 64 percent of the variation. Now
pH, daily insolation and clay contribute to all three factors. Bare ground, slope, water
penetration, plant hits and tree basal area contribute to factor 1. Factor 2 also in-
cludes variation in water capacity, maximum water penetration and slope. Factor 3
concentrates on plant hits, bare ground, exposure and tree basal area.
Conclusions
In discussing the present soil situation, it is useful to know what the original
soil overlying the area was before stripmining in 1949-1951. The Soil Survey of Ver-
Soil and Atmospheric Sciences
591
WACIMFERPGDTHJSQLNBK WACIMEHTFRPGDQLJNSBK
10-
20-
30-
40-
IIWli
T
i '
jyju \
V
J
■ry—
-
PHYSICAL
SITE
FACTORS
PHYSICAL
SfTE
FACTORS
19b4 '
1981
Figure 10. Dendograms clustering sites on basis of physical factors at each site in
1964 and 1981.
million County, Indiana (14), based on fieldwork completed from 1972 to 1975 describes
the adjacent land as a mosaic of Hennepin loam on the steeper slopes and Russell
silt loam on the flatter terrain. Their soil map outline superimposed on an aerial photo
places sites W and A on these soil types respectively. These soil types projected by
the Soil Survey are based on air photos, a necessarily sparse sampling program and
knowledge of local soil genesis. The texture values of the mapped soils is confirmed
CORRELATION SCHEME
Environmental Characteristics 1964/1981
,PH
-0.48/^0.57
I Bare ground.
-0.68/-0.43
I
+0.84/+0.66
■| Water Capacity-
+0.57/+0.06
I
-0.82/-0.63
i I Maximum Water Penetration <
-0.40/-0.77
Ir Silt
-0.737-0.29
' Clay . i
+0.54/+0.05
+0.29/+0.63
1 Plant Hits-
+0.14/+0.04
0.577-0.14
-0.72/-0.21
+0.73/+0.39
-0.33/-0.49
■Tree Basal Area.
-0.38/-0.43
+0.23/l+0.36
■Annual Solar Radiation 40°N— J
+0.44*
■Daily Solar Radiation June 40 N-
g1 np°
-0.89*
I
Figure 11. Correlations between selected characteristics of the research sites in
1964/1981. Note that the sign of correlation remains unchanged. Some values were
considered unchanged between the two studies (*).
592
Indiana Academy of Science
Vol. 94 (1985)
SITE
1964
Figure 12. Principal components analysis of 1964 sites on physical site factors.
Soil and Atmospheric Sciences
-3
593
Figure 13. Principal components analysis of 1981 sites on physical site factors.
594 Indiana Academy of Science Vol. 94 (1985)
by both the 1964 and 1981 textures measured on sites W and A in this study. There
is little reason not to believe that such soils covered the whole research area prior
to mining.
After stripping, the unleveled spoilbanks are classified as Orthents and consist
of variable amounts of ". . . loam glacial till, rock and shale fragments, and small
fragments of clay and coal." (14) This is confirmed by the mining soil samples taken
in this study. When deposited near the surface and subjected to weathering, the spoils
undergo chemical reactions releasing an excess of H+ ions (17). According to the
laboratory work of VonDemfange and Warner (19), ". . . there is sufficient neutraliz-
ing potential in only three feet of spoil to neutralize the acid that is likely to be pro-
duced by the spoils." Yet, they acknowledge that more acid is initially produced than
is neutralized by calcium and mangesium minerals in the spoilbank. This discrepancy
is likely due to the oxidation rate of pyrite proceeding more rapidly than the slower
neutralizing reactions. In this study, for the first time we follow specific sites and detect
neutralization directly over time.
Soil acidity appears to be "evening out" and approaching a median pH between
5.8 and 6.2. This probably involves the leaching of calcareous banks and the depletion
of sulfur compounds in acidic banks. This is suggested strongly by both Table 2 and
Figure 4. Two enigmas appear in the latter figure however: site D becomes slightly
more acidic and site P recovers with a dramatic shift twice that of any other site.
Site D is the only barren site in 1981, an impressive ridge over 60 feet high. In
June of 1980 a half-buried iron reinforcement rod was found in the sediment at the
base of this ridge. According to Munsee, that was one of the few rods left on site
in 1964 and its original location was at the top of the bank. This provides one possible
reason why site D remains acid and unrevegetated today. If a four-foot reinforcement
rod, buried for much of its length in a clay-slate bank, has been undercut and washed
to the base of the hill, there is grounds to believe that over 17 years, erosion has
been sufficient to remove weathered material and expose fresh sulfur compounds. This
could also prevent long-term rooting of perennials. Confirmation of this would re-
quire erosion pin techniques.
Soil at site P undergoes an unexpected rise in pH from 4.2 to 6.4 with a relatively
small variance in each sample series. Site P also shows an equally phenomenal change
in texture. Therefore it is valid to ask if we mislocated the site in 1981. However,
site P is very distinct. It is the south face of a small ridge with outstanding small
scale features. To the east is the flat basin of L-O; to the west the ridge bends and
levels out and becomes lower swampy territory never covered in 1964. Site P in 1964
was more barren than site D. Yet in 1981 all of the area that could conceivably be
site P is revegetating. This is a biological confirmation that site P has advanced and
supports the measured changes in pH and texture. Site P in 1981 has to be on site
P in 1964 and we believe the pH and texture values are real.
Is there anything unique about this site that might account for major shifts in
soil features that presumably take centuries to change? Site P is unique in having a
stagnant pool of water lapping at its base eight feet or so below the trapsites. This
pool has been present since mining and according to general observations by Munsee,
has risen perhaps six inches since 1964. This moisture undoubtedly accounts for species
of moss and fungi present at site P. The constant presence of soil moisture could
perhaps be a factor that exhausts the pyrite, either by permitting continuous oxidation
or by improving conditions for bacteria that can live in the presence of sulfur com-
pounds. Water percolation appears to be the main factor in removing sodium ions
from the surface foot of spoils in reclaimed coal spoils in the Northern Great Plains
(12). Deeper and more rapid penetration of water is attributed to enlarged root systems
Soil and Atmospheric Sciences 595
from revegetation. Although this is sodium (rather than sulfur) on a non-acid mine,
the process suggests that leaching is accelerated on spoils once vegetation is established.
Heterogeneity within each site was apparent, especially in 1964 (Table 2). Therefore,
the lack of a high number of subsamples as recommended by Berg (1) is a valid criticism.
Measurements of plant-available phosphorus and nitrogen, shown to be a critical factor
on some spoils of this type (18), were not made.
The changes in soil factors described here are admittedly not beyond dispute.
We could be more comfortable if a more extensive grid of cores were sampled, if
these core sites were permanently marked, and if they were repeatedly sampled at perhaps
five-year intervals. While such a study was not anticipated by Munsee in 1964 or possi-
ble in a faunistic study in 1981, we feel our data suggest rapid changes in some spoilbanks
over time and provide a basis for further studies aimed at confirming this.
The Clinton Chapter of the Isaac Walton League graciously granted permission
for the 1964 and 1981 studies. Without their ongoing efforts over the last 34 years,
this specific site of naturally revegetating spoil banks would not have existed undisturbed.
Mr. Don Post, Department of Agronomy, Purdue University, was of valuable assistance
with hydrometer tests in 1964. Dr. George Byers, Dr. William J. Bell and Dr. William
Gordon, all of the K.U. Department of Entomology, provided laboratory equipment
for some of the soil tests. Dr. Curtis Sorenson of the Department of Geography of
the University of Kansas, also supplied laboratory materials, equipment and space for
soil texture tests at the Soil Analysis Laboratory. In addition, his advice on procedures
and the reliability of soil test results was very helpful. Mr. Douglas West provided
housing and support during the 1981 field research. And finally, we would like to
thank our wives who were financially supportive throughout the research period. We
are indebted to Dr. Leland Chandler, Purdue University, and Dr. Edward Martinko,
University of Kansas, who served as major professors in the respective studies. The
Office of Surface Mining provided funds permitting the updating and presentation
of these results.
Literature Cited
1. Berg, W.A. 1978. Limitations in the use of soil tests on drastically disturbed
lands. In Schaller, F.W. and P. Sutton (eds.) Reclamation of Drastically Disturbed
Lands. Amer. Soc. Agron., Crop Sci. Soc. Amer. and Soil Sci. Soc. Amer.,
Madison, Wisconsin. 742 p.
2. Bouyoucos, G.J. 1936. Directions for making mechanical analyses of soils by
the hydrometer method. Soil Science. 42:225-228.
3. Brady, N.C. 1974. The Nature and Properties of Soils. 8th edition. Macmillan
Publishing Co., New York. 639 p.
4. Buffo, J., L.J. Fritschen, and J.L. Murphy. 1972. Direct Solar Radiation on
Various Slopes from 0 to 60 Degrees North Latitude. Pacific Northwest Forest
and Range Exp. Station Res. Paper 142. Portland, Oregon. 74 p.
5. Byrnes, W.R. and J.H. Miller. 1973. Natural revegetation and cast overburden
properties of surface-mined coal lands in southern Indiana. In Hutnik, R.J. and
G. Davis (eds.) Ecology and Reclamation of Devastated Land. Gordon and Breach,
New York.
6. Gee, G.W., A. Bauer and R.S. Decker. 1978. Physical analyses of overburden
material and mine land soils. In Schaller, F.W. and P. Sutton (eds.) Reclamation
of Drastically Disturbed Lands. Amer. Soc. Agron., Crop Sci. Soc. Amer. and
Soil Sci. Soc. Amer., Madison, Wisconsin, 742 p.
7. Greenslade, P.J.M. 1973. Sampling ants with pitfall traps: digging-in effects. Insects
Sociaux. 20:343-353.
596 Indiana Academy of Science Vol. 94 (1985)
8. Joosse, E.N.G. and J.M. Kaptejin. 1968. Activity-stimulating phenomena caused
by field disturbance in the use of pitfall traps. Oecologia. 40:385-392.
9. Meyer, B.S. and D.B. Anderson. 1952. Plant Physiology. D. Van Nostrand Co.
Inc., Princeton, New Jersey. 785 p.
10. Munsee, J.R. 1966. The Ecology of Ants of Stripmine Spoil Banks. Ph.D. Disser-
tation. Purdue University, West Lafayette, Indiana. 243 p.
1 1 . Munsee, J.R. and J.R. Schrock. 1982. Comparison of ant faunae from unreclaimed
coal stripmines in Indiana in 1964 and 1981. Proc. Ind. Acad. Sci. 92:257-261.
12. Richardson, B.Z. and E.E. Farmer. 1982. Changes in Sodium Adsorption Ratios
Following Revegetation of Coal Mine Spoils in Southeastern Montana. Intermoun-
tain For. and Range Exp. Stat. Res. Paper. INT-287. 4 p.
13. Riley, C.V. 1973. Chemical alterations of stripmine spoil by furrow grading-
revegetation success. In Hutnik, R.J. and G. Davis (eds.) Ecology and Reclama-
tion of Devastated Land. Vol. 2. Gordon and Breach, Inc., New York.
14. Robbins, J.M. and M.H. Robards. 1978. Soil Survey of Vermillion County,
Indiana. U.S.D.A. Soil Conservation Service and Purdue Univ. Agric. Exp. Station,
Lafayette, Indiana. 124 p.
15. Schrock, J.R. 1983. The Succession of Insects on Unreclaimed Coal Strip Mine
Spoil Banks in Indiana. Ph.D. Dissertation. University of Kansas, Lawrence,
Kansas. 207 p.
16. Smith, R.M. and A. A. Sobek. 1978. Physical and chemical properties of over-
burden, spoils, wastes and new soils. In Schaller, F.W. and P. Sutton (eds.)
Reclamation of Drastically Disturbed Lands. Amer. Soc. Agron., Crop Sci. Soc.
Amer. and Soil Sci. Soc. Amer., Madison, Wisconsin, 742 p.
17. Stumm, W. 1966. Oxygenation of ferrous iron properties of aqueous iron as related
to mine drainage pollution. Symposium on Acid Mine Drainage Research, Ohio
River Valley Sanitary Commission.
18. Vogel, W.G. and W.R. Curtis. 1978. Reclamation research on coal surface-mined
land in the humid East. In Schaller, F.W. and P. Sutton (eds.) Reclamation of
Drastically Disturbed Lands. Amer. Soc. Agron., Crop Sci. Soc. Amer. and Soil
Sci. Amer., Madison, Wisconsin. 742 p.
19. VonDemfange, W.C. and D.L. Warner. 1975. Vertical distribution of sulfur forms
in surface coal mine spoils. In Third Symposium on Surface Mining and Reclama-
tion, Vol. I. National Coal Association, Louisville, Kentucky.
ZOOLOGY
Chairperson: Thomas Fogle
Department of Biology
St. Mary's College
Notre Dame, Indiana 46556
(219) 284-4675
Chairperson-Elect: James R. Litton, Jr.
Department of Biology
St. Mary's College
Notre Dame, Indiana 46556
(219)284-4669 or 4671
ABSTRACTS
The Adaptive(?) Significance of Brood Reduction in the Red-winged Blackbird (Agelaius
phoeniceus). James D. Hengeveld, Department of Biology, Indiana University, Bloom-
ington, Indiana 47405. In hatching synchrony experiments on redwings from 1981-3,
results indicated that synchronous hatching was not a necessary precursor for brood
reduction. Starvation was essentially the same among nests with synchronous hatching
and those with the natural degree of hatching asynchrony. When coupled with the
finding that young from nests in which one or more chicks starved tended to fledge
at lower weights than young from starvation-free nests, these results led me to ques-
tion the significance of the brood reduction process itself. The critical issue, however,
is not a comparison between young from reduced broods and those in which all chicks
have survived but rather a consideration of whether survivors of reduced broods do
better than they would have if their sibling(s) had not starved. During the 1984 breeding
season, I attempted to address this question by substituting healthy young of the
appropriate age for starved young in half of the nests in which brood reduction occurred.
I then monitored the growth rates, starvation, and fledging weights of chicks from
the experimental and control groups. Chicks from experimental nests (nests with substitu-
tions) fledged at lower weights than chicks from the control nests seeming to indicate
that the sacrifice of a chick is beneficial to its siblings. However, sample sizes thus
far are too small to make any definitive conclusions. (Supported in part by a grant
from the Indiana Academy of Science to J.D. Hengeveld).
Patterns of Relative Fecundity in Snakes. John B. Iverson, Department of Biology,
Earlham College, Richmond, Indiana 47374. Interspecific brood size-body length
comparisons for snakes indicate that the two variables are positively correlated. Three
factors (reproductive mode, taxonomic group, and habitat type) are identified as signifi-
cant correlates of relative fecundity. Snakes with the highest relative fecundities are
typically viviparous, tend to be from certain taxonomic (phylogenetic?) categories
(primarily the Natricinae, Xenodontinae, and Viperinae), and/or are most often aquatic,
semi-aquatic, or semi-fossorial.
Light Microscopic and Ultrastructural Features of the Gut of the Balsam Woolly Aphid,
Adelges piceae Ratz. Mohinder S. Jarial, Center for Medical Education, Ball State
University, Muncie, Indiana 47306. The structure of the gut of the balsam woolly
aphid was studied by dissection, light microscopy and electron microscopy. The gut
is a simple but slightly coiled tube and falls into three distinct regions, namely, foregut
597
598 Indiana Academy of Science Vol. 94 (1985)
comprising mouth, sucking pump, pharynx and esophagus; midgut consisting of stomach
and intestine; and hindgut representing the rectum. The horseshoe shaped sucking pump
and adjoining pharynx is connected to the stomach by the esophagus which is a long,
slender tube lined by cuticular intima. The stomach or the first part of the midgut
appears very dark in the dissected specimens that have over-wintered. The stomach
leads into the intestine or the second part of the midgut. The intestine is longer and
relatively smaller in diameter, bends on itself and empties into the rectum which nar-
rows to end in the anus. The rectum has extremely thin wall containing few mitochon-
dria and is lined with cuticular intima.
The cells in the stomach and intestine present a similar structure, except in the
former the epithelial cells are larger and contain an abundance of fine particulate material
and membrane bound, magnesium and calcium rich granules of various shapes and
forms, and crystaline rods. Isolated muscles with trachea form the outer covering.
The basal plasma membrane rests on a basement membrane, and is thrown into
numerous infoldings that penetrate deep into the cell. The cells contain well developed
nuclei and numerous mitochondria. The striated border exhibits closely packed microvilli
projecting into the lumen.
The accumulation of granules in the cells and their occasional release into the
lumen of the stomach appears to be related to the process of storage excretion in the
absence of Malpighian tubles.
Parental Investment in the Bee Ceratina calcarata Robertson (Hymenoptera:
Xylocopidae): A Preliminary Study. Michael D. Johnson, Department of Biological
Sciences, DePauw University, Greencastle, Indiana 46135. Ceratina calcarata nests,
each a linear series of brood cells in the hollowed out twig, were examined in a
preliminary study of parental investment in this solitary bee. I examined 71 nests, weighed
475 immatures and their provisions, placed each in a gelatin capsule for rearing, and
weighed the 200 males and 167 females that emerged.
Preliminary analysis showed that 1) for any nest, females were typically supplied
heavier provision masses (X* = 22.27±6.29 mg vs. X* = 16.67±2.95 mg), 2) wet
weight of females exceeded that of males (X* = 1 1.62 ±3.26 mg vs. X* = 8.73±2.89
mg) even if reared from similar provision masses, 3) females occurred more commonly
in the innermost cell, but 4) a comparison of the provision weights did not reflect
this sex ratio.
Ceratina females are larger (and more valued?) and should receive relatively greater
parental investment. Thus the mother typically puts a female in the first brood cell
after the investment of hollowing a twig and provisioning the cell. Then, the provi-
sioning of each subsequent cell influences the mother's "decision," such that female
eggs are typically laid on larger provision masses.
Territorial Behavior in the Prothonotary Warbler, Protonotaria citera, Between- and
Within-season Territory Relocations. Michael P. Kowalski, 5690 Kings Road, Bloom-
ington, Indiana 47401. In April of 1983 a study of the population dynamics of
a color-banded population of Prothonotary Warblers was begun on the North Fork
of Salt Creek in eastern Monroe Co., IN, and was continued in 1984. In 1983 the
study area consisted of a 3 km stretch of river, and in 1984 this was expanded to
8.9 km. Territory abandonment and relocation by males was more common than
previously suspected. In 1984 a total of 47 males took up residence on the river, 45
of which were captured and color-banded. Nine of these (20%) abandoned their ter-
ritories and were not seen again during the season. Eight males (18%) abandoned their
territories and relocated to a new site on the study area. These new territories averaged
Zoology 599
603 m from the first, with the range being between 229 m and 2364 m. Most aban-
donments and relocations took place after 15 June 1984, and in 5 cases (63%) the
males had failed to attract a mate to the first territory. One male who relocated to
a territory 2364 m from his first territory eventually returned to the original territory
and succeeded in attracting a mate.
In 1983 seven males were marked, and 4 (57%) returned to the study area in
1984. Two of these returned to their 1983 territories, while the other 2 relocated. The
average distance relocated by these males was 1000 m, with a range of 782 m to 1220
m. One of the males who returned to his 1983 territory simultaneously held a second
territory located 763 m away.
It is suggested that adult dispersal is more common in passerines than is generally
thought, and so-called surplus males may in fact be wandering or relocating birds.
A Record of the Freshwater Nemertean Prostoma graecense (Bohmig) in Indiana. James
R. Litton, Jr., Department of Biology, Saint Mary's College, Notre Dame, Indiana
46556. Prostoma graecense (Bohmig) is a freshwater nemertean that has a very
spotty distribution in the United States. Specimens were collected in benthic grab samples
(mud) in Lake Marian, a concrete impoundment on the Saint Mary's College campus,
during March, April, and May of 1983 and the early spring of 1984 (February, March
only). Collections at other times of the year in 1983 or 1984 produced no individuals.
All individuals were small (< 5 mm in length; < 0.7 mm diameter), whitish, pale
yellow or vermillion, and sexually immature. No gut contents were ever noted or iden-
tified. Specimens were also found in two locations in Juday Creek, Saint Joseph County,
during May and June, 1984 while collecting benthic samples with a sweep net. Subse-
quent collections produced no individuals. At both locations the Prostoma were
associated with rooted aquatic vegetation. All of these individuals were large (> 10
mm in length; > 1.0 mm in diameter), yellowish-red or deep red in color, and sexually
mature. Gut contents of the mature individuals showed an almost exclusive diet of
small oligochaetes.
Seasonal Abundance of the Psammic Rotifers of Spicer Lake, Indiana. James R. Litton,
Jr., Department of Biology, Saint Mary's College, Notre Dame, Indiana 46556. The
psammic rotifer community inhabiting the substrate of the littoral zone of Spicer Lake
was studied from March to November, 1983. Thirty-nine species of rotifers were found,
but abundances were extremely low (range 0.0 to 7.8 cm"3) when compared with
other freshwater psammic habitats. Total rotifer density and densities of major genera
differed significantly over time, and between depths in the sediment, but not between
sites (p < 0.01). Highest densities occurred in the top 2.0 cm of sediment in early
spring and late fall. Total rotifer density and densities of Cephalodella spp. and
Trichocerca spp. were positively correlated with alkalinity (p < 0.01). Densities of
all taxa were negatively correlated with variability in water chemistry. Rotifer species
were found to be randomly distributed in time and space, with no evidence of com-
petitive exclusion. The density and diversity of the Spicer Lake rotifer community is
comparable to that found for highly variable and unpredictable environments.
Visual Signals in Sticklebacks: A Reexamination and Extension of Some Classic
Experiments. William J. Rowland, Department of Biology, Indiana University, Bloom-
ington, Indiana 47405. Fifty years ago Tinbergen and his coworkers at Leiden,
Netherlands began an investigation of the factors that influence the reproductive behavior
of the threespined stickleback {Gasterosteus aculeatus). By presenting territorial males
with simple dummies that varied with respect to only a single feature, these workers
600 Indiana Academy of Science Vol. 94 (1985)
discovered that a few simple stimuli were of paramount importance in eliciting stickleback
social behavior. These classic experiments thus provided much of the evidence for con-
cepts such as sign stimuli, releasers, and the innate releasing mechanism. However,
recent research suggests that the results of these experiments and their interpretation
may have been oversimplified.
Using fish from Dutch and American populations I reexamined and extended
the dummy presentation experiments. I found that territorial males attacked nuptially
colored (red undersides) or headdown dummies less than nonred or horizontal dummies,
respectively. This may be because red undersides and/or headdown posture are in-
dicative of rival males and thus elicit avoidance as well as aggression in a subject.
I also found that males court (and attack) dummy females in headup "courtship"
posture less than those presented horizontally. This suggests that the primary function
of headup posture in both male and female sticklebacks in appeasement of aggression
and that such appeasement has a suppressing effect even on the male's courtship. Results
confirmed the importance of the gravid shape of females in eliciting male courtship.
Furthermore, it was found that males preferentially court supernormally large and super-
normally gravid dummy females over normal size, gravid dummy females.
Venom Antigens in Oral Secretions of Colubrid Snakes. Sherman A. Minton, Depart-
ment of Microbology and Immunology, Indiana University School of Medicine,
Indianapolis, Indiana 46223. Venom in colubrid snakes is secreted by Duvernoy's
gland which lies in the postocular region. It may or may not be associated with enlarged,
grooved posterior maxillary teeth. Duvernoy's gland secretion was collected from 11
species of colubrid snakes, some with enlarged grooved or ungrooved teeth, and some
with more or less uniform maxillary teeth. Oral secretions were also obtained from
two species of boas which lack Duvernoy's gland. These secretions were reacted with
15 commercial antivenoms in Ouchterlony immunodiffusion preparations. Three
antivenoms, a polyvalent cobra and a polyvalent and monovalent mamba antivenom,
reacted with all colubrid oral secretions; four other reacted with at least five. Reac-
tions consisted of one to three precipitin lines. Boa oral secretions gave weak reactions
with polyvalent cobra and mamba antivenoms.
Ouchterlony preparations in which oral secretion and plasma of the same snake
species were reacted in alternate wells against antivenoms indicated some reacting antigens
were present in both and hence are probably widespread in snake tissues; other antigens
were present in oral secretion and not plasma. Immunoelectrophoresis of some col-
ubrid oral secretions indicated the reacting antigens evidently do not correspond to
the major toxins of cobra and mamba venoms.
Colubrid venoms share at least three antigens with venoms of African elapid snakes
(cobras and mambas) and two with saw-scaled vipers, another genus probably of African
ancestry. There is little evidence of shared antigens between colubrids and other vipers,
pit vipers, or coral snakes, but one antigen is shared with Australian elapids.
Physiology of Vocalization by an Echolocating Bird. Roderick A. Suthers, School
of Medicine and Department of Biology, Indiana University, Bloomington, Indiana
47405. Oilbirds (Steatornis caripensis; Steatornithidae) have a bilaterally asym-
metrical bronchial syrinx with which they produce echolocating clicks and a variety
of social vocalizations. Vocalizations are initiated by contraction of the sternotrachealis
muscles which stretch the trachea, reducing the tension across the syrinx and causing
the cartilaginous bronchial semi-rings supporting the cranial and caudal edges of the
external tympaniform membranes (ETM) to hinge inward, folding the ETM into the
syringeal lumen. Sonar clicks are terminated by rapid contraction of a previously
Zoology 601
undescribed intrinsic syringeal muscle, the broncholateralis, which inserts on the semi-
ring supporting the anterior edge of the ETM. Tracheal airflow at first increases as
expiratory effort increases subsyringeal pressure. The initial high rate of airflow drops
at the onset of phonation due to the increased syringeal resistance. In the case of a
double click, airflow momentarily ceases during the intraclick interval when the ETM
temporarily closes the syrinx. Air sac pressure rises to its maximum level at this time.
Expiratory airflow rapidly increases as the ETM is abducted from either its closed
or phonatory position to its open, resting position. Each sonar click requires about
1 cc of air; a typical agonistic squawk may use about 27 cc of air.
An Experimental Study of Biparental Care in the Dark-eyed Junco. Licia Wolf, Depart-
ment of Biology, Indiana University, Bloomington, Indiana 47405. It commonly
is assumed that males of monogamous birds care for their young because without that
care success in reproduction would be impossible or reduced. This study examines the
significance of male care in the monogamous, double-brooded Dark-eyed Junco {Junco
hyemalis) by quantitative comparison of reproductive success of females with and without
the help of a male. In contrast to male parental care on the survivorship of the young
to independence as well as the season-long reproductive success of females. Over two
summers, males of 24 breeding pairs were captured at the time their eggs hatched and
were held for the remainder of the breeding season; the nests of their mates (experimen-
tals) were subsequently monitored, as were the nests of unmanipulated pairs (controls,
n = 49). Earlier studies that have addressed this problem report no significant dif-
ferences in growth rate of experimental and control young in the nest. My preliminary
results support these findings, and further, indicate no differences in several other
variables that were considered important factors affecting the reproductive success of
females: weight of young at fledging, weight loss of nesting females, and interclutch
interval. However, fewer young raised by unaided females survived to independence
relative to young raised by two parents. Further, fledging success (at least one young
that fledged) of experimental broods was slightly higher than that of control broods
in both years of the study.
Rediscovery of the Spotted Darter, Etheostoma maculatum,
in Indiana Waters: Blue River; Crawford, Harrison and
Washington Counties; Ohio River Drainage, USA
Claude D. Baker, Bill J. Forsyth and Tom Wiles
Division of Natural Sciences
Department of Biology
Indiana University Southeast
New Albany, Indiana 47150
and
D. Brian Abrell
Indiana Natural Heritage Program
Indiana Department of Natural Resources
Indianapolis, Indiana 46204
The spotted darter, Etheostoma maculatum, originally inhabited the faster, deeper
riffles of medium-sized rivers in a triangular range extending from northwestern In-
diana to northwestern Pennsylvania to southern Tennessee. Indiana Heritage Program
records reveal that, prior to 1900, Jordan may have collected spotted darters from
several locations in the northern half of the state (3, 4); however, the only verifiable
museum specimens (U.S. National Museum No. 69233) are two spotted darters col-
lected by P. Kirsch on 25 August 1899 near Delong in Fulton County (6). Despite
earlier records, the spotted darter was not collected during Gerking's 1945 statewide
survey (2), and it was excluded from an annotated key to the fishes of Indiana published
in 1968 (5). The purpose of this paper is to report an extant spotted darter population
in Indiana waters, and the first collections ever for a stream located in southern Indiana.
Our initial collections were taken from the Blue River near White Cloud in Har-
rison County downstream from an abandoned bridge and rock dam (Figure 1, Table
1). All specimens were taken by seining a smaller side branch of the main stream which
coursed around the right side of an island located below the dam. Unfortunately, chang-
ing stream bed morphology and a new canoe access in the center of the dam have
resulted in the elimination of this particular habitat. Swift mainstream riffles were
suspected to be the darter's preferred habitat, but our collection attempts in the river
Table 1 . Spotted Darter Records for the Blue River, a Southern Indiana Tributary of
the Ohio River.
Location
1. White Cloud below dam
2. Near Wyandotte Cave
3. Below Rothrock's Dam
4. Below Milltown Dam
5. Below Fredericksburg
6. Above Fredericksburg
Date
24 SEP 1976
1
20 JUL 1977
1
11 JUL 1978
3
25 SEP 1977
3
10 JUL 1979
5
17 JUL 1980
1
10 OCT 1983
5
13 SEP 1984
5:
25 JUL 1984
5
14 AUG 1984
1
14 AUG 1984
2
Latitude-Longitude
38°13'45",86°13'37'
38°13 '23",86°15 '14"
38°16'17",86°16'20"
38°20'26",86°16'27"
38°25'45",86°11 '50"
38°26'02",86°11 '31"
"Several additional specimens returned to stream
603
604
Indiana Academy of Science
Vol. 94 (1985)
SCALE IN MILES
Figure 1. Map of Blue River with Spotted Darter Collection Locations. Numbers
1-6 are identified in Table 1.
proper were unsuccessful until an extremely low discharge period in 1983 when we
took five additional specimens in a series of mainstream riffles near the entrance to
Wyandotte Cave (Figure 1, Table 1).
Zoology 605
We hypothesized that the spotted darters apparently prefer the riffles located
downstream from old rock dams where the velocity is normally high and the water
well-oxygenated. In 1984, we substantiated this hypothesis by locating significant popula-
tions below Rothrock's Dam and Milltown Dam (Figure 1, Table 1). In addition, we
found specimens in two locations near Fredericksburg which is almost 50 river miles
above the Wyandotte Cave location (Figure 1).
At present, the spotted darter is regarded by the Department of Natural Resources
as a species of special concern. Since E. m. maculatum, the nominate subspecies which
our specimens represent, is very difficult to capture throughout most of its former
range, and may do well in only a few locations in the upper Ohio drainage of Penn-
sylvania (1), we feel that aggressive action should be taken to locate this darter in
remaining ideal habitats.
Acknowledgments
Research was supported by a grant-in-aid from Indiana University Southeast, and
in 1983 by a travel grant from the Indiana Heritage Program. Dr. David A. Etnier
of the University of Tennessee verified the identifications. This publication is dedicated
posthumously to Dr. William M. Clay who suggested that we initiate this study.
Literature Cited
1. Etnier, D. Personal communication. Zoology Dept., Univ. of Tennessee, Knox-
ville, Tennessee.
2. Gerking, S.D. 1945. Distribution of the fishes of Indiana. Investigations of Indiana
Lakes and Streams. 3: 1-137.
3. Jordan, D.S. 1877. A partial synopsis of the fishes of upper Georgia; with sup-
plementary papers on fishes of Tennessee, Kentucky, and Indiana. Ann. N.Y.
Lye. Nat. Hist. 1874-77. 11: 307-377.
4. Jordan, D.S. 1890. Report of explorations made during the summer and autumn
of 1888, in the Allegheny region of Virginia, North Carolina, and Tennessee,
and in western Indiana, with an account of the fishes found in each of the river
basins of those regions. Bull. U.S. Fish Comm. 1888. 8: 97-173.
5. Nelson, J.S. and S.D. Gerking. 1968. Annotated key to the fishes of Indiana.
Dept. of Zoology, Indiana Univ., Bloomington, IN. 83 p.
6. Zorach, T. and E.C. Raney. 1967. Systematics of the percid fish Etheostoma
maculatum Kirtland, and related species of the subgenus Nothonotus. Amer. Midi.
Nat. 77(2): 296-322.
Food Habits of Urban American Kestrels, Falco sparverius
Virgil Brack, Jr.,* Ted T. Cable** and Daniel E. Driscoll
Department of Forestry and Natural Resources
Purdue University,
West Lafayette, Indiana 47907.
Introduction
American Kestrels {Falco sparverius) have been noted on the campus of Purdue
University, West Lafayette, Indiana, since 1932 (14), and have nested there since at
least 1979 (3). We noted kestrels nesting on the Purdue campus from 1982-1984.
Similarly, nesting in an urban environment has been observed in downtown Oklahoma
City (2), and on the University of Oklahoma campus (13). Information on the diet
of urban kestrels is limited. On the University of Oklahoma campus, capture of House
Sparrows {Passer domesticus) was observed. In a residential area of Nevada, Missouri,
kestrels ate 5 types of prey; grasshoppers (Orthoptera), Six-lined Race Runners
{Cnemidophorus sexlineatus), House Sparrows, a Horned Lark {Eremophila alpestris),
and an American Robin {Turdus migratorius) (7). In contrast to the little studied urban
American Kestrel, quantitative food habits data are available for the European Kestrel
{Falco tinnunculus) which was studied on the University of Manchester campus (16).
The purpose of this study was to examine the food habits of urban American
Kestrels and the habitats used for hunting. It was of particular interest to see if the
abundant House Sparrow population was preyed upon, and whether the kestrels traveled
from the urban campus setting to hunt short-grass habitats in surrounding areas. These
data are compared with those of more frequently studied rural kestrels.
Materials and Methods
Food Habits Analysis
Pellets and prey parts were routinely collected from beneath a Douglas fir {Pseudot-
suga menziesii) and an American sycamore {Platanus occidentalis) used by American
Kestrels as feeding stations on Purdue University campus from winter 1983 through
spring 1984. Pellet contents were examined under a disecting microscope. Prey parts
and pellets were identified to major taxa, and to species when possible. Pellet contents
were estimated to the nearest 5% of pellet volume if greater than 5%, or 1% if less
than 5%. Each prey part was treated as one pellet. For each sample date, pellet con-
tents were combined and averaged, providing a percent volume of the total for each
prey type.
In 1982-83 the kestrel's diet contained significant quantities of House Sparrows,
and bird-feeder seeds (millet and cracked corn) were consistently associated with these
remains. Therefore, millet was dyed with green food coloring and placed daily at an
existing bird feeding area from 16 to 28 April 1984. Pellets collected after 16 April
were checked for green seed to see if sparrows were being captured on campus.
Habitat Analysis
Areas within a 1 .64 km radius (maximum radius for American Kestrel pairs ranges
from 0.4102 km to 1.8871 km [4]) of the Douglas fir feeding station were cover-mapped
Present address: WAPORA, Inc., 511 Old Lancaster Road, Berwyn, Pennsylvania 19312.
Present address: Department of Forestry, Kansas State University, Manhattan, Kansas 66506.
607
608 Indiana Academy of Science Vol. 94 (1985)
to determine the relative availability of various habitat types as an indicator of the
availability of various prey types. Vegetation was classified as follows: rank grass (bush-
hogged occasionally), shrubland, pasture and cropland, large lawns, forest, and residen-
tial (including roads, parking lots, buildings, and surrounding small lawns). An
Intergraph Interactive Graphics Design systems computer was used for analysis and
graphic plotting.
Kestrel Watches
Coordinated observations of potential kestrel feeding areas were conducted. Some
observers were stationed in study area habitats, while one observer remained on top
of a parking garage with a view of the feeding station and large areas of campus.
Observations of kestrel movements between areas were coordinated between observers
with walkie-talkies; the times and directions of flight were recorded. Time, location,
activity, and sex data were analyzed to see if males and females hunted different habitats.
Results
Food Habits Analysis
During 1983, the kestrel's diet consisted largely of voles, which were present on
all sample dates (Table 1). All parts identified to species were Prairie Voles {Microtus
ochrogaster), but frequently parts could not be identified to species. In winter (February)
and spring (March through May), the diet consisted largely of small mammals, mainly
voles, although Short-tailed Shrews {Blarina brevicauda), a House Mouse {Mus
Table 1 . The diet of American kestrels on Purdue University campus, West Lafayette,
Indiana, in 1983. (Data from pellet analysis by % volume technique). (T indicates < 0.5%).
Other
Other
Date
Microtus
Blarina
mammals
Passer
birds
Insects
Comment
17 Feb.
54
44
2
Mus, Araneida
22 Feb.
95
5
17 Mar.
96
1
4
Lepid., Coleop., Scarab.
24 Mar.
28
72
T
28 Mar.
51
49
30 Mar.
91
1
1
1
Curcul., Formic.
Homop., Scarab.
31 Mar.
48
9
32
10
1
Coleoptera
Orthoptera
5 Apr.
38
10
52
T
Coleoptera
7 Apr.
28
72
11 Apr.
89
3
8
18 Apr.
38
20
40
2
T
Coleoptera
21 Apr.
60
40
22 Apr.
23
20
30
26
1
Synaptomys
Coleoptera
26 Apr.
99
1
Coleoptera
28 Apr.
98
2
Carabidae
Cicindelidae
2 May
26
49
19
6
24 Jun.
24
10
30
35
1
Carduelis
Passerina
5 Jul.
13
20
27
40
Spermophilus
6 Jul.
6
6
10
74
2
1
Colepotera
12 Jul.
64
31
4
Curculionidae
Coleoptera
13 Jul.
60
40
Zoology 609
musculus), and a Southern Bog Lemming (Synaptomys cooperi) were also eaten. No
birds were present in the winter diet. The spring diet contained fewer House Sparrows
than other avian species. Insects were present on 69% of the pellet collection dates,
although they usually were a very small part of the diet. Beetles (Coleoptera) comprised
most of the insects in the winter and spring diet. Ground beetles (Carabidae), scarab
beetles (Scarabaeidae), snout beetles (Curculionidae), and tiger beetles (Cicindelidae)
were also present. Other insects eaten included representatives of the orders Lepidoptera
Homoptera, Orthoptera, and Hymenoptera (ants; Family Formicidae). Spiders
(Araneida) were present in the winter diet.
The 1983 summer (June and July) diet was largely birds. House Sparrows were
eaten most frequently. An American Goldfinch (Carduelis tristis) and an Indigo Bunting
(Passerina cyanea) were also eaten. Small mammals in the diet were largely voles,
although Short-tailed Shrews and a Thirteen-lined Ground Squirrel (Spermophilus
tridecemlineatus) were also eaten. Beetles, including snout beetles, made up the insect
portion of the summer diet.
The diet in 1984 contained mostly birds; House Sparrows were most abundant
(Table 2). Winter and early spring diets consisted almost totally of House Sparrows.
From 13 April throughout spring, voles were present on every sample date, although
birds still dominated the diet. Birds were present in all samples; House Sparrows were
present in 87.5% of the samples. Three partially eaten Mourning Dove nestlings (Zenaida
macroura) were recovered. Dark-eyed Juncos (Junco hyemalis) were found in the diet
on five separate occasions. A European Starling (Sturnus vulgarus) was also eaten.
Insects were few.
Habitat Analysis
Residential areas comprised 50% of the study area. The remaining area was 15%
large lawns, 10% pasture and cropland, 10% forest, 5% shrubland, 5% rank grass,
and 5% water (Figure 1). Kestrels hunted two areas off campus, designated as Areas
1 and 2, which were both rank grass habitats. Most vegetation in Area 1 was greater
than 1 m tall, and included teasel (Dipsacus sylvestris), fescue (Festuca sp.), smart-
weed (Polygonum sp.), mint (Menthus sp.), goldenrod (Solidago sp.), queen anne's
Table 2. The diet of American kestrels on Purdue University campus, West Lafayette,
Indiana, in 1984. (Data from pellet analysis by % volume technique.)
Other
Other
Date
Microtus
mammals
Passer
birds
Insects
Comment
21 Feb.
99
1
24 Feb.
60
40
12 Mar.
100
16 Mar.
98
2
26 Mar.
100
9 Apr.
100
Sturnus
13 Apr.
60
40
16 Apr.
41
19
41
Zenaida
17 Apr.
64
12
24
Junco
19 Apr.
47
6
2
45
Junco
20 Apr.
30
20
50
Junco
21 Apr.
15
65
20
Zenaida
24 Apr.
44
56
25 Apr.
63
36
1
26 Apr.
7
59
33
1
Junco
1 May
50
44
6
Junco
610
Indiana Academy of Science
Vol. 94 (1985)
mnniD
RESI DENTI AL
FOREST
RANK GRASS
PASTURE AND CROPLAND
ROAD ®
LAWN
SHRUBLAND
WATER
KESTREL FEEDING
STATION
Figure 1. Habitat types within a 1.64 km radius of the Douglas fir feeding station
on Purdue University campus, West Lafayette, Indiana. Five known hunting areas
are labeled with numerals.
lace (Daucus carota), and a few scattered smooth sumac (Rhus glabra) and tree-of-heaven
(Ailanthus altissima). Vegetation in Area 2 was less than 1 m tall, and included fescue,
clover (Trifolium sp.), mint, and dandelion (Taraxacum sp.).
Zoology 611
Kestrels were observed capturing small mammals in Area 1, a small waste area
between bridge ramps, and the surrounding residential lawns. In Area 2, many attempted
captures of small mammals were observed. Area 2 contained many vole runways and
burrow entrances 3.5 cm. in diameter. To the east of Area 2 was a large lawn — an
extension of the university airport supporting a large population of Thirteen-lined Ground
Squirrels.
Kestrels were also regularly sighted in three areas along the campus periphery.
Area 3 was a rank grass area, Area 4 a football field, and Area 5 a golf course and
associated athletic fields. Though kestrels from campus were frequently observed heading
towards these areas, we cannot be certain whether the birds observed hunting there
were campus birds.
Kestrels were observed attempting to capture House Sparrows at a bird feeding
station, and at other places on campus. As noted previously, millet and cracked corn
were usually present in pellets containing House Sparrow remains, although no green
dyed seed was found.
Discussion
The American Kestrel preys upon a variety of small mammals, birds, insects,
and herptiles (6, 8); the diet varies considerably by season and locality (1). House Spar-
rows are sometimes eaten in an urban environment (14). In a rural Indiana environ-
ment kestrels ate the following: 78% insects and spiders, 14% mammals, 6% herp-
tiles, and 3% birds (9). It is evident from the present urban study, despite a lack of
data from mid-May through mid-June, that birds and mammals are more heavily preyed
upon. It is likely that our method of pellet analysis underestimates the number of
insects in the diet, because of their small proportion of non-digestible parts, but our
data surely indicate that insects are minor.
Theoretically, predation by an individual raptor will tend to reflect local prey
densities of the species within its range that it is adapted to catch (4). Numerous vole
runways and burrows in Area 2, and observed captures of mammals in Area 1, testify
to the abundance of small mammals in these areas. Small birds were frequently pre-
sent in Area 2, but the kestrels were never seen attempting to capture them. Small
birds, especially House Sparrows, were abundant on Purdue's campus and surround-
ing residential areas. Pellets containing sparrow remains also contained seeds typically
supplied at bird feeders. Kestrels were several times observed attempting to capture
sparrows on campus, but were never seen attempting to catch birds in rank grass habitats.
Outside the study area, on the northern edge of Lafayette, kestrels were seen flying
into town and returning with sparrows. These kestrels regularly hunted a city playground
where they were observed catching sparrows on two occasions.
Area 1 was heavily hunted for rodents. Thus small tracts of "waste areas" pro-
vide important habitat for urban kestrels. As urbanization continues to reduce the
acreage of rural and wild lands, the importance of urban wildlife increases. The pro-
portion of Americans living in urban areas is expected to increase in the future. The
major contact many of these people will make with wildlife will be in this urban setting.
Unlike many species, the kestrel is relatively tolerant of urban pressures, providing
an easily observable and enjoyable urban wildlife experience. Thus, the wise manage-
ment of resources to benefit kestrels and the urban wildlife enthusiast likely includes
those management techniques which provide "waste areas" for kestrel use.
Behavior
As in previous studies (15, 10, 11), the kestrels were often observing caching and
retrieving prey. Unlike the silence noted by Sutton and Tyler (13), frequent calling
612 Indiana Academy of Science Vol. 94 (1985)
accompanied exchange of prey between male and female. Although it has been reported
that kestrels do not use the same nest site in consecutive years (5), the kestrels at Purdue
have nested in the same cavity for three consecutive years.
On 21 April 1984, in area 2, the male and female alternately hovered and pounced,
"leap-frogging" over one another five times while hunting. It is assumed these actions
flushed insects or small mammals which were then pursued. Although male kestrels
have been found to hunt areas with vegetation taller than 1 m, while females prefer
vegetation shorter than 1 m (11), no correlation of sex with vegetation height was
found in this study.
Summary
Several prey items found in this study have not been previously cited in the
literature. These included the following: scarab beetle, snout beetle, tiger beetle, Dark-
eyed Junco, American Goldfinch, Indigo Bunting, and Southern Bog Lemming. The
food habits of urban American Kestrels differed in several ways from the food habits
of rural American Kestrels; fewer mammals and insects were eaten, while more birds
were eaten. Our data, like the Craighead's (4) suggests that prey is taken as available.
It appeared that different habitats frequently were used for capturing different prey
types. Birds, mainly House Sparrows, were readily available in most urban areas, while
mammals were hunted in rank grass and short grass areas. In an urban situation, areas
frequently considered "waste areas" are valuable to this small falcon.
Acknowledgments
We would like to thank individuals who helped with the retrieving of pellets,
gave us data on sightings, and participated on kestrel watches, especially Toni Rogers,
Charles Rosenburg, and Karen Andreef. Russell E. Mumford proved invaluable for
discussion and suggestions throughout the project, and he reviewed the manuscript.
Bobby Witcher was a source of inspiration to one and all.
Literature Cited
1. Bent, A.C. 1938. Life Histories of North American Birds of Prey. Part 2. Bull.
U.S. Nat. Mus. 170:1-482.
2. Black, E.A. 1979. American Kestrel possibly two-brooded in central Oklahoma.
Bull. Oklahoma Ornith. Soc. 12:29.
3. Burr, I.W. 1979. The birds of Tippecanoe County, Indiana. Indiana Aud. Quart.
57:1-43.
4. Craighead, J.J. and F.C. Craighead, Jr. 1969. Hawks, Owls, and Wildlife. Dover
Publications, New York, N.Y. 433 pp.
5. Hamerstrom, F., F.N. Hamerstrom, and J. Hart. 1973. Nest boxes: an effective
management tool for kestrels. J. Wildl. Manage. 37:400-403.
6. Heintzelman, D.S. 1964. Spring and summer Sparrow Hawk food habits. Wilson
Bull. 76:323-330.
7. Lamore, D.H. 1963. Prey of a Sparrow Hawk family when raising young. Wilson
Bull. 75:461.
8. Mills, G.S. 1976. American Kestrel sex ratios and habitat selection. Auk 93:740-748.
9. Phillips, R.E. and CM. Kirkpatrick. 1969. Hawks and Owls of Indiana. Indiana
Department of Natural Resources, Division of Fish and Game. Bull. 8:1-38.
10. Roest, A.I. 1957. Notes on the American Sparrow Hawk. Auk 74:1-19.
11. Stendell, R. and L. Waian. 1968. Observations on food caching by an adult female
Sparrow Hawk. Condor 70:187.
12. Stinson, C.H., D.L. Crawford, and J. Lauthner. 1981. Sex differences in winter
Zoology 613
habitat of American Kestrels in Georgia, J. Field Ornith. 52:29-35.
13. Sutton, G.M. and J.D. Tyler. 1979. On the behavior of American Kestrels nesting
in town. Bull. Oklahoma Ornith. Soc. 12:25-29.
14. Test, L.A. and F.H. Test. 1932. Birds of Tippecanoe County, II. Proc. Ind. Acad.
Sci. 41:465-481.
15. Tordoff, H.B. 1955. Food storing in the Sparrow Hawk. Wilson Bull. 67:139-140.
16. Yalden, D.W. 1980. Notes on the diet of urban kestrels, Falco tinnunculus. Bird
Study 27:235-238.
Parasitic Endohelminths from Fishes of Southern Indiana
Richard L. Buckner
Division of Natural Sciences and Mathematics
Livingston University
Livingston, Alabama 35470
Melvin W. Denner
Indiana State University Evansville
8600 University Boulevard
Evansville, Indiana 47712
Daniel R. Brooks
Department of Zoology
The University of British Columbia
6270 University Boulevard
Vancouver, B.C., Canada V6T 2A9
and
Shareen C. Buckner
Division of Natural Sciences and Mathematics
Livingston University
Livingston, Alabama 35470
Introduction
There are numerous surveys of the parasites of freshwater fishes; however, there
is still little information available on the parasites of fishes from certain areas. One
such area is Indiana. The most extensive listing of parasites of Indiana fishes occurs
in Dolley's 1933 article on the biology of the St. Joseph River. He reported 10 parasites
of which only three were identified to species. A list of fish parasites reported from
Indiana fishes by Dolley (7) and other authors is given in Table 1 . The present study
provides additional information on the parasites of fishes from this poorly studied area.
Table 1 . Helminths known from Indiana, their piscine hosts, sites of infection, counties
where collected, and bibliographic references.
Helminth
Host
Site
County
Ref.
Monogenea
Gyrodactylus bairdi
Cottus bairdi
gl
St. Joseph
18
Woods & Mizelle, 1957
Gyrodactylus limi
Umbra limi
"
"
14,18
Woods & Mizelle, 1957
Digenea
Azygia sp.
Amia calva
st
"
7
Azygia acuminata
"
/<
?
10
Goldberger, 1911
(Lost Lake)
Azygia bulbosa
a
"
Marshall
10
Goldberger, 1911
Clinostomum complanatum
Umbra limi
bs
St. Joseph
14
(Rudolphi, 1819)
Cyathocotyloides sp.
Ictalurus punctatus
in
Tippecanoe
10
Hassallius hassalli
Ambloplites rupestris
st
Marshall
10
Goldberger, 1911
Holostephanus ictaluri
Ictalurus punctatus
in
Tippecanoe
17
Vernberg, 1952
Holostomid cyst
Micropterus dolomieu
615
fl
St. Joseph
7
616 Indiana Academy of Science
Table 1. — Continued
Vol. 94 (1985)
Helminth
Host
Site
County
Ref.
Leuceruthrus micropteri
Amia calva
St
Marshall
10
Marshall and Gilbert, 1905
Micropterus dolomieu
Micropterus salmoides
-
«
«
Microphallus opacus
Amia calva
in, st
St. Joseph
7
(Ward, 1894)
Neascus sp.
Ambloplites rupestris
Chaenobrittus gulosus
vs
«
"
Neochasmus umbellus
Morone mississippiensis
?
Gibson and
12
Van Cleave & Mueller, 1932
Monroe
Phyllodistomum brevicecum
Umbra limi
ub
Tippecanoe
15
Steen, 1938
"
"
St. Joseph
14
Phyllodistomum undulans
Cottus bairdi
"
Tippecanoe
15
Steen, 1938
Pristotrema manteri
Scaphirhynchus
in
"
4
Cable, 1952
platorhynchus
Prohemistomum chandleri
Micropterus dolomieu
vs
"
17
Vernberg, 1952
Micropterus salmoides
"
"
"
Cestoda
Cestodarian
Catostomus commersoni
Cyprinus carpio
in
lv,st
St. Joseph
7
Glaridacris catostomi
Catostomus commersoni
in
"
"
Cooper, 1920
Proteocephalus sp.
Esox lucius
in
"
"
Proteocephalus ambloplites
Morone mississippiensis
?
Gibson and
12
(Leidy, 1887)
Monroe
Trianophorus sp.
Perca flavescens
Iv
St. Joseph
7
Acanthocephala
Pomphorhynchus bulbocolli
Ameirus melas
st
St. Joseph
"
Linkins in Van Cleave, 1919
Catostomus commersoni
in
"
"
Cyprinus carpio
vs
"
"
Nematoda
Nematode
Ambloplites rupestris
in
"
"
Philometra sp.
Morone mississippiensis
7
Gibson and
Monroe
12
bs = body surface, fl = flesh, gl = gills, in = intestine.lv = liver, st = stomach, ub = urinary bladder, and vs = viscera.
Materials and Methods
From the fall of 1976 through the spring of 1978 fishes were collected from small
streams of Vanderburgh and Posey Counties, Indiana, and examined for endohelminths.
Fishes were collected by seining and maintained alive or kept on ice until necropsied,
usually within 24 hours of capture. The parasites were fixed in AFA, stained with
Mayer's carmalum, and mounted in Canada balsam. Nematodes were fixed in acetic
acid or hot 70% ethanol and examined as temporary mounts in lactophenol. Represen-
tative specimens have been deposited in the University of Nebraska State Museum,
Harold W. Manter Laboratory (HWML), Lincoln, Nebraska.
Results and Discussion
A total of 386 fish representing 10 families and 25 species were examined for
parasites. Of the total number of fishes examined 215 (56%) were found to be infected.
Helminths were found in all species except Aphredoderus sayanus (8 specimens ex-
amined). Seventeen species of helminths were collected — 3 digeneans, 5 cestodes, 5
nematodes, and 4 acanthocephalans — none of which have been previously reported
from Indiana. A listing of these helminths and their hosts is given in Table 2.
Zoology
617
Table 2. Prevalence and intesity of endohelminths collected in Posey and Vanderburgh
Counties, Indiana, 1976-1978.
Helminth
Host
HWML
No.
Preval. lnten. County
Digenea
Allocreadium lobatum
Wallin, 1906
Alloglossidium corti
(Lamont, 1921)
Pisciamphistoma stunkardi
(Holl, 1929)
Cestoda
Biacetabulum biloculoides
Mackiewicz & McRae, 1962
Bothhocephalus formosus
Mueller & Van Cleave, 1932
Corallobothrium fimbriatum
Essex, 1927
Megathylacoides intermedia
(Fritts, 1959)
Proteocephalus pinguis
LaRue, 1911
Nematoda
Camallanus ancylodirus
Ward & Magath, 1916
Camallanus muttilineatus
Kung, 1948
Dichelyne sp.
Philometra nodulosa +
Thomas, 1929
Spiniteclus micracanthus
Christian, 1972
Acanthocephala
Acanthocephalus dirus
(Van Cleave, 1931)
Gracilisentis gracitisentis
(Van Cleave, 1913)
Neoechinorhynchus cylindratus
(Van Cleave, 1913)
Semotilus atromaculatus
—
5/64
1-2
V.
Ictalurus natalis
21464
1/8
1
V.
Lepomis cyanellus
21465
1/47
1
V.
*Carpiodes velifer
*Fundulus notatus
* Lepomis cyanellus
•Notropus umbratilus
* Phenacobius mirabilis
*Pimephales notatus
Ictalurus melas
Ictalurus nebulosus
Esox americanus
"Lepomis cyanellus
*Lepomis macrochirus
*Carpiodes velifer
Ictalurus natalis
Lepomis cyanellus
Carpiodes velifer
Lepomis macrochirus
21469
2/3
21472
3/34
1,3
—
4/26
1
P.
—
1/47
1
P.
—
3/22
1-4
P.
—
1/3
1
P.
21487
10/31
1
P.
21466
1/6
1
P.
21468
2/8
1,5
V.
21467
1/8
1
P.
21474
2/47
1
P.
21475
1/34
1
V.
21470
1/3
1
p.
—
1/8
1
V.
21473
1/47
1
V.
21471
1/3
1
p.
14
P.,V.
Aplodinotus grunniens
22855
2/2
2,6
P,V
Campostoma anomalum
22853, 22854
5/12
1-9
P,V
* Carpiodes velifer
22847
3/3
1-22
P.
Cyprinus carpio
22859
1/2
18
V.
*Ericymba buccata
22848
15/29
1-44
P.,V
Esox americanus
22846
3/8
1-4
P,V
*Etheostoma squamiceps
22843
9/9
2-28
V.
*Fundulus notatus
22845
19/26
1-9
P.,V
Ictalurus melas
—
2/6
2,42
P.V
Lepomis cyanellus
22838, 22861
30/47
1-15
P..V
Lepomis macrochirus
22851
10/34
1-15
P..V
Lepomis megalotis
22842
7/12
1-123
P.,V
Micropterus salmoides
22844, 22852
3/3
2-15
P.,V
Notemigonus crysoleucas
22856
1/14
1
V.
*Notropus atherinoides
22857
3/10
1-4
P..V
Notropus spilopterus
22860
7/12
1-13
P.,V
Notropus umbratilis
22849
13/22
1-11
p.
* Phenacobius mirabilis
22839, 22840
3/3
1-4
P..V
Pimephales notatus
22858
15/31
1-23
p.
*Pomoxis annularis
—
1/13
15
V.
Semotilus atromaculatus
2284!
54/64
1-50
P.,V
Dorosoma cepedianum
—
2/6
5/7
p.
Micropterus salmoides
22837
1/3
Indiana Academy of Science
Vol. 94 (1985)
Table 2. — Continued
Helminth
Host
HWML
No. Preval. Inten. County
Neoechinorhynchus cylindratus Micropterus salmoides
(Van Cleave, 1913)
Neoechinorhynchus notemigoni Notemigonus crysoleucas
Dechtiar, 1967
Pomphorhynchus rocci Aplodinotus grunniens
Cordonnier & Ward, 1967
22837
22850
21476
21477
1/3
1/4
1/1
1/1
V.
V.
p.
V.
* = new host record, + = occurred in cheek galleries, P. = Posey County, and V. = Vanderburgh County
Acanthocephalus dims (Van Cleave, 1931) was the most frequently found parasite,
occurring in 21 species (8 families). Heavy fish predation on isopods may account
for the prevalence of this parasite. Isopods, many of which were found to be infected
with A. dims, occurred in large numbers in most of the areas where fish were collected
and were commonly found in the intestines of the fish examined. Because of the advanced
state of maturity of A: dims cystacanths, it is often difficult to distinguish definite
hosts from accidental hosts. However, in the present collection gravid worms occurred
in all 21 species of fish.
Amin (1) analyzed species of the genus Acanthocephalus occurring in fishes of
North America. He concluded that A. jacksoni Bullock, 1962 and A. parksidei Amin,
1975 represent geographic variants of A. dims. He noted marked variation between
northern and southern populations. The number of proboscus hooks per longitudinal
row was one of the more prominent differences. Specimen from the present collection
represent a deme situated between the populations examined by Amin (1). A wide
range of variation, especially regarding proboscus armature, was observed within this
collection.
Three specimens of Neoechinorhynchus notemigoni Dechtiar, 1967 were recovered
from one Notemigonus crysoleucas. This species was described from Lake Ontario,
Canada (6). In 1983 Buckner (3) reported it from N. crysoleucas of Alabama and
Mississippi. This is the third report of Neoechinorhynchus notemigoni. These specimens
are in agreement with those reported by Buckner (3).
Bothriocephalus formosus Mueller and Van Cleave, 1932 occurred in five species
but was more prevalent in Pimephales notatus. Specimens from the present collection
agree with the original description except in number of testes and lengths of scolices.
These specimens possess 50 to 70 testes and scolices up to 600 mm long whereas Mueller
and Van Cleave (13) reported 30 to 45 testes and scolices up to 475 mm long.
A procedure for discrimination between Pomphorhynchus bulbocolli Linkins in
Van Cleave, 1919 and P. rocci Cordonnier and Ward, 1967 was provided by Huffman
and Nickol (11) and refined by Gleason and Huffman (9). Specimens of P. rocci were
identified from two Aplodinotus grunniens according to the procedure outlined by
these authors. Lengths of proboscis hooks in the 60 to 80% position region ranged
from 46 to 62 um (54) and the ratio of anterior hook lengths to most massive hook
lengths ranged from 0.81 to 1.00. Specimens were sent to Dr. Brent Nickol, the University
of Nebraska-Lincoln, for verification. He concurred with the identification, as deter-
mined by the above procedure. This is the first identification of P. rocci from a
freshwater fish collected in a non-coastal habitat.
Megathylacoides intermedia (Fritts, 1959) Befus and Freeman, 1973 was iden-
tified from two Ictalurus nebulosus. This cestode was originally described from /.
Zoology 6 1 9
nebulosus of Idaho by Fritts (8) as Corallotaenia intermedia. This is the first report
of this species east of the Mississippi River.
Camallanus ancylodirus Ward and Magath, 1916 was collected from two species
of centrarchids, Lepomis cynallus and L. macrochirus. These occurrences may be
accidental since, with the exception of a report in Stizostedion vitreum by Sutherland
and Holloway (16), C. ancylodirus is known only from catostomids (2).
Immature C. multilineatus Kung, 1948 was collected from Carpiodes velifer. This
nematode is previously known only from its original description from a North American
frog that died in a London zoo.
Literature Cited
1. Amin, O.M. 1984. Variability and redescription of Acanthocephalus dims (Acan-
thocephala: Echinorhynchidae) from freshwater fishes in North America. Proc.
Helminthol. Soc. Wash. 51:225-237.
2. Baker, M.R. 1979. Redescription of Camallanus ancylodirus Ward and Magath
1916 (Nematoda: Camallanidae) from freshwater fishes of North America. J.
Parasitol. 65:389-392.
3. Buckner, R.L. 1983. Occurrence of two species of Neoechinorhynchus (Acan-
thocephala) in golden shiners of Alabama and Mississippi. Proc. Helminthol. Soc.
Wash. 50:176-178.
4. Cable, R.M. 1952. On the systematic position of the genus Deropristis, of
Dihemistaphanus sturionis Little, 1930 of a new digenetic trematode from a
sturgeon. Parasitol. 42:85-91.
5. Cable, R.M. and W.B. Vernberg. 1949. The occurrence of an adult holostome
(Trematode: Cyathocotylidae) in the intestine of a fish. J. Parasitol. 35: Sup.
pg. 20 (Abst. 33).
6. Dechtiar. A. 1967. Neoechinorhynchus notemigoni n. sp. (Acanthocephala:
Neoechinorhynchidae) from golden shiner of Lake Ontario. Can. J. Zool.
45:155-159.
7. Dolley, J.S. 1933. Preliminary notes on the biology of the St. Joseph River. Am.
Midi. Nat. 14:193-227.
8. Fritts, D.H. 1959. Helminth parasites of the fishes of northern Idaho. Trans.
Am. Microsc. Soc. 78:194.
9. Gleason, L.N. and D.G. Huffman. 1981. Meristogram analysis of a collection
of Pomphorhynchus bulbocolli from south-central Kentucky. J. Parasitol.
67:133-134.
10. Goldberger, J. 1911. Some known and three new endoparasitic trematodes from
American freshwater fish. Hgy. Lab. U.S. Pub. Health and Mar-Hosp. Serv.
Bull. 71:7-35.
11. Huffman, D.G. and B.B. Nickol. 1978. Meristogram analysis of the acan-
thocephalan genus Pomphorhynchus in North America. J. Parasitol. 64:851-859.
12. McReynolds, M. and J.D. Webster. 1980. Parasites of the yellow bass from two
southern Indiana lakes. Proc. Indiana Acad. Sci. 89:154-158.
13. Mueller, J.F. and H.J. Van Cleave. 1932. Parasites of Oneida Lake fishes. II.
Descriptions of new species and some general taxonomic considerations, especially
concerning the trematode family Heterophyidae. Bull. New York State Coll. Forest.
5(2c). Roosevelt Wild Life Ann. 3:79-137.
14. Peckham, R.S. and C.F. Dineen. 1957. Ecology of the central and minnow Um-
bra limi (Kirtland). Am. Midi. Nat. 58:222-231.
15. Steen, E.B. 1938. Two new species of Phyllodistomum (Trematoda: Gorgoderidae)
from Indiana fishes. Am. Midi. Nat. 20:201-210.
620 Indiana Academy of Science Vol. 94 (1985)
16. Sutherland, D.R. and H.L. Holloway, Jr. 1979. Parasites of fish from the Missouri,
James, Sheyenne, and Wild Rice Rivers in North Dakota. Proc. Helminthol. Soc.
Wash. 46:128-134.
17. Vernberg, W.B. 1952. Studies on the trematode family Cyathocotylidae Poche,
1926, with the description of a new species of Holostephanus from fish and the
life history of Prohemistomum chandleri sp. nov. J. Parasitol. 38:327-340.
18. Woods, R.A. and J.D. Mizelle. 1957. Studies on monogenetic trematodes. XXI.
North American Gyrodactylinae, Dactylogyrinae and a new host record for
Urocleidus dispar (Mueller, 1936). Am. Midi. Nat. 57:183-202.
The Present Distribution and Status of the
Eastern Woodrat, Neotoma floridana, in Indiana
Wynn W. Cudmore
Department of Life Sciences
Indiana State University
Terre Haute, Indiana 47809*
Introduction
Although the genus Neotoma is best represented in southwestern United States,
the eastern woodrat, Neotoma floridana, has a rather widespread distribution in both
western and eastern United States. The species occurs as far west as Colorado and
in the east from extreme southern Illinois, Indiana, Ohio and New York south to Loui-
siana, Alabama, Georgia and Florida. It is absent along the coast from New Jersey
to central South Carolina and Georgia. The presence of N. floridana in Indiana was
not documented until 1930 (Hickie and Harrison 1930) although there were references
to what were probably woodrats as early as 1872 (Cope 1872, Packard 1888, Blatchley
1897, Lyon 1936). Cave deposits from Missouri (Parmalee and Jacobson 1959, Par-
malee 1967), Illinois (Parmalee, et al. 1961, Parmalee 1967), Indiana (Bader and Hall
1960, Richards 1972, Parmalee, et al. 1978) and Ohio (Goslin 1955) indicate a historic
distribution of TV. floridana much further northward than the current distribution.
In Indiana the species has been taken only from locations in Harrison County (Hickie
and Harrison 1930) and Crawford County but the limits of its distribution in the state
had not been determined. Neotoma floridana is currently included on the threatened
list for Indiana (McReynolds, Whitaker and Gammon 1979) and the endangered list
for Illinois (Nawrot and Klimstra 1976). Climatic factors have been proposed to ex-
plain the restricted range of the species in Indiana and Illinois (Richards 1972, Nawrot
and Klimstra 1976).
The purpose of the present study was to determine the distribution of N. floridana
in Indiana and to investigate limiting factors by comparing characteristics of active
and inactive woodrat locations with those of areas lacking woodrats. Population estimates
were conducted to evaluate the status of the species in the state.
Methods and Materials
To determine the distribution of N. floridana in Indiana, 100 potential sites were
located on topographic maps (scale 1:24,000) of Perry, Crawford, Harrison, Floyd,
Clark and Jefferson Counties and were inspected for evidence of woodrats. Sign in-
cluded fresh cuttings, debris piles, fecal deposits and nests. Sites with questionable
or old sign were livetrapped to verify their occurrence. Characteristics of all sites, in-
cluding extent of cliff, number and quality of openings, rock type (limestone, sand-
stone or shale), and exposure, were recorded and comparisons made between active,
inactive, and uninhabited sites to determine habitat requirements.
The availability of potential den sites was assessed at six Neotoma localities. Open-
ings and crevices were judged to be potential den sites if they were similar in
characteristics to active den sites. Active and potential den sites were counted and
described for lengths of cliff from 480 to 1470 m (X" = 767 m). The relationship
between potential den site density and Neotoma density was tested by calculating Pear-
son's correlation coefficient for these two variables.
♦Present address: Zoology Department— Wildlife Biology, Washington State University, Pullman, Washington
99164-4220
621
622
Indiana Academy of Science
Vol. 94 (1985)
Neotoma populations were estimated at six sites using a mark-recapture method.
Forty-four livetraps set for three consecutive nights and one week later for two addi-
tional nights would generally trap all woodrats within a trapping site. Traps were baited
with peanut butter and rolled oats and set at or near all Neotoma dens, fecal deposits
and cutting piles within the area. Population estimates are given as number of animals
per 1000 m of cliff for each site. A statewide estimate of the Neotoma population
was attempted by determining the amount of available habitat in the state and apply-
ing the most appropriate population estimate. Ohio River bluffs within the range of
Neotoma were measured on topographic maps and categorized as being most similar
in site characteristics to one of the six sites where estimates had been made. Lengths
of cliffs in each category were totalled and the density estimate applied to the total
length. Population estimates for all categories were then totalled for the state estimate.
Results and Discussion
A total of 100 sites was inspected for evidence of woodrats (Figure 1). Over-
lapping of sites that were too close to be represented by more than one point accounts
for the discrepancy between the number of sites checked and the number of points
on the map. A more detailed view of Neotoma distribution in Perry, Crawford and
Harrison Counties is given in Figure 2. Twenty-four sites had evidence of woodrats;
twenty sites harbored active populations while four were inactive sites where only old
sign was observed. Active sites were restricted primarily to the limestone bluffs along
the Ohio River from the Little Blue River in Crawford Co. east to Evan's Landing,
Harrison Co. (Figure 2).
Examination of active Neotoma sites with respect to cliff extent, exposure, number
and types of openings and rock type resulted in a characterization of sites that are
suitable for woodrats in Indiana. All active cliff sites (N = 19) have a southern com-
ponent to their exposure ranging from SW to ESE. One active site is a cave not associated
Figure 1. Locations of Sites Inspected for Evidence of Neotoma floridana in Southern
Indiana.
Zoology
623
10 kilometers
Figure 2. Distribution of Neotoma floridana in Indiana (dashed line corresponds
to Blue River).
with cliffs. All but two sites (nNIO and nN17) are associated with extensive bluffs
along the Ohio River. These two sites are dry caves without running water. Most caves
not associated with cliffs along the Ohio River that were inspected tended to be wet
caves and did not harbor woodrats. Moisture may be an important factor on cave
sites not associated with cliffs. Caves or cave-like openings were present on 13 of 20
active sites (65.0%) and all active sites had moderate to abundant numbers of crevices
or openings suitable for woodrat dens. The relationship between number of potential
dens sites and woodrat densities was examined for seven Neotoma sites (Figure 3).
The two variables are highly correlated (r = 0.932) indicating that the availability
of den sites may be an important factor determining the density of woodrats. All active
sites were located in limestone bedrock. Sandstone sites that satisfied exposure and
cliff extent requirements generally lacked sufficient numbers of openings suitable as
den sites. Appropriate characteristics for a site to harbor woodrats therefore, appear
to be extensive limestone cliffs with a southern component to their exposure and suitable
rock formations for den sites. Dry caves not associated with cliffs and abandoned
buildings provide possible alternative habitats.
Seventy-six sites harboring no woodrats were examined to test the hypothesis that
the above factors are in fact determining the distribution of woodrats. Seventy-four
of 76 sites could be judged unsuitable as Neotoma habitat based on extent of cliff,
abundance of openings, exposure and isolation. Two remaining sites that satisfied these
requirements, but did not harbor woodrats may be lacking an additional factor that
was not examined.
Woodrat populations were estimated by mark and recapture on six sites. A total
of 1253 trap-nights resulted in 335 captures of 115 animals. Fifty-three males (46.1%)
624
Indiana Academy of Science
Vol. 94 (1985)
80-i
NO. NEOTOMa/
1000 m
NO. POTENTIAL DEN SITEs/lOOOm
Figure 3. Relationship between Neotoma Density and Density of Potential Den Sites.
and 61 females (53.0%) were captured for a sex ratio of 0.87:1. The sex of one animal
could not be determined. There were 143 captures of males (2.7 captures per male)
and 191 captures of females (3.1 captures per female). This difference is due to the
greater tendency for males to wander away from den sites and has been reported
previously by Rainey (1956). Population estimates for individual sites are given in Table
1. Woodrat densities averaged 27.5 animals per 1000 m of cliff and ranged from 8.3
to 71.9 animals per 1000 m. Variation in densities between sites is probably most closely
related to the availability of den sites (Figure 3). The density of juniper (Juniperus
virginiana), however, was also positively correlated with woodrat densities (r = 0.768)
and may be an important factor influencing population levels. Interestingly, woodrat
densities were not strongly correlated (r = -0.478) with the density of Tree-of-heaven
{Ailanthus altissima), the major food of Indiana woodrats. Site 82 harbored the greatest
density of woodrats (71.9 per 1000 m) and also yielded the greatest number of cap-
Table 1 . Estimated woodrat densities on six Ohio River bluff sites in extreme southern
Indiana.
Site ft
tt Neotoma
Length of
Cliff Censused (m)
tt Neotoma/
1000 m Cliff
tt Captures/
100 tn
30
82
15
930
16.1
30.6
12
780
15.4
17.4
4
480
8.3
5.5
22
1470
15.0
20.2
41
570
71.9
50.5
21
545
38.5
28.2
X" = 27.5
Zoology 625
tures per unit effort. Over half of the trapping effort resulted in captures on this site
(Table 1). The lowest densities were obtained on site 8 where only four woodrats were
trapped for an estimated density of 8.3 per 1000 m of cliff. The site also produced
the smallest return per unit trapping effort with only 5.5 captures per 100 trap-nights.
Measurements from topographic maps yielded an estimate of 43,855 m of Ohio
River bluff present within the range of N. floridana in Indiana. All Ohio River bluffs
occurring from the Little Blue River in Crawford Co. east to Evan's Landing in Harrison
Co. were included except those areas that were surveyed and found to lack woodrats.
This figure is considered to be an estimate of habitat available to woodrats in the
state although some caves occurring away from the Ohio River probably harbor small
populations. If the mean value of 27.5 Neotoma per 1000 m of cliff is applied to
this figure a statewide estimate of 1206 animals results. This is probably an overestimate
of the state population due to the uneven distribution of available cliff as being most
similar in site characteristics to one of the sites where population estimates had been
made and then to apply that estimate to the total length in each category. Lengths
of cliff assigned to each site category were as follows:
Site ft Length of Cliff
1 3855 m
2 7373 m
8 9181 m
30 17422 m
82 1084 m
98 4940 m
A state estimate of 781 woodrats were obtained using this method. Neotoma
floridana is currently included on the threatened list for Indiana (McReynolds, Whitaker
and Gammon, 1979); however, based on this estimate and the restricted range of the
species in the state (Figure 2) it is recommended that the species be elevated to
"endangered" status.
Previously known localities for N. floridana in Indiana include Tobacco Landing
(Hickie and Harrison 1930), Harrison State Forest (Kirkpatrick and Conaway 1948)
and a site 3 mi. SE of Wyandotte Cave (Mumford and Whitaker 1982) in Harrison
Co. and Wyandotte Cave (Mumford and Whitaker 1982) in Crawford Co. Mumford
(1969) discounted a report of Neotoma occurring 5 mi. SW of Bloomington (Wayne
1960) as being Rattus norvegicus. Rose (1982) reported the occurrence of a single
Neotoma skull among 630 prey items in Barn owl pellets and a woodrat colony in
the Coal Knobs area of Spencer County. The skull has been lost, however, and I have
since inspected the Coal Knobs site and found no evidence of woodrats. Whitaker
(unpublished) also examined this area and additional Barn owl prey items from the
same area and found no evidence of woodrats. Although the area has some sandstone
outcrop, it is generally unsuitable for Neotoma.
It is clear from "Recent" fossil evidence that the Eastern woodrat once had a
much more extensive distribution in Indiana than it presently does. Bader and Hall
(1960) reported N. floridana remains "probably less than a few hundred years old"
from Sullivan's Cave, 1.5 mi. W of Springville in Lawrence County. Richards (1972)
reported woodrat bones from ten sites in Indiana as far north as western Monroe County
and as far east as Jennings County. Based on fauna associated with these deposits
he estimated their age to be a few thousand years old. Parmalee, Munson and Guilday
(1978) found woodrat remains of late Pleistocene age at Harrodsburg Crevice in Monroe
County. The present and historic distributions of N. floridana in Missouri, Illinois
626 Indiana Academy of Science Vol. 94 (1985)
and Ohio parallel the situation in Indiana with present distributions restricted to extreme
southern regions and more widespread historic distributions.
As indicated from "Recent" fossil evidence, TV. floridana probably occupied most
of the karst region of southern Indiana prior to the Wisconsin glaciation. Parmalee,
Munson and Guilday (1978) proposed that extreme periods of climatic fluctuation during
the late Wisconsin exterminated northern populations and that woodrats have failed
to repopulate since that time. The southern-most advance of the glacial boundary lies
immediately north of Monroe County and brought boreal conditions there. Richards
(1972) indicated, however, that temperature alone would not seem to play an impor-
tant role in the distribution of the species since it ranges through the cooler Appalachian
Mountains. He proposed that some other ecological factor, possibly an indirect result
of a mild climatic change, caused the depopulation, There is some evidence, however,
that woodrats may be sensitive to temperature changes. Brown and Lee (1969)
demonstrated a "Bergmann's Response" (i.e. large body size in northern latitudes)
in four species of woodrats (Neotoma lepida, TV. cinerea, TV. albigula and TV. fuscipes).
Only those homeotherms in which environmental temperature has profoundly influenced
reproduction and mortality would be expected to show such a response. Nawrot and
Klimstra (1976) and Fitch and Rainey (1956) attributed population declines to below-
average temperatures and above-average snowfall in winter in Illinois and Kansas,
respectively.
During depopulation of northern areas in the late Wisconsin, woodrats were prob-
ably able to survive due to the milder winters of extreme southern Indiana. Current
populations occupy cliff sites with southern exposure and abundant openings suitable
for den sites and it is likely that these sites were also occupied at this time. Southern
exposure cliffs provide a microhabitat that is significantly warmer than surrounding
microhabitats. Herbaceous plant species flowered as early as 19 February (Cardamine
parviflora) on southern exposure cliffs in Harrison County, Indiana and many early
spring species on these sites were flowered one to two weeks earlier than in adjacent
woodlands. Crim (1961) reported that 95% of 370 sites of woodrat activity were situated
near SW facing rock formations in southern Illinois. Strong positive correlation be-
tween den site availability and population density (Figure 3) indicates that the presence
of suitable rock formations for den sites is an important limiting factor. In Illinois,
woodrats were eliminated from marginal habitat with only a few crevices in severe
winter weather but not from sites with many crevices, faults and ledges (Nawrot and
Klimstra 1976). Deep crevices and especially caves may serve to moderate temperature
extremes during periods of severe weather. Several authors (Heisler 1941, Sands 1951,
Crum 1961) have reported the availability of suitable rock shelters as the most impor-
tant limiting resource for TV. floridana at the northern limits of its range.
Woodrats have been unable to repopulate the karst area in Indiana probably due
to their slow migration (Richards 1972) and reproduction rates (Worth 1950). Nawrot
and Klimstra (1976) indicated that dispersion from present populations in Illinois may
be hindered by the obstruction of natural dispersal routes by man. These barriers in-
clude reservoir inundation of extensive outcrops, isolation of outcrops by agricultural
lands, residential development, stone quarries and highways. These authors felt that
these barriers would be sufficient to prevent natural repopulation of the former range
of the woodrat in Illinois. Similar conditions may be contributing to the hindrance
of migration in Indiana.
Literature Cited
1. Bader, R.S. and J.S. Hall. 1960. Mammalian remains from an Indiana cave. J.
Mammal. 41:111-112.
Zoology 627
2. Blatchley, W.S. 1897. Indiana caves and their fauna. 21st Ann. Rep. Dept. Geol.
Nat. Res. Indiana for 1896:121-212.
3. Brown, J.H. and A.K. Lee. 1969. Bergmann's rule and climatic adaptation in
woodrats (Neotoma). Evolution 23:329-338.
4. Cope, E.D. 1872. Observations on Wyandotte Cave and its fauna. Amer. Nat.
6:406-422.
5. Crim, J. A. 1961. Habitat of the woodrat in southern Illinois. M.S. Thesis, So.
111. Univ., Carbondale, 111. 83 pp.
6. Fitch, H.S. and D.G. Rainey. 1956. Ecological observations on the woodrat,
Neotoma floridana. Publ. Mus. Nat. Hist. Univ. Kansas 8:499-533.
7. Goslin, R.M. 1955. Animal remains from Ohio rock shelters. Ohio J. Sci.
55:358-362.
8. Heisler, W.T. 1941. Allegheny woodrat populations. M.S. Thesis, Penn. State
College. 77 pp.
9. Hickie, P.F. and T. Harrison. 1930. The Allegheny woodrat in Indiana. Amer.
Midi. Nat. 12:169-174.
10. Kirkpatrick, CM. and C.H. Conaway. 1948. Some notes on Indiana mammals.
Amer. Midi. Nat. 39:128-136.
11. Lyon, M.W., Jr. 1936. Mammals of Indiana. Amer. Midi. Nat. 17:1-384.
12. McReynolds, H.E., J.O. Whitaker, Jr. and J.R. Gammon. 1979. Development
of a proposed list of endangered and threatened vertebrate animals for Indiana.
Proc. Ind. Acad. Sci. 88:166-170.
13. Mumford, R.E. 1969. Distribution of the mammals of Indiana. Monog. No. 1,
Ind. Acad. Sci., Indianapolis. 114 pp.
14. Mumford, R.E. and J.O. Whitaker, Jr. 1982. Mammals of Indiana. Indiana
University Press, Bloomington. 537 pp.
15. Nawrot, J.R. and W.O. Klimstra. 1976. Present and distribution of the endangered
southern Illinois woodrat, Neotoma floridana illinoensis. Nat. Hist. Misc. Chic.
Acad. Sci. No. 196:1-12.
16. Packard, A.S. 1888. Cave fauna of North America, with remarks on the anatomy
of the brain and the origin of the blind species. Mem. Nat. Acad. Sci. 4:3-156.
17. Parmalee, P.W. 1967. A recent bone deposit in southwestern Illinois. Nat. Spel.
Soc. Bull. 29:119-147.
18. Parmalee, P.W., R.A. Bieri and R.K. Mohrman. 1961. Mammal remains from
an Illinois cave. J. Mammal. 42:119.
19. Parmalee, P.W. and K.W. Jacobson. 1959. Vertebrate remains from a Missouri
cave. J. Mammal. 40:401-405.
20. Parmalee, P.W., P.J. Munson and J.E. Guilday. 1978. The Pleistocene mam-
malian fauna of Harrodsburg Crevice, Monroe County, Indiana. Nat. Spel. Soc.
Bull. 40:64-75.
21. Rainey, D.G. 1956. Eastern woodrat, Neotoma floridana: life history and ecology.
Publ. Mus. Nat. Hist. Univ. Kansas 8:535-646.
22. Richards, R.L. 1972. The woodrat in Indiana: Recent fossils, Proc. Ind. Acad.
Sci. 81:370-375.
23. Rose, R.K. 1982. Small mammals of southern Indiana. Proc. Ind. Acad. Sci.
91:217-225.
24. Sands, D.E. 1951. A study of the Allegheny woodrat in central Pennsylvania.
M.S. Thesis, Penn. State College, University Park. 73 pp.
25. Wayne, W.J. 1960. Range extension of the Allegheny woodrat {Neotoma magister)
in Indiana. Proc. Ind. Acad. Sci. 69:311.
26. Worth, C.B. 1950. Observations on the behavior and breeding of captive rice
rats and woodrats. J. Mammal. 31:421-426.
Occurrence of Swimmers' Itch In Northeast Indiana
David L. Daniell
Department of Zoology
Butler University, Indianapolis, Indiana 46208
Introduction
Swimmers' itch or schistosome cercarial dermatitis is a discomfort experienced
by many persons each summer. It involves the development of red papular eruptions
and intense itching on skin that has been exposed to water inhabited by snails that
are infected with larval avian schistosomes. When avian schistosome cercariae accidentally
penetrate the outer layers of human skin, they die near the point of entry, and a
dermatitis reaction occurs around each parasite.
In recent years there have been many reports of outbreaks of swimmers' itch
in North America and many studies of the prevalence of schistosomes in both in-
termediate and definitive hosts. None of the studies, however, have included Indiana.
The abundance of lakes in northeast Indiana and their heavy use for recreation
during the summer months would suggest that cercarial dermatitis will occur if snails
infected with schistosomes are present in the water. This study was undertaken in order
to determine if infected snails are present, and if so, the prevalence and identification
of, schistosomes in various snail species.
Methods and Materials
At regular intervals during June, July, and August of 1984, snails were collected
from the following sites on 5 natural lakes in northeast Indiana: (1) the southwest
corner of Tippecanoe Lake in Kosciusko Co., (2) the Crooked Lake Biological Station
on the north side of Crooked Lake in southern Noble Co., (3) the northwest corner
of Loon Lake in Steuben Co., (4) the south side of Crooked Lake in Steuben Co.,
and (5) the south side of Jimmerson Lake in Steuben Co. Snails of all sizes were col-
lected by hand in shallow water along the shore.
The following criteria were used in the selection of collecting sites: (1) the presence
of humans engaged in activities that expose them to water, (2) the presence of bird
species known to harbor dermititis-producing schistosome adults, and (3) the presence
of snails from each of three pulmonate families (Physidae, Lymnaeidae, and Planor-
bidae) known to serve as intermediate hosts for avian schistosomes.
Snail species examined were Physa gyrina (Physidae), Lymnaea sp. and Pseudosuc-
cinea columella (Lymnaeidae), and Gyraulus sp., Promenetus exacuous, and Planor-
bula armigera (Planorbidae). Snails were placed in small culture jars (5-10 per jar)
of filtered lake water and kept for 2 days under normal light conditions. Water was
checked 3 times each day for the presence of emerged cercariae. If cercariae were found,
snails from that jar were isolated in small vials. Numbers of snails shedding cercariae,
sizes of snails, and all cercarial types were recorded.
Schistosome cercariae were examined while alive and also after being fixed in
hot 5% formalin. Measurements were taken of 10 formalin-fixed cercariae from each
snail infected with schistosomes. Identification was made on the basis of these
measurements and the morphology and behavior of living cercariae.
To determine if the cercariae isolated from snails were able to produce a skin
reaction (cercarial dermatitis), several from each infected snail were placed on the skin
of the author's arm. This technique works quite well because the author is sensitized
to most species of avian schistosomes.
629
630
Indiana Academy of Science
Vol. 94 (1985)
Results
Of the 7,835 snails examined during the summer of 1984, only 21 (0.27%) harbored
patent schistosome infections (Table 1). These occurred in Physa gyrina (13), Pseudosuc-
Table 1 . Patent infections of avian schistosomes and all trematode species in pulmonate
snails from northeast Indiana.
Total No.
Snail Host
Collected
Physa gyrina
2326
Lymnaea sp.
688
Pseudosuccinea
columella
962
Gyraulus sp.
2317
Planorbula
armigera
1042
exacuous
500
TOTALS
7835
No. of Patent Infections (°7o)
Avian Schistosomes
All Trematode Spec.
13 (0.56%)
0 (0.0 %)
7 (0.73°7o)
0 (0.0 %)
0 (0.0 %)
1 (0.20%)
21 (0.27%)
78 (3.35%)
9 (1.31%)
74 (7.69%)
126 (5.44%)
5 (0.48%)
34 (6.80%)
326 (4.16%)
cinea columella (7), and Promenetus exacuous (1). No schistosome infections were found
in the other 3 snail species. Patent infections of all trematode species occurred in 4.16%
of the snails examined. Pseudosuccinea columella had the highest total infection rate
(7.69%), as well as the highest schistosome infection rate (0.73%).
Snails harboring patent schistosome infections were collected at only 3 of the
5 study sites (Crooked Lake Biological Station, Tippecanoe lake, and Loon Lake).
The author experienced cercarial entry into the skin (dermatitis) while collecting at
each of these localities and also while collecting at Jimmerson Lake on one occasion.
Infected snails were not collected at this site, however.
Four species of schistosomes were recovered during this study (Table 2). The species
most frequently recovered was Gigantobilharzia huronensis from 9 individuals of P.
gyrina. Another species of this genus, G. elongata, was found in 1 P. exacuous. One
species of Trichobilharzia (probably T. physellae) was recovered from 4 P. gyrina,
and a second species of Trichobilharzia occurred in 7 P. columella. All 4 of these
schistosome species produce cercarial dermatitis.
Prevalence of infection for each schistosome species collected during the summer
of 1984 was determined for every collection period (Table 2). These data suggest that
patent infections of each species may be limited to specific, relatively short time periods.
G. elongata in P. exacuous and Trichobilharzia sp. in P. gyrina were found only during
the first half of June. P. gyrina infected with G. huronensis were collected from mid-
June to mid-July, while Trichobilharzia sp. infections in P. columella were limited
to the last 3 collections of the summer (July 24, August 8, and August 23).
Discussion
Although prevalence of schistosomes in snails of northeast Indiana, as determined
by this study, is quite low (0.27%), the presence of infected snails at 3 of the 5 collec-
ting sites and the contraction of dermatitis by the author at 4 sites indicate that per-
sons using these lakes for recreation could develop swimmers' itch.
Prevalence of schistosomes is frequently low, even in surveys done at the time
of outbreaks of dermititis. Lane et al. (9) found 1.1% of Physa sp. and 2.3% of Gyraulus
Zoology 63 1
Table 2. Prevalence of infection for each schistosome species found in snails collected
t T i „ a a .* ino^ / number of snails infected .
uuniig juuc, Juiy.
i cuiu rvu;
guai 17
"" v number
of snails collected }'
Schistosome
June
July
August
Totals
and host
4
14
23
2
12
24
8
25
Gigantobilharzia
elongate: in
0
96
1
84
0
26
0
51
0
72
0
51
0
100
0
20
la
500
P. exacuous
Gigantobilharzia
huronensis in
0
105
2
368
1
431
1
465
5
422
0
184
0
173
0
178
9b
2326
P. gyrina
Trichobilharzia sp.
in P. gyrina
2
!05
2
368
0
431
0
465
0
422
0
184
0
173
0
178
4C
2326
Trichobilharzia sp.
in P. columella
0
21
0
47
0
79
0
102
0
108
2
186
3
224
2
195
7a
962
all from Biological Station.
3 from Biological Station, 6 from Loon Lake.
3 from Biological Station, 1 from Tippecanoe Lake.
sp. infected after a dermititis outbreak in Alameda Co., California. Likewise, Howard
and Walden (6) found 3% of Lymnaea emarginata infected with schistosomes after
a dermititis outbreak at Cultus Lake, British Columbia.
In two recent studies done in southwest Michigan on large populations of snails,
0.19% of Physa integra from Gunn Lake in Barry Co. were infected with schistosomes
(7), and 2.54% of Gyraulus parvus from Wintergreen Lake in Kalamazoo Co. were
infected (8). The higher infection rate at Wintergreen Lake might be expected since
this lake is located at the Kellog Bird Sanctuary, where many species of waterfowl
reside. None of the 2,317 Gyraulus sp. in the present study was infected even though
waterfowl were seen at every collecting sight.
The schistosome cercariae found in P. columella collected at Crooked Lake
Biological Station were not identified to species. In structure and behavior they closely
resemble certain species of Trichobilharzia. Adult worms derived from experimental
infections using these cercariae would be useful. Attempts to produce experimental
infections in chickens exposed to these cercariae were unsuccessful, however.
Trichobilharzia cercariae have not previously been reported from P. columella.
Rankin (13) found schistosome cercariae in this snail in western Massachusetts, but
did not describe them. Both species of mammalian schistosomes (Heterobilharzia
americana and Schistosomatium douthitti) found in the United States may use P.
columella as a definitive host (10, 11).
Trichobilharzia sp. found in P. gyrina closely resembles T. physellae described
by Talbot (15). There are a number of species of this genus that use physid snails
as intermediate hosts (3), however, and experimental exposures to determine the adult
are necessary.
G. elongata cercariae were described by Brackett (1) from Gyraulus parvus col-
lected near Madison, Wis. Grodhaus (4) found this species in Gyraulus sp. in Califor-
nia. The occurrence of G. elongata in P. exacuous constitutes a new host record. Adults
of this species are found in pied-billed grebes (5).
G. huronensis, which was found in this study in P. gyrina, was described by
Najim from this same host species from the Huron River near Ann Arbor, Michigan
(12). The adult worms are found in a number of species of passerine birds (2, 12,
632 Indiana Academy of Science Vol. 94 (1985)
14). In the present study, part of each collecting site was near a stand of cattails where
red winged blackbirds were nesting in early summer. They were probably the source
of infection for the snails.
Acknowledgments
This study was funded through a Butler University Fellowship. Use of the facilities
at the Crooked Lake Biological Station, Indiana University-Purdue University at Fort
Wayne was appreciated greatly.
Literature Cited
1. Brackett, S. 1940. Two new species of schistosome cercariae from Wisconsin.
J. Parasitol. 26:195-200.
2. Daniell, D.L. 1979. Biology and host-parasite relationships of Gigantobilharzia
huronensis (Trematoda: Schistosomatidae). Dissertation Abstracts 39(B):3188-3189.
3. Farley, J. 1971. A review of the family Schistosomatidae: excluding the genus
Schistosoma from mammals. J. Helminthol. 45:289-320.
4. Grodhaus, G. 1960. Some schistosome cercariae from Gyraulus. J. Parasitol.
46(suppl.):33.
5. Grodhaus, G. 1965. Laboratory rearing and natural occurrence of Gigantobilharzia
elongata (= Cercaria elongata). J. Parasitol. 5 1(4): 680-681.
6. Howard, T.D. and C.C. Walden. 1965. An ecological study of the snail hosts
for the agent of schistosome dermititis at Cultus Lake, British Columbia. J. Appl.
Ecol. 2:121-135.
7. Kulesa, M.W., H.D. Blankespoor, and K.E. Roney. 1982. Prevalence of avian
schistosomes in Physa integra from southwestern Michigan. Proc. Helminthol.
Soc. Wash. 49:14-18.
8. Laman, T.D., D.L. Daniell, and H.D. Blankespoor. 1984. The role of Gyraulus
parvus as an intermediate host for avian schistosomes. Proc. Helminthol. Soc.
Wash. 51:267-269.
9. Lane, R.S., E.W. Mortenson, and G. Grodhaus. 1976. Control of schistosome
dermatitis at Shadow Cliffs Lake, Alameda County, California. Vector Views
23:15-20.
10. Lee, H.F. 1960. The life history of Heterobilharzia americana. J. Parasitol.
46(suppl.):35.
11. Malek, E.A. 1977. Geographical distribution, hosts, and biology of
Schistosomatium douthitti (Cort, 1914) Price, 1931. Can. J. Zool. 55:661-671.
12. Najim, A.T. 1956. Life history of Gigantobilharzia huronensis Najim, 1950. A
dermatitis-producing bird blood fluke (Trematoda-Schistosomatidae). Parasitol.
46:443-469.
13. Rankin, J.S., Jr. 1939. Ecological studies on larval trematodes from western
Massachusetts. J. Parasitol. 25:309-328.
14. Strohm, B.C., H.D. Blankespoor, and P.G. Meier. 1981. Natural infections of
the dermatitis-producing schistosome Gigantobilharzia huronensis Najim, 1950
in passerines in southeastern Michigan. Proc. Helminthol. Soc. Wash. 48(l):80-82.
15. Talbot, S.B. 1936. Studies on schistosome dermatitis. II. Morphological and life
history studies on three dermatitis-producing schistosome cercariae, C. elvae Miller,
1923, C. stagnicolae n.sp., and C. physellae n.sp. Am. J. Hyg. 23:272-284.
Cottonmouth, Agkistrodon piscivorus, Records from
the Blue River and Potato Run in Harrison County, Indiana
(Ohio River Drainage, USA)
Bill J. Forsyth, Claude D. Baker, Tom Wiles and Charles Weilbaker
Department of Biology
Indiana University Southeast
New Albany, Indiana 47150
Introduction
In May 1983, the first Indiana population of cottonmouth, Agkistrodon piscivorus,
was located in a swampy area known as Buffalo Bottom, northeast of Jasper in Dubois
County, Indiana (2). We report two additional cottonmouth records which represent
an easternmost range extension within the Ohio River drainage for the western subspecies,
A. p. leucostoma.
Localities
One adult specimen was taken by B.J. Forsyth and G. Wilson on April 27, 1968
from the lower Blue River near the confluence with the Ohio River at the entrance
to Stygeon River Cave in Harrison County (Lat. 38°10 '46", Long. 86°18 '37"). In April
1970, a subadult was taken by Forsyth from Potato Run which enters the Ohio River
about two miles upstream from the mouth of the Blue River in Harrison County (Lat.
38°10'15", Long. 86°18 '21 "). We have both specimens in our collections at Indiana
University Southeast. The locations of these localities and localities for other recently
verified records in Indiana and Kentucky are given in Figure 1.
ILLINOIS
MILES
Figure 1. Map of Kentucky and Indiana Indicating Verified Cottonmouth Records
in Dubois and Harrison Counties in Indiana, and in Daviess County in Kentucky.
633
634 Indiana Academy of Science Vol. 94 (1985)
Discussion
Both specimens probably were taken immediately following hibernation in near-
by limestone caves and crevices. Later in the year, the western subspecies is essentially
nocturnal which may account for our failure to locate additional individuals. Our first
inclination was that perhaps they were transported up the Ohio River on barges (which
accounts for the delay in reporting the records). Alternatively, the specimens could
represent a native relict population similar to the one in Dubois County which is only
about 35 miles northwest of the Harrison County records (Figure 1). In addition, a
Kentucky population, recently verified by MacGregor (1) in August 1984 near Owensboro
in Daviess County, Kentucky in the Panther Creek portion of the Green River drainage,
is situated about 50 miles southwest of our localities (Figure 1).
Acknowledgments
We acknowledge Dr. Sherman A. Minton of the Indiana University School of
Medicine who alerted us regarding the significance of these findings. Mr. Michael J.
Lodato verified the identifications.
Literature Cited
1. MacGregor, J. 1984. Personal communication from Kentucky's Non-Game
Biologist.
2. Minton, S.A., List, J.C., and M.J. Lodato. 1982. Recent records and status
of amphibians and reptiles in Indiana. Note added in proof. Proc. Indiana Acad.
Sci. 92:489-498.
Dental Anomalies in Three Species of Shrews from Indiana
Thomas W. French*
Department of Life Sciences
Indiana State University
Terre Haute, Indiana 47809
Introduction
Dental anomalies in shrews are not common. Hall (1940) found no cases of miss-
ing or extra teeth in 1,837 specimens of North American shrews and Jackson (1928)
reported only 5 specimens (0.05%) with dental anomalies out of 10,431 Sorex and
Microsorex examined. Information on dental anomalies in shrews has been best sum-
marized by Choate (1968). Most reported dental anomalies result from a reduction
in the number of upper, and to a lesser extent, lower unicuspid teeth (12, 13, C, P2,
P3, 13, and c of Choate 1968). No case of genetically deleted molariforms or first
incisors has been reported in shrews. Subnumerary dentitions have been reported from
Sorex araneus (Reinwaldt 1961), S. cinereus ohionensis (Bole and Moulthrop 1942),
5. minutus (Reinwaldt 1961), S. obscurus longicauda (Jackson 1928), S. tundrensis
(Pruitt 1957), Blarina brevicauda and B. b. carolinensis (= B. carolinensis) (Choate
1968), B. adamsi (from the upper Pliocene — Hibbard 1953), Cryptotis goodwini, C.
mexicana, C. nigrescens, and C. parva (Choate 1970), and Microsorex hoyi (Jackson
1928).
Supernumerary dental formulas have resulted most often from extra upper
unicuspids and have been reported from 1 Sorex o. obscurus (Jackson 1928), 1 S.
s. saussurei (Hooper 1946), 3 Blarina carolinensis (Choate 1968), 1 Crocidura cyanea,
2 C. hirta (Meester 1953), and 3 C. marquensis (Dippenaar 1978). Extra unicuspids
in the lower jaw have been reported only from Blarina brevicauda (Hibbard 1953).
Supernumerary molariform teeth in shrews are especially scarce. Small molariform
teeth located behind the lower third molar have been described from 3 (1 unilateral,
1 bilateral) of 514 (0.4%) Crocidura marquensis (Dippenaar 1978), 1 (unilateral) of
145 (0.7%) Blarina b. brevicauda (Choate 1968) and 1 (unilateral) of 111 (0.9%) B.
b. kirklandi (Choate 1968). Small extra molariforms posterior to the third upper molars
are known from the white-toothed shrews Crocidura olivieri (1 unilateral) (Setzer 1957)
and C. hirta (1 unilateral) (Meester 1959). Choate (1968) indicated that T.E. Lawlor
would discuss an instance of supernumerary molars in Blarina brevicauda from specimens
now housed in the Cleveland Museum. This data, however, has never been reported
(T.E. Lawlor, personal communication).
Knowledge of dental variation in Soricidae is important because cranial and dental
morphological features are considered the most useful key characters (Junge and Hoff-
man 1981). Both types of characters preserve well and are considered to be rather
dependable although few few data are available to suggest just how dependable. This
paper gives an assessment of rates of occurrence of rather pronounced morphological
variations within populations. These as well as more subtle variations, if genetically
based, are presumably part of the raw material upon which natural selection may act
and may help elucidate evolutionary trends in the Soricidae.
♦Current Address: Nongame and Endangered Species Program, Massachusetts Division of Fisheries and
Wildlife, 100 Cambridge Street, Boston, Massachusetts 02202.
635
636 Indiana Academy of Science Vol. 94 (1985)
Materials and Methods
During a study of the Southeastern Shrew (Sorex I. longirostris) and the Masked
Shrew (5. cinereus lesueuhi) in Vigo County, Indiana (French 1980), skulls of 125
Southeastern Shrews and 214 Masked Shrews were examined for dental anomalies.
In addition, skulls of 4 Southeastern Shrews and 115 Masked Shrews were examined
from other areas of Indiana and 95 Southeastern Shrews were examined from Alabama.
From Vigo and adjacent Clay counties, Indiana 385 Short-tailed Shrews, Blarina
brevicauda were examined for dental anomalies, along with 37 from Alabama, 20 from
Georgia, 5 from Maine, and 3 from Massachusetts.
Results
Six of 125 (4.8%) Southeastern Shrews from Vigo County exhibited subnumerary
dental formulas. In four cases the upper fifth unicuspid was missing (3 unilateral,
1 bilateral) and in two cases the upper fourth unicuspid was missing (both bilateral).
Four of these specimens include an adult female and her three nearly weaned offspring
which were trapped in a pitfall as a family unit. The tooth missing in the mother
is the left upper fifth unicuspid and in the offspring include a right upper fifth unicuspid,
the upper fifth unicuspids bilaterally, and what appears to be the upper fourth unicuspids
bilaterally. This is the first case in which a family of S. longirostris have been captured
together away from a nest and the first evidence that dental anomalies in shrews may
be inherited and not the result of spontaneous mutation. Specimens in this study also
represent the first reported examples of dental anomales in S. longirostris.
In Parke County, Indiana two of four Southeastern Shrews had upper unicuspids
missing bilaterally, the fourth unicuspids in one and the fifth in the other case. Of
ninety-five specimens of 5. longirostris from Alabama examined, one (1.1%) had the
left upper fourth unicuspid missing and two others had missing teeth that appeared
to be the result of injury. In one case the right upper fourth unicuspid was missing
and in the other the right upper third, fourth and fifth unicuspids were missing. In
both of these latter cases the teeth were not crowded and large gaps were present where
the teeth should have been located.
In S. cinereus two of 214 (0.9%) from Vigo County had subnumerary dental
formulas. In one, the upper fifth unicuspids were missing bilaterally and in the other
specimen this tooth was missing on the right and was peg-like on the left. One of
15 specimens from Wabash County, Indiana was missing the upper fifth unicuspids
on both sides. No cases were found in either species which involved molariform teeth.
The only case of supernumerary dentition in Sorex was a Southeastern Shrew
from Vigo County in which there were six upper unicuspids on the left side. This
is a 0.8% occurrence. The extra tooth appeared to be between the third and fourth
unicuspid (see Figure 1).
Palmer (1937) noted a greater incidence of subnumerary dentitions in smaller
subspecies of the broad-footed mole, Scapanus latimanus, than in larger subspecies,
a relationship that he considered due to crowding of the teeth in the smaller skulls.
Choate (1968) found the same relationship between two forms of Blarina that are now
considered closely related species, B. brevicauda and the much smaller B. carolinensis
and in his study only two of 145 (1.4%) specimens of B. b. brevicauda had subnumerary
complements of unicuspids, displaced unicuspids, or diminutive unicuspids; abnormalities
that Choate also considers related to tooth crowding.
The same relationship seems to exist between S. cinereus and S. longirostris. In
Vigo County 4.8% of the 5. longirostris and 0.9% of the 5. cinereus displayed sub-
numerary dental complements. As Miller (1895) noted, the palate of S. longirostris
Zoology
637
Figure 1. Dental patterns of Sorex longirostris and S. cinereus. Patterns of 5.
longirostris are on the left and include specimens with the fifth unicuspid missing and
the fourth smaller than the third (A), the fifth unicuspid missing and the third and
fourth about the same size (B), a sixth unicuspid between the third and fourth (C),
and the typical dental pattern (D). Patterns of 5. cinereus are on the right and include
specimens with the fifth unicuspid missing and the other unicuspids of typical propor-
tions (E), the fifth unicuspid displaced and unusually large (F), the typical dental pat-
tern of S. cinereus ohioensis with the third unicuspid smaller than the fourth (G),
and the typical dental pattern of other races of S. cinereus (H).
is "remarkably broad and short," a character which results in crowding and thus an
increased frequency of related tooth abnormalities.
Seventeen of 384 (4.4%) Vigo and Clay County Blarina had reduced, displaced,
or missing unicuspid teeth. Thirteen (3.4%) specimens had anomalies in the upper
and 4 (1.0%) in the lower unicuspid tooth rows. In at least 6 of the 14 skulls with
missing teeth these teeth appeared to have been lost due to injury. In the remaining
eight cases (2.1%) the teeth apparently were genetically deleted, in five cases the fifth
upper unicuspids were missing (3 unilateral, 2 bilateral), and in one case each, the
third and fourth upper unicuspids and first lower unicuspid were missing unilaterally.
Choate (1968) reported subnumerary complements of unicuspids, displaced unicuspids,
or diminutive unicuspids in two of 145 (1.4%) of the Short-tailed Shrews B. brevicauda
from the University of Kansas Museum of Natural History collection.
Two especially unusual cases of supernumerary dentitions were found in Vigo
County Blarina. On 8 April 1977 an adult pregnant specimen (ISU#5315) with heavily
worn teeth was trapped at Coal Creek and Indiana Highway 63. This specimen has
two extra molariform teeth positioned bilaterally and posterior to the upper third molars
(Figure 2A). On 14 October 1978 a juvenile female (ISU#5316) from this locality was
638
Indiana Academy of Science
Vol. 94 (1985)
Figure 2. A — Posterior end of Blarina brevicauda (ISU#5315) palate showing two
extra molariform teeth positioned bilaterally and posterior to the upper third molars.
B — Same condition as in 1A in a younger specimen (ISU#5316).
C — Posterior end of B. brevicauda (ISU#5316) lower tooth rows showing
one extra molar posterior to the left lower third molar.
discovered with the same condition of bilateral extra molariforms posterior to the up-
per third molars (Figure 2B) and also with an extra tooth posterior to the left lower
third molar (Figure 2C). The ages and length of time between captures of these two
specimens eliminates the possibility that the second specimen was the offspring of the
first but they well may have been closely related individuals. These two Blarina appear
to represent the first reported cases of bilateral supernumerary upper molariform teeth
in shrews. Seventy-two Blarina specimens were collected at this locality between 1977
and 1979 but no other supernumerary dentitions were discovered.
Of 65 Blarina examined from outside Indiana no additional anomalies involving
molariform teeth were found but one specimen from Massachusetts (ISU#5317) had
six upper right unicuspids. The extra unicuspid in this specimen is lingual to the fourth
unicuspid (P2 of Choate 1968).
Several other abnormalities probably not directly related to crowding also were
found. Two Southeastern Shrews, one from Vigo County and one from Alabama had
asymmetrical rostrums. The rostrums were curved to the left and displayed a noticeable
reduction in the size of several of the unicuspid teeth on the shorter side. One Masked
Shrew from Wabash County, Indiana had a steeply sloping rostrum resulting in very
uneven wear of the teeth. Another from Vigo County had an abnormally large upper
left fifth unicuspid which was separated from the fourth by a noticeable gap. This
normally diminutive tooth was as large as the fourth unicuspid and just as well pigmented
Zoology 639
(see Figure 1). It is not known if this abnormality was present on both sides because
the skull was crushed and part of the right maxilla was missing.
Normally the upper fifth unicuspid is not pigmented, even in very young individuals,
but 10 Masked and one Southeastern Shrew from Indiana had this tooth pigmented.
Four Southeastern Shrews from Alabama also had pigment on this tooth. In young
specimens the upper third unicuspid is usually pigmented but two young Southeastern
Shrews from Indiana lacked pigment on this tooth. Two of the 214 Vigo County S.
cinereus displayed unusually light colored dental pigment but no complete absence of
pigment was found. Bole and Moulthrop (1942) reported a series of 5. c. ohioensis
from Cuyahoga County, Ohio that "show practically no pigmentation in the teeth,
2 specimens from this locality being absolutely without dental pigment."
Hall (1940) had no instances of diseased teeth in 1837 Sorex but in this study
four of 132 (3.0%) Southeastern Shrews and five of 260 (1.9%) Masked Shrews from
Indiana had decayed teeth. Two of 95 (2.1%) Southeastern Shrews from Alabama
also had decayed teeth. All but one specimen with decayed teeth (a Vigo County S.
longirostris) were old shrews with heavy tooth wear.
Discussion
Within the Soricidae the basic dental formula consists of 32 teeth (i 3/1, c 1/1,
pm 3/1, m 3/3 x 2 = 32), including 5 upper unicuspids and 2 lower unicuspids. In
North America this full dental complement is found in the genera Sorex, Microsorex,
and Blarina although 2, rather than the usual 1 unicuspids are vestigial in Microsorex.
Reduced dentitions are typical of Cryptotis (30 teeth) and Notiosorex (28 teeth), each
the result of losses of upper unicuspids. In this study, reductions in upper unicuspid
numbers was the most frequent anomaly encountered and appears to be a genetically
inherited character and increases in frequency in species with shorter, more crowded
toothrows.
The only variation in the normal mandibular dentitions of recent Soricidae is
an extra unicuspid between the second and third tooth (c and p 4) in the African genus
Myosorex and rarely in its geographically and phylogenetically near relative Surdisorex
norae. No anomalies resembling this condition were found in this study but one case
has previously been reported from Blarina (Hibbard 1953). No examples of variations
in molariform teeth in recent species of Soricidae normally occur although other members
of the order Insectivora, such as moles, golden moles, elephant shrews, and tenrecs,
do possess greater molariform complements.
Acknowledgments
Appreciation is extended to G.S. Jones, D.D. Pascal, Jr., and J.O. Whitaker,
Jr. for the loan of Vigo and Clay county Blarina skulls in their collections. I thank
J.O. Whitaker, Jr. for his encouragement and suggestions concerning the manuscript.
ISU numbers refer to specimens in the Indiana State University mammal collection,
Terre Haute, Indiana.
Literature Cited
1. Bole, B.P. and P.N. Moulthrop. 1942. The Ohio recent mammal collection in
the Cleveland Museum of Natural History. Sci. Publ. Cleveland Mus. Nat. Hist.
5:83-181.
2. Choate, J.R. 1968. Dental abnormalities in the short-tailed shrew, Blarina
brevicauda. J. Mammal. 49:251-258.
3- . 1970. Systematics and zoogeography of middle American shrews of the
genus Cryptotis. Univ. Kansas Publ., Mus. Nat. Hist. 19:195-317.
640 Indiana Academy of Science Vol. 94 (1985)
4. Dippenaar, N.J. 1978. Dental abnormalities in Crocidura mariquensis (A. Smith
1844) (Mammalia: Soricidae). Ann. Transv. Mus. 31:165-168.
5. French, T.W. 1980. Ecological relationships between the southeastern shrew {Sorex
longirostris Bachman) and the masked shrew (5. cinereus Kerr) in Vigo County,
Indiana. Unpubl. Ph.D. dissertation, Indiana State Univ., Terre Haute, 54 pp.
6. Hall, E.R. 1940. Supernumerary and missing teeth in wild animals of the orders
Insectivora and Carnivora with some notes on disease. J. Dent. Res. 19:103-143.
7. Hibbard, C.W. 1953. The insectivores of the Rexroad Fauna, Upper Pliocene
of Kansas. J. Paleont. 27:21-32.
8. Hooper, E.T. 1946. Extra teeth in a shrew. J. Mammal. 27:394.
9. Jackson, H.H.T. 1928. A taxonomic review of the American long-tailed shrews.
North Am. Fauna 51:1-238.
10. Junge, J. A. and R.S. Hoffmann. 1981. An annotated key to the long-tailed shrews
(genus Sorex) of the United States and Canada, with notes on Middle American
Sorex. Occas. Pap. Mus. Nat. Hist., University of Kansas, No. 94, 48 pp.
11. Meester, J. 1953. The genera of African shrews. Ann. Transv. Mus. 22:205-214.
12. . 1959. Dental abnormalities in African shrews. Ann. Transv. Mus. 23:411-412.
13. Miller, G.S., Jr. 1895. The long-tailed shrews of the eastern United States. North
Am. Fauna 10:35-56.
14. Palmer, F.G. 1937. Geographic variation in the mole Scapanus latimanus. J.
Mammal. 18:280-314.
15. Pruitt, W.O., Jr. 1957. Tooth reduction in the tundra shrew. J. Mammal. 38:121.
16. Reinwaldt, V.E. 1961. Uber Zahnanomalien und die Zahnformel der Gattung
Sorex Linne. (On tooth anomalies and the tooth formula of the genus Sorex Linne.)
Arkiv For Zoologi 13:533-539 (in German).
17. Setzer, H.W. 1957. An extra tooth in Crocidura. J. Mammal. 38:258-259.
Reproduction and Age Structure of Three Indiana Shrews
Thomas W. French*
Department of Life Sciences
Indiana State University
Terre Haute, Indiana 47809
Introduction
Seasonal patterns of reproduction and age structure have been studied in several
individual species of shrew (Hamilton 1940, Pearson 1945, Conaway 1952, Jameson
1955, Clough 1963, Dapson 1968, French 1980a), but the relationship of these patterns
between species in the same geographical areas is poorly known. The purpose of this
study is to compare the seasonal patterns of reproduction and age structure of the
Southeastern Shrew (Sorex longirostris), Masked Shrew (5. cinereus) and Short-tailed
Shrew (Blarina brevicauda) in the vicinity of Terre Haute, Indiana.
Materials and Methods
A study of the Southeastern and Masked Shrews was conducted in Vigo County
between 1976 and 1979 (French 1980b). During this study 145 Southeastern, 214 Masked
and 216 Short-tailed Shrews were trapped. These specimens, plus previously existing
Vigo County museum specimens and 107 specimens of Blarina from adjacent Clay
County, were used to plot age structure. Reproductive data were gathered from these
specimens, and because of the natural scarcity of pregnant shrews (Jameson 1955,
Dapson 1968), the data were supplemented with data from 4 Masked and 14 Short-
tailed Shrew females from other parts of Indiana.
Four major age classes were recognized, and each class was broken into three
subgroups following the procedure of Rudd (1955). Each age class covered about four
months of life and was based on tooth wear rather than reproduction status. In Rudd's
study breeding was initiated in age class 2. In this study a representative specimen
of each species was selected from Vigo County to represent each age class. Subsequent
shrews were compared and assigned the number of the standard that they most closely
resembled. Plus or minus designations were used if tooth wear was more or less ad-
vanced than the standard. Thus, the youngest possible designation is 1 - and the oldest
4+.
Results
In Indiana, 13 pregnant Southeastern Shrews averaged 4.55 (4 to 6) embryos,
9 pregnant Masked Shrews averaged 6.10 (4 to 7) embryos, and 18 pregnant Short-
tailed Shrews averaged 5.39 (2 to 8) embryos. Dates of pregnancy ranged from 8 April
to 25 September, 28 April to 23 August, and 29 February to 11 September, respectively
(Table 1). Similar earliest dates of pregnancy, and first appearance of young of the
year suggested that the onset of breeding probably occurred at about the same time
in both species of Sorex, with the onset about 3 weeks earlier in S. longirostris in
this study (Figure 1). The onset of reproduction in Blarina, however, was 5 weeks
earlier than 5. longirostris and a full 8 weeks earlier than S. cinereus in this study.
*Current Address: Nongame and Endangered Species Program, Massachusetts Division of Fisheries and
Wildlife, 100 Cambridge Street, Boston, Massachusetts 02202.
641
642 Indiana Academy of Science Vol. 94 (1985)
Table 1 . Monthly distribution of pregnant and lactating individuals of Sorex longirostris,
S. cinereus and Blarina brevicauda near Terre Haute, Indiana.
Feb
Mar
Apr
May
June July Aug
Sept
Oct
Nov
Dec
Total
Sorex longirostris
pregnant
0
0
4
1
2 1 4
1
0
0
0
13
lactating
0
0
0
0
1 1 1
Sorex cinereus
0
1
0
0
4
pregnant
0
0
2
2
0 4 1
0
0
0
0
9
lactating
0
0
0
1
0 1 0
Blarina brevicauda
0
4
0
0
6
pregnant
1
1
8
5
1 0 1
1
0
0
0
18
lactating
0
0
2
4
2 3 0
3
4
1
2
21
Previously it has been shown that several species of shrews are capable of reaching
sexual maturity and reproducing during their first year of life (Pearson 1945, Con-
away 1952, Clothier 1955, Short 1961, Clough 1963, Dapson 1968, French 1980a, and
others). In this study, two S. longirostris in age class 1 were found to be pregnant,
three were lactating, and four other females showed signs of sexual maturity. No 5.
cinereus in age class 1 were pregnant or lactating, but 4 females had enlarged uteri,
indicating the approach of sexual maturity. At least one Blarina in age class 1 had
placental scars, one had enlarged teats but a small uterus and 4 had enlarged uteri.
Other specimens of each species in age class 2 also showed signs of reproduction in
their first year of life. Fifteen females of 5. longirostris and 20 females of S. cinereus
in age class 1 were examined, suggesting maturity of female 5. longirostris in their
first year is more common than in S. cinereus in Indiana. In Vigo County, S. longirostris
was trapped consistently with less frequency than 5. cinereus (1.02 and 5.65 per 100
trap nights in hardwood floodplain habitats, respectively), suggesting lower popula-
tion densities of S. longirostris. These results are consistent wi.th Stein's (1961) sugges-
4
4
4
+
3
3
CO *■
A A* 41 A A 11 A a 1
AAA* A ••* AAA A A A A A*f • A
1 1 AAA
tSfllglttt **•'
*tt* "•*•;* £?*£& r •
••
A A AA A A
****** A A* |4ft* A*** A *A
9— Am
-•••*** •#•
• AltA *** * • »A A ***** • **
1 - AAAAA AAAAA AA • AAAAAAAA AAA
1 - . A.AAJ. AA. AA*A
_l I I I I I I I I I I
Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec
Figure 1 . Monthly changes in age class composition of Sorex longirostris (triangles)
and S. cinereus (circles) from Vigo County, Indiana.
Zoology 643
tion that reproduction in the first year of life is related to low population densities
in Sorex. Blarina population densities were intermediate with trap results yielding 1.20
per 100 trap nights when taken with S. longirosths and 1.16 per 100 trap nights when
taken with S. cinereus in floodplain hardwoods. Individuals of each of these shrew
species have previously been found, simultaneously pregnant and lactating (Hamilton
1949, French 1980a), indicating successive litters in rapid succession during part of
the breeding season.
Although most breeding occurs in the spring, Dapson (1968) reports that under
certain conditions Blarina can produce litters at any time of the year. Several authors
(Seton 1909, Hamilton 1929, Lyon 1936, Blair 1940) have suggested that there are
two breeding peaks in Blarina during the spring and fall with a reduction occurring
in mid-summer. In this study, however, there seemed to be only one major peak in
Blarina reproduction in the spring (late March to early June — Figure 2) which con-
4+
•
• • •
4
4~
-
• MM • • • —
3+
•
MM MM M • • •
3
_ •
M • •
«•• • M* «M M •
a,3"
-
• —
• • M • M •
£V
M« • • • M|M M •
2
2~
1+
_ •
• • M M«« • • MM M j MM •• •
"B jl rWPH
1
• M • M*M tfff fffffff *** f* 1*
f
I
1 1
M
1 1 . 1 1 1 1 1 1 1 .....
Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec
Figure 2. Monthly changes in age class composition of Blarina brevicauda (circles).
tinued to drop off through the late summer and fall. This is most evident when com-
paring the proportion of young shrews just entering the trappable population (age
class 1) to shrews in age class 2 (Table 2). The proportion of younger shrews is high
from April to July and then steadily drops to zero by December. Both species of Sorex
do appear to have two reproductive peaks, in the spring and again in the fall, with
a reduction of reproduction during the mid-summer (Table 2). The major peak in the
appearance of young in the population occurs in June and July, with the proportion
of younger shrews (age class 1) dropping to zero in August, and a lesser peak appear-
ing in October and November before again dropping to zero in December.
Acknowledgments
Appreciation is extended to G.S. Jones, D.D. Pascal, Jr. and J.O. Whitaker,
Jr. for the loan of Blarina skulls in their collections. I thank J.O. Whitaker, Jr. for
his encouragement and suggestions concerning the manuscript.
644 Indiana Academy of Science Vol. 94 (1985)
Table 2. Monthly changes in the proportion of very young (age class 1) and older (age
class 2) shrews still in their first year of life from Vigo and Clay counties, Indiana.
May June July Aug. Sept. Oct. Nov. Dec.
age class 2
age class 1
percent in age 1
age class 2
age class 1
percent in age 1
age class 2
age class 1
percent in age 1
Sorex longirostris
0
0
1 13
4
9
1
3
1
6
9 0
1
14
3
0
100
100
90 0
Sorex cinereus
20
61
75
0
0
0
9 8
2
6
16
5
0
6
9 0
1
11
6
0
0
100
50 0
33
65
27
0
Blarina brevicauda
2
3
5 5
23
83
28
10
16
34
36 7
7
24
5
0
89
92
88 58
23
22
15
0
Literature Cited
1. Blair, W.F. 1940. Notes on home ranges and populations of the short-tailed shrew.
Ecology 21:284-288.
2. Clothier, R.R. 1955. Contribution to the life history of Sorex vagrans in Montana.
J. Mammal. 36:214-221.
3. Clough, G.C. 1963. Biology of the arctic shrew, Sorex arcticus. Am. Midi. Nat.
69:69-81.
4. Conaway, C.H. 1952. Life history of the water shrew {Sorex palustris navigator).
Am. Midi. Nat. 48:219-248.
5. Dapson, R.W. 1968. Reproduction and age structure in a population of short-
tailed shrews {Blarina brevicauda). J. Mammal. 49:205-214.
6. French, T.W. 1980a. Natural history of the Southeastern Shrew, Sorex longirostris
Bachman. Am. Midi. Nat. 104:13-31.
7. French, T.W. 1980b. Ecological relationships between the southeastern shrew {Sorex
longirostris Bachman) and the masked shrew (5. cinereus Kerr) in Vigo County,
Indiana. Unpubl. Ph.D. dissertation, Indiana State Univ., Terre Haute. 54 pp.
8. Hamilton, W.J., Jr. 1929. Breeding habits of the short-tailed shrew, Blarina
brevicauda. J. Mammal. 10:125-134.
9. Hamilton, W.J., Jr. 1940. The biology of the smoky shrew {Sorex fumeus fumeus
Miller). Zoologica 25:473-492.
10. Hamilton, W.J., Jr. 1949. The reproductive rates of some small mammals. J.
Mammal. 30:257-260.
11. Jameson, E.W., Jr. 1955. Observation on the biology of Sorex trowbridgei in
the Sierra Nevada, California. J. Mammal. 36:339-345.
12. Lyon, M.W., Jr. 1936. Mammals of Indiana. Am. Midi. Nat. 17:1-384.
13. Pearson, O.P. 1945. Longevity of the short-tailed shrew. Am. Midi. Nat.
34:531-546.
14. Rudd, R.L. 1955. Age, sex and weight comparisons in three species of shrews.
J. Mammal. 36:323-339.
15. Seton, E.T. 1909. Life histories of northern animals. Charles Schribner's Sons,
New York, vol. 1, p. 677-1267.
16. Short, H.L. 1961. Fall breeding activity of a young shrew. J. Mammal. 42:95.
17. Stein, G.H.W. 1961. Beziehungen szischen Bestandsdichte und Vermehrung bei
der Waldspitzmaus, Sorex araneus, und weiteren Rotzahnspitxmausen. Z.
Saugetierhunde 26:13-28 (in German).
Canine Dirofilariasis in Central Indiana
Neil J. Parke and Charles E. Mays
Department of Biological Sciences
DePauw University, Greencastle, Indiana 46135
Introduction
The heartworm, Dirofilaria immitis, is primarily a parasite of the domestic dog
(10). The incidence of heartworm disease (dirofilariasis) was at one time thought to
have been confined primarily to southern regions of the United States (11). However,
there is increasing concern of its prevalence in northern parts of the country (16).
Dirofilaria immitis (microfilariae) are transmitted to dogs by mosquitoes when
they feed on or near the muzzle, eye, or pelvic regions. Microfilariae feed on blood
and circulate to the heart where they reach maturity in about nine weeks. Approx-
imately fourteen weeks later, the adults release new microfilariae (1).
Initial stages of dirofilariasis generally occur in the right ventricle or the pulmonary
artery followed by intrusion into the lungs and liver (5). Among the symptoms associated
with the disease are loss of stamina and body weight, vomitting, poor haircoat, and
dehydration (1). In advanced stages, the infection can be fatal (3, 4, 5).
The literature on heartworm disease is somewhat spotty. Most studies have been
conducted in southern and eastern coastal states (5, 14, 15, 17). The present study
was made to determine the extent of canine dirofilariasis in Central Indiana. Among
the issues addressed are: (1) the current status of the disease in different locations,
(2) the effect of the breed and living environment of the dog, and (3) the effect of age.
Materials and Methods
Blood samples analyzed in this study were obtained from dogs in the four Cen-
tral Indiana communities of Greencastle, Greenfield, Indianapolis, and Lafayette. All
dogs surveyed were one year of age or older. Distinctions regarding sex, age, breed,
year of occurrence and location of occurrence were made.
Most of the heartworm analyses were made using direct blood smears and a
modified Knott technique (13). A few samples were analyzed by a micropore filter
technique (1). Differentiation of D. immitis microfilariae from those of the other primary
canine-infecting filariid of the United States, Dipetalonema reconditum was based on
previously described characteristics (9, 10, 17).
The Chi-square method was used for statistical analysis of the data; p < 0.05
was considered to be significant.
Results and Discussion
A total of 3424 dogs in Central Indiana was surveyed for dirofilariasis in 1983
and 1984. Of this total, 1654 specimens were from American Kennel Club recognized
breeds, and 770 were from dogs of mixed breeding. The average size of the D. immitis
microfilariae encountered in this study ranged from 286 to 340 microns in length and
6.1 to 7.2 microns in width. These dimensions are in a general size range with those
previously reported (2).
An analysis was made regarding geographic location and the incidence of heart-
worm disease. The four localities selected for this study represent rural (Greencastle),
semi-rural (Greenfield), small city (Lafayette), and large metropolitan (Indianapolis)
communities. There is a definite trend between the type of community surveyed and
the incidence of dirofilariasis (Table 1). It is significantly higher in the Greencastle
645
646 Indiana Academy of Science
Table 1. Incidence of dirofilariasis in Central Indiana.
Vol. 94 (1985)
Locality
No. dogs
No. dogs
examined
infected
% infected
181
14
7.8
200
5
2.5
2021
37
1.8
1022
24
2.3
Greencastle
Greenfield
Indianapolis
Lafayette
Totals
3424
80
2.3 fx)
area (7.8%) than the other three localities. Indianapolis (1.8%) had the lowest occurrence
of the disease, whereas Greenfield (2.5%) and Lafayette (2.3%) were intermediate.
The mean incidence of dirofilariasis in the four communities is 2.3%, which is in general
agreement with that reported for neighboring states (2, 7, 8, 18, 19).
The occurrence of infection in relation to breed and type of living environment
was determined. Breed categories were modified from those listed in another study
(3). Dogs were classified according to the environment in which they lived. Dogs that
were allowed out-of-doors only for exercise and elimination were classified as "in-
side" dogs; those spending approximately equal time in the house and out-of-doors,
as "inside-outside" dogs; and those which were kept exclusively out-of-doors as "out-
side" dogs.
The data indicate that hounds have the highest incidence (4.8%) of heartworm
disease, followed by mixed breeds (3.8%), working dogs (2.7%), and sporting dogs
(1.6) (Table 2). No infection was recorded among the miscellaneous breeds (e.g., Bichon
Frise, Lhasa Apso, Chow Chow, etc.), which are typically inside dogs.
The data regarding environment shows that outside dogs have a significantly higher
incidence of dirofilariasis than either inside dogs or inside-outside dogs. (Table 2).
This significant trend is similar to that reported in studies done in Georgia (15) and
Louisiana (14, 16). Mixed breeds show the highest incidence in each environmental
category.
The prevalence of dirofilariasis relative to age was analyzed. Results indicate that
the incidence of heartworm disease increases at age 4 and reaches a peak between ages
9 and 11. After that period, the infection rate seems to level off. The incidence at
age 10 was statistically significant (Table 3).
Table 2. Incidence of dirofilariasis as related to breed and environment.
Environment
1
Inside dogs
Inside
outside dogs
Outside dog;
Breed or
No.
No.
°7o
No.
No.
%
No.
No.
%
Tot.
Tot. %
breed type
exam.
infec.
infec.
exam.
infec.
infec.
exam.
infec.
infec.
exam.
infec.
Mixed
194
2
1.0
387
14
3.6
189
13
6.9
770
3.8
Working
0
0
0
179
3
1.7
538
16
3.0
717
2.7
Sporting
0
0
0
153
1
0.7
457
9
2.0
610
1.6
Hound
0
0
0
0
0
0
331
16
4.8
331
4.8
Nonsporting
84
0
0
161
1
0.6
76
1
1.3
321
0.6
Toy
289
2
0.7
0
0
0
0
0
0
289
0.7
Terrier
140
0
0
127
2
1.6
0
0
0
267
0.8
Miscellaneous
119
0
0
0
0
0
0
0
0
119
0
Totals
826
4
0.5 (x)
1007
21
2.1 (x)
1591
55
3.4 (x)
3424
2.3
Zoology
647
Table 3. Incidence of dirofilariasis as related to age.
Age in
No. of
No. of
<7o of
years
dogs
dogs
dogs
(approx.)
examined
infected
infected
1
346
5
1.5
2
355
5
1.4
3
329
6
1.8
4
300
9
3.0
5
423
13
3.1
6
409
10
2.5
7
275
9
3.3
8
250
0
0
9
151
5
3.3
10
164
9
5.5
11
122
3
2.5
12
117
2
1.7
13
91
3
3.3
14
57
1
1.8
15
35
0
0
The incidence of heartworm disease has been shown to be highest in regions which
favor the breeding of mosquitos, such as the southern and southeastern sections of
the United States (10). Infection rates in excess of 40% have been recorded in these
areas (5, 15). Rural communities tend to have numerous breeding areas, and often
do not have regular private and municipal spraying programs found in metropolitan
areas (personal communication, Jerry Rud, Biologist, Indiana State Board of Health).
This may partly account for the significant difference in the prevalence of dirofilariasis
between Greencastle (7.8%) and Indianapolis (1.8%). A Michigan study recorded an
incidence of D. immitis infection of 1.6% in the Detroit area, but noted a sharp in-
crease in infection rate along the marsh areas east of the city (19).
There is also a tendency for rural areas to have a greater percentage of outside
dogs than to urban areas. Such dogs would be more susceptible to mosquito bite.
Various studies have shown that larger outdoor breeds, especially hunting and work-
ing dogs have a significantly higher rate of infection than other breeds (3, 6, 12).
The difference in dirofilariasis between rural and urban areas may also have a
socioeconomic basis. A number of surveys have shown a significantly higher prevalence
of the infection in pound dogs than in privately owned dogs (6, 16).
In animals under 4 years of age, the incidence of D. immitis infection is low
(Table 3). It then rises somewhat and peaks at age 10 before declining. This trend
is similar to that reported in other studies (8, 17).
The results of this study support the growing impression that D. immitis infec-
tion is spreading (16, 17). Although the incidence of 2.3% in Central Indiana is in
close agreement with studies done in neighboring states, the prevalence of dirofilariasis
appears to vary depending on geographic location, breed, living environment, and age
of the dog. Furthermore, the incidence of infection in the Greencastle area (7.8%)
is similar to that reported for several southern and coastal regions of the country (3,
6, 16, 17). This suggests that dirofilariasis may pose a particular health problem for
rural areas.
Acknowledgments
We wish to thank Drs. Jeffrey Hanssen (Westwood Veterinary Clinic, Green-
field, Indiana), James Albrecht (Northside Animal Hospital, Indianapolis, Indiana),
648 Indiana Academy of Science Vol. 94 (1985)
Phillip Watson, Kathryn Carter, and Todd Wheeler (16th Street Veterinary Clinic,
Indianapolis, Indiana), Donald Brattain and John Scamahorm (Greencastle Veterinary
Clinic, Greencastle, Indiana, and John Blair (Blair Animal Clinic, Lafayette, Indiana)
for their assistance in collecting the data.
Literature Cited
1. Ettinger, S.J. and P.F. Sutter. 1970. Canine cardiology. W.B. Saunders Co.,
Philadelphia. 616 pp.
2. Groves, H.F. and F.R. Koutz. 1964. Survey of microfilariae in Ohio dogs. J.
Amer. Vet. Med. Assoc. 144:600-602.
3. Hirth, R.S., H.W. Huizinga, and S.W. Nielsen. 1966. Dirofilariasis in Con-
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7. McKinney, R.E. 1962. The prevalence of Dirofilaria immitis and Dipetalonema
sp. microfilaria in dogs in Champaign County, Illinois. 111. Vet. MEd. 5:43-44.
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Canine microfilariasis in eastern United States. J. Parasitol. 47:661-665.
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University Press, Ames. 274 pp.
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infections of dogs in Atlanta, Georgia. J. Amer. Vet. Med. Assoc. 153:1059-1063.
17. Wallenstein, W.L. and B.J. Tibola. 1960. Survey of canine filariasis. J. Amer.
Vet. Med. Assoc. 137:712-716.
18. Worley, D.E. 1964. Helminth parasites of dogs in southeastern Michigan. J.
Amer. Vet. Med. Assoc. 144:605-608.
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156:890-891.
Ectoparasites of Pine Voles, Microtus pinetorum,
from Clark County, Illinois
D. David Pascal, Jr.
and
John O. Whitaker, Jr.
Department of Life Sciences
Indiana State University
Terre Haute, Indiana 47809
Introduction
In studies on pine voles, only Hamilton (1938) and Benton (1955), both working
in New York, attempted to collect and identify external parasites. Hamilton noted
the mite, Laelaps kochi Oudemans, 1936 and lice of the genus Hoplopleura to be par-
ticularly numerous. Benton reported Hoplopleura acanthopus (Burmeister, 1839) and
Androlaelaps fahrenholzi (Berlese, 191 1) as the most abundant pine vole ectoparasites.
Ferris (1921) reported H. acanthopus from pine voles in New York and Iowa. Another
louse, Polyplax spinulosa (Burmeister, 1839), was found to infest pine voles in Georgia
(Morlan, 1952), although Rattus norvegicus and R. rattus are the true hosts for this
louse (Pratt and Karp, 1953; Wilson, 1961). The occurrence on pine voles was accidental.
Mite records (other than chiggers) on Microtus pinetorum have been summarized
by Whitaker and Wilson (1974), but include LAELAPIDAE: Androlaelaps fahrenholzi
(Berlese, 1911), Laelaps kochi Oudemans, 1936, Haemogamasus longitarsus (Banks,
1910), H. liponyssoides Ewing, 1925, H. ambulans (Thorell, 1872), Laelaps alaskensis
Grant, 1947, and Eulaelaps stabularis (Koch, 1836); GLYCYPHAGIDAE: Glycyphagus
hypudaei (Koch, 1841) and Orycteroxenus soricis (Oudemans, 1915)
LISTROPHORIDAE: Listrophorus pitymys Fain and Hyland, 1972; MYOBIIDAE
Radfordia ensifera (Poppe, 1896) and R. lemnina (Koch, 1841); CHEYLETIDAE
Eucheyletia bishoppi Baker, 1949.
In addition, Smiley and Whitaker (1979) reported the following pygmephorid mites
from Microtus pinetorum from Indiana: Pygmephorus equitrichosus Mahunka, 1975,
P. hastatus Mahunka, 1973, P. scalopi Mahunka, 1973 and P. whitakeri Mahunka, 1973.
Pine voles have numerous chiggers (Trombiculidae). Neotrombicula goodpasteri
(Brennan and Wharton, 1950), TV. lipovskyi (Brennan and Wharton, 1950), TV. microti
(Ewing, 1928), TV. whartoni (Ewing, 1929), Euschoengastia peromysci (Ewing, 1929),
E. ohioensis Farrell, 1956, E. carolinensis Farrell, 1956, and E. diversa Loomis, 1956
(Brennan and Wharton, 1950; Farrell, 1956; Kardos, 1954; Loomis, 1956; MacCreary,
1945; Manischweitz, 1966).
Although once thought to be primarily a bat chigger, Leptotrombidium myotis
(Ewing, 1929) has been reported from pine voles (Loomis, 1956; Manischewitz, 1966).
However, this record may be the more recently described L. peromysci Vercammen-
Grandjean and Langston, 1976.
Ctenophthalmus pseudagyrtes Baker, 1904 is the most commonly reported flea
from the pine vole, and has been collected in many areas (Benton and Cerwonka,
1960; Benton and Krug, 1956; Ellis, 1955; Geary, 1959; Holland and Benton, 1968;
Jameson, 1943; Jameson, 1947; Jordan, 1928; Layne, 1958; MacCreary, 1945;
Mathewson and Hyland, 1964; Morlan, 1952; Whitaker and Corthum, 1967; and Wilson,
1961).
Other fleas reported from pine voles are Stenoponia americana (Baker, 1899),
Atyphloceras bishopi (Jordan, 1933), Rhadinopsylla orama Smit, 1957, Doratopsylla
blarinaeC. Fox, 1914 (probably accidental), Opisodasys pseudarctomys (Baker, 1904),
649
650 Indiana Academy of Science Vol. 94 (1985)
Orchopeas leucopus (Baker, 1904), Orchopeas howardi (Baker, 1895), and Peromyscop-
sylla catatina (Jordan, 1928), (Benton, 1955; Ellis, 1955; Geary, 1959; Holland and
Benton, 1968; Jameson, 1947; MacCreary, 1945; Morlan, 1952; Poorbaugh and Gier,
1961; and Wilson, 1957).
The pine vole is known to harbor the tick Dermacentor variabilis (Say, 1821),
(Clifford, Anastos, and Elbl, 1961; Mellot and Connell, 1965; Sonenshine, Atwood,
and Lamb, 1966; Wilson, 1961).
Mumford and Whitaker (1982) presented more recent information on average
numbers of ectoparasites per host and percent of hosts parasitized for 28 pine voles
from Indiana. Included were many of the species mentioned above but also
MACRONYSSIDAE: Ornithonyssus bacoti (Hirst, 1913); PYGMEPHORIDAE: Baker-
dania sp., Pygmephorus equitrichosus Mahunka, 1975, P. hastatus Mahunka, 1973,
P. scalopi Mahunka, 1973, P. whitaker i Mahunka, 1973; MYOCOPTIDAE: Myooptes
musculinus (Koch, 1844), M. japonensis canadensis Radford, 1955;
CYRTOLAELAPIDAE: Cyrtolaelaps sp; and Anoetidae sp. In addition, they record-
ed the myobiid, Radfordia hylandi Fain and Lukoschus, 1977, probably the same species
earlier recorded as R. lemnina.
The purpose of this study was to determine the ectoparasite community from
a population of pine voles from Clark County, Illinois, and to determine whether their
abundance varied with age or sex of the voles, and if parasite abundance or frequency
varied with season.
Materials and Methods
A 450 x 399 meter tract in a woodlot in Clark County, Illinois, was divided into
3x3 meter plots, of which 125 were randomly selected for trapping. Plots in stream
beds were omitted, leaving 116 sample plots for study (Pascal, 1974). Plots were sampled
from April-December, 1968. Twelve traps were set per plot, underground across the
floor of burrows when burrows were present, at the surface when not. Traps were
baited with peanut butter and checked each day for one week. Pine voles were placed
in individual plastic bags in the field. Voles and plastic bags were examined in the
laboratory with a 10-30X zoom dissecting microscope for external parasites. The fur
was brushed with dissecting needles. Parasites were preserved in alcohol and later
transferred to Nesbitts Solution with acid fuchsin stain added. After a few days they
were mounted in Hoyers Solution and the cover slips were ringed with Euparal.
Results
A total of 1948 ectoparasites and other associates was found on 80 (93.0%) of
the Clark County pine voles examined (Table 1). The average number of ectoparasites
per vole was significantly higher (36.4) for voles collected during the spring and summer
months than for those taken in the fall and winter (19.1) (chi square = 164.77**,
1 df). There was no significant difference between the frequency of infested voles from
the spring-summer period (93.6%) and that of the fall-winter period (92.6%) (chi square
= 0.001, 1 df)- Subadult voles had a significantly lower average number of ectoparasites
per vole (15.1) than adults (21.9) (chi square = 35.92**, 1 df). The only juvenile vole
trapped during the study, a male taken in June, harbored 232 ectoparasites. The average
number of ectoparasites per vole was significantly higher for males (27.2) than for
females (17.6) (chi square = 83.06**, 1 df).
The ectoparasites are listed in order of decreasing abundance, and both the average
number per vole of each parasite and the average number per infested vole are given
(Table 1). Further discussion occurs below concerning some of the ectoparsites.
Zoology
651
Table 1. Ectoparasites of 86 pine voles, Microtus pinetorum, examined from Clark
County, Illinois.
No.
Av. no.
Av. no. per
Parasites
voles
Percent
No.
per vole
infested vole
Mites
Androlaelaps fahrenholzi
45
52.3
703
8.2
15.6
Euschoengastia ohioensis
63
73.3
473
5.5
7.5
Glycyphagus hypudaei
15
17.4
416
4.8
27.7
Haemogamasus longitarsus
22
25.6
85
1.0
3.9
Laelaps kochi
26
30.2
54
0.6
2.1
Pygmephoridae
22
25.6
40
0.5
1.8
Myocoptes japonensis
7
8.1.
26
0.3
3.7
Hypoaspis sp.
15
17.4
21
0.2
1.4
Haemogamasus liponyssoides
5
5.8
16
0.2
3.2
Euschoengastia diversa
7
8.1
12
0.1
1.7
Haemogamasus harperi
7
8.1
11
0.1
1.6
Eulaelaps stabularis
9
10.5
10
0.1
1.1
Neotrombicula whartoni
6
7.0
10
0.1
1.7
Wichmannia sp.
3
3.5
8
0.1
2.7
Neotrombicula lipovskyi
5
5.8
7
0.1
1.4
Radfordia lemnina
4
4.7
5
0.1
1.3
Dermacarus hylandi
1
1.2
1
0.01
1.0
Euschoengastia peromysci
1
1.2
1
0.01
1.0
Eutrombicula alfreddugesi
1
1.2
1
0.01
1.0
Haemogamasus ambulans
1
1.2
1
0.01
1.0
Eucheyletia bishoppi
1
1.2
1
0.01
1.0
Cheyletidae
1
1.2
1
0.01
1.0
Ornithonyssus bacoti
1
1.2
1
0.01
1.0
Fleas
Ctenophthalmus pseudagyrtes
17
19.8
32
0.04
1.9
Peromyscopsylla hamifer
I
1.2
1
0.01
1.0
Epitedia wenmanni
1
1.2
1
0.01
1.0
Doratopsylla blarinae
1
1.2
1
0.01
1.0
Stenoponia americana
1
1.2
1
0.01
1.0
Ticks
Dermacentor variabilis
9
10.5
9
0.1
1.0
Laelapidae
Androlaelaps fahrenholzi (Berlese, 1911), a laelapid mite, was the most abundant
ectoparasite, with 349 females, 91 males, and 263 nymphs collected. Adults and nymphs
were found together on 33.3% of the infested voles, females and nymphs on 24.4%
and females only on 22.2%. Androlaelaps fahrenholzi was taken from pine voles
throughout the study, but was significantly more abundant in the spring and summer
months (17.5 per vole) than in the fall and winter (6.0 per vole) (chi square = 209.09**,
1 df). The percentage of voles infested with A. fahrenholzi during the spring-summer
period (68.8%) was not significantly different from that of the fall-winter period (48.6%)
(chi square = 0.98, 1 df)- Adult mites did not vary significantly in infestation frequency
between spring-summer (62.5%) and fall-winter (44.3%) (chi square = 0.93, 1 df).,
but nymphs were taken at significantly higher frequencies in the spring and summer
(62.5%) than in the fall and winter (28.6%) (chi square = 4.25*, 1 df). The higher
infestation frequency of nymphs during this period may indicate an increase in mite
reproduction at this time. Adult pine voles had a significantly higher average number
of individuals per vole (8.1) than subadult voles (3.3) (chi square *> 52.54**, 1 df).
Male voles showed a higher average number of individuals per vole (9.9) than did
females (4.4) (chi square = 86.4**, 1 df). The higher abundance of both life stages
652 Indiana Academy of Science Vol. 94 (1985)
on male voles may reflect mere mobility by males, resulting in more opportunities
to pick up unattached mites.
Fourth and fifth in abundance were two additional laelapid mites, Haemogamasus
longitarsus (Banks, 1910) (76 females, 5 males, and 4 nymphs) and Laelaps kochi
Oudemans, 1936 (38 females, 9 males, 4 protonymphs, and 3 deutonymphs). Adults
of H. longitarsus were significantly more abundant during the spring and summer (1.7
per vole) than during the fall and winter (0.8 per vole) (chi square = 11.35**, 1 df).
The average number per vole of adult H longitarsus on subadult voles was 0.6 and
on adult voles it was 0.9, a difference that was not significant (chi square = 2.51,
1 df). Male voles showed a significantly higher average abundance of adult mites per
vole (1.3) than females (0.7) (chi square = 7.26**, 1 df)- Adults of L. kochi showed
no significant difference in the average number per vole for the spring-summer (0.8)
and fall-winter (0.50) (chi-square = 1.53, 1 df)- There was, however, a significantly
higher incidence of infestation during the spring and summer (50.0%) when compared
to fall-winter (21.4%) (chi-square = 3.91*, 1 df). Subadult voles had the same average
number of adult L. kochi per vole (0.5) as adults. An average number per vole of
0.6 was found for male and female voles alike.
Some other laelapid mite species taken were Haemogamasus liponyssoides Ewing,
1925 (3 females and 13 nymphs), Haemogamasus harperi Keegan, 1951 (3 females,
2 males, and 6 nymphs), Eulaelaps stabularis (Koch, 1836) (10 females), and
Haemogamasus ambulans (Thorell, 1872) (1 male).
Chiggers (Trombiculidae)
The parasite second in abundance was the chigger, Euschoengastia ohioensis Farrell,
1956. It had the greatest frequency of any parasite and was the only parasite found
to have a significantly higher average abundance during the fall and winter months
(6.6 per vole) than during the spring-summer period (0.9 per vole) (chi-square = 76.45**,
1 df). Frequency of infestation was also significantly higher during the fall- winter months
(84.3%) than during spring and summer (25.0%) (chi square = 6.23**, 1 df). This
species also had a significantly higher average number per vole on subadults (6.9) than
on adults (5.1) (chi-square = 9.04**, 1 df)- Whereas most parasites encountered were
more abundant on male voles, E. ohioensis had a significantly higher average abun-
dance per vole on females (70) than on males (4.3) (chi-square = 27.57**, 1 df). These
results indicate more active reproduction in fall and winter, and that infestation prob-
ably occurs in the nest since female and subadult voles are more heavily infested.
Other chiggers found were Euschoengastia diversa Loomis, 1956, Euschoengastia
peromysci (Ewing, 1929), Neotrombicula whartoni (Ewing, 1929), Neotrombicula
lipovskyi (Brennan and Wharton, 1950), and Eutrombicula alfreddugesi (Oudemans,
1910). The specimen of E. alfreddugesi was taken in July; the others in October through
December.
Glycyphagidae
Adults and tritonymphs of Glycyphagus hypudaei (Koch, 1841) (Glycyphagidae)
have been taken in Europe (Turk and Turk, 1957; Rupes, 1967), but have not been
reported from North America. The hyopial form (deutonymph), however, has been
found on a variety of North America mammals (Fain and Whitaker, 1973). Adults
of each sex, tritonymphs and deutonymphs, apparently all of this species, were found
on pine voles during the present study, although the adults and tritonymphs do not
precisely fit the previous descriptions. This was the third most abundant ectoparasite
found on the pine voles. A total of 107 adult males, 102 adult females, 181 tritonymphs,
and 26 deutonymphs was collected. Adults of both sexes were taken from each of
Zoology 653
five voles (5.8%), with an average abundance of 21.4 and 20.4 per infested vole for
males and females, respectively. Tritonymphs were found on eight voles (9.3%), five
of which also harbored adults. The average number of tritonymphs per infested animal
was 22.6. Hypopi were recovered from eight voles (9.3%), with an average of 3.3
per infested vole. All three life stages were found together on only one vole. The average
number of individuals per vole (all life stages included) was significantly higher during
the spring-summer period (10.3) than during the fall and winter (3.6) (chi-square =
121.80**, 1 df). The abundance of adult mites was also significantly higher in spring
and summer than during fall and winter, with average numbers per vole of 3.4 and
2.2 respectively (chi-square = 7.20**, 1 df). The tritonymphs showed a similar pattern,
with an average of 6.8 per vole in the spring-summer and 1.0 in the fall-winter (chi-
square = 206.74**, 1 df). The deutonymphs showed no significant seasonal difference.
The frequency of voles infested with G. hypudaei (all life stages included) also was
significantly higher in the spring-summer period (37.5%) than during the fall-winter
period (12.9%) (chi square = 4.50*, 1 df). The average number of G. hypudaei per
vole (all life stages included) for the subadult voles (1.8) was significantly lower than
for adults (4.4) (chi square = 29.13**, 1 df). Adult mites averaged 2.9 per vole on
the adult voles, but none were taken from subadult voles, again a significant difference
(chi square = 61.36**, 1 df). Deutonymphs had a significantly higher abundance on
subadults (0.6 per vole) than on adults ((0.2 per vole) (chi-square = 7.22**, 1 df).
The one juvenile pine vole caught during the study harbored one deutonymph, 75
tritonymphs, and 22 adults. Male pine voles had a significantly higher average number
of G. hypudaei (all life stages included) per vole (8.4) than females (2.1) (chi-square
= 150.23**, 1 df). The deutonymphs, however, had a significantly higher abundance
on females (0.5 per vole) than on males (0.1 per vole) (chi-square = 11.65**, 1 df).
The significance of the numbers of life stages other than deutonymphs in this sample
is not understood at this time.
Another glycyphagid mite collected during this study was a single deutonymph
of Dermacarus hylandi Fain, 1969, a species that has been reported only from Tamias
striatus (Fain et al., 1971; Fain and Whitaker, 1973) and Clethrionomys gapperi (Fain,
1969).
Other mites
A few other kinds of mites were found (Table 1). Some deserve further mention.
All anoetids appeared to be Wichmannia. One of two cheyletids taken appears to be
Eucheyletia bishoppi Baker, 1949. The second could not be identified because of its
poor condition. Radfordia hylandi Fain and Lukoschus, 1977 was the only myobiid
species found (five specimens taken, all females). Myocoptes japonensis Radford, 1955
was the only myocoptid mite recovered (20 females, two males, and four immatures.)
Several species of Pygmephoridae were collected. Species identified from this
material by S. Mahunka are Bakerdania jonesi Mahunka, 1975, B. plurisetosa Mahunka,
1975, Pseudopygmephorus quadratus (Ewing, 1917), Pygmephorus equitrichosus
Mahunka, 1975, P. hastatus Mahunka, 1973, P. scalopi Mahunka, 1973, and P.
whitakeri Mahunka, 1973.
It is interesting that no listrophorid mites were collected. Listophorid mites,
particularly Listrophorus mexicanus, are often abundant on microtine rodents (Fain
and Hyland, 1974; Whitaker, 1982), and Listrophorus pity my s was described from
Microtus pinetorum from Rhode Island. We have since seen specimens of Listrophorus
pitymys from pine voles from Georgia and Kentucky (Whitaker, unpublished) and New
York (Whitaker and French, unpublished). However, no listrophorids were taken from
this host from Illinois during the present study nor from Indiana (Whitaker, 1982).
654 Indiana Academy of Science Vol. 94 (1985)
Fleas
Ctenophthalmus pseudagyrtes Baker, 1904, the normal flea of the pine vole, was
the only species collected from more than one animal (16 of each sex taken). Other
fleas taken (one male each) were Peromyscopsylla hamifer (Rothschild, 1906), Epitedia
wenmanni (Rothschild, 1904), Doratopsylla blarinae C. Fox, 1914, and Stenoponia
americana (Baker, 1899).
Ticks
Dermacentor variabilis (Say, 1821) was the only tick found (5 larvae).
Discussion
Species not previously taken on this host are the laelapid mite Haemogamasus
harperi, the pygmephorid mites Bakerdania jonesi, B. plurisetosa, and
Pseudopygmephorus quadratus, and the fleas Peromyscopsylla hamifer and Epitedia
wenmanni. Mites other than chiggers not previously taken from Illinois are
GLYCYPHAGIDAE: Glycyphagus hypudaei, Dermcarus hylandi; LAELAPIDAE:
Eulaelaps stabularis, Haemogamasus harperi, H. longitarsus; MYOBIIDAE: Radfordia
hylandi; PYGMEPHORIDAE: Bakerdania jonesi, B. plurisetosa, Pseudopygmephorus
quadratus.
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15. Holland, G.P., and A.H. Benton, 1968. Siphonaptera from Pennsylvania mammals.
Amer. Midi. Natur., 80: 252-261.
16. Jameson, E.W., Jr. 1943. Notes on the habits and siphonapterous parasites of
the mammals of Welland County, Ontario. J. Mammal., 24:194-197.
17. Jameson, E.W., Jr. 1947. Natural history of the prairie vole (Mammalian genus
Microtus). Univ. Kans. Publ., Mus. Natur. Hist., 1:125-151.
18. Jordan, K. 1928. Siphonaptera collected during a visit to the eastern United States
of America in 1927. Novit. Zool., 34:178-188.
19. Kardos, E.H. 1954. Biological and systematic studies of the subgenus Neotrom-
bicula (Genus Trombicula) in the central United States (Acarina, Trombiculidae).
Univ. Kans. Sci. Bull., 36:69-123.
20. Layne, J.N. 1958. Records of fleas (Siphonaptera) from Illinois mammals. Natur.
Hist. Misc., Chicago Acad. Sci., 162:1-7.
21. Loomis, R.B. 1956. The chigger mites of Kansas (Acarina, Trombiculidae). Univ.
Kans. Sci. Bull., 37:1195-1443.
22. MacCreary, D. 1945. Some ectoparasites, excluding Ixodoidea, of Delaware mam-
mals. J. Econ. Entomol., 38:126-127.
23. Manischewitz, J.R. 1966. Studies on parasitic mites of New Jersey. J. N.Y. En-
tomol. Soc, 74:189-197.
24. Mathewson, J. A., and K.E. Hyland, Jr. 1964. The ectoparasites of Rhode Island
mammals. III. A collection of fleas from nondomestic hosts (Siphonaptera). J.
Kans. Entomol. Soc, 37:157-163.
25. Mellot, J.L., and W.A. Connell. 1965. A preliminary list of Delaware Acarina.
Trans. Amer. Entomol. Soc, 91:85-94.
26. Morlan, H.B. 1952. Host relationships and seasonal abundance of some southwest
Georgia ectoparasites. Amer. Midi. Natur., 48:74-93.
27. Mumford, R.E., and J.O. Whitaker, Jr. 1982. Mammals of Indiana. Indiana
Univ. Press, Bloomington. 537 p.
28. Poorbaugh, J.H., and H.T. Gier. 1961. Fleas (Siphonaptera) of small mammals
in Kansas. J. Kans. Entomol. Soc, 34:198-204.
29. Pratt, H.D., and H. Karp. 1953. Notes on the rat lice Polyplax spinulosa
(Burmeister) and Hoplopleura oenomydis Ferris. J. Parasitol., 39:495-504.
30. Rupes, V. 1967. The mites of the subfamily Labidophorinae Zachvatkin, 1941
(Acarina, Acarinae) from the nests of small mammals. Vest. Ceskoslov. Spolec-
nosti Zool., Acta Soc. Zool. Bohemoslov., 31:68-79.
31. Smiley, R.L., and J.O. Whitaker, Jr. 1979. Mites of the genus Pygmephorus
(Acari: Pygmephoridae) on small mammals in North America. Acta Zool. Acad.
Scient. Hungaricae 25:383-408.
32. Sonenshine, D.E., E.L. Atwood, and J.T. Lamb, Jr. 1966. The ecology of ticks
transmitting Rocky Mountain spotted fever in a study area in Virginia. Ann.
Entomol. Soc. Amer., 59:1234-1262.
33. Turk, E., and F. Turk. 1957. Systematik und okologie der p. Tyroglyphiden
Mitteleuiopas. In: H.J. Stamer, Beitrage zur systematik und okologie Mitteleuropas,
Acarina, 1:1-231.
34. Whitaker, J.O., Jr. 1982. Ectoparasites of mammals of Indiana. Ind. Acad. Sci.
Monograph No. 4. Indianapolis. 240 p.
35. Whitaker, J.O., Jr., and K.W. Corthum, Jr. 1967. Fleas of Vigo County, Indiana,
Proc Ind. Acad. Sci., 76:431-440.
36. Whitaker, J.O., Jr., and N. Wilson. 1974. Host and distribution lists of mites
656 Indiana Academy of Science Vol. 94 (1985)
(Acari), parasitic and phoretic, in the hair of wild mammals of North America,
north of Mexico. Amer. Midi. Natur., 91:1-67.
37. Wilson, N. 1957. Some ectoparasites from Indiana mammals. J. Mammal.,
38:281-282.
38. Wilson, N. 1961. The ectoparasites (Ixodides, Anoplura, and Siphonaptera) of
Indiana mammals. Unpubl. Ph.D. thesis, Purdue Univ. 527 p.
Quaternary Remains of the Spotted Skunk,
Spilogale putorius, in Indiana
Ronald L. Richards,
Indiana State Museum and Historic Sites
202 North Alabama Street
Indianapolis, Indiana 46204
Introduction
There is no physical evidence (skin or skull) of a live spotted skunk (Spilogale
putohus) in Indiana. However, "civet cats" were known to fur dealers in extreme
southwestern Indiana prior to about 1922. Evermann and Butler (5) suggested that
the spotted skunk might occur in Indiana, and Hahn (9) reported the trapping of a
"civet cat" between Bicknell and Bruceville in Knox County. He also cited a fur dealer
from Vincennes who had obtained a few "civet cats" with "several curved white stripes
and spots on the body" from the southern part of the State. Hahn also related that
a St. Louis fur dealer had received a few civet cat skins from Indiana.
Lyon (15) noted three occurrences of the animal in the Mount Vernon, Posey
Co. area, one of which (Lynn Township) he regarded as the best documented observa-
tion of the skunk in Indiana to date. Lyon also noted that some authors had listed
the spotted skunk for Indiana (eg. Howell (12) for Posey Co. and Cory (4) for southern
Indiana) without presenting any evidence. Mumford (16) summarized previous reports
and Mumford and Whitaker (17), in treating only extant fauna, excluded the spotted
skunk.
It appears that the spotted skunk was a rare inhabitant of at least extreme
southwestern Indiana in historic times and is now extirpated from the state. The spotted
skunk presently occurs no nearer to Indiana than Missouri on the west, and Tennessee
and eastern Kentucky on the south.
The spotted skunk inhabits prairies and brushy or sparsely wooded areas (3, 14).
It is an agile climber and in eastern Kentucky is found among the cliffs and rocks
in the rugged terrain where it usually dens in crevices at the base of a cliff or among
boulders (2). It is a common cave fossil within its modern range in Missouri (21, 24, 25).
In 1958 a left dentary of S. putorius was recovered from a woodrat den in Sullivan's
Cave, Lawrence County, Indiana (1), demonstrating that the animal once had a wider
distribution in Indiana. More recent references to spotted skunk include remains from
south-central Indiana caves (26, 29, 31, 32) and the Pleistocene occurrence of the animal
in the Harrodsburg Crevice fauna of Monroe County (23).
Descriptive Paleozoology
Most S. putorius remains were associated with rather extensive faunas. The species
that were extinct, extralocal (occurring out of their modern range), or extirpated (ex-
terminated within Indiana) are listed in Table 1. All references to a locality or any
of its fauna are cited. Abbreviations: B.P., Before Present (1950 A.D.); MNI, minimum
number of individuals; L, R, left, right; C, c, M, m, upper and lower canines and
molars respectively; cm, centimeter; mm, millimeter; m, meter.
Remains of at least 25 individuals are known from 9 localities in Monroe, Lawrence
and Harrison Counties (Figure 1).
1. Freeman's Pit, Monroe County, Indiana. OCCURRENCE: Bones occurred in
the upper 30 cm. of a 60 cm. deep laminated silt/clay deposit in a corner alcove
of the entrance room of the 29.6 m. (97 foot) deep pit. Bones of other individuals
were found in the silty sediments of a small chamber ("Attic") located off the
657
658 Indiana Academy of Science Vol. 94 (1985)
Table 1. Important Associations on Indiana Spilogile putorius bone localities.'
FP HC IR
sc ccc
Ledge Floor
KLC
PPC FMW
Extinct:
Dasypus bellus, Beautiful Armadillo
Canis cf. C. dirus, Dire Wolf
Smilodon fatalis, Sabertooth
Panthera onca augusta, Pleistocene Jaguar
Equus cf. E. complicatus, Pleistocene Horse
Mylohyus sp., Long-nosed Peccary
Platygonus cf. P. vetus ( = P. cf.
cumberlandensis), Leidy's Peccary
X
X
X
X
X
X
X
Extralocal:
Parascalops breweri, Hairy-tailed Mole
Spermophilus tridecemlineatus, Thirteen-
lined Ground Squirrel
Geomys cf. G. bursarius. Plains Pocket
Gopher
Oryzomys plaustris. Rice Rat
Neotoma floridana, Eastern Woodrat
Clethrionomys gapperi, Boreal Red-backed
Vole
Phenacomys cf. P. intermedius, Heather
Vole
X X
X
X X
X
XXX X X X X (local) X
X
X
Extirpated:
Ectopistes migratorius. Passenger Pigeon
Erethizon dorsatum, Porcupine
Ursus americanus. Black Bear
Cervus elaphus, Wapiti
X
X XXX
X X X X X
X3 X
1 . FP, Freeman's Pit, Monroe Co.; HC, Harrodsburg Crevice, Monroe Co.; IR, Indun Rockshelter, Monroe Co.;
SC, Sullivan's Cave, Lawrence Co. (note ledge and floor deposits); CCC, Carcass Crypt Cave, Lawrence Co.; KLC,
King Leo Cave, Harrison Co.; PPC, Passenger Pigeon Cave, Harrison Co.; FMW, Fair-to-Middlin Well, Harrison, Co.
2. Nomenclature of Kurten and Anderson (14).
3. Noted by Bader and Hall (1) only.
main room some 18.3 m. above the cave floor. MATERIALS: 2 complete, 1
fragmented skulls; 2L, 3R additional maxillae; 11L (2 are small fragments), 9R
dentaries; 4L, 4R fragmented scapulae; 4L, 8R humeri; 5L, 5R ulnae; 3L, 8R
radii, 5L, 7R fragmented innominates; 6L, 6R femora; 4L, 5R fragmented tibiae;
3L, 4R calcanea; 3L, 3R astragali: (MNI = 11). "Attic": RC; Lc; skull fragment;
2LM1 and 2LM2 (MNI ■ 2). COMMENTS: A radiocarbon date of 2,315 ± 65
years B.P. was determined for the base of deposits containing S. putorius (30).
PUBLISHED RECORDS: (26, 30, 31).
Harrodsburg Crevice, Monroe County, Indiana. OCCURRENCE: Remains from
an "undisturbed mass of bone embedded in a beige clay-limestone, detritus-
travertine matrix" (23) from the floor deposit of a collapsed, filled in cave sec-
tioned during highway construction. MATERIALS: L humerus, distal end
(MNI = 1). COMMENTS: A warmer and drier-than-present short-grass
prairie/forest edge environment of Sangamonian (last interglacial) age was pro-
posed for the fauna. PUBLISHED RECORDS: (18, 23, 36).
Indun Rockshelter, Monroe County, Indiana. OCCURRENCE: Remains recovered
from Levels 1 and 2 in sediments of a sandstone shelter. MATERIALS: L radius
Zoology
659
Figure 1 . Modern (lettered), fossil and subfossil (numbered) localities for the Spotted
Skunk, Spilogale putorius, in Indiana. A. Knox County (Hahn, 1909). B. Posey County
(Lyon, 1936). 1. Freeman's Pit, Monroe County. 2. Harrodsburg Crevice, Monroe County.
3. Indun Rockshelter, Monroe County. 4. Sullivan's Cave, Lawrence County. 5. Car-
cass Crypt Cave, Lawrence County. 6. King Leo Cave, Harrison County. 7. Passenger
Pigeon Cave, Harrison County. 8. N. Jim Cave, Harrison County. 9. Fair-to-Middlin
Well (pit), Harrison County.
and L ulna, proximal section (MNI = 1). COMMENTS: Because a fauna ranging
in age from Late Pleistocene to modern age was mixed in the disturbed deposit
any associations are unclear. PUBLISHED RECORDS: None.
4. Sullivan's Cave, Lawrence Co., Indiana. OCCURRENCE: Bones recovered from
sediments of an ancient woodrat den on a ledge a couple meters above the cave
floor and within a 23 cm. deep sand, gravel and organic silt deposit in a floor
channel below and adjacent to the ledge some 600 m. inside the cave.
MATERIALS: "Ledge:" L dentary (collected by Bader and Hall in 1958); RC;
L, R c; LM1; Rm3; L radius (collected by author in same deposit in 1971).
"Floor:" RM2 from upper 8 cm. of sediment. RM2 collected in gravely sediments
below this. (MNI = 3). COMMENTS: Temporal association of the fauna is unclear.
Further excavation will be undertaken. PUBLISHED RECORDS: (1, 26, 31).
5. Carcass Crypt Cave, Lawrence County, Indiana. OCCURRENCE: Bones found
in the upper 5 cm. of carbon-rich silt/clay in floor deposits ca. 25.9 m. inside
the pit cave (21.6 m. entrance drop). A calcaneum had been transported by a
woodrat to a cache 2.4 m. off the floor of the room. MATERIALS: Complete
skull; R dentary; R innominate; L ulna; 2L, 1R femora; 2L, 1R tibiae; R calcaneum
(MNI =2). COMMENTS: Much of the fauna, primarily of deciduous woodlands,
660 Indiana Academy of Science Vol. 94 (1985)
is believed to be of Holocene age, though no C-14 dates are available. PUBLISHED
RECORDS: (26, 28, 31).
6. King Leo Cave, Harrison Co., Indiana. OCCURRENCE: Bones occurred in the
upper 23 cm. of a ca. 33 cm. deep sand/silt deposit in a small passageway perhaps
6 m. from the top of the 19.8 m. (65 foot deep) pit entrance. MATERIALS:
L dentary; LM1; RM2; R humerus (MNI = 1). PUBLISHED RECORDS: (30).
7. Passenger Pigeon Cave, Harrison Co., Indiana. OCCURRENCE: Bone recovered
from within the top 30 cm. of a loose, dusty silt deposit a couple meters inside
a shallow shelter-like limestone cave on an Ohio River bluff. MATERIALS: RM1
in maxilla section; R dentary, rodent gnawed (MNI = 1). PUBLISHED RECORDS:
(27). COMMENTS: The teeth are of large size (Table 2); identification is ten-
tative (skunk cf. Spilogale putorius).
8. N. Jim Cave, Harrison Co., Indiana. OCCURRENCE: Bone found in winnowed
gravely sediments on surface, ca. 21 m. inside pit cave (10.7 meter deep pit).
MATERIALS: Dentary (MNI = 1). COMMENTS: Isolated dentary with most of
the dentition was identified by author in the field, and later lost in transit to
the lab. PUBLISHED RECORDS: (26).
9. Fair-to-Middlin Well (Pit), Harrison County, Indiana. OCCURRENCE: Skull
noted in gravely sediments on surface associated with bear skeleton in a water
washed area at corner of room below the 27.4 m. (90 foot) entrance drop.
MATERIALS: Complete skull; L dentary; L, R humeri; L, R innominates; R
femur; L, R tibiae (MNI = 2). PUBLISHED RECORDS: (32).
The complete S. putorius skulls are illustrated in Figure 2. Measurements are given
in Table 2.
The Indun Rockshelter remains are on file at the Glenn A. Black Laboratory
of Archaeology, Bloomington, Indiana, the Fair-to-Middlin Well bones are at the Indiana
State Museum, Indianapolis, and the Harrodsburg Crevice remains are at the Dept.
of Anthropology, University of Tennessee, Knoxville. All other remains, largely from
undescribed faunas, are presently on file with the author.
Discussion
Two subspecies of S. putorius are present in eastern United States (exclusive of
Florida), the easterly S. putorius putorius and westerly S. p. interrupta; both intergrade
in the southern part of their ranges (35). Though the largest cranial measurements
are of 5. p. putorius from Hale Co., Alabama, the slightly larger of the two subspecies,
the great overlap in size makes subspecific separation of individual fossils difficult
(35). The Indiana material (Harrodsburg Crevice bone not examined) was of relatively
large size, with some dimensions (especially toothrow lengths) similar to those from
Hale Co., Alabama. This suggests that S. p. putorius, the subspecies typically indicated
for Indiana (10, 35), is represented by the Indiana fossils. Van Gelder (1959) notes
that average cranial measurements of S. putorius are in general larger in the northern
part of its range. Existing metric data, however, is difficult to utilize in interpreting
fossil gradients of size, as measureable skulls are not commonly recovered from fossil
localities, and there is a lack of published postcranial measurements. For this reason
selected measurements of Indiana S. putorius have been presented (Table 2). S. putorius
displays a sexual dimorphism in size with male crania 5% to 9°7o larger than those
of females in various measurements (35). This dimorphism is apparent in especially
the Indiana postcranial material.
Several species of extralegal and some extinct mammals have been associated with
5. putorius on the Indiana fossil localities (Table 1). This suggests that environmental
conditions at that time were different than today. None of the localities (except, perhaps
Zoology
661
Figure 2. Complete skulls of the Spotted Skunk, Spilogale putorius, recovered from
Indiana pit caves. A. Freeman's Pit, Monroe County (dorsal view). B. Fair-to-Middlin
Well, Harrison County (dorsal view). C. Freeman's Pit, Monroe County (palatal view).
D. Carcass Crypt Cave, Lawrence County (right lateral view). E. Right dentary, Car-
cass Crypt Cave, Lawrence County (labial view). Scale in centimeters.
Sullivan's Cave) appear to have full-glacial boreal faunas. All Indiana S. putorius remains
have been found in interglacial (Harrodsburg Crevice), very late glacial (?Sullivan's Cave),
or post-glacial deposits. The most common extralocal associates are the Woodrat, Plains
Pocket Gopher and Hairy-tailed Mole, with the Thirteen-lined Ground Squirrel, Rice Rat,
Red-backed and Heather Voles at one locality each. Excepting the Hairy-tailed Mole, Red-
662
Indiana Academy of Science
Vol. 94 (1985)
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664 Indiana Academy of Science Vol. 94 (1985)
backed and Heather Voles, all are southern or western in distribution today, and the Plains
Pocket Gopher and Thirteen-lined Ground Squirrel are indicative of open vegetation
and well-developed soils. Geomys in particular once had a much more extensive range
in Illinois, Indiana, Kentucky and Tennessee which Parmalee and Klippel (22) suggest
indicates drier conditions than today with extensive prairies or open parklands throughout
the region. The Hairy-tailed Mole, Red-backed and Heather Voles suggest cooler, more
mesic conditions. Most of the associates, however, including the reptiles and amphi-
bians, are within their modern range. Graham (letter, November 15, 1984) suggests
that the presence of western species in "eastern" faunas indicates an "environmental
mosaic, with patches of open vegetation and well developed soils," that could be created
by a slight change in climate or edaphic conditions.
The Illinois records of Spilogale putorius are similar to those in Indiana. There
is one unreliable observation for 1910 in southeast Illinois (11), and two Holocene
bone localities. The Modoc Rock Shelter, Randolph County, produced extralocal Plains
Pocket Gopher remains and bones of at least 9 Spotted Skunks from the 4500-6500
B.P. levels (20). Meyer Cave, Monroe County, Illinois had an extensive fauna with
northern, southern and western extralocals that included the Plains Pocket Gopher
and 25 Spotted Skunks. The presence of the western animals was thought to have
dated from the post-Wisconsinan Xerothermic Interval, characterized by a warm, dry
climate (19). At that time the ranges of forest animals are thought to have been
fragmented and the ranges of grassland, forest-edge and aridity-tolerant forest animals
shifted eastward (33, 34).
Guilday et al. (8) in discussing the "Prairie Peninsula," noted that the evidence
for a late-glacial eastern movement of western species was much stronger than that
for the hypothesized Xerothermic Interval thousands of years later.
The assignment of western extralocal species in a fossil fauna to either of these
two periods should be undertaken with caution and in context with the entire fauna.
This is true of the Sullivan's Cave, Indiana fauna.
Graham (7) and King and Graham (13) suggested that each species responds to
environmental change individually, with its distribution controlled by the interaction
of many variables such as habitat type, competitive advantage, and physiological
tolerances interfacing with climatic parameters. Climatic reconstructions based solely
upon modern temperature and humidity extremes in the distribution of a species without
regard to other limiting factors could be oversimplified. Until limiting factors are better
known for modern species, it will be difficult to speculate upon the cause of many
range extensions. It seems productive at present to establish a chronology of common
faunal associates, making inferences from particular species with better known limiting
factors. The common extralocal associates of S. putorius tend to be southern and western
species, and Geomys in particular suggests more extensive tracts of open vegetation.
This accords well with interglacial, late-glacial and postglacial range extensions of the
spotted skunk. The Late Pleistocene in general is thought to have been more equable
than at present, with greater overlapping of many presently separated animal ranges (6).
As indicated by the Freeman's Pit, Monroe County, Indiana Radiocarbon date
of 2,315 ± 65 years B.P., the spotted skunk was still present in south-central Indiana,
long before the Caucasoid settlement of the region when its extirpation from the lower
Wabash Valley was recorded. The lack of historic accounts from south-central Indiana
where its bones are common suggest that it had been undergoing a general range reduc-
tion since at least 2300 years ago, in response to environmental change.
Acknowledgments
Russel W. Graham, Illinois State Museum, kindly reviewed this manuscript, and made
Zoology 665
helpful notation on range limiting factors of western species. Dave Rieger and Donald
B. Haddix produced Figures 1 and 2 respectively, and Deborah Randolph, Indiana State
Museum, typed the manuscript. A thanks is extended to cavers too numerous to name
for help in the field recovery of specimens.
Literature Cited
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Mammal., 41:111-112.
2. Barbour, R.W. and W.H. Davis. 1974. Mammals of Kentucky. The University
Press of Kentucky. 322 p.
3. Burt, W.H. and R.P. Grossenheider. 1964. A field guide to the mammals.
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4. Cory, C.B. 1912. The mammals of Illinois and Wisconsin. Field Mus. Nat. Hist.
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5. Evermann, B.W. and A.W. Butler. 1894. Preliminary list of Indiana mammals.
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6. Graham, R.W. 1976. Late Wisconsin mammalian faunas and environmental gra-
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15. Lyon, M.W., Jr. 1936. Mammals of Indiana. Amer. Midi. Natur., 17(1): 1-384.
16. Mumford, R.E. 1969. Distribution of the mammals of Indiana. Indiana Acad.
Sci. Monogr. 1. Indianapolis, 114 p.
17. Mumford, R.E. and J.O. Whitaker, Jr. 1982. Mammals of Indiana. Indiana Univ.
Press, Bloomington. 537 p.
18. Munson, P. J., P.W. Parmalee and J.E. Guilday. 1980. Additional comments on
the Pleistocene mammalian fauna of Harrodsburg Crevice, Monroe County,
Indiana. Nat'l Speleol. Soc. Bull. 42:78-79.
19. Parmalee, P.W. 1967. A recent cave bone deposit in southwestern Illinois. Nat'l.
Speleol. Soc. Bull. 29(4): 119-147.
20. Parmalee, P.W. and D.F. Hoffmeister. 1957. Archaeozoological evidence of the
spotted skunk in Illinois. J. Mammal. 38(2):261.
21. Parmalee, P.W. and K.W. Jacobson. 1959. Vertebrate remains from a Missouri
cave. J. Mammal. 40(3): 40 1-405.
666 Indiana Academy of Science Vol. 94 (1985)
22. Parmalee, P.W. and W.E. Klippel. 1981. A late Pleistocene population of the
Pocket Gopher, Geomys cf. busahus, in the Nashville Basin, Tennessee. J. Mam-
mal., 62(4): 83 1-835.
23. Parmalee, P.W., P.J. Munson and J.E. Guilday. 1978. The Pleistocene mam-
malian fauna of Harrodsburg Crevice, Monroe County, Indiana. Nat'l Speleol.
Soc. Bull. 40(2):64-75.
24. Parmalee, P.W. and R.D. Oesch. 1972. Pleistocene and Recent faunas from the
Brynjulfson Caves, Missouri. Illinois State Mus. Rept. Inv. No. 25, 52 p.
25. Parmalee, P.W., R.D. Oesch and J.E. Guilday. 1969. Pleistocene and Recent
vertebrate faunas from Crankshaft Cave, Missouri. Illinois State Mus. Rept. Inv.
No. 14, 37 p.
26. Richards, R.L. 1972. The woodrat in Indiana: Recent fossils. Proc. Indiana Acad.
Sci. 81:370-375.
27. Richards, R.L. 1980. Rice Rat (Oryzomys cf. palustris) remains from southern
Indiana caves. Proc. Indiana Acad. Sci., 89:425-431.
28. Richards, R.L. 1981. Vertebrate remains from Carcass Crypt Cave, Lawrence
County, Indiana. Proc. Indiana Acad. Sci., 90:442.
29. Richards, R.L. 1982. Hairy-tailed Mole (Parascalops breweri) remains from south-
central Indiana caves. Proc. Indiana Acad. Sci., 91:613-617.
30. Richards, R.L. 1983. Quaternary records of the Pygmy and Smoky Shrews from
southern Indiana caves. Proc. Indiana Acad. Sci., 92:507-521.
31. Richards, R.L. 1984. The Pleistocene Vertebrate collection of the Indiana State
Museum, with a list of the extinct and extralocal Pleistocene vertebrates of Indiana.
Proc. Indiana Acad. Sci., 93:483-504.
32. Richards, R.L. 1984. It's the pits: explorer discovers black bear bones. Outdoor
Indiana, vol. 49 (7):25-27. October.
33. Smith, P.W. 1957. An analysis of post-Wisconsin biogeography of the prairie
peninsula region based on distributional phenomena among terrestrial vertebrate
populations. Ecology, 38(2):205-218.
34. Smith, P.W. 1965. Recent adjustments in animal ranges, pp. 633-642 in: H.E.
Wright, Jr. and D.G. Frey, (eds.), The Quaternary of the United States, Princeton
Univ. Press.
35. Van Gelder, R.G. 1959. A taxonomic revision of the spotted skunks (Genus
Spilogale). Bull. Amer. Mus. Nat. Hist., 117(5):233-392.
36. Volz, S.A. 1977. Preliminary report on a Late Pleistocene death-trap fauna from
Monroe County, Indiana. Proc. Indiana Acad. Sci., 86:293-307.
Extinct Woodland Muskox, Symbos cavifrons, Cranium
from Miami County, North Central Indiana
Ronald L. Richards, Indiana State Museum and Historic Sites
202 North Alabama Street, Indianapolis, Indiana 46204
and
William R. Wepler, Miami County Historical Museum
4th Floor, Courthouse, Peru, Indiana 46920
Introduction
During a survey of natural history specimens at the Miami County Historical
Museum in the spring of 1984 the senior author noted a Symbos cavifrons skull among
the exhibited materials. The junior author, director of the museum, provided the follow-
ing information: Catalogue No. 170.56; found in muck on the farm of ex-sheriff Homer
Fenters, south of Macy; collected by C.F. Fite; donated by Pearl M. Fite, August
29, 1919. Examination of county plat records indicated that this locality was in the
south 1/2 of Section 24 and NE corner of Section 25, T29N, R3E, Allen Township,
Macy Quad., Miami County. Our search of the literature failed to locate any descrip-
tion or published record of a Symbos skull from Miami County, with the exception
of an excellently preserved skull from adjacent Union Township now in the U.S. National
Museum (10). Jerry N. McDonald, now actively studying muskoxen, confirmed that
two different specimens were involved, and that the Allen Township skull had been
previously unrecorded (personal communication, August, 1984). The Union Township
skull (USNM 8574), also collected by C.F. Fite in 1916 had been sold for $10.00 and
sent to the U.S. National Museum by O.P. Hay, received on February 20, 1917 as
Accession No. 60856 (personal communication, August, 1984, Ed Ducco, Dept. of
Vertebrate Paleontology, U.S. National Museum).
Description
From dorsal aspect, most of the skull is present, with an excellently preserved
exostosis. Ventrally, most of the sphenoid area and much of the palate (except the
left alveolar area) is missing including the rostral section of the left maxilla. Also absent
are the zygomatic arches, caudal wall of the right orbit, and both premaxillae. Only
the roots of the left first and second molars are present; the crowns have broken off
at the level of the alveoli. No other teeth are present. Though several sutures are obscured
by the exostosis and horn core bases (frontal, fronto-parietal and parieto-occipital
sutures), the basioccipital, nasofrontal and caudal nasomaxillary sutures are obliterated,
suggesting an adult animal. The parieto-temporal suture is visible and horizontally
oriented as in other 5. cavifrons (12).
Many of the thin extremities of bone (tips of horncores, tips of nasals, rostral
maxillaries, orbital rims) appear have been heavily gnawed by rodents. This might
indicate exposure for some time before burial. Several other marks on the skull might
not have been made by rodents and are under further study. The dark brown skull
has peat with minor inclusions of pond weed seeds and sand embedded in many of
its foramina and interior cavities. The skull lacks the abrasion that is present on many
apparently stream tumbled muskoxen skulls (2, 6, 15). It had been coated with varnish
some years ago, though this has not prevented some cortical bone from peeling. The
skull is illustrated in Figure 1, with selected measurements presented in Table 1.
667
668
Indiana Academy of Science
Vol. 94 (1985)
IIIIIIIIIII1III
Ficure 1. Woodland Muskox, Sumbos cavifrons, skull from Allen Township, Miami
County, Indiana in the collection of the Miami County Historical Museum (Catalogue
No. 170.56). A. Dorsal view. B. Left lateral view. C. Palatal view. Scale in centimeters.
Zoology 669
Table 1. Symbos cavifrons, Cranial Measurements (mm.)1
Exostosis length:
Exostosis width, anterior to horncores:
Exostosis width, across anteroinferior flanges:
Greatest depth of concavity between horn cores:
Anteroposterior diameter of horncore at base
Dorsoventral diameter of horncore at base:
Circumference of horn core at base:
Width between horncore tips (as preserved):
Tip of (preserved) horn core to sagittal plane:
Height from ventral margin of occipital condyle to dorsal surface
of cranium:
Greatest height of occipital region: basion-nuchal line:
Height from upper lip of foramen magnum to midline on dorsal
surface of cranium:
Height from upper lip of foramen magnum to top of nuchal crest:
Minimum height, occipital: ophisthion-nuchal line:
Minimum height, occipital: opisthion-dorsal edge nuchal ligament
insertion:
Height of foramen magnum: basion — opisthion:
Width of foramen magnum (at rim of condyle):
Height of skull above alveolar border to front of exostosis:
Transverse width of cranium at base of horn cores:
Width of cranium at constriction between horncores and orbits:
Greatest breadth of bases of paraoccipital processes
Greatest breadth of basioccipital:
Greatest breadth across occipital condyles:
Mastoid width:
Face width between supraorbital foramina:
Zygomatic width (on malars, ventral to orbits):
Angle, foramen magnum plane with occipital plane:
Angle, basioccipital plane with foramen magnum plane:
Greatest length, front of nasal (gnawed) to rear of skull, measured
along dorsal suface:
From front of nasals (gnawed) to anterior end of exostosis:
Greatest diameter of orbit (dorsoventral):
Width of orbit (anteroposterior):
1. Measurements primarily after McDonald, 1984; Harington, 1975, and Semken, Miller and Stevens, 1964.
*Fragmented or gnawed
**Calipers difficult to position along curving contours of skull
Discussion
Next to remains of the Mastodon (Mammut americanum) and Jefferson's
Mammoth (Mammuthus jeffersonii), remains of muskoxen are among the most com-
monly recovered of large extinct mammals in Indiana. The most recent descriptive
summary of Indiana muskoxen fossils was Lyon (10), though Kitts (6) provided some
additonal records. There are now 20 known specimens, comprising 2 or 3 species.
One fossil of the living muskox (Ovibos moschatus) has been recovered from Wayne
Co. (3, 10). One specimen of Bootherium is known from a gravel pit in Gibson Co.
(USNM 24885; information and casts sent to Indiana State Museum by Robert W.
Purdy, Dept. of Paleobiology, U.S. National Museum, March, 1984). Semken, Miller
and Stevens (17) suggest that Bootherium bombifrons is a distinct genus, and that
B. sargenti may be a female of Symbos (5, 21). An unidentified ovibovine tooth was
identified from the Prairie Creek Locality, Daviess County, by John Sparling (specimen
in Glenn A. Black Lab. of Archaeology, Indiana University, Bloomington). All other
iami Co., IN.
243
117
160
25
L = 108**
R =
115**
L= 74
R =
78
L= ca. 298**
R =
ca. 277
494*
L = 251*
R =
243*
208
136
169
110
111
81
37
47
204
136
138
156 + *
69
125*
182+*
101
228
ca. 127°
ca. 142°
485+*
235*
L= ca. 68
R =
ca. 67
L = ca. 64
670 Indiana Academy of Science Vol. 94 (1985)
muskoxen fossils have been referred to Symbos cavifrons. These include specimens
from: Bartholomew Co. (4); LaGrange Co. (14); LaPorte Co. (9); Miami Co. (10);
Montgomery Co. (11); Newton Co. (1, 10); Porter Co. (10); Randolph Co. (10) and
St. Joseph Co. (8). There are seven other skulls now under study by Patrick Munson
and Russell Graham. These include specimens from Knox (3 individuals), Koscuisko,
Marion, Owen and Parke counties.
S. cavifrons skulls are readily identified by the bony, pitted exostosis between
the horn cores, and the absence of the median groove (sulcus) normally present in
O. moschatus (7).
Other characters for the genus are noted in Semken, Miller & Stevens (17),
Harington (2) and Kurten & Anderson (7). McDonald (12) has examined additional
characters and suggests the possibility that Symbos may not be a monotypic genus
as it is usually considered. Semken, Miller & Stevens (17) and McDonald and Bartlett
(13) give detailed descriptions of the Symbos skeleton.
Symbos cavifrons was taller and more slender than the living muskox Ovibos
moschatus (2). Symbos is thought to have inhabited steppe grasslands or parklands
(2) as well as woodlands (6, 17), living in more temperate regions than the living muskox
which inhabits some tundra regions today (7). Symbos is known from late Irvingto-
nian through Rancholabrean times when the most recent Radiocarbon date available
is 10,370 ± 160 B.P. (7).
The Allen Township and presumably also the Union Township specimens appear
to have been recovered from sediments within or above the Packerton Moraine (16).
The Packerton Moraine was formed at the maximum advance of the Saginaw lobe
during the Cary Substage of the Wisconsinan glaciation (19, 20). Deposition of the
remains would have occurred in post-Cary times some 13,000-14,000 years ago (18,
19). The Allen Township specimen appears to have been deposited in the peat layers
that would have resulted from the natural infilling of a kettle lake which was left
behind the retreating Wisconsinan ice front.
We wish to thank Jerry N. McDonald, Department of Geography, Radford Univer-
sity, Radford, Virginia for reviewing this manuscript and for other information supplied
during the study. Patrick J. Munson, Department of Anthropology, Indiana Universi-
ty, Bloomington, kindly provided unpublished information on seven muskox skulls
under his study. Dave Rieger produced Figure 1, and Deborah Randolph, Indiana State
Museum, typed the manuscript.
Literature Cited
1. Bradley, F.H. 1870. Geology of Kankakee and Iroquois Counties. Geol. Survey
Illinois, Vol. 4, p. 229.
2. Harington, C.R. 1975. Pleistocene Muskoxen (Symbos) from Alberta and British
Columbia. Canad. J. Earth. Sci. 12(6):903-19.
3. Hay. O.P. 1912. The Pleistocene Period and its vertebrata. 36th Ann. Rept. In-
diana Dept. Geol. Natur. Res., p. 541-784.
4. Hay, O.P. 1923. The Pleistocene of North America and its vertebrated animals
from the states east of the Mississippi River and from the Canadian province
east of longitude 95°. Publ. Cam. Inst. Wash. 322:1-532.
5. Hibbard, C.W. and F.J. Hinds. 1960. A radiocarbon date for a Woodland Muskox
in Michigan. Paps. Michigan Acad. Sci. Arts, and Letters, Vol. XLV: 103-108.
6. Kitts, D.B. 1953. A Pleistocene Musk-Ox from New York and the distribution
of the musk-oxen. Amer. Mus. Novitates, No. 1607, p. 1-8.
7. Kurten, B. and E. Anderson. 1980. Pleistocene mammals of North America.
Columbia University Press, New York. 443 p.
Zoology 671
8. Lyon, M.W., Jr. 1926. A specimen of the extinct musk-ox Symbos cavifrons (Leidy)
from North Liberty, Indiana. Proc. Indiana Acad. Sci., 35:321-324.
9. Lyon, M.W., Jr. 1931. A small collection of Pleistocene mammals from LaPorte
County, Indiana. Amer. Midi. Natur., 12:406-410.
10. Lyon, M.W., Jr. 1936. Mammals of Indiana. Amer. Midi. Natur., 17(1): 1-384.
11. Lyon, M.W., Jr. and F.T. Hall. 1937. Skull of Musk-Ox, Genus Symbos, from
Montgomery County, Indiana. Amer. Midi. Natur., 18:608-611.
12. McDonald, J.N. 1984. A record of Symbos (Artiodactyla; Bovidae) from Kauf-
man County, Texas (in press).
13. McDonald, J.N. and C.S. Bartlett, Jr. 1983. An associated muskox skeleton from
Saltville, Virginia. J. Vert. Paleontol. 2(4):453-470.
14. Rarick, W.D. and W.J. Wayne. 1969. The Wolcottville Skull. Outdoor Indiana,
vol. 34 (1):10-11.
15. Ray, C.E., D.L. Wills and J.C. Palmquist. 1968. Fossil muskoxen of Illinois.
Trans. Illinois State Acad. Sci., 61(3):282-292.
16. Schneider, A.F. and G.H. Johnson. 1967. Late Wisconsinan Glacial history of
the area around Lake Maxinkuckee. Proc. Indiana. Acad. Sci., 76:328-334.
17. Semken, H.A., B.B. Miller, and J.B. Stevens. 1964. Late Wisconsinan Woodland
Muskoxen in Association with pollen and invertebrates from Michigan. Jour.
Paleontol. 38(5):823-835.
18. Wayne, W.J. 1963. Pleistocene formations in Indiana. Indiana Dept. Conserv.,
Geol. Surv. Bull. No. 25. 85 p.
19. Wayne, W.J. 1966. Ice and Land: A review of the Tertiary and Pleistocene history
of Indiana, pp. 21-39, in: E. Lindsay (ed), Natural Features of Indiana, Indiana
Acad. Sci., Indianapolis.
20. Wayne, W.J. and J.H. Zumberge. 1965. Pleistocene geology of Indiana and
Michigan, p. 63-83, in: H.E. Wright, Jr. and D.G. Frey (eds). The Quaternary
of the United States. Princeton Univ. Press. Princeton, N. Jersey.
21. Wilson, R.L. 1967. The Pleistocene vertebrates of Michigan. Paps. Michigan Acad.
Sci., Arts and Letters, Vol. LII: 197-234.
Survey of the Fishes of the Kingsbury State Fish and
Wildlife Area, LaPorte County, Indiana
David M. Sever
Department of Biology, Saint Mary's College
Notre Dame, Indiana 46556
and
Douglas Duff
Department of Biology, Indiana University
South Bend, Indiana 46634
Introduction
The Kankakee River is the northernmost tributary of the Mississippi River in
Indiana. The source of the Kankakee is just west of South Bend, Indiana, and the
river flows southwesterly until joining the Illinois River in eastern Illinois. Major Indiana
tributaries of the Kankakee River are the Yellow and Iroquois Rivers. In total, the
Kankakee River and its major and minor tributaries drain nearly 8100 sq km of nor-
thwest Indiana (3). The basin has been extensively drained, and most of the original
channel of the Kankakee River has been replaced by a series of ditches. The land
bordering the Kankakee River is largely agricultural. One area where the channel and
the surrounding land have been least modified is the Kingsbury State Fish and Wildlife
Area (KSFWA), a tract of 1820 ha in LaPorte County, Indiana. The northern shore
of the Kankakee River forms the KSFWA' s southeastern border. The KSFWA is a
natural swampy lowland where construction of drainage ditches by the Indiana Depart-
ment of Natural Resources is ongoing to accommodate further development of the
area for deer and rabbit management.
The main purpose of this study was to make regular collections of fish from
three different stream habitats in the KSFWA to see if there were differences in species
diversity, The habitats chosen were a relatively unmodified stream, an old ditch, and
a new drainage ditch. Another purpose of this study was to compare the types of
fish we collected to species records compiled for the Kankakee River basin by earlier
workers. Previous reports on fish on the Kankakee River basin include reports by Hay
(2), Robertson and Ledet (3) as well as records listed in the state reports by Gerking
(1) for Indiana and Smith (4) for Illinois. None of these reports contain any records
specifically from the KSFWA.
Methods
Samples were taken by seine on a monthly basis from March through November,
1983, inclusive. Dates were: 18 March; 22-23 April; 12 May; 8 June; 13 July; 15 August;
9 September; 14 October; and 18 November.
Three localities were chosen. Locality 1 (at KSFWA parking lot 4B) was a relatively
undisturbed tree-lined stream passing through cultivated fields and brushland. The stream
bed was sandy-bottomed, and current was relatively swift. Maximum depth was 1.5
m (although mid-channel depth averaged about 0.5 m), and width varied from 3-4.5
m. Locality 2 was a section of Breckenridge Ditch from 1-1.5 km upstream of its
confluence with the Kankakee River. This ditch was clogged with tree limbs and aquatic
vegetation. The sides of its channel were extremely silty, but the middle of the channel
was sandy. Average depth at the middle of the channel was about 0.8 m, and maximum
depth of pools was 1.8 m. Width of the stream was 4-6 m, and current was moderately
swift . The stream for locality 1 is a minor tributary of Breckenridge Ditch about
673
674 Indiana Academy of Science Vol. 94 (1985)
2 km from the collecting site for locality 2. Locality 3 was a newly constructed ditch
parallel to and directly across a dirt road (River Road) from locality 2. This ditch
was constructed to drain a large swampy wetland. Flow was controlled by a gate operated
by the DNR at parking lot 5F. The ditch was about 3.5-4 m wide and varied from
1.8-2.3 m maximum depth above the gate at all seasons. Below the gate, average depth
of the channel was nearly 1 m in the spring, but only pools (maximum depth 0.7 m)
connected with shallow (> .10 m) rivulets existed in the summer and fall. The bottom
of the entire channel was extremely silty, there was little aquatic or emergent vegeta-
tion, and, depending upon water level, flow was moderately swift. In the spring months
(March-June), we could collect from the source of the ditch to its confluence with the
Kankakee River, a distance of about 1 km. In the remaining months, however, our
collections from locality 3 were limited to pools near KSFWA parking lot 5F.
All collections were made between 1000-1500 hrs, and average time for a collec-
tion was about 90 min. About 0.5 km usually was sampled per collection except as
noted above for locality 3. Specimens were preserved upon collection in 10% formalin.
After sorting and identification, specimens were placed in 60% isoporpanol. Specimens
are currently stored at Saint Mary's College.
Results and Discussion
Analysis of Collections
From our samples, we preserved a total of 2849 individuals from 33 species
representing 13 families (Tables 1-3). All specimens were saved except for some Cyprinus
carpio, Catostomous commersoni, Notemigonus crysoleucas, Ictalurus melas, and /.
nebulosus. Specifically, we saved 663 individuals of 12 species from locality 1, 552
individuals of 28 species from locality 2, and 1 634 individuals of 25 species from locality 3 .
Only eight species were found in all three localities. These were: Esox americanus,
C. carpio, C. commersoni, Erimyzon sucetta, I. melas, Lepomis cyanellus, L. gibbosus
and Perca flavescens. Only single individuals of E. sucetta and L. gibbosus were ob-
tained, however, from locality 1, and only one C. commersoni came from locality 3.
Each locality had at least one species not found at other localities, but such species
were represented by single specimens except in two instances. Nocomis biguttatus was
collected only at locality 1 (April — 1). Moxostoma macrolepidotum (August — 1),
Table 1. Fish preserved from locality 1. An asterisk (*) indicates that not all of the
fish collected were preserved.
Species MONTH TOTAL
3 4 5 6 7 8 9 10 11
Esox americanus 1 4 4 1 3 13
Cyprinus carpio
Nocomis biguttatus
Rhinichthys atratulus
Catostomous commersoni
Erimyzon sucetta
Ictalurus melas
Cot t us bairdi 109
Lepomis cyanellus
Lepomis gibbosus
Perca flavescens
Etheostoma nigrum 32
2*
2
4
1
1
3
2
12
19
6
4
1
2*
1
8
3
2
3*
11
26
2
3
34
35
36
12
32
65
47
48
408
3
1
1
2
1
1
2
2
10
1
2
10
17
7
10
27
28
16
27
174
Zoology
675
Table 2. Fish preserved from locality 2. An asterisk (*) indicates that not all of the
fish collected were preserved.
Species
MONTH
TOTAL
3
4
5
6
7
8
9
10
11
Amia calva
2
1
3
Umbra limi
1
1
Esox americanus
2
1
7
6
9
3
5
33
Esox lucius
3
2
1
2
8
Cyprinus carpio
1
1
3
3*
7
Notemigonus crysoleucas
1
4
4
18
5
56
14
7
9
118
Semotilus atromaeulatus
4
1
- '
5
Rhinichthys atratulus
1
1
Ictiobus bubalus
1
1
Moxostoma macrolepidotum
1
1
Catostomous commersoni
1
5
6
4
3*
6
7
1
1
34
Minytrema melanops
1
1
2
4
Erimyzon sucetta
1
1
2
2
6
Ictalurus melas
5
18
7
12*
1
7
9
10
69
Ictalurus natalis
1
2
4
8*
3
2
5
5
30
Ictalurus nebulosus
2
7
3
6
3
6
2
29
Noturus gyrinus
1
1
2
Apredoderus sayanus
1
2
1
4
Fundulus dispar
2
2
Cottus bairdi
3
1
4
Micropterus salmoides
2
2
Lepomis cyan el I us
5
1
1
1
15
2
6
26
57
Lepomis gibbosus
2
1
4
2
5
2
16
Lepomis macrochirus
1
1
6
40
4
3
5
60
Pomoxis nigromaculatus
1
1
Perca flavescens
1
7
7
16
Percina maculata
1
1
Etheostoma nigrum
1
3
12
9
2
6
4
37
Minytrema melanops (June through August — 4), Fundulus dispar (April — 2), and Per-
cina maculata (June — 1) were found only at locality 2. Four species found only at
locality 3 were each represented by single specimens from the May or June collections:
Dorosoma cepedianum, Notropis spilopterus, Pimephales notatus, and Labidesthes sic-
culus. Water levels in the ditch at locality 3 were high during May and June, providing
a continuous channel at least 1 m deep from the gate at parking lot 5F to the Kankakee
River.
Three species were found in localities i and 2, but not locality 3. These were,
with numbers of specimens (locality l:locality 2) in parentheses: Rhinichthys -atratulus
(19:1), Cottus bairdi (408:4), and Etheostoma nigrum (174:37). There were no species
common to localities 1 and 3, but not locality 2.
There were, however, 13 species found at localities 2 and 3 but not locality 1.
These were, with number of specimens (locality 2:locality 3) in parenthesis: Amia calva
(3:2), Umbra limi (1:24), Esox lucius (8:2), N. crysoleucas (118:950), Semotilus
atromaeulatus (5:3), Ictiobus bubalus (1:6), Ictalurus natalis (30:1), /. nebulosus (29:66),
Noturus gyrinus (2:1), Apredoderus sayanus (4: 1), Micropterus salmoides (2:30), Lepomis
machrochirus (60:153), and Pomoxis nigromaculatus (1:33).
The dominant fish at locality 1 were Cottus bairdi and Etheostoma nigrum, the
only species collected each month and in any numbers. Only 2-6 species were collected
in any month at locality 1 except for July (8) and August (10). The limited fish fauna
locality 1 is probably representative of the local natural diversity in a shallow, relatively
676
Indiana Academy of Science
Vol. 94 (1985)
Table 3. Fish preserved from locality 3. An asterisk (*) indicates that not all of the
fish collected were preserved.
Species
MONTH
TOTAL
3
4
5
6
7
8
9
10
11
Amia calva
1
1
2
Dorosoma cepedianum
1
1
Umbra limi
8
11
4
1
24
Esox americanus
13
1
1
2
1
18
Esox lucius
1
1
2
Cyprinus carpio
57*
16*
16
9
21
3
3
1
6
132
Notemigonus crysoleucas
797
14
83
12
37
7
950
Semotilus atromaculatus
3
3
Notropis spilopterus
1
1
Pimephales notatus
1
1
Ictiobus bubalus
6
6
Catostomous commersoni
1
1
Erimyzon sucetta
1
2
1
12
1
2
2
21
Ictalurus melas
2
5
31
16
12
9
18
5
10
108
Ictalurus natalis
1
1
Ictalurus nebulosus
1
6
9
4
14
13
10
3
6
66
Noturus gyrinus
1
1
Apredoderus sayanus
1
1
Labidesthes sicculus
1
1
Micropterus salmoides
4
1
1
2
2
3
2
5
10
30
Lepomis cyanelius
4
4
3
11
Lepomis gibbosus
19
5
1
13
7
4
0
3
58
Lepomis macrochirus
58
33
33
3
4
9
8
153
Perca flavescens
3
4
2
9
Pomoxis nigroma culatus
16
1
3
7
6
33
swiftly flowing, sandy-bottomed stream. There are a few Lepomis, and some Esox
americanus, Cyprinus carpio, and Catostomous commersoni moved into the area during
the summer, but usually the stream is dominated by two small carnivores, one benthic
(C. bairdi) and the other nektonic (E. nigrum).
The dominant fish at locality 2 were: eoscids (Esox americanus and E. lucius),
the cyprinid Notemigonus crysoleucas, catastomids (especially Catostomous commer-
soni), ictalurids (especially Ictalurus melas), centrarchids (especially Lepomis cyanelius
and L. macrochirus) and percids (Perca flavescens and Etheostoma nigrum). Species
diversity was rather constant throughout the year at locality 2 with 11-15 species col-
lected each month except March (8 species).
Species composition of localities 2 and 3 were more similar to each other than
either was to locality 1. Certain habitat similarities between localities 2 and 3, therefore,
must have been more important to species diversity than the fact that localities 1 and
2 are directly connected while locality 3 is separate from the other two streams. The
most important similarity between localities 2 and 3 is probably closeness to the river.
The collecting site at locality 2 was only 1-1.5 km upstream from its confluence with
the Kankakee River, and at locality 3, we were able to collect from the gate at parking
lot 5F to the Kankakee River during high water periods. The fish fauna of both localities
2 and 3, therefore, were more directly influenced by movement of fish from the main
channel of the river than the fish community at locality 1, 2 km upstream of locality 2.
Although species composition was similar between localities 2 and 3, there were
some differences in the dominant forms. Umbra limi was abundant in the spring at
Zoology 677
locality 3 and absent at this time at locality 3. C. carpio was a more abundant species
at locality 3 than locality 2, especially in the March-July collections while water levels
were high at locality 3. Suckers generally were not as dominant at locality 3 as at
locality 2. C. commersoni was represented by only a single specimen at locality 3,
and Minytrema melanops and Moxostoma macrolepidotum were absent at locality 3.
Ictiobus bubalus and Erimyzon sucetta, however, were more abundant at locality 3
than locality 2. Among ictalurids, Ictalurus melas was a dominant form at both locales,
but /. natalis was represented by just one specimen at locality 3 while it was frequently
collected at locality 2. Among centrarchids, L. cyanellus was less common and Micropterus
salmoides was more common at locality 3 than locality 2, and Pomoxis nigromaculatus
was represented by 33 specimens at locality 3, but by only one specimen at locality 2.
During the high water periods (March-July), diversity was similar at locality 3
(12-15 species) to that previously noted for locality 2. Since the diversity and species
groups involved were similar for localities 2 and 3, why were there such differences,
as noted above, in the dominant forms between the two streams? Locality 2 was the
older ditch. It had a good growth of aquatic and emergent vegetation, numerous snags
and fallen limbs on the bottom, and a relatively constant water depth — seasonal fluc-
tuations were >0.5 m. The passing through or stranding of fish, especially larger esocids
and catastomids, resulted in the minor, but noticeable changes in seasonal diversity
at locality 2. We believe, therefore, that our collections from locality 2 represent natural
associations of fish resulting from their residence in or seasonal movements through
the collecting area.
Locality 3 was freshly dug and had little aquatic or shore-line vegetation. With
the sluice gate at parking lot 5F controlling the draining of the swamp upstream, cur-
rent and water-level were dependent upon manipulation by the DNR of the situation.
These factors, coupled with the heavy siltation, made locality 3 seemingly poor perma-
nent fish habitat. When the sluice was opened during high water in spring, a greater
diversity and number of species were taken. The May collection yielded 214 individuals
of 15 species. The large numbers of C. carpio and N. crysoleucas collected in spring
may simply indicate an influx of individuals from the swamp or river during periods
of high water. When the sluice was closed in July, water depth above the gate became
too deep for seining, while below the gate, the stream generally dired over the summer
and fall months into a series of pools connected by rivulets accessible only to very
small fish. Only 31 individuals from 7 species were collected from these pools in October.
Most likely the larger individuals of species collected from these pools in summer months,
such as some Amia calva and M. salmoides, were trapped when decreasing water levels
made escape to the river or the swamp impossible.
The composition of the collections from locality 3, therefore, cannot be considered
representative of a stable fish community. The differences in dominant species bet-
ween localities 2 and 3 is likely the result of the chance movement or stranding of
individuals at locality 3 due to shifting water levels. Individuals from locality 3 were
collected while in a connecting link of rather inhospitable habitat during movements
either from the swamp or the river.
Relation to Previous Records
The only species collected not previously recorded by Gerking (1) or Smith (4)
from any portion of the Kankakee River drainage is Ictiobus bubalus. Based on Gerk-
ing (1), four species were recorded for the first time from Indiana portions of the
Kankakee drainage: Dorsoma cepedianum, M. macrolepidotum, E. lucius, and M.
salmoides. All of these species, however, have been recorded from Illinois portions
of the Kankakee system, and their presence in our samples is not surprising. Eleven
species had been recorded from other portions of the Kankakee River in Indiana by
678 Indiana Academy of Science Vol. 94 (1985)
Gerking (1), but not previously from LaPorte County. These new LaPorte County
records are: A. calva, E. sucetta, M. melanops, C. carpio, I. melas, I. nebulosus,
I. natal is, Fundulus dispar, L. macrochirus, Pomoxis nigromaculatus, and Labidesthes
sicculus. Again, the occurrence of these forms in our samples could be expected.
Some 53 species have been reported by Gerking (1) and Smith (4) from the
Kankakee River drainage but were not collected at KSFWA in our study (Table 4).
In addition, a fishery survey by Robertson and Ledet (3) on the Kankakee River revealed
three species, Anguilla rostrata, Ictiobus niger and Carpiodes velifer, not recorded in
the system by Gerking (1), Smith (4), or the present study. Our collections reveal,
therefore, that the streams sampled at KSFWA contain >40% of the total number
of species known from the Kankakee River system. Also, there are some other species
of probable occurrence in the Kankakee River basin that have not been recorded as
of yet in the scientific literature. Such species include, for example, Ichthyomyzon
castaneus, Lepiosteus oculatus, Nocomis micropogon, Notropis atherinoides, Pylodic-
tis olivaris, and Etheostoma blennoides, among others known from connecting river
basins. Confirmation of their occurrence in the Kankakee River system awaits further
collecting.
The Kankakee River system thus has a recorded fish diversity of about 90 species
with a potential for probably a dozen more. The absence of many of these species
from the streams sampled at KSFWA can most obviously be related to lack of ade-
quate habitat. The complete absence of many cyprinid genera and the representation
of Notropis by only one specimen of N. spilopterus, however, is clearly quite striking.
We do not believe that the lack of these forms is an artifact of collection, but that
it does, indeed, represent the actual situation. It would seem an over-simplification
to blame the absence of so many characteristic stream fish of northwestern Indiana
(such as N. cornutus or N. texanus) on inadequate habitat, but we can offer no fur-
ther explanation at this time. It will be interesting to sample other small tributaries
of the Kankakee River in nearby areas to see if their minnow fauna is as depauperate
in species diversity as that of the streams we sampled at KSFWA.
Acknowledgments
We thank R. Haney of the KSFWA for his permission to conduct this study.
The collections were supported by a grant from the Indiana Academy of Science. We
thank the following persons for aid in making one or more collections: D. Deery,
K. Higgs, K. Hoban, J. Litton, E. Mould, C. Patricoski, P. Patricoski, and L. Weber.
Literature Cited
1. Gerking, S.D. 1945. The distribution of the fishes of Indiana. Ivest. Ind. Lakes
Streams 3:1-137.
2. Hay, O.P. 1896. On some collections of fishes made in the Kankakee and Illinois
Rivers. Field Columbian Museum Zool. Ser. Publ. 12, Vol. l(4):85-97.
3. Robertson, B. and N. Ledet. 1981. A fisheries survey of the Kankakee River
in Indiana. Fisheries Sec, Ind. Depart. Nat. Res., Indianapolis. 58 pp.
4. Smith, P.W. 1979. The fishes of Illinois. Univ. Illinois Press, Urbana. 314 pp.
Zoology 679
Table 4. Species reported from the Kankakee River drainage from LaPorte County,
other Indiana Counties, and from Illinois but not represented in the Kingsbury State
Fish and Wildlife Area Collections. Indiana records from Gerking (1949), and Illinois
records from Smith (1979).
LaPorte Other
Species County Indiana Illinois
Ichthymoyzon fossor X
Ichthyomyzon unicuspis X
Lampetra appendix X
Lepiosteous osseus X
Salmo gairdneri X
Salmo trutta X X
Hybopsis amblops X
Phenacobius mirabilis X X
Notropis chalybaeus X X
Notropis cornutus XXX
Notropis dorsalis X
Notropis emiliae X
Notropis heterodon X X
Notropis heterolepis X X
Notropis hudsonius X X
Notropis lutrensis X
Notropis rubellus X X
Notropis stramineus XXX
Notropis texanus XXX
Notropis umbratilus XXX
Notropis volucellus XXX
Ericymba buccata XXX
Phoxinus erythrogaster X
Pimephales vigilax X X
Campostoma anomalum XXX
Ictiobus cyprinellus X
Capriodes cyprinus X X
Moxostoma anisurum X
Moxostoma carinatum X
Moxostoma duquesnei X X
Moxostoma erythrurum X X
Hypentilium nigricans X X
Erimyzon oblongus X X
Ictalurus punctatus X X
Noturus flavus X X
Fundulus diaphanus X
Fundulus notatus XXX
Morone chrysops X
Micropterus dolomieui X X
Lepomis gulosus X X
Lepomis humilis X X
Lepomis megalotis XXX
Lepomis microlophus X
Lepomis punctatus X
Ambloplites rupestris XXX
Pomoxis annularis X
Stizostedion vitreum X
Percina caprodes X X
Percina phoxocephala X X
Etheostoma caeruleum X X
Etheostoma microperca XXX
Etheostoma spectabile X
Etheostoma zonale X
Heat Loss from Avian Integument:
Effects of Posture and the Plumage
Marcus D. Webster
Department of Biology
Franklin College
Franklin, Indiana 46131
Introduction
Heat loss from the body surface of birds is reduced through physiological
adjustments of heat flow to the skin surface and by the presence of an insulating feather
coat. Maintenance of a high and stable body core temperature is made possible by
high avian metabolic rates coupled with behavioral, physiological, and morphological
adaptations to reduce heat loss in the cold and to increase heat dissipation when air
temperature is high. Previous studies of avian thermoregulation have provided a strong
theoretical and empirical framework for studies of energy expenditure in birds, but
relatively few investigators have focused on the role of the plumage in impeding heat
loss through the integument (5).
The fibrous coat of mammals and birds, like human clothing, provides a barrier
to heat flux by trapping a layer of still air next to the skin. The effectiveness of the
insulating coat can be reduced by wind penetration or by free convection due to
temperature gradients within the coat (4). Cutaneous tissue beneath the coat also pro-
vides some insulation, depending on the amount of bloodflow (hence heat flux) to
different body regions. The relative importance of these and other factors in heat transfer
relations of birds can be evaluated using relatively simple biophysical models that in-
corporate measurements of animal and environmental variables (3, 5, 9).
Sensible heat flux at the surface of an animal may be treated, as a first approx-
imation, as a linear diffusion process in which radiative and convective heat losses
occur across a thermal resistance and down a temperature gradient. Thermal resistance
for a whole animal is given by:
rhb = [ PCP (Tb - Ta>l / <M - XE>' <*>
where rrh is in s/m, PC is the volumetric heat capacity of air (1200 J m~ 3 K~ '), Tb
is the body core temperature (C), Ta is air temperature (C), M is metabolic rate in W/m2,
and XE is the total evaporative heat flux (W/m2) [3, 5, 8, 9].
The whole-body thermal resistance (Eq. 1) can be illustrated and partitioned into
tissue, coat, and boundary layer components using electrical circuit analogues (8, 9).
Temperature differences between points in the animal-environment interface are treated
as voltages and heat fluxes across the interface as current sources. The plumage,
boundary layer, and subcutaneous tissues form three series resistances to heat flow,
which can be quantified using equivalent circuit analysis (3, 8, 9).
Materials and Methods
Five male and five female pigeons (472.9 ± 12.4 g body weight) were obtained
from a local breeder and acclimated to standard environmental conditions prior to
experimentation. The birds were maintained under a 12:12 light-dark cycle, at 20C
and 40-50% relative humidity, and were fed and watered ad lib. Pigeons were selected
for docility and acceptance of training to tolerate experimental restraint.
681
682 Indiana Academy of Science Vol. 94 (1985)
To separate cutaneous from respiratory evaporative water loss I used a two com-
partment metabolic chamber constructed of plexiglass and placed in a temperature-
controlled cabinet. Air dried to less than 0.3 g/m3 water vapor density by columns
of Drierite was pumped into each compartment at 1.5-1.8 1/min, and led to a dew-
point hygrometer (EG&G 660) for humidity determination and to an oxygen analyzer
(Electrochem N-3) for measurement of oxygen consumption. Skin temperatures were
measured with fine wire Cu-Cn thermocouples closely affixed to the skin and glued
(methyl-cyanoacrylate adhesive) to the base of a contour feather to hold them in place.
Temperature transmitters (Minimitter Model M) surgically implanted in the birds allowed
continuous monitoring of intraperitoneal body temperature. Humidity in the chamber
was determined with a Weathertronics 5118 electrical hygrometer, and compartment
temperatures were measured with thermocouples and controlled to within ± 0.2 C.
Further details of the experimental apparatus are provided in (12).
Each of the pigeons used in this study was trained (for 20-25 h) to stand quietly
in the chamber, with its head protruding through a latex collar into the head compart-
ment and supported by a plexiglass pillory. Each experiment was conducted in darkness,
during the subjective night of the birds, on resting and postabsorbative animals. No
measurements were taken until the 95% equilibration time for body compartment water
vapor had expired (1 1/2 hrs.), and efflux air oxygen content and body temperature
remained stable for a minimum of five min before a measurement was taken. Data
from runs in which fecal water contaminated the chamber air were discarded.
Determinations were performed at six temperatures over the range of 0 C to 40
C. Each bird was used at each air temperature, and the number of replicates ranged
from 23 to 36 measurements at each chamber temperature. Statistical analyses relied
on stepwise comparisons between temperature groups using analysis of variance.
Thermal resistances to heat loss were computed using the methods of (9). Boun-
dary layer resistance was taken as the sum of free and forced convective resistances,
since windspeed in the chamber was low and constant (0.1-0.15 m/s) at all air
temperatures (13). For computing boundary layer resistance, the characteristic dimen-
sion of the birds was estimated as 10 cm, the approximate diameter of the pigeons sitting
parallel to chamber airflow.
Results
Body temperature remained approximately stable with increasing air temperature,
averaging 40.6 ± 0.1 C at 0, 10, and 20 C, but increased significantly (ANOVA, p
< 0.05) at 35 C air temperature, to 41.0 ± 0.1 C, and at 40 C increased (p < 0.05)
to 41.6 ± 0.15 C. Skin temperature was linearly related to chamber air temperature,
increasing from 22.6 ± 0.5 C at 0 C by 4.2 ± 0.5 C each 10 C increment until reaching
values near core temperature. At 40 C, skin temperature (40.9 ± 0.2 C) was not
significantly different from body temperature (12, 13).
Whole-body thermal resistance (rhb) was constant between 0 and 35 C air
temperature at 335 + /- 30 s/m (Figure 1). At 40 C, rhb was reduced to 192 + /-
22 s/m (p < 0.05). Comparison of these estimates to pigeon whole-body resistances
computed from available data (3) reveals that my restrained pigeons were able to in-
crease rhb to levels that were only 50-60% of those measured in unrestrained pigeons at
0 to 10 C.
Tissue thermal resistance was 140 +/- 14 s/m at 0, 10 and 20 C, but declined
significantly to 105 + /- 3.2 s/m at 30 and 35 C, and dropped to 76.5 + /- 1.8
s/m at 40 C (Figure 2). Feather coat thermal resistance did not change significantly
from 0-35 C, and averaged 139 + /- 25 s/m over that temperature range. At 40 C,
however, a sharp decrease in plumage thermal resistance was noted, to 30.5 + /-
Zoology
683
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o
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Calder & Schmidt -
Nielsen 1967
rhb in Restrained
and Unrestrained Pigeons
10
20 30
Tair(°C)
40
Figure 1 . Whole body thermal resistance (mean + / - SE) of restrained and unre-
strained pigeons as a function of chamber air temperatures. Data for the lower points
were obtained in this study; the upper curve was calculated from data reported in (2).
T400+
(0
a,30Q
o
c
*200
Q)
DC
1 100
Plumage Resistance rhc y
* * + t~,.\
Tissue Resistance rM si
0
10
20
Tajr(°C)
30
40
Figure 2. Tissue (rh ) and plumage (rhc) thermal resistance (mean + / - SE) in pigeons
at different chamber air temperatures. No significant differences were detected between
0 and 20 C. Tissue thermal resistance decreased significantly at 30 and 35 C (p <
0.05, ANOVA) and both tissue and plumage resistance decreased significantly (p <
0.05) at 40 C.
684
Indiana Academy of Science
Vol. 94 (1985)
5.3 s/m. Over the entire range of temperatures used, boundary layer thermal resistance
was 90 s/m.
Discussion
Comparison of my results on the temperature dependence of whole-body thermal
resistance of experimentally restrained pigeons to those of Calder and Schmidt-Nielson
(2) in unrestrained pigeons indicates that in pigeons free to adjust their posture in
the cold, a progressive increase in body insulation to twice that of thermoneutral levels
is possible. Because these two data sets were collected under experimental conditions
similar in all respects except restraint of the animal, differences in body resistance
at 0 and 10 C most probably result from the inability of restrained pigeons to alter
their exposed surface area by changing posture. If so, pigeons are evidently able to
reduce their effective surface area to about 50% of thermoneutral zone levels by tuck-
ing the head, withdrawing the feet, fluffing the plumage, and bringing the wings in
close to the body. The hypothesis that postural adjustments are an important means
of increasing thermal resistance is sound, but further experimental testing is needed
before these results can be confirmed.
Thermal resistances of the subcutaneous tissues and the feather coat in pigeons
are approximately equal from 0-35 C under the experimental conditions of this study
(Figure 2). My estimates of tissue heat loss resistance are slightly higher than those
summarized by Campbell (3) for mammals, but a marked decrease in tissue thermal
resistance at high air temperatures is apparently common to mammals and birds and
almost certainly results from increased cutaneous bloodflow.
Estimates of avian plumage thermal insulation have been made for excised pat-
ches of feathers using heat flux plates (1 1), on live birds by measuring plumage surface
temperatures with radiometers (6, 10), by removing the feathers and measuring changes
in metabolic rate (1), and by measuring or estimating important animal and environmen-
tal variables to solve heat balance equations as in this study and (9). Excised plumage
measurements may not accurately estimate coat thermal resistance in nature, although
such data can be useful for comparison of the insulative value of different animal
coats. Whole body measurements like those reported here reflect differences in coat
thickness over the body and incorporate heat loss from unfeathered regions.
Feather coat resistances of several species of birds are given in Table 1 . For com-
parison, the thermal resistance of a 1 cm thick layer of still air is 480 s/m (3), and
the resistance of heavy winter coats worn by humans is about 400 s/m (5). Estimates
Table 1. Thermal resistance of avian plumages. Resistances in the third column are
whole-body estimates; those in the fourth column are excised plumage measurements
or whole-body data divided by estimated average coat depth.
Coat
Coat Resistance
Species
Tair(C>
Resistance (s/m)
per cm depth (s/m)
Source
Goose (down)
227
(5)
California Quail
20
282
—
(1)
Pigeon
—
—
176
(11)
Gray Jay
-8
465
232
(10)
Black -capped Chickadee
10
427
214
(6)
White-crowned Sparrow
20
400
240
(9)
Zoology 685
of the thermal resistance of avian plumage per cm depth range from 176 s/m for pigeon
breast plumage (11) to 240 s/m in the White-crowned Sparrow (9). Feather coats are
evidently only 37-50% as effective as still air in impeding the loss of heat by convec-
tion and radiation from the skin surface. All available data indicate that avian plumages
and mammalian pelage have thermal resistance values that are approximately half those
of an equivalent depth of still air (5).
The sharp reduction of coat resistance in pigeons at high air temperatures ap-
parent in Figure 2 may result from sleeking of the plumage, from alternate erection
and depression of the plumage improving convective heat loss, or from free convec-
tion currents arising in the feathers due to the high temperature of the skin surface
(4, 14). Such a reduction in coat resistance coupled with lower tissue thermal resistance
could play an important role in increasing heat dissipation in heat stressed birds.
Conclusions
1) Whole-body thermal resistances at 0 and 10°C of the restrained pigeons used in
this study differed significantly from those measured in unrestrained pigeons under
identical conditions (2). These differences may reflect the inability of restrained birds
to adjust their posture and thus reduce exposed surface area. Postural adjustments
may therefore account for 40-50% of the increased thermal resistance of birds in cold
versus thermoneutral air temperatures.
2) Thermal resistances of the tissue and the plumage coat in pigeons are reduced during
mild heat stress, probably due to cutaneous vasodilation and ptilomotor adjustments
coupled with increased free convection within the plumage.
Acknowledgments
This study was supported by grants from the National Science Foundation (DEB
79-09806 and DEB 81-141590) to James R. King. A National Institute of Health predoc-
toral traineeship (GM-01276) and Washington State University E.O. Holland Fellowship
provided additional financial support during the course of the study. For helpful discus-
sions which improved the analysis, I thank G.S. Campbell and D.R. Webb. An
anonymous reviewer's perceptive comments are acknowledged gratefully.
Literature Cited
1. Brush, A. 1965. Energetics, temperature regulation and circulation in resting,
active and defeathered California quail, Lophortyx calif ornicus. Comp. Biochem.
Physiol. 15:399-421.
2. Calder, W.A. and K. Schmidt-Nielsen. 1967. Temperature regulation and
evaporation in the pigeon and Roadrunner. Amer. J. Physiol. 213:883-889.
3. Campbell, G.S. 1977. An introduction to environmental biophysics. Springer-
Verlag, New York. 159 pp.
4. Campbell, G.S., A.J. McArthur, and J.L. Monteith. 1980. Windspeed
dependence of heat and mass transfer through coats and clothing. Bound. Layer
Meterol. 18:485-493.
5. Cena, K. and J. A. Clark. 1979. Transfer of heat through animal coats and
clothing. Pp. 1-42 in D. Robertshaw, ed., International Review of Physiology,
Vol. 20. Environmental Physiology III. University Park Press, Baltimore.
6. Hill, R.W., D.L. Beaver and J.H. Veghte. 1980. Body surface temperatures
and thermoregulation in the Black-capped Chickadee (Pants atricapillus). Physiol.
Zool. 53:305-321.
7. Kaufman, W.C., D. Bothe and S.C. Meyer. 1982. Thermal insulating capabilities
of outdoor clothing materials. Science 215:690-691.
686 Indiana Academy of Science Vol. 94 (1985)
8. Mc Arthur, A.J. 1981. Thermal resistance and sensible heat loss from animals.
J. Therm. Biol. 6:43-47.
9. Robinson, D.E., G.S. Campbell, and J.R. King. 1976. An evaluation of heat
exchange in small birds. J. Comp. Physiol. 105:153-166.
10. Veghte, J.H. and C.F. Herreid. 1965. Radiometric determination of feather
INSULATION AND METABOLISM OF ARCTIC BIRDS. PHYSIOL. ZOOL. 38:267-275.
11. Walsberg, G.E., G.S. Campbell and J.R. King. 1978. Animal coat color and
radiative heat gain: a re-evaluation. J. Comp. Physiol. 126:211-222.
12. Webster, M.D. 1983. Temperature and humidity dynamics of cutaneous and
respiratory evaporation in the pigeon, Columba livia. Ph.D. dissertation,
Washington St. Univ.
13. Webster, M.D., G.S. Campbell and J.R. King. 1985. Resistance to cutaneous
water-vapor diffusion in pigeons and the role of the plumage. Physiol. Zool.
58:58-70.
The Freshwater Naiads, Bivalvia: Unionidae, of the Blue River, a Southern
Indiana Tributary of the Ohio River
Charles Weilbaker, Claude D. Baker, Bill J. Forsyth
and Carl M. Christenson
Department of Biology
Indiana University Southeast
New Albany, Indiana 47150
and
Ralph W. Taylor
Department of Biological Sciences
Marshall University
Huntington, West Virginia 25701
Introduction
Freshwater bivalve mussels of the family Unionidae have been in existence since
early Mesozoic times in the lakes and rivers of North America (2). In the last century
to the present time, these naiads have been exploited first by the pearl button industry
and more recently by the Japanese cultured pearl industry. This commercial exploita-
tion along with impoundments, clear-cutting, siltation, and pollution has resulted in
decreased bivalve diversity in many streams. Since many bivalve species are extinct
or virtually so, most environmental biologists feel that an immediate accounting is
essential if we are to preserve the remaining species.
Because of channelization and impounding of water, current naiad faunas of our
larger rivers are different from those present at the turn of the century (3). Some smaller
streams, however, may not have been so severely altered. Thus, a logical conservation
approach would be to concentrate on protecting smaller streams which may be serving
as refugia for rare and endangered species.
In Indiana, most published work dealing with unionids is concerned with the naiad
faunas of the larger rivers (1, 4). Very little is known about the molluscan faunas
in the smaller streams and rivers, especially those located in southern Indiana. The
purpose of this study was to inventory the freshwater bivalves inhabiting the Blue River,
a southern Indiana tributary of the Ohio River.
Description of Study Area
Situated within unglaciated middle and upper Mississippian bedrock of extreme
south central Indiana, the Blue River is unique because the main stream has few sur-
face tributaries (Figure 1). The watershed of the Crawford Upland and Mitchell Plain
physiographic regions is characterized by numerous sinkholes. Areas of sinkhole plains
collect the available surface water into subterranean systems which eventually re-emerge
near the Blue River as karst springs. The largest karst spring in the area is Harrison
Spring which empties into the Blue River near White Cloud (Figure 1). In addition,
solution of limestone over the years has produced a substantial number of cave systems.
Indiana's best known caves, Marengo Cave and Wyandotte Cave, are located within
the watershed.
The Blue River is a fourth order stream with typical alternating riffles and pools.
Most riffles have significant quantities of limestone boulders and rubble while the pools
have bottoms comprised of limestone boulders, rocks, gravel, sand and silt. Water
willow, Dianthera americana, commonly forms dense stands along many riffles during
the summer months. Riparian trees form a significant canopy with sycamore being
the most common streamside tree. In general, the stream water quality is relatively
687
688
Indiana Academy of Science
Vol. 94 (1985)
MILES
Figure 1. Map of the Blue River, a southern Indiana tributary of the Ohio River.
Bivalves were taken from almost every accessible location along the river. The area
from Wyandotte Cave to the Ohio River was collected via SCUBA and snorkeling.
Much of the remainder of the main stream was collecting during float trips. Collec-
tions were terminated at Salem on the West Fork and Pekin on the South Fork where
no specimens were found. Collections summarized in Table 1 included 9 collections
in the main stream from the mouth to Fredericksburg, 7 collections in the upper Blue
from Fredericksburg to Salem, and 9 collections in the South Fork from Fredericksburg
to Pekin.
Zoology
689
good with high oxygenation, but the water has exceptional clarity only in the fall during
periods of low discharge.
Methods
Bivalve specimens were taken from 25 stations using handpicking, snorkeling,
and SCUBA. Raccoon and muskrat middens were also reliable sources of fresh shells.
In addition, some shells were retrieved from the area of an old button factory located
near the mouth of the river. Normally, only fresh shells were retained. Weathered
or subfossil shells were noted and discarded. Fresh shells and living specimens were
transported to the laboratory, cleaned, and identified. Voucher specimens have been
placed in the museums at The Ohio State University and Marshall University. Dr.
David H. Stansbery of The Ohio State University Museum of Zoology verified the
identifications. A species list with number of specimens and locality descriptions for
each station has been placed on file with the Indiana Natural Heritage Program in
Indianapolis.
Results and Discussion
Thirty-seven species of unionids plus the exotic Asiatic clam, Corbicula, were
taken from the Blue River system (Table 1). Of these, 30 species were found in the
Table 1 . Summary of unionid mussels taken from the Blue River, a southern Indiana
tributary of the Ohio River. Based on 25 collections in 1984. A = abundant, C = com-
mon, R = rare, L = living specimen taken, WS = weathered shell, E = proposed en-
dangered by Indiana Department of Natural Resources, T = proposed threatened, SC
= proposed special concern.
Scientific name
(1)
Main Stream
Upper Blue
South Fork
Anodonta grandis
Strophitus undulatus
Alasmidonta marginata
Alasmidonta viridis
Simpsonaias ambigua
Lasmigona complanata
Lasmigona costata
Magnonaias nervosa
Tritogonia verrucosa
Quadrula quadrula
Quadrula metanevra
Quadrula pustulosa
Amblema plica ta
Fusconaia maculata
Fusconaia flava
Cyclonaias tuberculata
Pleurobema clava
Pleurobema cordatum
Pleurobema rubrum
Elliptio crassidens
Elliptio dilatata
Ptychobranchus fasciolaris
Obliquaria reflexa
Actionanaias carina ta
Obovaria subrotunda
Obovaria retusa
Truncilla truncata
Truncilla donaciformis
Leptodea fragillis
R
CL
—
R
R
—
CL
AL
R
CL
CL
R
—
CL
—
R
—
R
—
CL
—
AL
AL
R
CL
—
RWSE
—
RSC
—
RWSE
—
RL
R
CL
AL
CL
R
CL
—
—
R
RWS
R
—
A
RSC
AL
RT
AL
R
690
Indiana Academy of Science
Vol. 94 (1985)
Table 1. — Continued
Scientific name
(1)
Main Stream
Upper Blue
South Fork
Potamilus alatus
Potamilus ohiensis
Ligumia recta
Villosa iris
Villosa lienosa
Lampsilis r. luteola
Lampsilis fasciola
Lampsilis ventricosa
Corbicula = Asiatic clam
(2)
AL
—
RLE
—
R
—
CL
CL
R
R
AL
AL
R
—
AL
CL
AL
CL
c
R
AL
R
AL
Total
30
13
19
(1) Scientific names according to D.H. Stansbery, The Ohio State Univ.
(2) Exotic species widespread in Indiana waters not included in count.
main stream. Although the species composition gradually changed as discharge decreased,
the most abundant species in the main stream were Lampsilis ventricosa, L. radiata
luteola, and Potamilus alatus. Species diversity decreased in the forks of the river with
13 species taken from the upper Blue River-west fork and 19 species noted in the south
fork (Table 1). Amblema plicata, Alasmidonta viridis, Elliptio dilatata, and Villosa
iris were the most abundant forms in the branches of the river.
Several naiads taken during this study have been placed on the proposed en-
dangered, threatened, and species of special concern list recently published by the Indiana
Department of Natural Resources (designated E, T or SC in Table 1). Of the six
designated species, all except Potamilus ohiensis were represented by a single valve.
Several P. ohiensis shells including a living specimen found near Wyandotte Cave were
taken from the main stream. Since this species normally inhabits larger streams, the
Blue River may, in fact, be providing refuge for this endangered mussel.
The paucity of recent comparative mussel abundance data for Indiana indicates
the need for additional inventories of Indiana bivalves. In 1966 and 1967, Krumholz
et al. (2) covered over 500 miles of the Wabash and White Rivers and found only
30 species in 99 collections. It should be noted, however, that Krumholz was studying
commercial exploitation of large river species. This coupled with the fact that only
living specimens were recorded probably accounts for the low species number. In our
study, we recorded only 15 species of living shells (Table 1). More recently, Taylor
(4) in his collections of nearby Indian Creek found only 15 species at 7 locations.
In contrast to Krumholz et al. (2) species list which, as expected, is typically Wabash
in nature, our profile is more Ohioan in nature. The recorded diversity is similar to
the species composition of Kentucky streams such as the Salt River and Floyd's Fork
which are also tributaries of the Ohio River (3). As the largest southern Indiana tributary
of the Ohio River, the Blue River could conceivably harbor the largest mussel assemblage
of this type in the state of Indiana. For this reason and because the river may be
acting as a refuge for the endangered Potamilus ohiensis, the main portion of the
Blue River should be provided with continual protection so that the existing habitats
can be maintained.
Acknowledgments
This research was supported by a grant-in-aid from Indiana University Southeast
and a travel grant from the Indiana Natural Heritage Program. Dr. David H. Stansbery
of the Ohio State University Museum of Zoology verified our identifications.
Zoology 69 1
Literature Cited
1. Goodrich, C. and H. Van Der Schalie. 1944. A revision of the Mollusca of
Indiana. Amer. Midi. Nat. 32(2):257-326.
2. Krumholz, L.A., Bingham, R.L., and E.R. Meyer. 1970. A survey of the com-
mercially valuable mussels of the Wabash and White Rivers of Indiana. Proc.
Indiana Acad. Sci. 79:205-226.
3. Taylor, R.W. 1980. Mussels of Floyd's Fork, a small northcentral Kentucky
stream. The Nautilus. 94(1): 13-15.
4. Taylor, R.W. 1982. The freshwater mussels (naiads) of Big Indian Creek, a small
southern Indiana tributary of the Ohio River (Bivalvia: Unionidae). The Nautilus.
95(1):13-14.
INSTRUCTIONS FOR CONTRIBUTORS
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693
694 Indiana Academy of Science Vol. 94 (1985)
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Instructions For Contributors 695
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Revised July 31, 1974.
INDEX
Abrell, D. Brian, 245, 603
A Brief History of the Cell Biology Section,
Indiana Academy of Science*, 152
A Case of Tuberculosis in the University
Setting*, 401
A Checklist of the Aquatic Coleoptera of
Indiana, 357
Acid Rain: A Synopsis, 381
A Comparison of Soils on Unreclaimed
1949 Indiana Coal Stripmine Surfaces in
1964 and 1981, 579
A Competitive Ecotone between Hardwood
and Relict Hemlock Communities*, 210
A Compilation of Plant Diseases and
Disorders in Indiana — 1984, 145
Additions to the Flora of Indiana: II*, 426
Additions to the Flora of Pike and Gibson
Counties, Indiana, 455
Addresses and Contributed Papers, 80
A Description of Kenneth Chert*, 94
Ahlrichs, J.L., 518, 521, 565
Ahn, Myong-Ku, 174
Air Temperature Fluctuation in Alabama
During the Annular Solar Eclipse on 30
May 1984*, 517
Akhavan, Sepehra, 167
Aldrich, James R., 209, 245, 425, 457
Alwine, S., 426
Ambidentate Phosphine LLgands:
Phospine-amino and Phosphine-imidate
Complexes of Tungsten*, 167
An Analysis of the 28 March 1984 Tornado
Outbreak in the Carolinas, 555
Andersen, Douglas C, 213
Anderson, James M., 170
Andresen, J. A., 521
Anecdotal History of Entomology in In-
diana, 307
An Electron Spin Resonance Method for
the Measurement of Liposomal
Leakage*, 170
A New Amine as an Uncoupler of
Chloroplast Electron Transport, 113
A New and Challenging Science Program
from AAAS for Grades 7 and 8*, 483
A New Approach to Fostering Scientific
Literacy among Indiana's Secondary
School Students*, 485
An Examination of 495 Splice Junction
Sequences*, 403
An Experimental Study of Biparental Care
in the Dark-eyed Junco*, 601
An Introductory Titration for First Year
Chemistry Students: A Comparison of
Antacid Effectiveness, 487
An Investigation of Aluminum Concentra-
tions in Water*, 174
Annual Changes in Flea Population on
Three Domestic Pets, 1978-1984, 329
Annual Financial Report, 59
Annual Report, Indiana Junior Academy
of Science, 64
Anslinger, C. Michael, 93
Anthropology, 93
A Preliminary Review and Multiple-entry
Key to the Rust Fungi on Cyperaceae
and Juncaceae in Indiana, 447
A Preliminary Survey of Phenolic Com-
pounds in Sympatric Populations of
Quercus shumardii and Q. Rubra in
Northern Indiana* 425
A Preliminary Survey of the Maumee River
in Allen County, Indiana*, 95
Apsley, David K., 215
A Rapid Method for the Determination of
Barley Seed Viability, 117
Araya, Jaime E., 303
A Record of the Freshwater Nemertean Pro-
stoma graecense (Bohmig) in Indiana*,
599
Armentano, Thomas V., 269
A SCC MO Calculation on the Tetracyano-
ethylene-benzine Complex, 181
Ash, Donald W., 388
A Simple, Reproducible High Performance
Liquid Chromatography Separation of
Amino Acids with Picomole Sensitivity*,
170
Assessing Variation in Mixed Oak Com-
munities: Evaluation of Multivariate
Analyses of Morphological Data*, 426
Assessment of Numbers of Striped
Cucumber Beetle Adults and Fequency
of Feeding Injury on Muskmelon
Cultivars*, 304
Astrophotography Using Celestron
Telescopes*, 420
A Study of the Coordination Compounds
of Some of the Transition Metals Using
2(2-Aminoethoxy)-Ethanol as a Ligand
697
698
Index
and l-Methyl-2-Pyrrolidinone as a
Solvent*, 173
A Summer Institute in Microcomputer Ap-
plications for Secondary School Science
Teachers, 499
A Superfund Risk Assessment in Indiana:
A Case Study of the Columbia City Site* ,
371
A System for Astronomical Photometry*,
419
Atomic Polarizations of Transition Metal
tris-3-Pentandedionates*, 173
A Trace Metal Analysis of Coal and Acid
Rain*, 172
Ault, Curtis H., 29
A Useful Morphological Characteristic of
Two Toed Sloth Hair*, 94
A Wind Tunnel Investigation of Roughness
Parameters for Surfaces of Regularly Ar-
rayed Roughness Elements, 571
Backs, Steven E., 227
Bacone, John A., 457
Bacterial Wilt Resistance in Commercial
Muskmelon Cultivars, 131
Baeske, Ericka, 193
Baker, Claude D., 373, 603, 633, 687
Banaszak, Konrad J., 387
Banking DNA for Future Diagnosis of
Hereditary Diseases*, 402
Barnell, Stasia A., 167
Barr, Rita, 113
Bartlett, Albert A., 417
Bartolucci, L.A., 519
Baumgardner, M.F., 517, 519
Beeson, Beth E., 167
Behforouz, Mohammad, 167, 168
Behforouz, Nancy C, 401
Bemis, Lynne, 113
Benson, Denise, 373
Beranek, William Jr., 371
Berry, James W., 209
Bhella, H.S., 99, 105
Biofiltration in Intensive Culture Systems:
Design Considerations*, 211
Blackwell, Will H., 391
Bloom, William W., 117
Bolan, Joseph L., 167
Borges, G., 97
Bossung, S., 402
Botany, 97
Bowden, W.W., 293, 295
Boyle, Jeffrey G., 117
Boyts, E., 98
Brack, Virgil Jr., 231, 607
Brekrus, S., 426
Brengle, Blair, 97
Brinker, Ruth, 93
Brooks, Daniel R., 615
Brooks, Richard, 155
Brookville Historical Tour, 13
Buckner, Richard L., 615
Buckner, Shareen C, 615
Burden, Stanley L., 169, 171
Burek, Kathy, 151
Burgin, Alex, 209
Burkett, Frank, 93
Bush, Christopher L., 177
Buskirk, William H., 30
Butler, M.G., 426
Byrnes, William R., 213
Cable, Ted T., 607
Cady, Marshall P. Jr., 417, 483, 487
Calmodulin Stimulation of ATP-
Dependent Ca2 + Uptake in Maize Root
Microsomes*, 154
Canine Dirofilariasis in Central Indiana,
645
Cantin, Mark, 93
Carpenter, Mary Ellen, 94
Carson, Catharine A., 94
Casebere, Lee A., 425
Castrale, John S., 239, 373
Cell Biology, 151
Centennial Address, 37
Chaney, William E., 303
Chaney, William R., 97, 213
Characteristics of Drumming Habitat of
Ruffed Grouse in Indana, 227
Characterization of Indiana Soils by
Porosimetry*, 518
Chaudhuri, N.N., 517
Checklist of Adult Carabid Beetles Known
from Indiana, 341
Chemistry, 167
Chen, B.H., 304
Chesak, David D., 293,
Chick Limb Duplications Produced by
Retinoic Acid Releasing Microimplants,
161
Christenson, Carl M., 687
Ciesla, Edward J., 171
Cisneros, Mark, 169, 181
Index
699
Clancy, Mary, 403
Clark, James A., 313
CLIMATE: A Microcomputer Program
Allowing Student Preparation of
Climatic Maps for Indiana, 489
Cochran, Donald R., 93,
Coggan, A.R., 153
Colglazier, Jerry M., 484
Color Vision: A Lecture Demonstration of
Afterimages*, 488
Comparison of Two Simple Methods for
Determining Lecithin/Sphingomyelin
(L/S) Ratios in Human Amniotic Fluid
Samples, 197
Competition for Ownership of Webs in the
Semi-social Spider Cyrtophora moluccen-
sis of Yap (Caroline Islands,
Micronesia)*, 209
Compression Strength Testing of the
Springfield Coal, Coal V, Pike County,
Indiana*, 387
Computer Aided Classroom Presentations
in Chemistry*, 487
Concanavalin A Inhibits Oral Regeneration
in Stentor coeruleus by Binding to the Cell
Surface*, 152
Conclusion of Acid Rain Monitoring in
Central Indiana*, 173
Conklin, Richard L., 417
Conn, Patricia S., 197
Contorl of Cell Growth by Transplasma-
lemma Redox: Stimulation of Hela Cell
Growth by Impermeable Oxidants, 407
Control of Vegetable Insects with Neem
Seed Extracts, 335
Cook, Della Collins, 94
Cooksy, Kevin, 170
cortwright, spencer a., 210
Cory, Walter, 483
Cottonmouth, Agkistrodon piscivorous,
Records from the Blue River and Potato
Run in Harrison County, Indiana (Ohio
River Drainage, USA), 633
Coulometric Titrations: Low Cost Alter-
natives for Computer Controlled
Titrations*, 169
Cragnulino, G., 293
Crane, Frederick L., 113, 407
Crovello, Theodore J., 380
Cudmore, Wynn W., 621
Curran, John T., 529
Daily, Fay Kenoyer, 18, 21, 69
Daniell, David L., 629
Davis, Grayson S., 161
de Almeida, Alfonso, 518
Debitage Classification Systems*, 93
deCassia, Rita, 97
Denner, Melvin W., 615
Density-dependent Mortality on Galls of the
Goldenrod Gall Fly, Eurosta solidaginis* ,
214
Dental Anomalies in Three Species of
Shrews from Indiana, 635
Determining Needs: First Step for Improv-
ing Science and Mathematics Instruction
in Rural High Schools in Northwestern
Indiana*, 484
Development and Analysis of a CFI Data
Base for Indiana*, 211
Development of a Model System for the
Study of Murine Leukocyte
Chemiluminescence, 404
Dhawale, S., 293
2,4-Dinitrophenylhydrazones: A Modified
Method for the Preparation of these
Derivatives and an Explanation of
Previous Conflicting Results*, 167
Dinner for Senior Academy Officers, 38
DiNoto, Vincent A. Jr., 417, 418
Dolph, Gary E., 483, 489
Do Tadpoles Die for their Siblings?*, 212
Dotterer, Sally K., 169
Downie, N.M., 357
Driscoll, Daniel E., 607
Duff, Douglas, 673
Duffy, Kim, 471
Dunn, Howard, 372
Dupuis, Roxane A., 425, 426
DuSold, Elizabeth, 371
Dyke, Jennifer L., 170
Ecology, 209
Ectoparasites of Pine Voles, Microtus
pinetorum, from Clark County, Illinois,
649
Edmondson, Frank K., 418
Effect of Acetylcholine Stimulation on
Cytosolic Chloride in Parotia Acinar
Cells*, 151
Effect of Barley Yellow-dwarf Virus Infec-
tion of Wheat and Oats on the Life Cycle
of Rhopalosiphum padi (L.)*, 303
Effect of Cyclosporine A on Leishmania
tropica*, 401
700
Index
Effect of Cytokinins on Erythritol
Permeability to Phosphatidylcholine
Bilayers*, 97
Effect of Viruliferous and Non-viruliferous
Rhopalosipum padi (L.) Aphids on
Winter Wheat*, 304
Efficiency of Pollen Traps with Various
Sized Trap Screens*, 303
Ellis, Bernice, 170
Engineering, 293
Engineering and Science Education's
Dilemma: Inadequate Science Programs
in the Public School System*, 293
Engineering Properties of Indiana Peats and
Mucks*, 518
Engle, Mike, 203
Entomology, 303
Environmental Quality, 371
Evaluation of Landsat Thematic Mapper
Data for Classifying Forest Lands, 297
Evaulation of Sample Pre-treatments as
Potential Methods of Enhancing
Phospholipid Extraction from Human
Amniotic Fluid, 193
Evaporation Rates of Organic Liquids at
Various Wind Speeds and
Temperatures*, 372
Evidence of Algal Source of Micrite in a
Saluda Coral Zone in Southeastern In-
diana, 391
Ewert, Michael A., 210
Executive Committee Meetings, 39
Extinct Woodland Muskox, Symbos
cavifrons, Cranium from Miami County,
North Central Indiana, 667
Farringer, L. Dwight, 418, 499
Faust, Kristen, 167
Favinger, John J., 307
Ferreira, A.M., 518
Ferson, Scott, 210
Field Biology: A Blow to Provincialism*,
486
Fischer, Burnell C, 211
Fischer, Robert, 471
Fisher, D., 426
Flynt, Michael S., 167
Food Habits of Urban American Kestrels,
Falco sparrerius, 607
Foos, K. Michael, 109, 503
Forsyth, Bill J., 373, 603, 633, 687
Foster, John E., 303, 304, 305, 404
Fowler, S.E., 518
Francq, G. Earle, 484
Frantz, Vonda, 98
Franzmeier, D.P., 533
French, Thomas W., 635, 641
FULLENKAMP, A.M., 471
Functionalized Crown Ethers*, 172
Furgason, E.S., 305
Furia, Edmond J., 94
Gallagher, Patrick, 170
Galloway, H.M., 533
G alloy, Ronald J., 381
Garcia, J.R., 401
G-banding in Lens culinaris and Vicia
faba*, 98
Geology and Geography, 387
Geology and Geomorphic History of the
Garrison Chapel Cave System, Monroe
County, Indiana*, 388
George, James, 485
Goebel, Edwin M., 151
Goff, Charles W., 154
Gommel, William R., 517
Gravel Hill Praries of Indiana, 457
Gray, Bonnie, 471
Greeman, Theodore K., 387
Green, Ralph J. Jr., 98
Gust Fronts in Doppler Radar Data, 547
Hamilton, Jodi, 419
Hamrick, Linda, 483, 485
Hardwood Tree and Ground Cover
Establishment on Reclaimed Mineland
and Unmined Reference Sites in
Indiana*, 213
Hartmann, Walter, 475
Harty, Harold, 485, 509
Heat Loss from Avian Integument: Effects
of Posture and the Plumage, 681
Heiser, Charles B., Jr., 88
Hengeveld, James D., 597
Hennen, J.F., 425
Hennen, M.M., 425
Herbicide (Alachlor, Atrazine, Linuron and
Paraquat) Residues in Deer Mice In-
habiting Conventional and Minimum
Tillage Row-crop Fields, 373
Heterosexual Social Interactions in the
Syrian Hamster*, 471
Hicks, Ronald, 95
Highlights of the Spring Meeting, 13
Index
701
Hill, Jonn R., 29
Hill, Maureen L., 170
Hindered Ligand Systems: Structure of the
cis, os-l,3,5,-Tris (pyridine-2-carboxaldi-
mine) cyclohexane Complexes of Fe(II)
and Ni(II) Ions*, 171
History of Science, 395
Hodes, M.E., 402, 403
Hoffer, Roger M., 297, 484
Holland Chert Quarries/Workshops Near
Huntingburg, Dubois County, Indiana*,
93
Holland, James P., 155
HOLLERMAN, ANDREW, 293
Homoya, Michael A., 245, 426
HOSKINS, JOANN, 402
Huffman, C.J., 171
Huffman, J.C., 171
Hughes, James P., 153
Ideas Concerning the Use of Computer
Data Acquisition Systems to Improve
Teaching Effectiveness within the
Laboratory*, 483
Identification of a Pectinase in Larvae of
the Hessian Fly, Mayetiola destructor
(Say)*, 305
Improving Efficiency of Iron Uptake by
Soybeans, 141
Improving the Results of Molecular Mass
Determination Experiments by Using a
Microelectronic Thermistor Device*, 486
Increased Binding of Growth Hormone
following Cleavage by Rabbit Liver
Plasmalemima*, 153
Index, 697
Indiana Gypsy Moth Survey — A History,
313
Indiana Junior Academy of Science, 64
Insect Pest Control in the Greenhouse:
Alternatives to Commercial Toxins*, 98
Insects and Other Arthropods of Economic
Importance in Indiana in 1984, 323
Instructions for Contributors, 693
Integer-valued Equivalent Resistance, 421
Interactions among Mast, Small Mammals,
and Insects, and their Implications*, 213
Interpretation of Glacial Geology and
Groundwater Problems in Eastcentral In-
diana Using Improved Compilations of
Water Well Driller's Records*, 388
Isolation of the Coprophilous Fungus,
Pilobolus, from Wayne County, Indiana,
109
Iverson, John B., 597
Jackson, Marion T., 425, 427,
Jackson, Misty, 95
Jacobson, John E., 297
Jarial, Mohinder S., 597
Jarrett, Harry W., 170
Jensen, Richard, 425, 426, 429
Jersild, Ralph A. Jr., 151, 152
Johnson, John G., 35
Johnson, Michael D., 598
Johnson, Susan M., 485
Jordan, David, 371
Joseph, Paul, 518
Kane, Barbara, 471
Kastelein, Nathan E., 171
Kelly, Sean T., 227
Kershaw, John A. Jr., 211
Kimmerle, Kenneth R., 173
Kirsch, Joe, 169, 173, 181
Kissinger, Paul B., 486
Kjonaas, Richard A., 172
Klansmeier, Michael E., 372
Kleinhans, F.W., 170
Klingle, Diana L., 547
Klingler, T.E., 555
Klunzinger, Phillip E., 171
Koenig, Thomas, 152
Kowalski, Michael P., 598
Kramer, Rosalie, 486
Kristof, S.J., 517
Krohne, David T., 209
Kroll, LeRoy, 172
Kudagamage, C, 304
Kuo, K.C., 387
Kurtz, Kristine S., 187
Lambert, Richard H., 401
Land Cover Classification of Rupgang
Thana Dhaka, Bangladesh Using Landsat
MSS Data*, 517
Landfills in Marion County — A Revisit*,
387
Langona, M., 401, 402
Larsen, Steven H., 402
Larter, Raima M., 177
Latin, Richard X., 145
Lawrence, Joseph D., 421
Leeds, Jonathan, 113
Legal Game Harvest by Indiana Land-
owners Hunting without a License, 239
702
Index
Leopold, Donald J., 212, 215
Libey, George S., 211
Licensing and Certification of Physics
Teachers by Examination: What are the
Dangers?, 419
Lieb, Shannon, 167, 181
Light Microscopic and Ultrastructural
Features of the Gut of the Balsam Wooly
Aphid, Adelges piceae Ratz*, 597
Linear Differentiation of A lluim cepa, Lens
culinaris and Viciafaba Chromosomes* ,
426
Lister, R.M., 404
Litton, James R., Jr., 597, 599
Lopez-F, R.M., 425
Lovell, C.W., 518
Lozano-Garcia, D. Fabian, 484
Macdonald, D.D., 293
McCain, John W., 447
McFee, W.W., 521
McMahan, Deborah A., 151
Macer, Charles B., 372
Macri, Jeff, 170
Madisen, Linda, 402
Major, P. Decker, 227
Male Mating Behavior in Hyla cinerea*, 213
Maloney, Michael S., 152
Mann Site Figurines*, 93
Marking in Submissive Male Gerbils after
Contact with a Dominant Male and His
Odors*, 471
Marks, Gayton C, 117
Marshall, Philip T., 98, 313
Mass Rearing the Bird Cherry Oat Aphid,
Rhopalosiphum padi (L.)*, 304
Mays, Charles E., 645
Mehra, Romesh C, 98, 426
Menges, Eric S., 269
Mennen, K.E., 168
Merrill, John, 371
Meunier, Gary, 471
Meyer, Robert W., 323
Microbiology and Molecular Biology, 401
Miller, Benjamin P., 372
Miller, Brian K., 227
Miller, Gary E., 211
Minton, Sherman A., 30, 600
Minutes of the Budget Committee Meeting,
57
Minutes of the Fall Meeting (Executive
Committee), 48
Minutes of the Fall Meeting (General Ses-
sion), 54
Minutes of the Spring Meeting (Executive
Committee), 31
Modrak, Gin a, 193, 197, 203
Mohow, James A., 95
Moreira, Nuno, 518
Morris, Molly, 212
Mosbo, John A., 170
Mueller, Paul W., 297
Mulkey, Timohty J., 154
Munford, John W., 152
Munro's Doctrines: A Forgotten Pioneer
in Holism and Hypnosis, 475
Munsee, Jack R., 329, 579
Nass, Lisa B., 164
Necrology, 69
Nelson, Craig E., 210, 212
Newman, John, 13
New Members for 1984, 76
Newman, Steve, 172
Noller, C.H., 518
Nonspecificity with Varied Effectivity in
Mycorrhizal Associations*, 97
Noon Luncheon, 41
Norris, F.H., 403
Oak "Leaf Tatters": A Malady of
Unknown Cause in Indiana*, 98
Oblander, Scott, 293
Occurrence of Swimmers' Itch in Northeast
Indiana, 629
Officers and Committees for 1984, 4
Ogle, M.E., 168
Ohm, H.W., 304
On the Measurement of Thermal Dif fusivities
with Bryngdahl Interferometry*, 417
Orr, M., 402
Orwell's 1984, Skinner's Walden II, Marx'
Classless Society and other Utopias: An
Exploration of Human Expectation and
the Psychological Factors in a "Perfect
Society", 472
Our Brookville Bond, 18
Pace, Robert E., 94, 95
Palbykin, J., 426
Parasitic Endohelminths from Fishes of
Southern Indiana, 615
Parental Investment in the Bee Ceratina
Index
703
calcarata Robertson (Hymenoptera:
Xylocopidea): A Preliminary Study*, 598
Parke, Neil J., 645
Parker, George R., 212, 215
Pascal, D. David Jr., 649
Patterns of Relative Fecundity in Snakes*,
597
PawU, K.T., 521
Pecknold, Paul C, 145
Perrill, Stephen A., 213
Personality Types and Perceptual-motor
Performance*, 473
Pfingsten, William J., 239
Phillips, T.L., 519
Physics and Astronomy, 417
Physiological Studies of Azospirillum
amazonese*, 151
Physiology of Vocalization by an
Echolocating Bird*, 600
Pictorial Highlights of the Fall Meeting, 35
Pipewort Pond, a Unique Wetland with
Atlantic Coastal Plain Elements in
Elkhart County, Indiana*, 209
Pires, Ana L., 565
Plant Taxonomy, 425
Plasma Progesterone, Blastocyst Steriod-
ogenesis and Blastocyst Survival in Rats
with Altered Thyroid Status, 155
Poad, Douglas W., 517
Pokorney, Laura, 173
Poorman, Lawrence E., 419
Pope, Phillip E., 97, 213
Population Studies of Threatened and En-
dangered Plants of Barker Woods Nature
Preserve, LaPort County, Indiana, 121
Poster Sessions, 42
Post, Thomas W., 245, 455, 457
Potter, CD., 388
Pre-burning Floral Inventory of Little
Bluestem Prairie, Vigo County, Indiana*,
427
Predator-determined Structure in Amphi-
bian Pond Communities*, 210
Prediction of the Variation of Azeotropic
Composition Using the Gibbs-Konovalov
Theorem*, 293
Preface to the Centennial Volume, 3
Preference of the Bird Cherry Oat Aphid,
Rhopalosiphum padi (L.) on Hessian Fly-
infested Wheat and Effects on its
Biology*, 305
Presidential Address: "Computers, Educa-
tion, and Artificial Intelligence", 80
Protein Degradation after Eccentric
Exercise*, 153
Psychology, 471
Psychovector Love Scale and its Differenti-
ability*, 472
Pugh, Tom, 403
Putnam, J.E., 407
Quaternary Remains of the Spotted Skunk
Spilogale putorius, in Indiana, 657
Ragatz, Barth H., 193, 197, 203
Reaction Sequence Alteration in the
Acetoacetic Ester Synthesis of Ketones*,
172
Rediscovery of the Spotted Darter,
Etheostoma maculatum, in Indiana
Waters: Blue River; Crawford, Harrison
and Washington Counties; Ohio River
Drainage, USA, 603
Reed, David K., 304, 335
Reed, Gary L., 131, 304, 335
Reed, Patricia Wiese, 121, 465
Regional Low Density and Extinction in
Populations of Peromyscus leucopus*,
209
Relationship between Symptomatic
Resistance and Virus Production in
Barley Cultivars Inoculated with Barley
Yellow-dwarf Virus*, 404
Relationship of Probing Behavior of Sito-
bion avenal (Fabricins) of Transmission
of Luteoviruses Causing Cereal Yellow-
dwarf Diseases*, 305
Reports from field trip leaders, 29
Reproduction and Age Structure of Three
Indiana Shrews, 641
Response of Forage Corps to Dolomitic
Lime, 565
Response of Muskmelon to Within-row
Plant Spacing, 99
Rhykerd, C.L., 517, 518, 565
Richards, Ronald L., 657, 667
Ricketts, John A., 486
rlemenschneider, victor, 425, 465
Rishaw, Claudia, 171
Robertson, Thomas H., 419
Robots in the Chemistry Laboratory, Part
I: A High Speed RS-232C Serial Commu-
nications Link for Controlling a HERO
704
Index
I Robot from an Apple II Plus Micro-
computer*, 171
Robots in the Chemistry Laboratory, Part
II: Software for Controlling a HERO I
Robot from an Apple II Plus Microcom-
puter via a High Speed RS-232C Com-
munications Link*, 171
RODIBAUGH, ROSEMAY, 141
Rolley, Robert E., 239
Rostek, Wayne F., 571
Roth, Cynthia L., 171
Rothrock, Paul E., 427
Rowland, William J., 599
Royer, Judith A., 109
Ruhl, Gail E., 145
Rust Species Diversity in Temperate and
Tropical Regions of the Americas*, 425
Rybarczky, James P., 172, 173
Samuelson, Alan C, 388
Sans, J.R., 388
Sayegh, Samir I., 421
Scabies, A Nosocomial Outbreak*, 402
SCHARMANN, LAWRENCE, 509
Scheller, H.V., 305
Schepper, Jeanette M., 153
schliessmann, roberta, 472
schlueter, annette j., 161
Schrock, John Richard, 341, 486, 579
Schuley, Richard E., 486
Schultz, Phillip W., 169
Schwartz, Eugene, 173
Science Education, 483
Science Training for the Industrial Environ-
ment (STIE)*, 486
Scircle, John, 173
Scott, Donald H., 145
Seasonal Abundance of the Psammic
Rotifers of Spicer Lake, Indiana*, 599
Seeley, Barbara A., 395
Seeley, Gerald R., 395
Semel, Brad, 213
Sensitivity Studies of a Computer Model
for the Peroxidase-oxidase Oscillating
Reaction, 177
Serum Hormone Levels in Germfree and
Conventional Rats: Effect of Dietary
Restriction*, 404
Sessions, Katharine, 487
Sever, David M., 673
Sexual Selection and Alternative Mating
Strategies in Hyla crucifer and Hyla
chrysoscelis, 212
Shea, Gerald J., 419
Shellhaas, James L., 404
Shimer, Stanley S., 487
Shipe, Albert P., 529
Shukle, R.H., 305
Siefker, Joseph R., 173
Skaria, M., 404
Smith, David R., 547, 555
Snow, John T., 571
Snyder, A.C., 153
Snyder, David L., 404
Software for Astronomical Photometry*,
419
Soil and Atmospheric Sciences, 517
Soils an Important Component in a Digital
Geographic Information System*, 519
Soil Survey in Indiana: Past, Present and
Future, 533
Sojka, Gary A., 21, 22
Some Late Archaic Manifestations in
Indiana*, 96
Sousa, Lynn R., 167
"Speaker of the Year" Address, 1984-85
"The Contributions of the Nightshade
Family (Solanaceae) to Human
Welfare", 88
Speaking of Sex — A Presentation on Ter-
minology for Students in Reproductive
Biology Classes*, 486
Special Acknowledgment, 44
Spectra and Equilibria of the Thiocyanate
Complexes of Copper (I) in Aqueous
Solution*, 169
Spectra and Photochemistry of the Chloro
Complexes of Cooper (I), 187
Squiers, Edwin R., 209, 289
Starcs, Helene, 425
Stark, Robert J., 151
St. Clair, Martin, 371
Steldt, F.R., 420
Stem Length as an Estimator of Muskmelon
Growth, 105
Stephenson, P. Ranel, 95
Steric and Electronic Effects upon cis:trans
Distributions in W(CO)4(L)(L ') Com-
plexes when L and L ' are Phosphorus
Ligands*, 170
Stevenson, Kenneth L., 169, 187
Stevenson, W.R., 131
Still well, William, 97
Stockton, Daniel D., 210
Index
705
Storhoff, Bruce, 167, 172
Strait, Rebecca A., 427
Streator, James T., 487
Streib, W.E., 171
Stress Corrosion Cracking of Sensitized
Austenitic Stainless Steels in Basic Acid
Solution Containing Sulfur Oxyanions*,
293
Successional Relationships of Pine Stands
at Indiana Dunes, 269
Sun, I.L., 407
Survey of the Fishes of the Kingsbury State
Fish and Wildlife Area, LaPort County,
Indiana, 673
Survey of the Mineral Composition of
Forage Corps in Portugal*, 518
Suthers, Roderick A., 600
Synthesis Experiments for High School
Chemistry, 485
Synthesis of /3-Carbolines Derived from
2-Amino-3-(3-indolyl)-butyric Acid ((3-
Methyltryptophan)*, 168
Taylor, Ralph W., 687
Temperature Dependent Infrared Studies
of the Hydrogen Bonding in Aliphatic
Alcohols*, 169, 173
Territorial Behavior of the Prothonotary
Warbler, Protonotaha citera, Between-
and Within- season Territory
Relocations*, 598
Test Excavations at the Smith Site,
(12-Vi-86), Viro County, Indiana*, 94
The Adaptive (?) Significance of Brood
Reduction in the Red-winged Blackbird
(Angelaius phoeniceus)*, 597
The Commissary Site (12-Hn-2) Revisited*,
93
The Complex Relationship of Embryonic
Development of Incubation Temperature
in Turtles*, 210
The Determination of the Removal Rate of
Specific Chemicals by the Indianapolis
Wastewater Treatment System*, 371
The Discovery of Native Rare Vascular
Plants in Northern Indiana*, 425
The Dynamics of the Population of the
United States*, 417
The Effect of Fasting on Sodium Pump Ac-
tivity in Rat Skeletal Muscle*, 152
The Effect of Illumination on the Rat Pineal
as Measured by MSH Activity*, 154
The Effects of Oligolysines and Polylysines
on Human Platelet Aggregation Induced
by Polylysines, Adenosine Diphosphate,
and Epinephrine, 203
The Foraging Ecology of Some Bats in In-
diana, 231
The Freshwater Naiads, Bivalvia:
Unionidae, of the Blue River, a Southern
Indiana Tributary of the Ohio River, 687
The Great Southern U.S. Geologic Uplift
Observed in the Early Months of 1984*,
419
The IAS Engineering Section: A Brief
History*, 293
The International Challenge: A Com-
parison of Science Education Models
from Four Nations*, 485
The Layered Classifier: A More Effective
Method for Studying Seasonal Changes
in Forest Cover Types Using Satellite
Data*, 484
The Making of David Starr Jordan, 22
The Manchester Interface Adapter for
Commodore and Apple Microcom-
puters*, 418
The National Optical Astronomy
Observatories*, 418
The National Weather Service Rainfall Data
Collection Network in Indiana, 529
The Natural Regions of Indiana, 245
The Physics of the Grist-mill*, 418
The Present Distribution and Status of the
Eastern Woodrat, Neotomafloridana, in
Indiana, 621
The PVT Behavior of Compressed
Liquids*, 295
The Ratio of PM-10 to TSP in the
Atmosphere*, 371
The Red and Black Oaks of Indiana, 429
The Regulation of S-Adenosylmethionine
Synthetase in Candida albicans*, 401
The Rich and Varied Past of the History
of Science Section, 395
The Roots of Ecology in Indiana, 289
The Several Themes of Adolescence*, 471
The Synthesis of a Crown Ether that May
Exhibit Metal Cation Enhanced
Fluorescence*, 167
The Year at Drombeg*, 95
Thirakhupt, V., 305
Three Cranial Tumors from Late Woodland
Sites: Diagnosis and Cultural
706
Index
Implications*, 94
Three-dimensional Patterns of Biotic Com-
position within the Cloudy Pass
Batholith, Washington*, 388
Three Plasmid Cloning Vectors for Mam-
malian Cells*, 402
Tomak, Curtis H., 96
Towards Predicting Loss of Archaeological
Resources from River Channel
Migrations*, 95
Transcriptional Regulation of the
Sporulation-specific Glucoamylase of
Saccharomyces cerevisiae*, 403
Tree Species Dynamics in an Old-growth
Deciduous Forest since 1926*, 212
Tree Species Response to Release from
Domestic Livestock Grazing, 215
Two-year College Biology Instructors'
Perceptions about their Role Expecta-
tions, 509
Tzeng, Oliver C.S., 472
Uhl, J. J., 153
Use of a Microcomputer to Enhance the
Coin Flip Probability Exercise in the
General Biology Laboratory, 503
Using the Microcomputer to Teach Science
in the Elementary Classrooms*, 487
Using Toys to Teach Physics to Middle
School Students*, 420
Valenzuela, C.R., 519
Vance, Robert A., 471
Vascular Flora of Grant County, Indiana:
Additions and Comments*, 427
Vascular Plants of Barker Woods Nature
Preserve, LaPorte County, Indiana, 465
Vaughan, Martin A., 154
Vayhinger, John A., 472
Venom Antigens in Oral Secretions of Col-
ubrid Snakes*, 600
Visual Signals in Sticklebacks: A Reex-
amination and Extension of Some Classic
Experiments*, 599
Walton, Rod, 214
Ware, Roger, 473
Watson, James Jr., 420
Watson, Nancy, 420
Weaver, Connie, 141
Webster, Marcus D., 681
Weidenbener, Erich, 155
Weilbaker, Charles, 633, 687
Welcome to Butler University, 35
Welcome to the Fall Meeting, 36
Wentworth, R.A.D., 171
Wepler, William R., 667
West, Dennis, 295
West, T.R., 387
Wet Atmospheric Deposition in Indiana,
521
Whitaker, John O. Jr., 649
White, Charles E., 357
Wicker, John W., 517
Wilcox, G.E., 105
Wiles, Tom, 603, 633
Williams, Albert A., 488, 499
Wilson, William L., 388
Wittig Reaction: Stable Ylides in the
Preparation of 7, 5 -unsaturated-/}-
Ketoesters*, 168
Wolf, Licia, 601
Womack, Henry C, 154
Woodland Sites and Ross Soils: A Correla-
tion in the Upper White River (West
Fork) Drainage, 95
Wostmann, Bernard S., 404
WUNDERLICH, DANIEL K., 174
Yahner, J.E., 533
Yess, Edward C, 529
Yokomoto, Charles, 473
Young, Frank N., 357
Zoology, 597
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